Progress in Oceanography: Joan E. Cartes, Vanesa Papiol, Beatriz Guijarro

Progress in Oceanography 79 (2008) 3754
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Progress in Oceanography
journal homepage: www.elsevier.com/locate/pocean
The feeding and diet of the deep-sea shrimp Aristeus antennatus off the
Balearic Islands (Western Mediterranean): Inuence of environmental factors
and relationship with the biological cycle
Joan E. Cartes a,*, Vanesa Papiol a, Beatriz Guijarro b
a
b
Institut de Cincies del Mar (CMIMA-CSIC), Psg. Martim de la Barceloneta 3749, Spain
IEO-Centre Oceanogrc de les Balears, P.O. Box 291, 07080 Mallorca, Spain
a r t i c l e
i n f o
Article history:
Received 14 February 2008
Received in revised form 25 July 2008
Accepted 25 July 2008
Available online 6 August 2008
Keywords:
Feeding intensity
Trophic dynamics
Deep Sea
Decapoda
Benthic-Planktonic coupling
Reproductive cycle
Water masses
Daily migrations
a b s t r a c t
Spatio-temporal variation of feeding intensity and diet in the red shrimp Aristeus antennatus was studied
at two locations around the island of Mallorca (Balearic Islands, Western Mediterraean) in August,
September, and November 2003, and in February, April and June 2004 at depths between 550 and
750 m. The two areas, with different oceanographic conditions, were respectively located in the northwest (Sller) and the south (Cabrera) of Mallorca. Off Sller, feeding intensity of A. antennatus showed
a signicant increase from February to April and June 2004 in all the three size-classes studied (small
shrimps: CL < 30 mm; medium: CL between 30 and 40 mm; large: CL P 40 mm). Off Cabrera, the highest
fullness was recorded in November 2003 among small and medium shrimp, while only large specimens
showed patterns similar to that found off Sller. Off Sller, the diet of both small (CL < 34 mm) and large
(CL P 34 mm) A. antennatus was mainly inuenced by season, with three dietary groups corresponding to
AugustSeptember 2003, to November 2003/February 2004, and to hauls from April to June 2004. Off
Cabrera, hauls (representing diets) were grouped by depth, never by season. The most remarkable seasonal shift in the diet of A. antennatus off Sller was the increase of mesopelagic prey in AprilJune relative to other months. In all size categories there was an increase off Sller in the energy intake of prey
ingested from February to June 2004, an increase not found off Cabrera. Degree of digestion of mesopelagic prey indicated nocturnal feeding on mesopelagic fauna. These prey probably have a shallower depth
distribution at night than found in our daylight sampling, and possible migratory movements among prey
and A. antennatus at night would explain the lack of correlation between prey abundance in guts and in
the environment found during daylight periods for most micronekton mesopelagic prey (euphausiids,
myctophids and sergestids). Off Sller, fullness and diet were signicantly linked to temporal changes
in water column productivity (e.g., Chl a readings, uorescence) and to changes in the shrimp biology
(lipid content of hepatopancreas, Gonado-somatic Index, GSI). Off Cabrera, we found a higher dependence
of fullness and diet with T and S, both variables in turn related to depth. The increase of stomach fullness
and dietary energy intake in pre-reproductive females from February to AprilJune 2004 found off Sller,
coupled with the consumption of mesopelagic prey, was parallel to a signicant increase of the gonad
weight (GSI, fecundity) in June. Most individuals attain gonad development in the period MayJune, after
two months of the peak of primary production at the surface. The strong link found between pelagic
resources and reproductive processes in a deep-sea species such as the shrimp Aristeus antennatus, situated near the top of the trophic web, suggests a rapid energy ow via mesopelagic fauna between surface
primary production and bathyal megabenthic communities at oligotrophic insular areas. In contrast to
mainland areas off the Catalan coasts submitted to the inuence of submarine canyons, around the island
of Mallorca the empoverishment of benthos biomass may enhance consumption of micronektonic prey
and a possible accumulation of pre-reproductive females of A. antennatus in areas (e.g., steep slopes
and persistent frontal systems found off Sller) with high zooplankton aggregations.
2008 Elsevier Ltd. All rights reserved.
1. Introduction
* Corresponding author.
E-mail address: jcartes@icm.csic.es (J.E. Cartes).
0079-6611/$ - see front matter 2008 Elsevier Ltd. All rights reserved.
doi:10.1016/j.pocean.2008.07.003
Aristeus antennatus (Risso, 1816) is a demersal deep-sea shrimp

distributed in the subtropical Atlantic from Cape Verde in the
south into Mediterranean waters (Crosnier and Forest, 1973;
38
J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754
Lagardre, 1977). It is a dominant species in Mediterranean

