the role of zenopsis spp. as a predator in seamount and
TRANSCRIPT
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ICES CM 2002/M:28
Theme Session on Oceanography and Ecology of Seamounts
Indications of Unique Ecosystems
The role of Zenopsis spp. as a predator in seamount and shelf habitats
Heike Zidowitz, Heino O. Fock, Hein v. Westernhagen
Alfred Wegener Institute for Polar and Marine Research, PO Box 12 01 61,
D- 27515 Bremerhaven, Germany [tel + 49 471 4831 1382, fax +49 471 4831
1425, email: initial first [email protected]]
Abstract
The genus Zenopsis Gill 1862 consists of three species. Zenopsis conchifer occurs in the
Atlantic and Indian Oceans whereas Zenopsis nebulosus has a wide distribution in the (Indo-)
Pacific Ocean. Zenopsis oblongus, described in 1989, is closely related to Z. nebulosus and
only known from the Nazca Ridge in the SE Pacific. Feeding studies are reviewed for all three
species. At the Great Meteor Seamount (GMR) (NE Atlantic), Z. conchifer shows a shift in
prey selectivity with ontogenetic development. Smaller specimens (<46 cm TL) preyed on
constituents of the sound scattering layer (SSL) (myctophids, stomiids). Diet of larger
specimens (>52 cm) predominantly consisted of bentho-pelagic Macroramphosus spp., which
was the most abundant fish species on that seamount. A similar ontogenetic shift was
observed for Z. conchifer on the Namibian shelf. However, prey for the larger specimens
consisted mainly of the pelagic species Trachurus trachurus and Synagrops microlepis. In the
Pacific, Z. nebulosus (oblongus) obtains a similar trophic position with regard to migrating
components of the SSL. However, though being also very abundant at the Nazca-Ridge
seamounts, no Macroramphosus spp. were eaten. Large specimens of Z. nebulosus were
found to prey on bentho-pelagic rockfishes (Sebastidae). Zenopsis spp. appears to be an off-
bottom pelagic predator with preference for mesopelagic food components in all areas
considered. It is suggested that larger specimens abandon the off-bottom pelagic feeding
mode and that body size thus determines the capabilities of Zenopsis spp. to prey on bentho-
pelagic species.
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Keywords: Zenopsis conchifer, Zenopsis nebulosus, Zenopsis oblongus, feeding ecology,
sound scattering layer,
Abbreviations: GMR – Great Meteor Seamount (follow Wilson and Kaufmann 1987), SSL –
sound scattering layer
Introduction: The genus Zenopsis spp. Gill 1862 consists of three species, Zenopsis conchifer, Zenopsis
nebulosus and Zenopsis oblongus. These species are very similar in body plan with laterally
flattened bodies with an upwardly pointing face and a silvery coloration (Swaby & Potts
1999).
Z. conchifer occurs in the Atlantic and Indian Oceans (fig.1). In the eastern Atlantic it
inhabits shelf areas from Ireland to South Africa, most abundant at 20° N (north-west Africa)
(Maurin & Quéro 1982). The distribution in north-western European areas expanded since the
1960ies reaching Ireland in the 1980ies (Swaby & Potts 1999). Besides the shelf habitats it is
also found around Islands (Canary islands) and on seamounts far off the continental rises e.g.
at the Great Meteor Seamount (GMR). In the western part of the Atlantic its distribution
ranges from Nova Scotia to North Carolina (Scott & Scott 1988) where it gets more abundant.
It is also known from Brazil (Haimovici et al. 1994) to Argentina (Quigley & Flannery 1995).
In the Indian Ocean Z. conchifer is found off the SW coast of India and off southern Africa
from Walvis Bay to Kenya (Smith & Heemstra 1986) up to Somalia and in Indonesia (Froese
& Pauly. Eds. 2002. FishBase).
Z. conchifer reaches a total length of 80 cm (Smith & Heemstra 1986) and a body weight up
to 3.2 kg (Robins & Ray 1986). Specimens of ca. 60 cm are estimated to be 12-14 years old
(this paper), larger specimens probably get older than 20 years.
