Journal of

Applied Ichthyology
J. Appl. Ichthyol. 26 (2010), 583–591 Received: April 22, 2009
 2010 Blackwell Verlag, Berlin Accepted: January 10, 2010
ISSN 0175–8659 doi: 10.1111/j.1439-0426.2010.01467.x

Age, growth and feeding habits of the bluemouth rockfish, Helicolenus dactylopterus
dactylopterus (Delaroche 1809) in the central Mediterranean (southern Tyrrhenian
Sea)
By P. Consoli1, P. Battaglia1, L. Castriota1, V. Esposito1, T. Romeo1 and F. Andaloro2
1
ISPRA (Superior Institute for the Environmental Protection and Research), Laboratory of Milazzo, Milazzo (ME), Italy;
2
ISPRA, Palermo, Italy

Summary by (Uiblein et al., 2003). It is considered selectively carnivo-

This paper provides data on age, growth and diet of the rous, feeding mainly on benthic crustaceans (decapods), fishes,
bluemouth rockfish, Helicolenus dactylopterus dactylopterus gelatinous plankton (pyrosomes) and incidental cephalopods
(Scorpaenidae), central Mediterranean Sea. Estimated by and polychaetes (Nouar and Maurin, 2000).
otolith readings, ages ranged from 0 to 21 years. Parameters Notwithstanding the several studies on diet and feeding
of the von Bertalanffy growth curves were Linf = 26.1 cm, behaviour in varous Mediterranean areas (Froglia, 1976;
k = 0.14 year)1, t0 = )1.92 years and the growth perfor- Wurtz, 1977; Macpherson, 1981; Sartor et al., 1993; Madurell
mance index was F = 1.98. The weight-length relationship, and Labropoulou, 2000), very little is known of its trophic
W = 0.016 TL2.99; R2 = 0.99, described an isometric growth spectrum at a specific level and none of these authors used a
of the species. This species is a carnivore focusing mainly on statistical approach.
the crustaceans Goneplax rhomboides and Lophogaster typicus. In light of these considerations, the aim of this paper was to
Statistical analysis showed an ontogenetic shift between small investigate some aspects of H. d. dactylopterus biology in the
(4.0–6.3 cm) and larger fishes. Significant differences were also southern Tyrrhenian Sea giving more detailed information on
found in relation to the sampling depth. age, growth, diet composition, niche breadth and also evalu-
ating how size and depth affect the observed feeding pattern.

Introduction
The bluemouth, Helicolenus dactylopterus dactylopterus (Del- Materials and methods
aroche 1809) (Pisces : Scorpaenidae), is a deep-sea scorpion- Specimens of H. d. dactylopterus were occasionally collected
fish living on the coarse and mud-sandy bottoms of the from January 2002 to December 2004, during several scientific
continental shelf and slope from 20 to 1000 m (Fischer et al., bottom trawl and longline surveys carried out in the southern
1987). It is widely distributed in the eastern Atlantic, from Tyrrhenian Sea (Fig. 1). Samplings were not specifically
Norway to the south-west coasts of Africa (Hureau and designed for the capture of this species, but were aimed at
Litvinenko, 1986), in the western Atlantic from Venezuela to estimating the abundance of demersal resources in the study
Nova Scotia, and in the entire Mediterranean. area on sea bottoms ranging from 100 to 600 m. Fishing
This teleost is a commercial species with an important surveys were uniformly performed on five depth ranges (100–
economic value targeted by long-lines and gillnets and which 200; 201–300; 301–400; 401–500; 501–600 m).
also appears regularly in the by-catch of bottom trawl and pots Each specimen (total length–TL) was measured by calliper
set for deep-water crustaceans (Fischer et al., 1987; Relini to the nearest 0.01 cm and weighed to the nearest 0.1 g.
et al., 1999). Stomachs were then removed and preserved in 70% ethanol.
In the Mediterranean, several authors investigated age and
growth of H. d. dactylopterus by means of modal progression
analysis and otolith readings. These studies were carried out in Age
the Alboran and Balearic seas (Massutı́ et al., 2000), Ligurian Sagittae were removed from the vestibular apparatus, cleaned,
Sea (Peirano and Tunesi, 1986; Ragonese, 1989), Sicily dried and stored with a code number. Before observation, they
Channel (Ragonese and Reale, 1992), Ionian Sea (DÕOnghia were immersed in a solution of glycerine, ethanol and water for
et al., 1992, 1996) and Adriatic Sea (Ungaro and Marano, 1 day and read the following day. Readings were carried out
1995; Romanelli et al., 1997), but no age data are available with a stereomicroscope under reflected light; whole otoliths
from the southern Tyrrhenian Sea. In spite of the differences in were observed in the above-mentioned mixture on a dark
maximum age and growth parameters, all authors seem to background, to better distinguish opaque and translucent
agree that bluemouth is a slow-growing and long-lived species. zones (Massutı́ et al., 2000). A growth increment including one
Sizes of this species are larger in the Atlantic Ocean, above opaque zone together with the adjacent translucent zone was
40 cm (Isidro, 1987; Morales-Nin, 1989), than in the Mediter- considered equivalent to 1 year of life (Beckman and Wilson,
ranean Sea where it reaches a maximum total length of 36 cm 1995), as suggested by several studies on this species in the
(Massutı́ et al., 2000). Mediterranean Sea (Ragonese and Reale, 1992, 1995; Roma-
H. d. dactylopterus is a typical sit-and-wait ambush predator nelli et al., 1997; Massutı́ et al., 2000) and the Atlantic Ocean
with a highly cryptic colouration, attacking prey as they swim (Isidro, 1987; Esteves et al., 1997; Abecasis et al., 2006).