megafaunal assemblages, occupying the middle and lower parts
of the slope between 450 and 2261 m (Cartes and Sard, 1992;
Cartes, 1993a), with a few specimens caught deeper, to 3300 m
(Sard et al., 2004). Other species in the genus Aristeus are also distributed at mid-bathyal depths (e.g., Aristeus varidens) in tropical
and subtropical waters (Crosnier and Forest, 1973). Aristeinae also
includes a number of large, often red-coloured shrimp such as Plesiopenaeus spp. and Aristaeaomorpha spp., all of them abundant in
deep waters and of high ecological and sheries interest.
The trophic role of A. antennatus has been studied both in the
Atlantic (Lagardre, 1971) and in Mediterranean waters(Relini-Orsi
and Wrtz, 1977; Cartes, 1994; Chartosia et al., 2005). Aristeus
antennatus has a highly diverse diet (H0 of 5.58: Cartes, 1994),
and together with some Macrouridae (Madurell and Cartes, 2006)
and other decapods (e.g., Crangonidae: Lagardre, 1977) it is
among the megabenthic predators whose diet is mainly based on
benthos in the deep Mediterranean (Cartes, 1994; Cartes and Carrassn, 2004). However, A. antennatus preys on a wide variety of
benthic and benthopelagic organisms with ontogenic changes in
the diet (Cartes and Sard, 1989). Small juveniles prey on small
crustaceans (amphipods, euphausiids and mysids), while among
adults there is an increasing importance of large prey (small sh,
the burrowing decapod Calocaris macandreae, mesopelagic decapods such as Pasiphaeidae and Sergestidae) in the diet. Size-related
dietary changes have largely been described among shes, but less
so for decapod crustaceans (Fanelli and Cartes, 2004). The diet of A.
antennatus also changes with depth and season (Cartes, 1994).
Although this has been attributed to changes in prey availability,
the simultaneous sampling of prey has rarely been accomplished.
Large specimens prey on infauna and epifauna (e.g., polychaetes,
ophiuroids) at the head of submarine canyons, where the biomass
of infauna and the stomach fullness (feeding intensity) of A.
antennatus increase, probably as a consequence of high local productivity in canyons (e.g., higher organic matter OM - content:
Cartes, 1994). In addition, A. antennatus occupies a high trophic level (elucidated by their isotopic composition: Polunin et al., 2001)
in deep-sea food webs, and it shows diel feeding rhythms and a relatively high food consumption rate (deduced from daily ration
models: Maynou and Cartes, 1997) linked to possible nocturnal
movements upward along the slope canyons (Cartes et al., 1993).
After 21 yrs of studies on the biology of this species, especially
off the coasts of Catalonia (Balearic Basin), important knowledge
gaps still remain. Aristeus antennatus shows a smaller-deeper trend
(Sard and Cartes, 1993), with smallest recruits (individuals between 7 and 16 mm CL) distributed below 1400 m (Cartes and
Demestre, 2003) in both the Balearic Basin (Sard and Cartes,
1993) and the Algerian Basin (Morales-Nin et al., 2003). Why these
recruits are linked to deep waters, expected a priori to be of low
productivity, is unknown. Relatively high levels of fresh OM have
been recorded below 1000 m in the Balearic Basin (Cartes et al.,
2002), At these depths, there is still a relatively high abundance
of suprabenthic crustaceans (e.g., amphipods, cumaceans), which
can exploit detritus (Madurell et al., 2008). These small crustaceans
are the prey of the smallest A. antennatus recruits (Cartes data in
Sard and Cartes, 1997). As the smallest recruits of A. antennatus
are practically absent from upper and mid-slope depths above
1000 m, possible coupling between potential prey availability
and shifts in the abundance of A. antennatus above 1000 m depth
must be related with the aggregation of reproductive specimens,
not with recruitment of juveniles.
The biological cycles of deep-sea macrofauna have been repeatedly suggested to derive from cyclic food availability, often linked
to periodic phytodetritus deposition (vertical ux), and to advective uxes in areas with mainland inuence (e.g. via submarine
canyons: Palanques et al., 2006). Among megafauna, links with
pulses of food also happens among detritivorous echinoderms, particularly holothurians (Ginger et al., 2001; Hudson et al., 2004) and
asteroids (Howell et al., 2004). Among deep-water predators,
including decapods (Cartes et al., 2007) occupying higher trophic
levels than echinoderms, we generally expect a delayed response
to peaks of primary production. Ortons rule (1920) suggested that
continuous (or asynchronous) gametogenesis should be the dominant reproductive pattern of deep-living species, permitted by the
increasing environmental stability at great depths. However, continuous and seasonal (or synchronous) reproduction has been documented at mid-slope depths (Gage and Tyler, 1991). In addition,
Thorsons rule (1950) predicted the prevalence of lecitotrophic larvae among deep water fauna. However, in the Atlantic Ocean at
temperate latitudes, both planktotrophic (e.g., Sergestidae) and
lecitotrophic (e.g., Pasiphaeidae) decapods can be found at mesopelagic levels (Omori, 1974). Lecitotrophic oplophorid shrimp are
dominant at bathypelagic depths, whereas at the abyssopelagic level we nd planktotrophic species such as Benthysicymus spp. or
Bentheogenemma spp., species with the same biological strategy
found in Aristeus antennatus. Penaeoidean shrimp are characterized
by shedding eggs into the water, a feature common in r strategists, not attaching them to their pleopods as do the rest of the
decapods. In summary, the data available for deep-water shrimp
are not consistent with a single expected pattern of reproduction,
probably because the energetic costs of both strategies (planktotrophy and lecitotrophy) are similar.
Complete understanding of reproductive success among deepsea invertebrates will require both better knowledge of their life
cycles and simultaneous environmental (physicalchemical) and
trophic data, among the most important probably being food availability and consumption. Recently, the joint study of food and feeding and of the reproductive cycle in the pandalid shrimp Plesionika
martia, a species inhabiting the mid-slope of the deep Mediterranean, indicates important shifts in the diet of this species depending on season, with a more energy-rich diet (based on the
consumption of energy-rich mesopelagic decapods) in spring coinciding with the beginning of gonad development in pre-reproductive females (Fanelli and Cartes, 2008). The congeneric species
Plesionika heterocarpus, distributed at shallower depths than P.
martia and showing a continuous reproductive cycle, did not exhibit such substantial changes in the diet as did P. martia. Lipids have
an important role in these cycles, and lipid biomarkers indicate different biological strategies for mesopelagic fauna (Ju and Harvey,
2004) and for deep-sea benthos, depending on the nature of lipids
accumulated by each species (Ginger et al., 2001). Fluctuations in
the lipid content in muscle, ovary and hepatopancreas have been
reported for A. antennatus (Rosa and Nunes, 2003a,b; Guijarro
et al., 2008), with shifts probably linked to its non-continuous
reproductive cycle (Arrobas and Ribeiro-Cascalho, 1987; Demestre
and Fortuo, 1992). The beginning of gonad development in prereproductive females occurs in May with a peak in summer
(Demestre and Fortuo, 1992) in the Balearic Basin.
In the present study, the trophic dynamics of Aristeus antennatus to the northwest and south of Mallorca (Balearic Islands) were
simultaneously studied in relation to suprabenthos-zooplankton
dynamics (see Cartes et al. (2008)) based on bi-monthly sampling.
The sampling covered two neighbouring regions (separated by
110 km) situated around the island of Mallorca. The areas were expected, before the sampling, to have different oceanographic conditions and clear differences in the population dynamics of the
red shrimp (Guijarro et al., 2008). The objectives pursued in this
study were: (1) to identify the importance of spatial patterns in
the diet and feeding of Aristeus antennatus; (2) to identify seasonal
variations; (3) to establish the main environmental variables controlling these trophic aspects; and (4) to establish possible links
between the trophic requirements of A. antennatus and its repro-
ductive cycle. We sought to demonstrate shifts in the diet and possible short-term coupling of general ecological processes with red
shrimp reproduction. We worked exclusively with gut contents,
excluding other methods used in trophic studies (e.g., isotopic
analyses) that are more focused on demonstrating average characteristics of the assimilated food.
2. Material and methods
2.1. Study area and sampling strategy
The study of changes in the feeding and diet of Aristeus antennatus was performed in two areas situated to the northwest and to
the south of Mallorca (Balearic Islands, Western Mediterranean:
Sller at 38980 N2570 E; 39140 N2760 E and Cabrera at
39680 N2180 E; 39810 N2370 E, (Fig. 1) The two sites are separated by ca. 110 km. The Sller site is in the Balearic Basin (between the coasts of Catalonia northeast Iberian Peninsula and
the Balearic Islands). The Cabrera site is close to the Cabrera Archipelago in the Algerian Basin. The Balearic and the Algerian basins
are separated by a ridge at ca. 600 m depth in the Mallorca channel.
The two sites have different oceanographic conditions (Millot,
1999; Lpez-Jurado et al., 2008), with different surface water
masses. Over the area as a whole there are two main surface water
types: Modied Atlantic Water (MAW) progressing northward
from the Algerian basin and Local Atlantic Water (LAW), a resident
water type in the Balearic basin (Lpez Garcia et al., 1994). Off
Cabrera, MAW predominates (particularly in springsummer),
and only weak frontal systems are formed. Off both Cabrera and
Sller, at mid-slope depths, where A. antennatus is distributed, variability is due to the occurrence of the Levantine Intermediate
water (LIW), which is present during the entire year with variable
seasonal inow. LIW seasonal inow was especially strong off
Sller at mid-bathyal depths (350550 m) in springsummer, after
circulating through the northern part of the Western Mediterranean from the Eastern Basin (Lpez-Jurado et al., 2008). The Western Mediterranean Intermediate Waters (WIW), which are formed
39
during winter in the Gulf of Lyons (Pinot et al., 2002), can arrive at
the Balearic Islands in early spring (Lpez-Jurado et al., 2008). WIW
is found just above the LIW, is not present every year and varies in
thickness yearly.
Regarding bottom character, the shelf is narrow and the occurrence of muddy terrigenous sediment is reduced due to low river
discharges. Muddy bottoms of biogenic origin dominate deeper,
between 350 and 750 m (own data, Cartes et al., 2008). The steepness of the slope was greater off Sller than off Cabrera (Fig. 1),
which could enhance differences in the distribution of some food
resources and their availability for red shrimp.
Six surveys were carried out to sample large megafauna, food
resources (macrofauna: suprabenthos and zooplankton) and
hydrographic features of the water column in August, September,
and November 2003, and in February, April and June 2004. Aristeus
antennatus were sampled from F/V Moralti Nou using a commercial
otter bottom trawl (Moranta et al., 2008). On each survey and in
each area, three hauls were carried out at 550, 650 and 750 m on
red shrimp shing grounds. All samplings were performed in daytime, comprising sunrise to noon (ca. 05.0012.30 GMT). Duration
of trawlings was 4560 min. Suprabenthos and meso-macrozooplankton were sampled from R/V Francisco de Paula Navarro. A total
of 46 Macer-GIROQ sledge and 461 m2-WP2 plankton net tows
were performed corresponding to six cruises (37 August 2003;
25 September1 October 2003; 1321 November 2003; 1420
February 2004; 713 April 2004; 2328 June 2004). During each
cruise, four stations (at depths ca. the 150, 350, 650 and 750 m isobaths) were sampled off both Sller and Cabrera (other details in
Cartes et al. (2008)). Distance between stations along the transects
ranged from ca. 6 to 18 km. The sledge sampled suprabenthos between 1040 and 5090 cm above the bottom. Macrozooplankton
in the water column was sampled using a WP2 net with a mouth
area of 1 m2, in horizontal-oblique hauls performed from between
13 and 90 m above the sea bottom to the sea surface. Both WP2
and sledges were equipped with 500 lm mesh size nets and were
trawled at ca. 1.5 knots. The durations of sledge hauls over the bottom were ca. 10 min; the duration of WP2 tows were 10 min close
Fig. 1. Map of the study area around the Balearic Islands indicating sampling locations and slope proles off Sller (NE of Mallorca) and off Cabrera Archiplago (S of
Mallorca). Proles constructed after calculating distance of trawls to the nearest coast.
40
to the bottom (horizontal haul) plus between 3 and 15 min during

the recovery of the net throughout the entire water column. The
same depth was maintained during all manoeuvres. Standard
2030 owmeters were attached to the mouth of nets to measure
the amount of ltered water and/or the distance/area covered by
each haul. All samples were collected during daytime.
2.2. Analyses of trends in abundance and in trophic data
Abundance of Aristeus antennatus was standardised to number
k m2 for each trawl (Guijarro et al., 2008). The centers of gravity
(COG: Stefanescu et al., 1992) for total abundance of small and
large (adult) females collected were calculated for each area and
cruise. This population COG is dened as
COG x1 z1 x2 z2 x3 z3 . . .=
xi ;
where xi is the calculated mean abundance of the species/population in the depth stratum i and zi is the mean depth of that stratum.
Specimens were classied in two/three size classes according to
previous information available on its diet (Cartes and Sard, 1989;
Cartes, 1994) and its biology (Demestre and Fortuo, 1992;
Demestre, 1995). To analyze feeding intensity and the energetic
content of diet we considered three size classes: (i) small with carapace length (CL) < 30 mm, which comprised all immature specimens and small males/females; (ii) mid-sized specimens, with CL
between 31 and 40 mm, comprising adult females and males, and
(iii) large, exclusively adult, females, with CL P 40 mm. For diet
only two groups were considered: (i) individuals smaller than
34 mm CL and (ii) individuals larger than or equal to 34 mm CL,
in the last case comprising only females. The diets of males and females were not considered separately after it was established that
diet depends mostly on size, not sex (Cartes and Sard, 1989), and
this increased the number of specimens analyzed per haul and size
classes for MDS analyses.
The following indices were calculated to measure feeding intensity and diet:
(1) Stomach fullness, F (stomach content weight/ predator
weight).
(2) W (weight of prey item/total gut content weight 100, in
wet weight).
These are traditional indices employed in the study of sh feeding (Hyslop, 1980) and used previously for decapods (e.g., in Cartes
(1994)).
The abundance of some dominant, broad taxa/species, key prey

in the diet of Aristeus antennatus, caught with sledges (e.g. polychates, bivalves and Calocaris macandreae among infauna, Boreomysis
arctica, and Natatolana borealis among suprabenthos) and WP2
nets (e.g., euphausiids, meso-bathypelagic decapods and Myctophidae among zooplankton/micronekton) and they were standardized to individuals/100 m2.
Degree of digestion was established for some key mesopelagic
prey (e.g., natantian decapods, Meganyctiphanes norvegica and
myctophids). Five degrees of digestion of food in stomachs were
considered, from undigested to very digested, similar to the scale
described by Aloncle and Dalaporte (1970) and adopted by the
authors (e.g., Stefanescu and Cartes, 1992) for deep water sh.
We counted prey of the two least digested values on that scale
appearing in stomachs.
Caloric content of the diet was also estimated, based on the values of caloric content of prey found in previous, similar studies
(Blaber and Bulman, 1987; Zagami et al., 1991; Madurell and
Cartes, 2005) performed on deep-sea fauna and summarized in
Table 1. Sum of caloric contents of prey found in stomachs was performed to estimate the whole caloric content of the diet. Caloric
content depends, among other variables, on the season, habitat,
and size of species and individuals (Hopkins, 1988; Zagami et al.,
1991). Therefore, we emphasized the data on caloric content from
a study of benthopelagic sh trophodynamics in a bathyal site in
the eastern Mediterranean (Madurell and Cartes, 2005), where
prey consumed by small benthopelagic sh (e.g., Macrouridae)
are similar in size to those consumed by A. antennatus.
2.3. Environmental and biological variables
The major part of environmental and biological variables is published in a monographic issue containing articles related with IDEA
project; however, these data were used in those articles for different objectives than in the current paper. There we detailed information as a function of depth and season on granulometry,
organic matter and Chl a satellite data (Cartes et al., 2008:
Madurell et al., 2008; Moranta et al., 2008), on T and S and uorescence (Lpez-Jurado et al., 2008) and on all biological variables
(Guijarro et al., 2008; see also Section 4). Only T, S (from 150 to
750 m) are again presented with some detail in this paper, though
only data from 550 to 750 m, simultaneously taken with red
shrimp, were considered for statistical analyses. To avoid, among
other things, duplicity in the publication of environmental/biological data, the rest of these variables are directly treated in the
Table 1
Caloric content (k cal/g DW) of the main prey found in Aristeus antennatus according to literature sources
Species/taxa
k cal/g DW
Source
Polychaeta
Crustacea
Gammaridae (Rhachotropis sp.)
Boreomysis arctica
Sergestes arcticus
Pasiphaea sivado
Plesionika martia
Euphausiacea
Cephalopoda
Teuthoidea
Myctophidae
Cyclothone braueri
Argyropelecus hemygymnus
Hygophum benoiti
Maurolicus muelleri
2.24
2.15
2.16
3.16
3.12
4.88
5.19
5.43
4.06
3.43
4.00
3.51
3.86
4.11
Madurell unp.
Madurell and Cartes (2005)
Blaber and Bulman (1987)
Blaber and Bulman (1987)
Zagami et al. (1991)
From polychaetes consumed by C. coelorhynchus