Z. nebulosus is known in the Indo-Pacific region, from Japan, Northwest shelf of Australia to
Broken Bay in New South Wales, New Zealand (Kailola et al. 1993), and elsewhere in the
region (Philippines (Anon 2001)) (fig.2). In the Eastern Pacific it is known off central and
southern California, USA (Eschmeyer et al. 1983) and from several seamounts in the western
Pacific (e.g. Hancock Seamount, Kammu Seamount (Hawaiian Ridge); Jumeau Seamount
(Richer de Forges 2000), Stylaster Seamount (Norfolk Ridge) (Froese & Pauly. Eds. 2002.
FishBase) Specimens caught at seamounts of the Nazca Ridge (SE Pacific) have been
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investigated by Parin et al. 1988. An analysis of otolith increments revealed an age of 13
years at a standard length of 46 cm. Different from Z. chonchifer, Z. nebulosus reaches only a
size up to 70 cm in total length and a maximum weight of 3.0 kg (Williams 1990), so that it
can be estimated that the species reaches an age of more than 20 years. These investigations
on the ecology of Z. nebulosus of the Nazca Ridge (Parin et al. 1988) are more likely to refer
to Z. oblongus.
Z. oblongus was first described in 1989 and is very closely related to Z. nebulosus, but differs
by a lower body and larger number of osseous scutella above the anal fin (Parin 1989). It is
only known from the Nazca Ridge in the Eastern Pacific Ocean as a probable endemic species
of the area (Parin et al. 1997) (fig.3).
Z. conchifer was caught in depths ranging from 50 – 730 m (Saldanha 1968) but mainly
occurs on the upper slope at 200 – 400 m (Quigley & Flannery 1995, Quéro 1998). It is
frequently encountered in coastal waters (Quéro et al. 1990) and found in midwater or near
the bottom (Swaby & Potts 1999) and it is probable that it occurs in schools (Berry 1978,
Whitehead et al. 1986). Z. conchifer is regarded as a bentho-pelagic species of the deep-water
(Froese & Pauly. Eds. 2002. FishBase), a mesopelagic species (Scott & Scott 1988) or
bathypelagic species (Quéro & Pariente 1977).
Z. nebulosus is considered as a bathydemersal deep-water species with occurrences in depths
ranging from 30 - 800 m (Froese & Pauly. Eds. 2002. FishBase).
Parin et al. (1997) consider Z. oblongus as a bentho-pelagic species in a wide sense inhabiting
depths between 180 – 330 m. They also indicate the species as an off-bottom pelagic species,
which can swim far away from the bottom, rising in midwater during diel vertical migrations.
Compared to fisheries of shelf areas, deepwater fisheries show a shift in families of fishes
which are commercially exploited. On the continental shelves primary families of exploited
fishes are Gadidae, clupeoids, salmonids, scombrids and Pleuronectidae whereas deepwater
fisheries are based on entirely different orders, such as Beryciformes, Zeiformes and
Scorpaeniformes (Koslow et al. 2000). Differences at this taxonomic level indicate
fundamental shifts in body plan and ecological strategy as well as in evolutionary lineage
(Koslow et al. 2000). Many deepwater species aggregate on seamounts and form a distinct
guild based on common features of their body plan, proximate composition, physiology and
metabolism, ecology and life history (Koslow 1996, 1997). They tend to be robust and deep-
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bodied in order to manoeuvre in the strong currents characteristic of this environment
(Koslow et al. 2000).
These fishes generally do not migrate vertically, but depend on the influx of meso- and
bathypelagic organisms past the seamount and on intercepting mesopelagic migrators on their
downward migration (Isaac and Schwartzlose, 1965; Genin et al. 1988; Koslow 1997). Some
predators developed an expectant hunting strategy along plateau margins and an increased
habitat dependent resource utilisation rate at locations of sound scattering layer interception
(Fock et al. 2002). To explain living conditions for seamount populations in often
impoverished nutritional conditions in the ambient oceanic regions, the sound scattering
layer-interception hypothesis (Isaacs and Schwartzlose 1965) has been developed. It implies a
primarily pelagic food utilisation for bentho-pelagic fishes, increased habitat dependent
utilisation rates at locations of interception with the sound scattering layer, diel changes in
utilisation rates due to availability of prey and sufficient resource partitioning among species
in order to avoid competitive exclusion (Fock et al. 2002b). It was suggested that it represents
a large enough prey source to maintain populations at seamounts (Hesthagen 1970, Rogers
1994, Parin et al. 1997).