U.S. Copyright Clearance Centre Code Statement: 0175–8659/2010/2604–0583$15.00/0

 14390426, 2010, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1439-0426.2010.01467.x by Cochrane Portugal, Wiley Online Library on [24/10/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
584 P. Consoli et al.

Fig. 1. Map of the southern

Tyrrhenian Sea

The count of annuli was performed on the concave side of Krug et al., 1998; White et al., 1998; Kelly et al., 1999; Allain
the sagitta, usually on the postrostral area, checking their and Lorance, 2000; Abecasis et al., 2006).
continuity all around the otolith to avoid adding any possible
fakes (Beckman and Wilson, 1995; Campana, 2001; Panfili and
Morales-Nin, 2002). Feeding habits
When otoliths were thick and not clearly readable, they were Gastric contents analysis was carried out under a binocular
made thinner using semi-automatic polishing equipment (LS2 microscope and prey items identified to the lowest possible
machine, Offshore srl, Bologna, Italy) on sandpaper (grid taxonomic level, counted and weighed to the nearest 0.1 mg,
1200–2000). Once the sagittal plan was close to the core, after removing excess water with blotting paper.
otoliths were polished with alumina 0.2 lm and the opaque To count the remains of digested prey, the criteria suggested
zones then counted. by Quiniou (1978) and Cherabi (1987) were used: (i) fragments
Each sagitta was read by two observers; only coincident of crustaceans and polychetes were considered as prey units;
interpretations were accepted. If readings had differing results, (ii) unidentified individuals or fragments of individuals of the
both readers discussed the age differences detected; the otolith same taxonomic group were counted and weighed; (iii) remains
was then read once more. When agreement was not possible, of individuals of the same species were counted and weighed
the otolith was considered unreadable and thus discarded. together.
In order to measure sample size sufficiency, the cumulative
number of new prey types was plotted against the cumulative
Growth number of non-empty stomachs analysed (Ferry and Caillet,
To describe H. d. dactylopterus growth, the weight-length 1996). The PRIMER statistical package v.6 (Clarke and
relationship was calculated using the exponential equation: Gorley, 2006 licensed to Pierpaolo Consoli) was utilized to
compute a prey species accumulation plot as an average of 999
W ¼ aLb curves based on different random orders of the stomachs.
where W is the total weight (g), L is the total length (mm), a is The logistic and the linear regressions were calculated and
the intercept of the regression line and represents a coefficient the goodness of fit coefficient (R2) compared: the sample size is
related to the body shape of the species, b is the regression considered sufficient when R2 for the logistic curve results is
coefficient and indicates the isometric growth when equal to 3 higher than R2 for the linear relation.
(Wootton, 1990; Anderson and Neumann, 1996). Estimation The contribution of each prey in the diet of H. d. dactyl-
of parameters a and b was carried out by transforming (ln) the opterus was estimated by calculating abundance (%N = num-
equation by linear regression. To check the theoretical ber of prey i ⁄ total number of prey* 100), weight
isometric (b = 3) or allometric growth (b „ 3) the StudentÕs (%W = weight of prey i ⁄ total weight of all prey*100) and
t-test (Snedecor and Cochran, 1967) was employed. frequency of occurrence (%F = number of stomachs contain-