Range size similar to that consumed by A. antennatus

TL 2.6 cm, similar to that found in WP2 sampling
TL 1.65.0
TL 5.35.6
TL 4.45.1
Data were selected based on studies performed in similar habitats, and on prey sizes similar to those found in A. antennatus.
respective statistical analyses performed or included in the

Section 4.
Environmental variables were collected along the sampling
transects using SBE911 and SBE25 CTD prolers from R/V FP Navarro and with an SB37-SM mounted on the mouth of the commercial otter trawl (see Lpez-Jurado et al. (2008)). From each prole
we obtained a measure of temperature and salinity at 5 m above
the bottom (T5 mab; S5 mab). Fluorescence was recorded throughout
the water column. Maximum uorescence and depth of that maximum were obtained from the raw data.
Sediment for granulometric and organic matter analyses were
collected with a Shipeck grab at each station and on each cruise,
obtaining the variables: (1) percentage of mud (silt + clay) using
Sedigraph techniques (see Cartes et al. (2008) and Moranta et al.
(2008) for additional details); and (2) total organic matter (%OM)
content, calculated as difference between dry weight (DW: 80 C
during 24 h until reaching constant weight) and ash weight
(500 C in a furnace during 2 h).
Phytoplankton pigment concentration (ppc, mg Chl a m3) obtained from satellite imagery (http://reason.gsfc.nasa.gov/
Giovanni) was used as an indicator of the productivity in the area.
Monthly averages of ppc for the positions of the bottom trawls
were used, considering periods simultaneous with and 1, 2 and 3
months before each shrimp sampling.
Biological variables: The following were measured:
(1) red shrimp density (individuals k m2);
(2) gonadosomatic index (GSI), calculated as GSI = 100 (GW/W),
the gonad wet weight divided by total individual weight;
(3) hepatosomatic index (HSI) calculated as HSI = 100 (HW/W),
the weight of hepatopancreas divided by individual weight;
(4) condition index (K n ; Le Cren, 1951) calculated as K n = OW/
EW, the observed weight divided by the expected weight
estimated from length-weight relationships based on the
whole data set; and
(5) total lipids from the hepatopancreas, as percentage of dry
weight. GSI, HIS, K n and total lipids, taken from Guijarro
et al. (2008).
2.4. Data treatment

Data treatment was based on a total of 1672 Aristeus antennatus,
collected in 36 trawls and dissected in the laboratory immediately
after capture. The stomach contents of 909 of these individuals
were examined (Table 2). Mean stomach fullness (feeding intensity) was calculated for 36 hauls (number of specimens per haul:
1323 for small males and females; 921 for large males/mid-sized
females; 1023 for large females). Individuals were measured
(cephalothorax length CL in mm) and weighed to the nearest
0.01 g. The stomach content was weighed to the nearest 0.0001 g
and prey identied to the lowest possible taxonomic level under
a stereomicroscope (1040). When it was not possible to obtain
a direct weight for each prey item/type, weight of prey consumed
(WW, g) was estimated after the percentage volume occupied by
each prey in stomach contents was estimated using the subjective
points methods (Swynnerton and Worthington, 1940). The whole
stomach content weight was thereby partitioned for each preytype.
Data on diet (per haul: all stomachs from a single trawl combined) were analyzed by MDS techniques by separate in Cabrera
and Sller. Based on the resemblance matrix obtained by raw matrix data, Multi-Dimensional Scaling (MDS) was performed. The
number of specimens analyzed for diet per haul ranged between
8 and 19 specimens, both for small shrimp and for large females.
The similarity index used in MDS ordination was 1 r (Pearson
41
correlation coefcient, after log-transformation of the data). The

aggregation algorithm was the UPGMA (Unweighted-paired group
method average). PERMANOVA analyses (one factor consisting in
time periods off Soller, in depth intervals off Cabrera; 9999 permutations) were performed on the resemblance matrix (Bray-Curtis
distance) to test signicance of groups formed in MDS analyses.
To relate feeding and diet of the red shrimp with biological and
environmental variables, we examined possible correlations between diet, and stomach fullness (the dependent variables) and
the following environmental and biological possible explanatory
variables:
Environmental variables: (1) depth (m); (2) temperature (C) and
salinity (psu) at 5 m above the bottom (T5 mab; S5 mab); (3) %
mud; (4) percentage of organic matter (OM) in sediments; (5)
phytoplankton pigment concentration (mg Chl a m3)
obtained by satellite imagery; (6) Maximum uorescence and
(7) depth of maximum uoresence taken in IDEA02 and IDEA04.
Biological variables: (1) density (individuals k m2); (2)
Gonadosomatic index (GSI); (3) Hepatosomatic index (HSI);
(4) K n ; and (5) Total lipids from the hepatopancreas.
Only T and S are presented in this paper, while the other environmental variables are already published (Cartes et al., 2008;
Guijarro et al., 2008; Lpez-Jurado et al., 2008).
We searched for possible correlations to identify the main
explanatory variables for feeding intensity and diet at two levels:
(1) by simple non-parametric Spearman rank correlations (feeding
intensity and diet); and (2) by linear multiple-regression models
(MLR; after log transformation of the data), only for feeding intensity. All the analyses were performed using PRIMER6 & PERMANOVA+ (Clarke and Warwick, 1995; Anderson, 2001) and STATISTICA
7.0 softwares.
3. Results
3.1. Abundance and size-depth distribution
There were some differences in the depth distribution (COG) of
total abundance and females between Sller and Cabrera. The most
remarkable trend was a displacement off Sller of the COG of females, especially large females, to shallower depths (on average
between 50 and 100 m, in a range of 300 m, between 550 and
750 m) during February and June (less in August). COGs reached
the shallowest depths observed in February 2004 (Fig. 2). Differences in COG between Cabrera and Sller transects increased from
April to June (Fig. 2), when COG of both females and total abundance were displaced to greater depths off Cabrera (Fig. 2).
3.2. Spatio-temporal changes in stomach fullness
The feeding intensity of A. antennatus in the depth interval 550
750 m, showed seasonal uctuations (Fig. 3), which differed
depending on site (Cabrera and Sller) and size class. Off Sller,
all size-classes showed signicant increases in stomach fullness
from February to April 2004 and June 2004 (Fig. 3). Minimum signicant values were always recorded in February 2004 (Fig. 3). Off
Cabrera, the highest fullness values were recorded in November
2003 (signicant in comparison to February 2004) among small
and medium shrimp, while only adult specimens showed patterns
similar to that found off Sller, i.e., an increase in fullness in April
June 2004. In summary, maximum feeding activity of Aristeus
antennatus was often found in springearly summer off Sller, with
lower values in late summerautumnwinter. Off Cabrera the pattern was opposite for small and mid-sized shrimp, with the highest
feeding rates found in autumn (SeptemberNovember).
42
Table 2
Diet (W: grams wet weight per 100 individuals) of small and large Aristeus antennatus
Small
CL <34 mm
Large
Sller 581738 m
Cabrera 576752 m
CL P34 mm
Sller 581738 m
Cabrera 576752 m
NovFeb
AprJun
Aug-Sep
NovFeb
AprJun
AugSep
NovFeb
AprJun
AugSep
NovFeb
75
84
91
61
86
89
71
72
64
58
92
AprJun
66
Porifera
Hydrozoa
Siphonophora (Chelophyes
appendiculata)
Stephanoscyphus sp.
Polychaeta
Glycera spp.
Aphroditidae
Eunicida
Nephthydae
Polychaeta (other)
Crustacea
Copepoda Calanoida
Euphausiacea
Meganyctiphanes non/egica
Other (Nematoscelis megalops)
Decapoda
Gennadas elegans
Aristeus antennatus
Sergestes arcticus
Sergia robusta
Pasiphaea multidentata
Plesionika martia
Natantia unid. (Processa sp.)
Calocaris macandreae
Macrura Reptantia
Decapoda (postlarvae)
Mysidacea
Lophogaster typicus
Eucopia hanseni
Boreomysis arctica
Other (Parapseudomma
calloplura)
Amphipoda Gammaridea
Rhachotropis spp.
Eusirus longipes
Urothoe corsica
Tryphosites spp.
Other Lysianassidae
Harpinia spp.
Oedicerotidae
Synopiidae (Bruzelia typica)
Amph.Gammaridea unid.
0.045
0.029
0.002
0.015
0.439
0.300
0.191
0.073
0.041
0.237
0.125
0.100
0.151
0.006
0.035
0.21
0.14
0.06
0.022
0.633
0.030
0.009
0.117
0.477
0.017
0.326
0.005
0.060
0.261
0.002
0.755
0.033
0.12
0.128
0.041
0.432
0.048
0.605
0.029
0.083
0.154
0.339
0.021
1.179
0.025
0.266
0.177
0.240
0.471
0.002
0.892
0.044
0.009
0.193
0.102
0.544
0.002
5.602
0.01
0.53
0.521
0.932
3.608
4.302
0.041
0.515
1.832
1.912
2.462
0.134
1.154
0.498
0.011
0.666
0.011
3.683
0.059
0.708
0.853
1.054
1.009
0.012
4.897
0.062
1.286
0.899
0.396
2.254
0.006
3.474
0.123
0.161
1.502
0.419
1.269
0.004
0.017
0.004
0.062
0.092
0.270
0.019
1.529
0.314
0.132
0.056
0.384
0.019
0.547
0.068
0.032
0.009
0.300
2.129
0.207
0.367
0.227
0.331
0.005
0.763
0.193
0.128
0.188
0.219
0.544
0.080
0.050
0.020
0.004
0.022
0.292
0.097
0.288
0.323
0.007
0.002
0.157
0.046
0.183
0.696
0.349
0.112
0.202
0.159
0.01
0.154
0.03
0.004
0.024
0.110
0.285
0.168
0.063
0.013
0.003
0.105
0.259
0.093
0.322
0.067
0.206
0.175
0.023
0.164
0.023
0.251
0.350
0.216
0.271
0.012
0.052
0.031
2.094
0.227
0.119
1.339
0.565
1.471
0.823
0.470
2.021
2.246
0.034
0.237
1.724
0.123
0.605
0.237
0.346
0.018
0.695
1.674
3.903
0.367
1.526
0.283
0.017
1.229
0.014
0.087
0.311
0.708
0.358
0.822
1.936
0.604
0.178
0.144
0.04
0.607
0.539
0.854
1.092
0.643
0.729
0.588
1.135
0.457
0.546
2.232
0.084
1.256
0.663
0.241
0.894
0.017
0.036
0.039
0.021
0.100
0.004
0.044
0.097
0.015
0.044
0.018
0.218
0.056
0.249
0.056
0.025
0.127
0.027
0.090
0.033
0.013
0.031
0.137
0.003
0.024
0.034
0.011
0.063
0.038
0.015
0.018
0.019
0.080
0.044
0.038
0.011
0.019
0.015
0.039
0.211
0.135
0.005
0.008
0.042
0.041
0.035
0.067
0.074
0.002
0.008
0.013
0.049
0.091
0.019