A very important reason for the success of the deepwater families is the far K-selected end of
the life-history spectrum (Koslow et al. 2000). It is characterised by longevity, slow growth
and delayed maturity. With these features the species fill a gap in the distribution of teleost
life-history patterns (Roff 1984). Because in deepwater habitats the recruitment appears to be
episodic and e.g. orange roughy and Sebastes spp. undergo extended periods (decade or more)
of very low recruitment to the adult population (Leaman and Beamish 1984), Murphy 1968
and Stearns 1976 hypothesised an evolutionary link between longevity and recruitment. The
adaptations on the life-history level could constitute the population’s ability to withstand
extended periods of poor recruitment (Koslow et al. 2000). Episodic recruitment can also be
assumed for commercially not exploited genera like Zenopsis, Antigonia, Capros.
Morids (Moridae), cusk-eels (Brotulidae) and hakes (Merlucciidae) are robust-bodied
Gadiformes and active predators (Koslow et al. 2000). Hakes form a small but widely
distributed family (Koslow et al. 2000). Species of Merluccius are voracious predators
inhabiting the continental shelf and upper slope (Froese & Pauly. Eds. 2002. FishBase) and
are often dominant piscivores over upper portions of the continental slope and typically
migrate vertically into the upper waters at night to feed (Bulman and Blaber 1986). Their
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productivity is thereby linked directly to the near-surface food web (Koslow et al. 2000)
which makes them less adjusted to seamount habitats. Fock et al. (2002) showed that the
Gadiformes contributed an extremely low share to overall catch at the Great Meteor
seamount although they dominate on the shallower shelf of the NE Atlantic.
Materials and methods Sampling
Sampling was carried out at GMR during R/V Meteor cruise M 42/3 in September 1998. The
GMR is located at 30°N, 28.5°W and is an isolated, flat-topped seamount. Sampling
procedures applied on 15 station were bottom trawls in depths between 286 and 435 m with
an ENGEL bottom trawl. Eight hauls were performed during daytime, two at dusk and dawn
and five during night time.
Biological standard measures
A total of 94 specimens of Zenopsis conchifer were caught on the GMR. Of these, 62
specimens were provided for stomach content analysis. Specimens caught were measured in
total length (TL) to the nearest centimetre and weighted in the whole, using an electronic
digital laboratory scale. After preparation single organs (stomach, liver, ovaries) were
weighted as wet weight. Organs and heads of specimens were then deep-frozen.
Stomach content analysis
To investigate a shift in prey selectivity with ontogenetic development, specimens were
subdivided in size classes. Small fish <36 cm, medium-size fish 36 – 46 cm, medium-size fish
47 – 52 cm and large fish >52 cm. The medium-size classes comprise 82 specimens (87% of
overall catch). 52 individuals of 36 – 46 cm in total length were caught (55% of overall
catch), of which 11.53% were male specimens. In size class 47 – 52 cm 32% of total catch
with 30 individuals were found (6.6% males). The size class >52 cm was represented by 11
specimens which were exceptionally females that made up 11.7% of the total catch. Only one
juvenile fish was caught and sent to the ichthyological collection of Madeira, so that the small
size class was abandoned for further investigations.
Stomach contents were accomplished by defrosting stomachs and adherent water was blotted
off with tissues. For comparison with the weight taken onboard, wet weight was taken to the
nearest 0.005 g on electronic digital scales.
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All containing food items and regurgitated items in the pharynx with clear signs of digestion
were counted and weighted. Specimens with totally regurgitated stomachs without any food
items were not considered for further analysis.
Stomach fullness, which is recommended as an index for feeding periodicity (Hyslop 1980),
was estimated by definitions shown in table 1. Only stomachs with no food item at all were
considered as empty. Cephalopod beaks were only counted but not weighted. Prey items were
identified to the lowest possible taxon. Freely occurring otoliths were used for identification
of already digested prey items or prey in an advance state of digestion by using otolith
catalogues or by comparison with otoliths of successfully identified fishes. Myctophids and
other mesopelagic fishes were identified by a catalogue of A. Kotthaus (1972) of previous
studies of GMR ichthyocoenosis. Trachurus sp. was identified by the catalogue of W.
Schmidt (1968).