0.098
0.090
0.010
0.038
0.023
0.079
0.007
0.008
0.036
0.007
0.002
0.005
0.005
0.005
0.015
0.007
0.007
0.018
0.054
0.007
0.123
0.051
0.039
0.011
0.028
0.092
0.009
0.042
0.011
0.013
0.051
0.020
0.007
Aug-Sep
0.019
0.005
0.021
0.066
0.026
0.526
0.054
0.318
0.734
0.32
0.050
0.638
0.057
0.056
0.080
0.094
0.017
0.010
0.136
0.045
0.053
0.342
0.164
0.008
0.022
0.054
0.024
1.181
1.998
0.122
0.050
0.628
0.053

0.051
0.013
0.009
0.395
0.481
0.089
0.004
0.213
0.003
0.037
0.036
0.032
0.003
0.270
0.002
0.002
0.003
0.007
0.450
0.008
0.009
0.030
1.220
0.050
0.016
0.007
0.028
0.002
0.096
1.188
0.031
0.011
0.005
0.012

0.071
0.001
1.377
0.076
0.033
0.015
0.041
2.757
0.027
0.004
0.103
3.704
0.002
0.014
0.003
3.439
0.009
0.009
0.043
5.999
0.013
0.002
0.007
0.055
8.309
0.005
0.021
0.002
0.055
6.097
0.512
0.007
0.035
0.038
0.003
0.003
0.033
0.010
0.048
0.006
0.054
0.038
0.017
0.065
0.010
0.012
0.003
0.002
0.077
0.006
0.001
0.020
0.066
0.002
0.172
0.009
0.010
0.005
0.054
0.029
0.031
0.006
0.030
0.115
0.088
0.004
0.023
0.143
0.002
0.013
0.009
0.083
0.007
0.095
0.048
0.010
0.002
0.056
0.008
0.179
0.086
0.004
0.024
0.026
0
0.027
0.098
0.073
0.043
0.009
0.017
0.005
0.107
0.052
0.035
0.026
0.017
0.004
0.012
0.048
0.159
0.010
0.010
0.075
0.023
0.013
0.089
0.007
0.023
0.065
0.140
0.076
0.056
0.006
0.158
0.057
0.017
0.004
0.073
0.023
0.007
0.094
0.019
0.012
0.002
0.012
0.001
0.004
0.001
0.076
0.018
0.064
0.252
0.572
0.286
0.016
0.079
0.014
0.216
0.002
0.318
0.383
0.136
0
0.02
0.129
0.335
0.289
0.033
0.007
0.069
0.592
0.010
0.022
0.023
0
0.013
0.016
0.123
0.309
0.117
0.005
0.004
2.733
0.373
0.138
0.011
0.030
0.345
0.165
0.019
0.011
0.010
0.060
0.078
0.141
0.078
0.005
0
0.008
0.094
0.302
0.169
0.004
0.005
0.078
0.023
1.469
0.006
0.029
0.083
0.010
0.022
0.191
0.418
0.034
0.016
0.012
0.469
0.012
0.358
0.125
0.288
0.159
0.051
0.075
0.029
0.033
0.072
0.002
0.046
0.097
0.008
0.158
0.121
0.022
0.241
0.022
0.055
0.013
0.023
0.187
0.216
0.116
0.231
0.077
0.189
0.606
0.061
0.023
0.296
0.027
0.078
0.006
0.224
0.339
0.263
0.083
0.021
0.804
0.030
0.008
0.398
0.011
0.339
0.013
0.003
0.026
0.016
0.006
0.011
0.061
0.348
0.017
0.001
0.167
0.073
0.226
0.061
0.320
0.034
0.002
0.087
0.003
0.330
0.013
0.143
0.055
0.074
0.013
3.345
0.564
1.433
0.129
0.126
0.532
0.006
0.027
0.015
4.336
0.252
0.227
0.028
0.004
1.189
2.926
0.275
1.755
0.051
0.022
0.448
0.277
0.515
1.265
0.032
0.007
1.408
0.477
0.010
3.564
0.067
0.930
0.095
0.002
0.254
2.062
0.143
1.322
0.102
0.236
0.369
Amphipoda Hyperiidea
Phronima sedentaria
Phrosina semilunata
Vibilia armata
Other Hyperiidea
Isopoda
Anthuridae
Natatolana borealis
Eurydice grimaldii
Gnathia sp.
Munnopsurus atlanticus
llyarachna sp.
Other (Desmosomatidae)
Tanaidacea (Apseudes sp.)
Cumacea
Leuconidae
Cyclaspis longicaudata
Platysympus typicus
Other Cumacea (Diastylidae)
Ostracoda (Cypridinidae)
Pycnogonida (Pallenopsis scoparia)
Mollusca
Caudofoveata
Escaphopoda
Bivalvia Taxodonta
Abra longicallus
Philine scabra
Cymbulia peroni
Alvania sp.
Benthonella tenella
Gastropoda unid.
Pteropoda
Cephalopoda (Teuthoidea)
Chaetognatha
Pyrosoma atlanticum
Thaliacea
Sipunculoidea
Echiurida
Echinodermata
Ophiuroidea
Echinoidea
Osteichthyes
Myctophidae
Cyclothone braueri
Osteychthyes unid. (remains)
Foraminifera
Nylon threats
Unidentied (gelatinous)
Plant remains (Posidonia)
Unidentifed
Other (e.g. Insecta)
Diet is expressed in base of seasonal groups deduced from the MDS analyses. n: number of individuals analyzed; (): <0.001 g.
43
44
Aug
2003
Set
2003
Nov
2003
Feb
2004
Apr
2004
Jun
2004
560
Total abundance
600
Cabrera
640
S
Sller
680
720
560
Small females
Depth (m)
600
Cabrera
640
Sller
680
720
560
Large females
600
640
Cabrera
Sller
680
720
Fig. 2. Center of gravity (COG) as a function of season for total abundance and abundance of small and large females of Aristeus antennatus off Cabrera and Sller.
3.3. Dietary analyses

A taxonomic range including 157 different prey items was identied in stomach contents of A. antennatus (98 categories identied
to genus/species level) (Table 2). Among small shrimp
(CL < 34 mm) the main prey off Sller were mesopelagic natantian
decapods (Gennadas elegans, Sergestidae, Pasiphaea multidentata),
sh (Myctophidae, Cyclothone braueri, sh remains) and the isopod

Natatolana borealis. Some other prey were important seasonally,
e.g., jellysh remains (probably salps) in AugSep and NovFeb,
polychaetes and Calocaris macandreae in NovFeb and AprJun,
and hyperiids and Meganyctiphanes norvegica in AprJun. Main
prey off Cabrera were polychaetes, and N. borealis. Bivalves were
also important prey there in comparison to Sller. Mesopelagic
0.01
CL
08, 02-06
0.008
30 mm
11, 02 11, 02
0.006
11
11
0.004
04,06
11
04,06
0.002
0
0.01
08, 02-06
08, 02
0.008
02
11
11
0.006
11
11
F
0.004
04,06
04
0.002
40 > CL > 30 mm
0
0.01
0.008
02
06
08
02
0.006
0.004
11, 04,06
0.002
CL
40 mm
Aug
2003
Sep
2003
Nov
2003
Feb
2004
Apr
2004
Jun
2004
Fig. 3. Fullness of Aristeus antennatus off Sller (---------) and Cabrera (

) as a
function of size (CL, cephalotorax length, mm) and seasonal periods. Numbers
above/behind each point means the periods (months, e.g., 08 = august) for which F
values were signicantly different in paired comparisons (1-way ANOVA; Tukeys
test; p < 0.05). In dark tone for Cabrera, in light tone for Sller. Parts of this gure
appeared previously in Cartes et al. (2008).
natantians (in AprJun) and sh (in Nov-Feb, myctophids) were

seasonally important off Cabrera, while siphonophores were
important in summer (AugSep).
Both off Cabrera and Sller dominant prey (in wet weight) in
the diet of large (Cl P 34 mm) females were polychates (mainly
Aphroditidae and Nephthydae). In numbers the most abundant
polychate taxon consumed by A. antennatus was Glycera spp. Other
dominant prey by weight in both areas were mesopelagic decapods, N. borealis, sh and jellysh (gelatinous) remains. Moreover,
off Sller euphausiids and hyperiids were important, especially in
Apr-Jun, while cephalopods were mainly consumed in AprJun/
NovFeb. In this same period off Cabrera, bivalves and echinoderms were among the dominant prey in the diet, and more Calocaris macandreae were consumed there all year round. The
mesopelagic decapod Sergia robusta was an especially signicant
food off Sller in Apr-Jun.
MDS analyses summarized trends in diets based on a multiprey
analysis. MDS were performed for small (CL < 34 mm) and large
specimes (CL between 34 and 62 mm) separately. The diet of A.
antennatus was primarily inuenced by a seasonal component off
45
Sller. Three (among small shrimps) and two (among large shrimps)
groups corresponding to AugustSeptember 2003 and to AprilJune
2004 (both in small and large shrimps) and November 2003
February 2004 (only among small shrimps) were identied in
MDS analyses in that area. Hauls from November 2003February
2004 were situated between these two groups (signicant differences in the distribution of AugustSeptember 2003, AprilJune
2004, and November 2003February 2004 hauls: PERMANOVA test:
Pseudo Fsmall = 7.44, p < 0.001; Pseudo Flarge = 4.74, p < 0.01; all posthoc tests signicant at p < 0.01 between all paired comparisons
excluding November 2003February 2004 vs. the two other groups
among large shrimps). So, a similar seasonal trend was observed
both for small individuals (Fig. 4a) and for adult females (Fig. 4b).
Off Sller we found the most remarkable seasonal shift in the diet
of small (CL < 34 mm) A. antennatus in AprilJune 2004, with a signicant increase (t test, p < 0.05) of mesopelagic prey (e.g. the
euphausiid Meganyctiphanes norvegica, the decapods Gennadas elegans, sergestids, and Pasiphaea multidentata, hyperiids and myctophids) (Table 2). Other prey hardly varied (e.g., polychaetes, N.
borealis, bivalves) or even decreased from AugustSeptember/
NovemberFebruary to AprilJune (e.g., gastropods, siphonophores,
jellysh remains, probably salps). Seasonal changes in the diet of
large shrimp (CL P 34 mm) were similar to those found in small
ones. The most remarkable trend was the increase of Sergia robusta
and of euphausiids in gut contents in AprilJune 2004. However,
other mesopelagic prey even decreased in this period (e.g., Gennadas
elegans, Pasiphaea multidentata, myctophids) in comparison to AugustSeptember/NovemberFebruary. Other prey hardly varied
(e.g., the whole polychaete groups, N. borealis) or they decreased
(e.g., bivalves, gastropods, siphionophores, jellysh remains).
Off Cabrera, by contrast, the MDS analysis segregated hauls by
depth, never by seasonal groups, again both for juvenile/males
(Fig. 4a) and adult females (Fig. 4b). Hauls performed between
581 and 662 m were grouped and separated from the deepest samples collected at 745752 m (signicant differences in the distribution of the two depth groups both among large and small shrimps:
PERMANOVA test: Pseudo Fsmall = 9.77, p < 0.001; Pseudo
Flarge = 9.03, p < 0.001). The diet both of small and large specimens
at 581662 m off Cabrera included substantial quantities of the
isopod Natatolana borealis (30% of the diet of large; 24.6% in small)
and also (in decreasing order) polychaetes (Eunicida, Aphroditida),
myctophids and natantian decapods. Sergia robusta and Plesionika
martia were especially consumed by adults. Deeper, at 750 m, diet
was not as dependent on a single prey as at 581662 m. Meganyctiphanes norvegica was the most important prey for small shrimp
(only 7.3% of diet weight), with polychaetes (mainly Nephthydae),
natantian decapods and bivalves. Among large shrimp unidentied
polychaetes (5.9% of diet) and eunicids were dominant, with significant amounts of brachyurans and myctophids.
Caloric content of the diet, based on values compiled in Table 1,
also showed differences between Sller and Cabrera (Fig. 5). In all
size categories there was an increase in the energy content of prey
ingested from February to June 2004 off Sller. Adults showed similar levels of energy content in their diet in SeptemberNovember
2003 than those found in February and June 2004. Off Cabrera
there was not a sharp increase of energy intake in these same periods, and the highest caloric content was recorded among medium
shrimp in November and among adults in June, while no important
changes were recorded among small specimens.
3.4. Distribution of prey
The abundance and depth ranges of the main prey consumed by
A. antennatus off Sller and Cabrera (Table 3) overlapped with the
distribution of A. antennatus. However, most prey-species showed
higher abundance in the other area than in the one where prey
46
S06-730
Sller
1.0
1.0
S06-659
S11-655
0.4 S08-581
S11-593
S04-657
S04-726 S06-601
S02-593
S09-653
Dim 2
Dim 2
S11-734
-0.2
C09-752
C11-659
S02-655
S08-734
C06-750
C11-745
Cabrera
S02-735
C04-657
C06-662
C08-659 C06-584
C11-583
C08-583 C04-586
C02-580
C09-661
C02-650
-0.5
C08-751
C02-750
C04-752
S04-605
S09-738
-0.8
S09-600
C09-584
-1.4
-1.4
-0.8
-0.2
0.4
1.0
-2.0
-1.0
1.6
-0.4
0.2
Dim 1
S04-605
S11-593
S11-655
1.0
S06-730
S08-581
Dim 2
Dim 2
S04-726
S09-653
-0.2
S06-659
S08-659
S09-738
S04-657
S11-734
C04-586
C09-661
C02-650
C04-657
C08-583C11-583C11-659
C06-662
C08-659
C06-584 C02-580
C08-751
-0.5
C04-752 C11-745
C06-750
C02-750
S02-655
-0.8
S06-601
C09-584
S08-734
-1.4
-1.2
1.4
Cabrera
S09-600
1.0
0.4
0.8
Dim 1
C09-752
S02-593
Sller
S = 0.171
S = 0.192
S08-659
-0.6
0.0
S = 0.221
0.6
1.2
Dim 1
-2.0
-1.2
-0.6
0.0
0.6
S = 0.131
1.2
1.8
Dim 1
Fig. 4. MDS analyses performed for (a) small (CL < 34 mm) and (b) large (CL P 34 mm) A. antennatus off Cabrera and off Sller. In haul codes C = Cabrera; S = Sller; numbers
before dashes are month, numbers after dashes are depth of haul. S is stress.
were mainly consumed. In fact only Natatolana borealis and Sergestes articus, mainly consumed by A. antennatus off Cabrera, and Gennadas elegans, more signicantly consumed off Sller, showed
coincidence between their abundance in stomach contents and in
the environment. Some prey were not well sampled with our gear
(sledges and WP2 net), the case of Sergia robusta with only two
specimens caught on all cruises.
The maximum size of some prey found in stomachs was higher
than that of specimens collected with the sledge-WP2. This was
the case of Natatolana borealis, Meganyctiphanes norvegica, Sergestes
arcticus and especially Sergia robusta, Pasiphaea multidentata and
Myctophidae (main species found in guts: Lampanyctus crocodilus,
Notoscopelus elongatus and Ceratoscopelus maderensis). This suggests net avoidance by large specimens of these prey consumed
by Aristeus antennatus.
Between 16% and 23% (in number) of identied prey among
undigested mesopelagic prey (e.g., decapods, myctophids) must
have been eaten by A. antennatus before (01 h) sunrise and to 4 h
after sunrise Fig. 6. The occurrence of somewhat digested and very
digested prey at these periods indirectly indicated nocturnal feeding
on these mesopelagic fauna by the red shrimp. The large proportion
of myctophids eaten at midday, 810 h after sunrise (appearing in
Fig. 6) was based in a low number (n = 5) of ingested sh.
3.5. Trends in environmental variables
Temperature (T) 5 m above bottom, varied with depth and site
and ranged between 12.94 C at 650 m to 13.47 C at 150 m. On
average, close to the bottom, there was an increase in T of 0.2

0.3 C from the shelf slope break (150 m) to 650750 m on the
slope. The seasonal pattern was determined by depth. On the
shelf slope break (the level of Winter Intermediate Water,
WIW) maximum T was recorded in February decreasing in
AprilJune to around 13 C. This decrease was recorded earlier
off Sller (April) than off Cabrera (Fig. 7a). Deeper at 550
750 m (a level including the lower depth distribution of Levantine Intermediate Water, LIW) there was a decrease of T from
November 2003 to April 2004 off Sller, with a further sharp increase in June 2004, more obvious toward 550650 m. Over the
deepest part of the slope (550750 m), more regular conditions
of T through the year were recorded off Cabrera in comparison
to Sller.
Salinity (S) 5 m above bottom ranged from 38.28 ppm at 150 m
(November 2003) to 38.51 ppm at 350 m (June 2004). In general S
increased from the shelf-slope break to bathyal depths to 550 m
(Fig. 8), then slightly decreasing. The seasonal pattern was different
depending on depth. S increased (above 38.4 psu) from February to
AprilJune at 350 m, and later (June) deeper (550750 m). Maximum S was regularly recorded in June 2004 both at 150 m and
on the slope. However, at 150 m maximum values were also recorded in November 2004. S was in general higher off Cabrera, except at 350 m depth where values were rather similar at both sites
(Fig. 7). In general T and S values decreased with depth (excluding S
at between 150 and 350 m), especially off Cabrera below 550 m.
Also both variables exhibited higher seasonal uctuations of Sller
than off Cabrera (Fig. 7).
47
1.0
0.8
CL < 30 mm
0.6
0.4
0.2
0.0
2.4
Kcal/g WW
2.0
1.6
1.2
0.8
40 > CL > 30 mm
0.4
0.0
6.0
5.0
4.0
3.0
2.0
CL > 40 mm
1.0
0.0
Aug
2003
Sep
Se
2003
Nov
2003
Feb
2004
Apr
Ap`
2004
Jun
2004
Fig. 5. Trends in the caloric content (in k cal/g WW) in the diet of Aristeus
antennatus off Sller (---------) and Cabrera (
) as a function of size and annual
period. CL = cephalotorax length (mm). Parts of this gure appeared previously in
Cartes et al. (2008).
3.6. Relationships between feeding/diet and biological/environmental

variables
Relationships were treated separately for Cabrera and Sller by
means of univariante (Spearman r) and multivariate (MLR models)
statistical methods. Off Cabrera, fullness (F) was positively corre-
lated (non-parametric Spearman rank; p < 0.05) with T close to