Age determination
Otoliths of Z. conchifer were prepared out of heads and sent to the Central Ageing Facility of
the Marine and Freshwater Resources Institute in Queenscliff, Australia for further
preparation and reading.
Whole otolith reading and sectioned otolith reading were accomplished. Each otolith was read
using a dissecting microscope at up to 40x magnification with reflected light. Otoliths were
distally ground on both sides to reveal growth increments and samples were viewed with
transmitted light. Increments were counted from the primordium to the edge of the largest
lobe and measured using an image analysis system.
Analysis of fish diet
The percentage of frequency of occurrence of each prey item (%F), the percentage of
abundance (%A) and percentage of weight (%W) were calculated (Hyslop 1980). %F was
determined as percentage of stomachs of fish with prey each item compared to all non-empty
stomachs. %A was calculated as percentage of abundance of prey item (N) compared to the
total abundance of all prey items (Ntot). %W was calculated respectively.
For overall comparison of prey utilisation the “relative importance index (RI)” (George &
Hadley 1979) was calculated for each prey item, which is based on the absolute importance
index (AI) as follows:
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AI = %frequency of occurrence + %total numbers + %total weight,
RI [%] = 100 AI/ AI ∑n
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Where n is the number of different food categories.
Results Size composition
94 specimens of Zenopsis conchifer were caught at GMR with a biomass of 104 kg. Z.
conchifer was tenth in position of the most abundant fishes and fifth in weight ranking of all
fishes caught (Zidowitz 2001).
The specimens caught were between 17.2 cm and 59 cm in total length (TL), but the fish were
predominantly between 35 and 59 cm (mean length 45.1 cm, mean weight 1109.2 g) There
was only one catch of a juvenile (17.2 cm) but no postlarvae or “fingerlings“ . Fig. 4 displays
the length-frequency of Z. conchifer with maxima in length class >40–42.5 cm with 28
individuals, and in length classes >45–52.5 cm with 13, 12 and 11 individuals. The maxima
probably attributes to cohorts of episodic recruitment. A Kolmogorov-Smirnov-Test showed
no normal distribution.
Age
Grinding and reading of otoliths of Z. conchifer revealed a lifespan of 12 – 14 years for the
largest specimens. Increments counted of whole otoliths were relatively clear from the
primordium to the edge although large otoliths were very opaque in the primordial region
(fig.5). Increments visible in transverse sections were very diffuse so otoliths were distally
ground in an attempt to increase increment clarity. A comparison of age estimates obtained
from whole and sectioned otoliths found that higher ages were derived from sectioned
otoliths. This may be because more increments were visible in the primordial region of
sectioned otoliths and because increments formed on the edge were also clearer in sectioned
samples. After examination of whole and transversely and distally sectioned otoliths, it was
evident that more accurate estimates of age were obtained from whole otoliths. Steward
(1992) found that increments formed in the otoliths of the mirror dory (Zenopsis nebulosus)
were also very diffuse when sectioned and that consistent age estimates were obtained from
whole otoliths. Similar age estimates were determined for Z. conchifer as for Z. nebulosus for
similar sized fish. Assuming that the counted increments were annual, age estimates obtained
from the Z. conchifer otoliths ranged from 4 to 14 years of age (table 2., fig. 6).
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Food items and diet composition
The food spectrum of Zenopsis conchifer comprise 14 food categories (table 3). The diet is
characterised by pelagic, mesopelagic and bentho-pelagic fishes. Seven genera of fishes could
be identified, of these four species. Due to an advanced state of digestion a lot of fishes could
not be determined to the lowest taxon but categorised as “unidentified fishes“.
Stomachs contained five genera of the family Myctophidae (Lampanyctus sp., Lepidophanes
sp., Ceratoscopelus sp., Diaphus sp., Hygophum sp.). Stomiiform fishes were represented by
families Stomiidae (Chauliodus danae), Phosichthyidae (Vinciguerria nimbaria) and
Sternoptychidae (Argyropelecus sp.) with a few individuals. Capros aper (Caproidae,
Zeiformes) and the genus Trachurus sp. (Carangidae, Perciformes) could be verified in
stomach contents. Most important food item in the diet of Z. conchifer was Macroramphosus
spp. (Macroramphosidae, Syngnathiformes).
Besides fishes molluscs (Cephalopoda, Decabrachia) could be identified by cutinisied jaws
and a rest of a buccal-apparatus. Crustacea in the diet belonged to the decapods.