the bottom for small and large shrimp (Table 4). Other variables
were correlated with F depending on shrimp size. Among small
shrimp, F was positively correlated with uorescence and negatively with shrimp density (higher F with lower density). Among
medium-sized shrimp, there were negative correlations between
F and Chl a readings 2 months before sampling (Table 4). In addition to the variables cited, depth was negatively correlated with F
among large shrimp.
Among small sizes off Sller, F was positively correlated with
Chl a readings taken simultaneously and 12 months before, also
with gonadosomatic index (GSI), and with shrimp biomass (Table
4). Among mid-sized shrimp, only a negative r between F and Chl
a 1 month before was found, while no variable was correlated with
F in large shrimp.
Regarding changes in the diet of A. antennatus (MDS dimensions: DIM1, DIM2), r was signicant between DIM1, DIM2 and a
number of environmental variables, though these correlations varied depending on the area and size of shrimp (Table 4). Among
small (CL < 34 mm) shrimp, DM1 off Cabrera was signicantly correlated with fullness (F), depth and T close to the bottom, whereas
DIM2 was only correlated with lipid content (n = 9; r = 0.717;
p = 0.03). Among large (CL P 34 mm) shrimp, DIM1 of diet was
correlated with F, depth and with T and S close to the bottom, without any signicant r in DIM2.
Off Sller the diet of small shrimp was signicantly correlated
with F, uoresence and Chl a (12 months before) and also with
density, while DIM2 was correlated with HSI (n = 12; r = 0.783;
p = 0.003). Among large females, DIM1 was signicantly correlated
with F, with Chl a (12 months) and with the hepatosomatic (HSI)
and K n indexes, while DIM2 was related with depth (n = 17;
r = 0.520; p = 0.03), mean S (n = 17; r = 0.576; p = 0.02), and the
gonadosomatic index (GSI) (n = 11; r = 0.630; p = 0.007).
From MLR models, Chl a in surface waters off Cabrera taken 2
months before sampling was the main explanatory variable of fullness for small Aristeus antennatus (Table 5), together with shrimps
density and GSI. All these variables showed, however, signicant
negative correlations. Salinity and mean size (W) explained fullness for mid-size shrimp (positive r) as did depth (negative r) for
large shrimp. Off Sller higher fullness was found among small
shrimp with decreasing density, and with increasing surface Chl
a taken 2 moths before. Among mid-sized Aristeus, higher F came
with increasing depth and decreasing surface Chl a taken simultaneously, whereas no variable signicantly explained fullness for
large specimens. The variance accumulated by MLR models ranged
between r2 = 0.418 and 0.549.
In summary we found a higher dependence of fullness and diet
with T and S off Cabrera than off Sller, and that is related to depth.
Table 3
Abundance (ind./100 m2) and depth ranges of main prey (in g wet weight, W/100 ind.) consumed by Aristeus antennatus off Sller and Cabrera
Prey-species
Boreomysis arctica
Natatolana borealis
Meganyctiphanes norvegica
Euphausia krohni
Nematoscelis megalops
Sergestes arcticus
Gennadas elegans
Sergia robusta
Pasiphaea multidentata
Calocaris macandreae
Myctophidae
Depth range m
Suprabenthos
Suprabenthos
Zooplankton
Zooplankton
Zooplankton
Zooplankton
Zooplankton
Micronekton
Micronekton
Infauna
Micronekton
335760
134748
363752
150753
156760
666760
347760
749
670752
365752
650753
COG m
N ind./100 m2
W g/100 ind.
Size (n) mm
Sller
Cabrera
Sller
Cabrera
Sller
Cabrera
Water column
699
465
713
569
661
728
660
727
613
711
682
590
692
599
657
676
698
749
732
715
703
18.2
0.11
0.24
2.03
4.74
0.06
0.57
0.01
0.28
0.07
22.4
0.28
0.89
1.96
6.14
0.21
0.51
0.08
0.04
0.69
0.11
0.21
3.61
1.44
0.20
0.25
0.20
1.72
2.43
1.43
0.55
3.77
0.13
8.06
0.84
0.12
0.19
0.77
0.45
1.32
0.90
0.45
2.24
1.37.4(1025)
1.821.2(48)
3.69.2(113)
1.16.4(330)
0.88.9 (693)
3.411.1 (27)
1.510.0(112)
6.39.3 (3)
2.613.8(13)
2.39.8 (57)
10.645.4(12)
(n): number of specimens used to calculate size range. Suprabenthos/infauna collected with sledges, zooplankton/micronekton with WP2 nets.
Guts
4.27.3(14)
4.024.5 (139)
3.210.6 (61)
4.85.3 (8)
2.66.9 (8)
5.912.9 (7)
9.9(1)
6.618.5 (19)
5.018.0 (13)
5.38.7(13)
15.052.6 (20)
48
40
Mesopelagic decapods
30
23
20
33
30
43
24
69
10
0
40
sunrise
0-2h
2-4h
4-6 h
6-8 h
8-10 h
Myctophidae
small meso-bathypelagic decapods and myctophids were more

important in the diet of the insular areas. Off the coasts of
Catalonia, benthic prey are particularly important around submarine canyon heads, where higher foregut fullness (more food consumption) was also recorded (Cartes, 1994; Fig. 8). Aristeus
antennatus, therefore, can eat such a wide variety of taxa that it
can exploit prey of distinct ecological habitats depending on local
availability. Both around the Catalan canyons and the Balearic islands, micronekton (euphausiids and small decapods) marked the
seasonal changes found in the diet of the red shrimp, to 1200 m
(Cartes, 1994), which is the assumed boundary of distribution of
mesopelagic fauna (Goodyear et al., 1972; Stefanescu and Cartes,
1992). Below this depth the inuence of the mesopelagic assemblages on the diet of A. antennatus and other demersal species is
low (Cartes, 1998).
30
%prey (n)
21
4.1. Spatio-temporal changes in the diet and prey availability:

inuence of environmental factors
20
22
23
39
10
11
0
sunrise
0-2h
2-4h
4-6 h
6-8 h
8-10 h
40
M. norvegica
30
28
20
16
28
40
10
22
16
0
sunrise
0-2h
2-4h
4-6 h
6-8 h
8-10 h
Fig. 6. Percentage undigested prey (by number) in gut contents of Aristeus

antennatus in hauls performed around sunrise and to 10 h after sunrise. E1
and E2
indicating two steps of undigested prey from fresh (E1) to slightly
digested (E2); numbers above bars indicate the total number of prey counted in
each period.
Depth was an important factor affecting diet off Cabrera (see

trends in MDS analyses), and in this area depth was in turn correlated with F among large females (higher fullness shallower:
r = 0.571; p = 0.02; n = 16) and T (lower T deeper: r = 0.825;
p = 2 105; n = 18). Off Sller, where seasonal forces were more
important in shifting the diet, variables explaining changes in fullness and diet were more strongly linked to temporal changes in
water column productivity (e.g., positive r with Chl a readings
and uorescence) and with shrimp biology (GSI, HSI, and kn).
4. Discussion
Aristeus antennatus has the most diverse diet found among
deep-sea megafauna in the Mediterranean (H0 = 5.58: Cartes,
1994), based on the large variety of small benthic/suprabenthic
and pelagic macrofauna and micronekton. Compared with other
decapods and sh cohabiting with it, this is the species in which
benthos (e.g., polychaetes, ophiuroids) contributes the most to its
diet (Lagardre, 1971; Relini-Orsi and Wrtz, 1977; Cartes,
1994). However, when comparing diets over the slope off the
Catalan coast with those around the Balearic Islands, euphausiids,
Off the Balearic Islands dependence upon mesopelagic prey was

more important off Sller than off Cabrera. This must be associated
with higher abundance of (meso)zooplankton off Sller (Cartes
et al., 2008) that is linked in turn to higher occurrence of oceanographic frontal systems and eddy formation to north of Mallorca
(Lpez-Jurado et al., 2008). Some differences between Sller and
Cabrera in the sources of food for suprabenthos (prey of small A.
antennatus) were evidenced based on the isotopic composition of
suprabenthos (Fanelli, 2007; Madurell et al., 2008). Correlations
between d13C and d15N of suprabenthos increased form February
to June 2004 off Sller, pointing to exploitation of a unique food
source after the main peak of primary production in surface occurring in FebruaryMarch (Cartes et al., 2008). Off Cabrera, by contrast, d13C vs. d15N did not follow the same pattern, d13Cd15N
correlations were lower than those recorded off Sller and the
highest d13Cd15N correlation were found in SeptemberNovember
2003 (Fanelli, 2007). This dependence upon mesopelagic prey
seems consistent with a longer duration for Chl a peaks (an indirect
indicator of surface production) off Sller in comparison to Cabrera,
which may probably favor higher coupling between primary production and zooplankton/suprabenthos in this area. Finally, a parallel increase in %OM (total organic matter) in sediments was
found in AprilJune 2004, also higher off Sller (Cartes et al.,
2008). All these factors explain the signicant correlations found
in MLR models between fullness/diet vs. Chl a in this area.
Oceanographic differences between Cabrera and Sller included
a higher ux and thickness of Levantine Intermediate Water (LIW
with salinity > 38.45 psu) off Sller at depths of 300 to 500 m
(Lpez-Jurado et al., 2008) and of Winter Intermediate Water
(WIW with temperature 613 C) distributed above LIW between
150350 m. WIW is formed during winter to the north of Balearic
Islands, in the Gulf of Lyons (Pinot et al., 2002). Winter water has
been recently linked to cascading events generated in the Gulf of
Lyons (Palanques et al., 2006), by which submarine canyons acts
not only as channels for discharges of short-torrential rivers found
along the coasts of Catalonia but for water masses down the slope.
As a consequence, both LIW and WIW ow was particularly strong
to the north of Balearic Islands (Sller) in late winterspring
(Lpez-Jurado et al., 2008), both increasing from February to April
and especially in June. These water masses can carry nutrients
from most productive areas in the Gulf of Lyons, enhancing phytoplankton and zooplankton production. LIW was also present off
Cabrera, and maximum inux (SeptemberNovember 2003:
Lpez-Jurado et al., 2008) again coincided with maximum changes
in the isotopic signal of suprabenthos there (Fanelli, 2007), hence
probably inducing changes in trophic web dynamics. Water masses
conditions (T and S) showed higher variability at shallower (150
49
b
13.20
13.60
150 m
Increasing depth
13.15
13.40
13.10
13.20
350 m
13.05
13.00
13.00
12.95
12.80
n2
0
04
4
Ju
pr
20
0
A
Fe
b2
0
04
3
00
ov
2
p2
0
Se
38.51
03
3
00
ug
2
n2
0
04
4
A
Fe
Ju
04
b2
0
00
pr
20
0
03
ov
2
N
Se
38.50
p2
0
ug
2
00
12.90
350 m
38.49
38.30
150 m
38.47
38.10
n2
0
04
4
Ju
pr
20
0
A
04
b2
0
Fe
ov
2
00
03
p2
0
Se
ug
2
00
04
n2
0
Ju
04
pr
20
0
Fe
b2
0
00
ov
2
N
03
p2
0
Se
ug
2
00
38.45
Fig. 7. Temperature (T) and salinity (S) of the water close to the bottom (5 m above bottom) at the three depth ranges studied off Sller (s) and Cabrera (--). At
150350 m (a) and at 550750 m (b). In general both T and S decreased with depth at 550750 m, therefore, lines at the top of plots represent the shallowest depth sampled,
and at the bottom the deepest. Redrawn from Cartes et al. (2008).
La Berenguera
Canyon (450 m)
0.04
600-650 m
0.02
D
ec
l
Ju
A
pr
M
ar
Fig. 8. Temporal changes of fullness (F CI 95%) in A. antennatus off the Catalan

coasts close to Barcelona during 1991 (April/December samples) 1992 (March/
July) at two stations situated in a submarine canyon (average depth, 450 m) and at
600650 m on the slope. Each point is the mean fullness calculated for >30
individuals/haul.
350 m) than at deeper (650750 m) depths. Variability was higher