For food categories the percentage of abundance (%A), percentage of frequency of occurrence
(%F), percentage of weight (%W) and the relative importance index (%RI) were calculated.
The most important food item was Macroramphosus spp. with 39 % relative abundance.
Myctophidae accounted for 32 %, „unidentified fishes“ 10 % of overall prey items. Other taxa
were only of minor relevance with one to five percent of relative abundance.
Prey item Macroramphosus spp. was found in 44 %, „unidentified fishes“ in 18% and
myctophids in 16 % of investigated stomachs. Capros aper was found in 10 % and Trachurus
sp. in seven percent of stomachs. Other prey categories accounted for two to five percent of
relative occurrence.
Looking at the relative weight of the prey categories of Zenopsis conchifer, Macroramphosus
spp. was again the dominant food with 44 %. The few individuals of Trachurus sp. Were the
second important in weight category with 38,6%, because of their large body size.
“unidentified fishes” were third in percentage of weight due to a few larger, and heavier
individuals with 10% of prey biomass. Capros aper, reaching a higher individual weight
compared to Macroramphosus spp., took eight percent of weight prey. The numerous
myctophids only made up two percent of biomass just like the decapos. The mesopelagic
fishes Chauliodus danae und Vinciguerria nimbaria accounted only for 0,4 % and 0,04 % of
weight.
The relative importance index shows that Macroramphosus spp. was the main prey item of
Zenopsis conchifer at GMR with 39% (fig.7).
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Myctophids were of relatively high significance with 18 % although of little weight (2 %) and
therefore the second important food category. Trachurus sp. Had a share of 17 %, due to its
size and weight but small number, in the overall comparisons, “unidentified fishes” 7%,
Capros aper 8 %. All other taxa, Chauliodus danae, V. nimbaria and decapods, were only
between 1 - 3 % of relative importance index.
Diurnal feeding rhythm
At GMR a habitat dependent resource utilisation was observed (Fock et al. 2002 b). Among
other fishes Z. conchifer shows a preference in consuming constituents of the sound scattering
layer. By partitioning habitats and the analysis of the consumed food in these areas, a
connection to the SSL-trap phenomenon was revealed.
The intensity of ingestion varied. Differences in feeding activity over course of time, were
acquired by degree of stomach fullness at different day and night times. The data revealed a
change in stomach fullness and periods of maximal feeding activity could be made visible
(fig. 9). At night time a high percentage of stomachs were empty and no or only little food
was consumed. At dawn and in the morning hours half of the stomachs were full or had a
moderate filling. This was the main feeding period of Z. conchifer. In the afternoon the filling
decreased and more intense digested food items could be observed. At dusk (18-21 h) a
second but weaker feeding period was determined.
At the Great Meteor seamount the analysis of the horizontal distribution of seamount fishes
over the plateau revealed that most of the populations were related to habitats at the edge of
the plateau (Fock et al. 2002). At plateau margins the probability of interception with the
horizontally advected SSL is increased and vertically migrating SSL ideally passes the
marginal habitats twice a day during its ascent and descent, while the summit plateau is only
supplied with advected planktonic prey during the descending phase of the SSL (Fock et al.
2002) (fig.10). Besides enhanced interception probability with the SSL, plateau margins are
further affected by topographically induced circular currents around the summit (Taylor-
Column) and local upwelling phenomena caused by these current anomalies, which are
typical at GMR (Meincke 1971, Mourino et al. 2001).
Ontogenetic shift in diet composition
The comparative presentation of relative importance indices in fig. 10 illustrates the
relevancy of single food categories in the nutrition of the different sized specimens.
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Specimens of 36 – 46 cm fed on six food categories, whereat no item extremely dominated in
the diet. Food items Trachurus sp., Myctophidae and Macroramphosus spp. took the main
body of prey, whilst cephalopods, Capros aper and “unidentified fishes” were of less
important trophic meaning. With increasing size a change in the diet composition could be
observed. In size class 47 – 52 cm the significance of Trachurus sp. decreased. Instead,
Macroramphosus spp. predominated the diet of Z. conchifer and “unidentified fishes” were
second in meaning. Myctophids still took an important role but by comparison with the
smallest size class (36 – 46 cm) were less important. In the prey spectrum of the moderate
size class other mesopelagic fishes turn up, but only in small numbers so that they were
summarised to “other fishes”. Decapods were found but with little importance. In size class
>52 cm a distinct taxonomic impoverishment of the prey spectrum occurs. Only two food
categories, Macroramphosus spp. and unidentified fishes were ascertainable though only two
specimens of Z. conchifer were available.
Discussion Individuals of Zenopsis conchifer reaching a maximum length of 59 cm were relatively large.
Because no larvae, postlarvae and small juveniles were caught in pelagic trawls above the
GMR and also very large specimens with total lengths over 60 cm were missing a regular
recruitment can be doubted. Ehrich (1974) suggests a self preserving recruitment of Z.
conchifer at GMR but there is no evidence that the population is independent from other
habitats. Irregular current situations may cause an influx of pelagic larvae from other areas
and therewith fill up the population at GMR. Maxima of several length classes could indicate
cohorts of episodic recruitment. This ability of withstanding episodic recruitment is one of the
factors for the ecological success of the genus Zenopsis and with this adaptation to prevail in
insecure recruitment phases is the causes for successfully inhabiting seamounts.
Like Z. conchifer, Z. nebulosus was one of the most important fishes of seamounts of the
Nazca Ridge. It occupied fourth to sixth place in lists of dominant species on these seamounts
and accounted for up to 5 % of the total catches (Parin et al. 1988). Catches of Z. nebulosus
were between 4 and 48 cm, with a mean length of 32-33 cm. They also had remarkably few
catches of small and juvenile specimens, in four years only two “fingerlings“ (postlarvae)
with standard lengths (SL) of 44 and 60 mm and only three juvenile females with standard
lengths of 140 to 160 mm. Parin et al. (1988) assume that the infrequency of the discovery of
the juveniles is explainable by their pelagic mode of life.
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The results of the analysed size classes revealed an increasing selectivity with ontogenetic
development from a diverse diet to a one-sided diet consisting of the most abundant species
Macroramphosus spp.. From an relatively even utilisation of all food categories Z. conchifer
shifts to a tighter utilisation with a predominant food category. It can be regarded as an
opportunistic feeder probably due to the possibilities of body size.
At GMR only Heptranchias perlo was the most competitive predator to Z. conchifer. H. perlo
fed predominantly on bentho-pelagic teleost fishes like the snipefishes Macroramphosus spp.,
boarfish Capros aper, congrid eels and other unidentified teleosts. Besides this cephalopods,
mainly octopods, a few elasmobranchs and remains of echinoderms and sponges were food
items of importance (Frentzel-Beyme & Koester 2002).
The diet of Z. conchifer (n=69) at the Namibian shelf consisted predominantly of fishes but
also of euphausiids. The major food component was represented by the family Myctophidae
and minor components were formed of Synagrops microlepis and Trachurus trachurus. The
larger the individuals were the less important became the crustaceans. In smaller individuals
(20 – 29 cm TL) the food were made up of 87.1 % of weight by myctophids mainly composed
of Diaphus dumerilli and D. taaningi. In larger specimens other fishes became more
important. In individuals of 30 – 39 cm TL crustaceans only accounted for 0.2 %, Trachurus
trachurus already reached 32.2 %, whereas the myctophids constituted only 67.7 % (29.2 %
D. dumerilli) of the food. Specimens larger than 40 cm only fed on Synagrops microlepis
(33.4 %), Trachurus trachurus (34.2 %) and the myctophid Symbolophorus boops (32.4 %)
but no crustaceans (MacPherson 1983). The stomach contents revealed a shift in diet
composition with ontogenetic development similar to the shift of Z. conchifer at GMR, from a
main share of mesopelagic food items to a more one-sided diet of a few pelagic and
mesopelagic components. At GMR the range of the size-shifting was in a different magnitude
because the specimens caught were larger in comparison and especially non-myctophid
components consisted of other species than those from the Namibian shelf. The food analysis
of Z. conchifer in the Northwest Atlantic (NOAA area Nova Scotia – South of Cape Hatteras)
showed a 100 % fish diet (Bowman et al. 2000). Of this a share of 32.1 % was identifiable as
Stenotomus chrysops (Sparidae), 67.9 % were unidentifiable osteichthyes. Because of the
small number of specimens (n=5) no shift could be pointed out.