off Sller and the strongest decrease of T found at 150 m occurred
earlier (in April) there than off Cabrera (see Fig. 7a). As main
changes in stomach fullness and diet of A. antennatus occurred
from February to April 2004, possible inuence of oceanographic
processes in the trophodynamics of red shrimp seems more likely
linked to ow of surcial water masses (WIW) than to deep waters
(LIW). Off Sller the COG of females was shallower than off Cabrera, closer to a possible inuence of water masses ow coinciding
with the period (AprilJune) of WIW and LIW occurrence in the
area (Lpez-Jurado et al., 2008). Deep mesopelagic prey consumed
by A. antennatus (on average distributed > 400 m depth: see below)
must trophically exploit zooplankton distributed at shallower

depths (150350 m) by means of daily migratory movements. In
any case, both WIW and LIW are found above the daylight distribution of A. antennatus (550750 m) on the Balearic slope, so hypothetical upward nocturnal migration by A. antennatus (discussed
below and suggested by Cartes et al., 1993 off Catalan canyons)
in search of prey could also explain a spatial coincidence between
the distribution of A. antennatus and zooplankton associated with
the seasonal inux of LIW and WIW.
Aristeus antennatus preferently consumed micronektonic prey
(e.g., Myctophidae, Sergia robusta, Meganyctiphanes norvegica. . .)
off Sller. However, most of these species were more abundant
in WP2 samplings off Cabrera. This lack of correlation between
prey in guts and in the environment could be explained by (i) A.
antennatus and their prey were not distributed at the same depth
range at the time of samplings (performed in daylight); (ii) possible
net avoidance by micronekton. Feeding rhythms have been reported in A. antennatus, consisting of a higher consumption of prey,
mainly suprabenthos, at night (Cartes, 1993b; Maynou and Cartes,
1997). Off the Balearic Islands Aristeus antennatus could also prey
on micronekton in synchrony with daynight movements of this
fauna. Only 3 of the 98 species identied in gut contents of A.
antennatus (two mysids: Boreomysis megalops, Lophogaster typicus
and one amphipod, Hippomedon massiliensis) showed a shallower
distribution than that of the red shrimp during daytime (unpublished data), and only one (L. typicus) was a relatively important
prey. However, most mesopelagic prey had a shallower maximum
peak of abundance in the environment than in the depth interval
where A. antennatus were caught. Thus, M. norvegica showed average weighted mean depths (abundance peaks) at 440500 m in the
NW Mediterranean (Sardou et al., 1996; Cartes, 1998), whereas we
sampled offshore, over bottoms of 576752 m. Myctophids probably followed a similar pattern. Because most mesopelagic species
50
Table 4
Relationships (Spearman rank correlation: r) between stomach fullness (F) and rst dietary dimension (DIM1) of Aristeus antennatus and environmental/biological variables
F
Cabrera
Sller
Small
Depth
T
S
OM(%)
% Mud
Redox
Depth uorescence
Fluorescence
Chl a (2 months)
Chl a (1 month)
Chl a sim
GSI
HSI
Kn
Lipids
D (ind./km2)
B (gl/k m2)
DIM1
Medium
Small
Medium
Large
18
17
17
11
10
11
7
7
18
18
18
17
12
18
9
18
18
0.401
0.495
0.233
0.455
0.297
0.627
0.546
0.764
0.384
0.271
0.086
0.319
0.007
0.059
0.200
0.558
0.515
ns
0.043
ns
ns
ns
0.039
ns
0.046
ns
ns
ns
ns
ns
ns
ns
0.016
0.029
18
17
17
11
10
11
7
7
18
18
18
17
12
18
13
18
18
0.402
0.455
0.104
0.627
0.624
0.700
0.400
0.691
0.489
0.393
0.148
0.270
0.175
0.096
0.516
0.218
0.218
ns
ns
ns
0.039
ns
0.016
ns
ns
0.040
ns
ns
ns
ns
ns
ns
ns
ns
16
15
15
11
10
11
7
7
16
16
16
16
11
14
9
16
16
0.571
0.670
0.488
0.373
0.467
0.818
0.036
0.327
0.217
0.160
0.171
0.168
0.245
0.389
0.017
0.347
0.347
0.021
0.006
ns
ns
ns
0.002
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
17
18
18
11
11
10
10
10
6
17
17
17
18
18
9
12
11
0.456
0.147
0.285
0.167
0.539
0.239
0.313
0.129
0.905
0.850
0.762
0.750
0.129
0.084
0.350
0.345
0.717
ns
ns
ns
ns
ns
ns
ns
ns
0.013
0.00002
0.0004
0.001
ns
ns
ns
ns
0.013
17
18
18
11
11
10
10
10
6
17
17
17
17
18
13
18
18
0.142
0.022
0.116
0.257
0.021
0.453
0.485
0.541
0.000
0.543
0.241
0.170
0.459
0.028
0.582
0.370
0.295
ns
ns
ns
ns
ns
ns
ns
ns
ns
0.024
ns
ns
ns
ns
0.037
ns
ns
17
17
17
10
10
9
5
5
16
16
16
17
11
17
13
17
17
0.446
0.412
0.409
0.042
0.297
0.250
0.462
0.564
0.244
0.339
0.373
0.025
0.309
0.456
0.148
0.130
0.100
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
Small
Large
Cabrera
F
Depth
T
S
OM(%)
% mud
Redox
Depth uorescence
Fluorescence
Chl a (2 months)
Chl a (1 month)
Chl a sim
GSI
HSI
Kn
Lipids
D (ind./k m2)
B (gl/k m2)
Large
Sller
Cabrera
Sller
18
18
17
17
11
10
11
7
7
18
18
18
17
12
18
9
18
18
0.515
0.661
0.514
0.320
0.291
0.455
0.536
0.600
0.709
0.068
0.085
0.211
0.027
0.182
0.344
0.050
0.141
0.148
0.029
0.003
0.035
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
18
18
18
18
10
10
9
6
6
17
17
17
18
12
18
9
18
18
0.480
0.125
0.238
0.418
0.176
0.103
0.467
0.618
0.971
0.585
0.566
0.341
0.273
0.301
0.294
0.533
0.496
0.350
0.044
ns
ns
ns
ns
ns
ns
ns
0.001
0.014
0.018
ns
ns
ns
ns
ns
0.036
ns
16
18
17
17
11
10
11
7
7
18
18
18
16
11
14
10
18
18
0.535
0.669
0.573
0.520
0.309
0.467
0.582
0.655
0.655
0.318
0.114
0.220
0.235
0.336
0.213
0.248
0.253
0.358
0.033
0.002
0.016
0.032
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
ns
17
17
17
17
10
10
9
5
5
16
16
16
17
13
17
13
17
17
0.686
0.074
0.086
0.230
0.455
0.055
0.517
0.821
0.718
0.602
0.595
0.324
0.147
0.808
0.608
0.451
0.216
0.380
0.002
ns
ns
ns
ns
ns
ns
Ns
Ns
0.014
0.015
ns
ns
0.0008
0.010
ns
ns
ns
Table 5
MLR models deduced for stomach fullness (F) of Aristeus antenntus as a function of environmental/biological explanatory variables
Small
MLR1
MLR2
Medium
MLR1
MLR2
Large
MLR1
MLR2
Cabrera
Sller
Cabrera
Sller
Cabrera
Sller
r2
Var.
17
0.508
Int
Chl a (2 months)
D
GSI
Int
D
Chl a (2 months)
18
12
17
16
17
0.647
0.546
0.325
0.375
0.374
Int
S
W
Int
Chl a (2 months)
Int
Depth
Int
Kn
p-level
0.0147
0.0008
0.0808
0.0005
0.0078
3.420
0.041
0.023
0.013
0.037
2.5E-04
0.018
0.033
0.038
3.1E-04
0.018
0.046
0.022
0.022
0.031
0.337
0.047
0.012
0.021
0.069
0.018
0.85
0.96
0.82
1.00
1.00
0.67
0.62
0.63
1.00
0.98
Models built for 3 size categories (small, medium, and large) and for the two areas (Sller, Cabrera) separately. GSI: gonoadosomatic-index; K n : condition index; D: density;
W: mean size (B/D). n: number of cases; r2: explained variance; int: intersect; B: beta coefcient.; T: tolerance (inverse to redundancy among explanatory variables).
often show a bigger-deeper size-with-depth relation (e.g., decapods: Hargreaves et al., 1984; sh: Stefanescu and Cartes, 1992)
small specimens found in guts of A. antennatus (TL between 2.2

and 4.8 cm) may have a shallower, more inshore distribution than
specimens of TL > 56 cm, which in the case of Lampanyctus

crocodilus are distributed over depths of 550750 m (Stefanescu
and Cartes, 1992). Other prey, by contrast, had similar depth distributions to that A. antennatus, as was the case of Gennadas elegans
with maximum concentrations at 660698 m and higher consumption and abundance off Sller. Net avoidance must be sizedependent (i.e. only smaller individuals of prey species are caught
with the 1 m2 WP2), and most mesopelagic species (e.g., M. norvegica, Sergestidae and Pasiphaeidae) found in gut contents were larger than specimens collected with nets. Therefore, it is possible
that the lack of coincidence between prey found in stomach contents of A. antennatus and their distribution in the environment
could be caused by a shallower distribution of prey sizes consumed
by the red shrimp during daylight when plankton sampling was
performed.
Off the Catalan coasts, it has been suggested that A. antennatus
migrates upward and inshore at night (Cartes et al., 1993). These
migrations would take place preferentially through canyon valleys,
especially among small shrimp and, suggested by the higher stomach fullness found within canyons (Cartes, 1994; Fig. 8). It is well
known the role of canyons enhancing secondary production especially at canyon heads (Vetter and Dayton, 1998) even favouring
recruitment success of demersal species (e.g. Vetter and Dayton,
1999). Hence, upward nocturnal movements by A. antennatus
would be done to feed in more productive depths. A. antennatus
is caught by trawling at night at only 100150 m off Calabrian
coasts (Matarresse et al., 1995). Off the Balearic Islands, these
migrations could point to the capture of mesopelagic prey, coinciding with upward-inshore migrations at night by mesopelagic fauna
as evidenced by Reid et al. (1991) off Hawaii. An important proportion of undigested mesopelagic prey (e.g., decapods, myctophids)
were mainly found in guts of A. antennatus before/close to sunrise,
and the occurrence at sunrise of digested decapods, euphausiids
and myctophids suggests feeding (previous to our sampling) on
these same prey at night. However, these migratory movements
by A. antennatus which are probably in search of food are only supported by indirect evidence and remain hypothetical.
4.2. Interactions between trophic requirements and the reproductive
cycle
Higher consumption of mesopelagic prey with a large energetic
content occurred off Sller and this might favour conditions for
reproduction in the red shrimp. Rosa and Nunes (2003b) evidenced
higher lipid, and probably energy content in muscle of Aristeus
antennatus females in MayJune off Portuguesse coasts. At midslope depths interactions between food availability and trophic
requirements may only inuence the reproductive cycling of A.
antennatus, and not recruitment, because early juveniles
(CL < 14 mm) are practically restricted to depths below 1400 m
(Cartes and Demestre, 2003). A common trend found in both areas
was a sharp increase in stomach fullness during AprilJune 2004,
after a decrease from November 2003 to February 2004. This trend
was more evident off Sller, where diet was based on mesopelagic
prey with a higher energy content also in AprilJune 2004. This
could have led to the signicant increase of the GSI (fecundity) observed off Sller in June and higher density of small females of A.
antennatus (Guijarro et al., 2008; Fig. 9). In addition, HSI, lipids in
hepatopancreas and K n among females were higher off Sller from
February to AprilJune 2003 (Guijarro et al., 2008; Fig. 8), previous
to and simultaneously with the period of gonad development. A last
consequence was the occurrence of smaller mature females off Sller (24.926.5 mm CL contrasting to 27.329.8 CL off Cabrera) in
this same period. Most individuals attain their rst gonad development in May off the Catalan coasts, when the previtellogenesis
phase begins (Demestre and Fortuo, 1992). However, in the NW
51
Mediterranean variability in the onset and the duration of the

reproductive cycle has been recorded between years (Carbonell
et al., 2006), which was attributable to environmental (maybe food)
variability. The phase of rapid oocyte growth must be incur high
metabolic costs. At the end of the reproductive period (August
and September), and coinciding with a decrease in the caloric content of the diet, GSI for females was similar both off Sller and Cabrera, while K n , HSI and the percentage of lipids from the
hepatopancreas reached minimum values, indicating that the organic reserves stored in the hepatopancreas are used for ovarian
development in the later part of the spawning period (Guijarro
et al., 2008). The beginning of gonad development occurs two
months after the peak of primary production in surface layers
(FebruaryMarch: Cartes et al., 2008). In this way, off Sller, we
found a positive relationship (in small shrimp) between fullness/
diet and temporal changes in water column productivity (e.g., Chl
a readings taken two months before sampling and uorescence).
Control of populations among abyssal detritivorous fauna must
be coupled with quality of organic matter arriving at the bottom
and to the ability of benthic fauna for selective feeding (Ginger
et al., 2001; Witbaard et al., 2001). The link between reproductive
strategy and temporal variations in food availability have been discussed for bathyal species, and synchrony between the release of
juveniles in peracarids and maximum food availability deriving
from spring phytoplankton blooms has been suggested for instance
among peracarids (Cartes et al., 2002; Richoux et al., 2004). Comparison of species with continuous and non-continuous reproduction is particularly interesting. It has been suggested that shifts in
the food consumption of some slope dwelling shes are stronger
among shes with peaks of gonad development (Madurell and
Cartes, 2005), and during pre-reproductive periods these species
base their diet on the consumption of mesopelagic prey. At abyssal
and at bathyal depths, macrofauna and detritivorous megabenthos
couple their biological cycles with phytodetritus deposition
(Hudson et al., 2004; Richoux et al., 2004). In the case of holothurians, the biological cycle types (continuous, synchronous) are related with changes in the food ingested, deduced by changes in
the amount of fatty acid biomarkers (Hudson et al., 2004) deposited under conditions before and after the spring phytoplankton
blooms at the surface. We suggest, in the case of the deep shrimp
Aristeus antennatus, that a highly energetic diet provided by high
prey availability can stimulate fecundity and/or allow an earlier
maturity. A similar trend has been found comparing the energy intake and the reproductive cycle of pandalids off the Balearic Islands
(Fanelli and Cartes, 2008).
In general, energy reserves (e.g., lipids) strongly affect fecundity
and reproduction among sh (e.g., Lloret et al., 2005; Adams,
1999), and also in deep water decapods (Rosa and Nunes,
2003b). The storage of lipids in the hepatopancreas of A. antennatus
begins before reproduction (Fig. 9), and storage must in turn be enhanced by a more energetic diet. Aggregation of micronektonic, pelagic prey in particular areas, hotspots (e.g., canyons, slope breaks)
for zooplankton biomass (Genin, 2004), as likely occurs over the
steep slope off Sller, can in turn favour aggregation of predators
for feeding. In the case of A. antennatus we hypothesize that prereproductive females may search for areas with accumulations of
highly energetic prey.
Reproductive aggregations of the red shrimp have been suggested to occur around submarine canyons off the Catalonian
coasts (Sard et al., 1994). Secondary production of macrofauna locally increases at canyon heads (Vetter and Dayton, 1998). In
Catalan canyons, there was also an increase of gut fullness in A.
antennatus females in MarchApril (Fig. 8: Cartes et al., 2006), prior
to the beginning of the reproductive period (May: Demestre, 1995).
In a pattern that is different from that observed in the Balearic
Islands, the increase of fullness off the Catalonian coast was not
52
16
1.10
Kn-F
1.08
12
1.06
1.04
1.02
1.00
0.98
0
0.96
16
A03
S03
N03
F04
A04
J04
N03
F04
A04
J04
Nov
2003
Feb
2004
Apr
2004
Jun
2004
11
HSI
12
% GSI
10
%
8
4
7
6
A03
16
S03
80
70
12
%lipids
60
50
40
30
20
10
20
40
60
80
CL (mm)
Aug
2003
Sep
S
2003
Fig. 9. GSI (left), and K n , HIS and % lipids (right) of Aristeus antennatus as a function of time off Sller ( (---------); sh) and Cabrera ((
); ). Individual data was considered
for GSI, mean monthly data for K n , HIS and % lipids. Plots of K n , HIS and % lipids re-drawn by Guijarro et al. (2008). GSI was signicantly higher in Soller in June 2004 (Mann
Withney test, p < 0.05) Part of this gure revised from a published gure in Guijarro et al. (2008).
synchronized with a higher consumption of mesopelagic prey, but

with higher consumption of benthic prey (e.g., large polychaetes
and especially the macruran Calocaris macandreae) that are more
abundant in canyons in that area (Cartes, 1994). Calocaris macandreae is a key prey in trophic webs around submarine canyons,
and some of their potential predators (e.g., Trachyrhynchus scabrus
in the insular part of the Balearic Basin: Stefanescu et al., 1993),
can decrease or even disappear coinciding with its absence. This
even induces local differences in assemblage composition and spe-
cies distribution between mainland and insular areas (Stefanescu,

1991; Cartes et al., 2004).
Local differences in the source of energy exploited by A. antennatus in pre-reproductive periods should also be considered for
long-term uctuations of red shrimp populations. Based on results
from the Balearic Islands (Cartes et al., 2008), it has been suggested
that landings of A. antennatus increase off the Catalan coasts (a
mainland area) in years of higher secondary production by zooplankton blooms, which are in turn correlated with years of posi-
tive NAO (North Atlantic oscillation) index, (Maynou, 2008). However, patterns are different around the Balearic Islands and in Catalonian canyons. While zooplankton is important in the diet of prereproductive A. antennatus females in the insular area, the benthic
pathway covers the energetic requirements of red shrimps in the
mainland area.
4.3. Conclusions
The increase of food consumption by A. antennatus in pre-reproductive periods can derive from different prey, depending on
changes in local prey availability. Around the island of Mallorca
the expected impoverishment of benthic biomass in comparison
to canyon heads off the mainland area of the Catalan coasts led
to enhanced consumption of micronektonic prey and possibly to
accumulation of pre-reproductive females of A. antennatus in areas
(e.g., steep slopes, persistent frontal systems) likely to be home to
dense zooplankton aggregations (Genin, 2004). The occurrence of
zooplankton/micronekton in stomach contents of deep-sea sh
and decapods has been widely documented (e.g., Mauchline and
Gordon, 1991; Cartes, 1993a,b). Also, coupling between reproductive processes and inputs of organic matter originating in surface
waters has been documented for deep-sea detritivorous megafauna (Hudson et al., 2004). However, in the case of the deep-sea
shrimp Aristeus antennatus, we found a strong link in a predator
at a high trophic level between pelagic resources and reproductive
processes. This suggests, together to some other similar cases
(Fanelli and Cartes, 2008), a rapid link via mesopelagic fauna between surface primary production and gonad development that
is available to much of the megabenthic, bathyal community.
Acknowledgements
The authors thank to all participants in the F/V Moralti Nou
crews and to Lpez Jurado (IEO) who provided some oceanographic
data and conducted oceanographic surveys. We especially thank
the help of Drs. Joan Moranta and E. Massut, the last main researcher of IDEA project.
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Aristeus antennatus (Risso, 1816) is a demersal deep-sea shrimp

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

Lagardre, 1977). It is a dominant species in Mediterranean

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

to the bottom (horizontal haul) plus between 3 and 15 min during

The abundance of some dominant, broad taxa/species, key prey

From polychaetes consumed by C. coelorhynchus

Range size similar to that consumed by A. antennatus

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

respective statistical analyses performed or included in the

2.4. Data treatment

correlation coefcient, after log-transformation of the data). The

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

3.3. Dietary analyses

sh (Myctophidae, Cyclothone braueri, sh remains) and the isopod

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

Fig. 3. Fullness of Aristeus antennatus off Sller (---------) and Cabrera (

natantians (in AprJun) and sh (in Nov-Feb, myctophids) were

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

average, close to the bottom, there was an increase in T of 0.2

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

3.6. Relationships between feeding/diet and biological/environmental

lated (non-parametric Spearman rank; p < 0.05) with T close to

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

small meso-bathypelagic decapods and myctophids were more

4.1. Spatio-temporal changes in the diet and prey availability:

Fig. 6. Percentage undigested prey (by number) in gut contents of Aristeus

Depth was an important factor affecting diet off Cabrera (see

Off the Balearic Islands dependence upon mesopelagic prey was

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

Fig. 8. Temporal changes of fullness (F CI 95%) in A. antennatus off the Catalan

350 m) than at deeper (650750 m) depths. Variability was higher

must trophically exploit zooplankton distributed at shallower

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

small specimens found in guts of A. antennatus (TL between 2.2

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

specimens of TL > 56 cm, which in the case of Lampanyctus

Mediterranean variability in the onset and the duration of the

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

synchronized with a higher consumption of mesopelagic prey, but

cies distribution between mainland and insular areas (Stefanescu,

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

J.E. Cartes et al. / Progress in Oceanography 79 (2008) 3754

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