In investigations of the upper slope demersal ichthyofauna in Brazil Zenopsis conchifer was
caught in 34 hauls. Stomach content analysis revealed mesopelagic Maurolicus muelleri,
Diaphus dumerilii and Euphausia similis as main food components (Haimovici et al.1994).
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Bigelow & Schroeder (1953) reported that they found two butterfish, 15 – 18 cm long, and
one squid in the stomach of a large specimen about 50 cm long captured in a trawl off the
American Atlantic coast.
In terms of optimal foraging (Hart 1986) the SSL-interception hypothesis firstly implies
reliance on pelagic prey. Secondly, an increased resource utilisation rate as cause for the
aggregation of fishes in plateau margins as locations of potential interceptions is predicted.
Thirdly, the SSL-interception hypothesis predicts either cessation of feeding during periods of
absence of migrating prey or a switch between diets depending on their diel availability (Fock
et al. 2002 b)..
Z. conchifer shows in all investigated areas an intense food utilisation of pelagic and
mesopelagic food items. At GMR and a positive relationship to plateau margins (Fock et al.
2002 b) and an increased resource utilisation rate at the plateau margins and a diurnal feeding
rhythm which corresponds to the availability of migrating prey could be determined. This
preference were suggested to be attributed to differential feeding modes with respect to the
SSL-interception hypothesis (Fock et al. 2002 b).
Seamount-associated predators such as Zenopsis spp. are less mobile and have a body plan
constituted for a different hunting strategy than the fast swimming neritic hakes. Zenopsis spp.
is considered as a feeble swimmer (Wheeler 1969) and profits of a expectant strategy
(Zidowitz 2001). Zenopsis spp. showed in all regions of occurrence a selectivity for
mesopelagic items during SSL decent with a therefore adapted diurnal feeding rhythm.
Smaller specimens of Z. conchifer at GMR, on the Namibian shelf and smaller and medium-
size specimens of Z. oblongus show furthermore an adaptation in ontogenetic development for
this favoured feeding strategy. Larger specimens abandon preference for the SSL and the off-
bottom pelagic feeding mode and turn to a more one-sided diet by feeding on the most
abundant species. Diet of larger specimens of Z. conchifer at GMR constitutes almost
exclusively of Macroramphosus spp., the most abundant fish on the seamount, probably
caused by body size enabling to prey on bentho-pelagic species. MacPherson (1983)
appraised the presence of fast swimmers like Trachurus trachurus in several stomachs, that it
shows the great efficiency of the hunting strategy of Zenopsis conchifer.
Z. nebulosus (oblongus) reaches a maximum size of 70 cm and therefore has a lower average
size compared to Z. conchifer. Like Z. conchifer the diets consists of the most abundant
species, whether “thalassobathyial bentho-pelagic” fishes (in particular Maurolicus), “oceanic
mesopelagic” animals (euphausiids, myctophids and squids) and “pseudoneritic and oceanic
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nektonic” fishes (Trachurus, Emmelichthys, Cubiceps) (Parin et al. 1988) thus making them to
opportunistic feeders with preferences depending on body size. Though being also very
abundant at the Nazca Ridge seamounts (Parin et al. 1997), no Macroramphosus spp. were
eaten here. In Californian waters Z. nebulosus was found to prey on rockfishes (Sebastidae).
Conclusions The species of the genus Zenopsis spp. are similar types of predators in the investigated
habitats. They fill the same ecological niche benefiting in seamount habitats of their typical
zeid body plan and life history with special adaptations. Besides a few sharks they play an
important role in food web structures as higher-level carnivores, whereas in shelf habitats
other fast swimming nektonic species dominate.
Literature Anon. (2001). Fish collection database of the National Museum of Natural History
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17
Figures and Tables
Fig
Fig
Fig
. 1: Distribution of Zenopsis conchifer (FishBase mapper).
. 2: Distribution of Zenopsis nebulosus (FishBase mapper).
. 3: Distribution of Zenopsis oblongus (FishBase mapper).
18
Table 1: Definitions for estimation of stomach fullness, estimated values.
degree of stomach fullness Definition
0 empty
< ¼ little
¼ moderate
½ half full
4/4 full
Empty
A few, or single remains <10%
Filled up to 30 %
Filled between 30 % and 70 %
Filled between 70 % and 100 %
13
8
28
9
13 12 11
3 3 4
0
5
10
15
20
25
30
17 18,5
>35 -
37,5
>37,5
- 40
>40 -
42,5
>42,5
- 45
>45 -
47,5
>47,5
- 50
>50 -
52,5
>52,5
- 55
>55 -
57,5
>57,5
- 60
length [cm]
num
ber [
n]
Fig. 4: Size composition (TL) of Zenopsis conchifer at GMS.
Ageing Transect
Fig. 5: A whole Z. conchifer otolith viewed with a dissecting microscope, reflected light and a black background. Magnification = 15.75x
19
Table 2. Sample information of otoliths. Total Length Otowght Age
40 0.001 440 0.001 440 0.002 440 0.002 4
642 0.001 542 0.001 643 0.001 643 0.001 649 0.004 950 0.003 850 0.003 853 0.004 957 0.006 1059 0.005 1459 0.007 1159 0.006 12
42
0
10
20
30
40
50
60
70
0 2 4 6 8 10 12 14 16
Whole age estimates (yrs)
Tota
l Len
gth
(cm
)
Fig. 6: The relationship between estimated age (determined from whole otoliths) and total length of Z. conchifer.
20
Table 3: Number of prey items, frequency of occurrence (%F), percentage of abundance (%A), percentage of weight (%W) and “relative importance index (RI)” for food categories of Zenopsis conchifer.
Taxon number prey item[n]
percent abundance (A) [%]
percent frequency (F) [%]
percent weight (W) [%]
rel. importance index (RI) [%]
Pisces Syngnathiformes Macroramphosidae Macroramphosus spp. Zeiformes Caproidae Capros aper Stomiiformes Stomiidae Chauliodus danae Phosichthyidae Vinciguerria nimbaria Sternoptychidae Argyropelecus sp.* Myctophiformes Myctophidae Lampanyctus sp.* Lepidophanes sp.* Ceratoscopelus sp.* Diaphus sp.* Hygophum sp.* Perciformes Carangidae Trachurus sp. Unidentified fishes
142 59 7 6 2 1 48 15 13 14 5 1 4 15
94,6 39,3 4,7 4 1,3 0,7 32 10 8,6 9,3 3,3 0,6 2,7 10
43,5 9,7 3,2 1,6 1,6 16,1 9,7 4,8 11,3 4,8 1,6 6,5 17,7
40,3 7,5 0,4 0,04 -- 2,2 -- -- -- -- -- 38,6 9,7
39 7,8 2,7 1 -- 17,9 -- -- -- -- -- 17,1 7,1
Mollusca Cephalopoda Decabrachia+
3
2
4,8
--
--
Arthropoda Crustacea Decapoda
5
3,3
1,6
1,8
2,4
* identification bay otoliths, no weight data available
+ no cephalopod beaks weighted
7 %
17 %
18 %
1 %3 %
8 %
39 %
2 % Macroramphosus spp.
Myctophidae
Capros aper
Chauliodus danae
V. nimbaria
Trachurus sp.
Pisces indet
Decapoda
Fig. 7: relative importance index (George & Hadley 1979) of food components in the diet of Zenopsis conchifer.
21
0%
20%
40%
60%
80%
100%
0 - 3
h3 -
6 h
6 - 9
h
9 - 12
h
12 - 1
5 h
15 - 1
8 h
18 - 2
1 h
21 - 2
4 h
Time
empty full
little moderate half full
other fishes
Cephalopodaunident. fishes
Chauliodus sp.Capros aperMacroramphosus spp.
Fig. 8: Diurnal feeding rhythm based on degree of stomach fullness.
0%
10%
20%
30%
40%
50%
60%
70%
80%
90%
100%
36 - 46 cm 47 - 52 cm > 52 cm
size classes TL [cm]
rel.
Impo
rtan
ce in
dex
[%] Decapoda
Trachurus sp.
Myctophidae sp.
Fig. 9: Comparison of food composition of the 3 size classes of Z. conchifer based on the relative importance indices. 36 – 46 cm: n = 32, 47 – 52 cm: n = 24, > 52 cm: n=6.
22
F
ig. 10: DSL (SSL) trap at Great Meteor Seamount (from Hesthagen 1970).