Moreover, growth parameters (Linf, k, t0) were estimated by ing prey i ⁄ total number of stomachs containing prey*100).
fitting age and related length data into the theoretical growth These values were used to calculate the index of relative
model of von Bertalanffy (1938): importance (IRI) for each taxonomic category, using mass
  instead of volume (Pinkas et al., 1971; Hyslop, 1980; Hacunda,
Lt ¼ Linf 1  ekðt  t0 Þ 1981): IRI = (%N + %W)(%F).
The %IRI was also calculated as: %IRI = (IRI ⁄ S
where Lt is the total length at age t (years), Linf is the predicted IRI) · 100.
asymptotic length, k is the instantaneous growth coefficient, To evaluate the rate of feeding activity, the vacuity
describing how rapidly this length is achieved, t0 is the coefficient (%V = percent of empty stomachs) was estimated.
theoretical age at zero length (years). Diet breadth was calculated by LevinÕs standardized index
The growth performance index (F = 2 log Linf + log k) (Hulbert, 1978; Krebs, 1989) for prey biomass:
(Munro and Pauly, 1983) was also calculated in order to !
compare the values of the growth parameters obtained in the 1 1
Bi ¼ P 2 1
present paper with those reported by other authors for the n1 j pij
same species in the Mediterranean Sea (Peirano and Tunesi,
1986; DÕOnghia et al., 1992, 1996; Ragonese and Reale, 1992; Bi = LevinÕs standardized index for predator i; pij = propor-
Ungaro and Marano, 1995; Massutı́ et al., 2000) and Atlantic tion of diet of predator i that is made up of prey j; n = number
Ocean (Isidro, 1987; Morales-Nin, 1989; Esteves et al., 1997; of prey categories.
 14390426, 2010, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1439-0426.2010.01467.x by Cochrane Portugal, Wiley Online Library on [24/10/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Age, growth and feeding habits of the bluemouth rockfish 585

The index ranges from 0 to 1; low values indicate diets 250

dominated by few prey items (specialist predator), high values
indicate generalist diets (Krebs, 1989). 200 N = 498
To assess the feeding behaviour of H. d. dactylopterus, a
modified Costello graphical method (Costello, 1990; Amund- 150

N. ind
sen et al., 1996) was applied to the data set of prey taxa
identified at species level. In the new graphical method, the 100
prey-specific abundance (Pi) is plotted against the frequency of
occurrence (Fi), providing a two-dimensional graph 50
(Amundsen et al., 1996). In mathematical terms Fi and Pi
can be described by the equations: 0
0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30
X X  TL (cm)
Fi ¼ ðNi =N Þ and Pi ¼ Si = Sti 100
Fig. 2. Length frequency distribution (2 cm TL classes) of bluemouth
Helicolenus d. dactylopterus specimens caught in the study area, 2002–
Where Ni is the number of predators with prey i in their 2004. N = 498
stomach, N is the total number of predators with empty
stomachs, Si is the stomach content (as a number) comprised
of prey i, and Sti the total stomach content in only those fishes
interpret. Otoliths showed the characteristic pattern of teleosts,
with prey i in their stomachs.
with hyaline and opaque rings laid down around an opaque
To verify the existence of differences in the diet of
nucleus due to slow and fast growth phases. As reported by
bluemouth in relation to the predator length and depth of
Massutı́ et al. (2000) the first 4–6 pairs of rings were wider than
capture a two-way ÔAnalysis of SimilaritiesÕ (ANOSIM;
the following rings, which were laid down with decreasing
Clarke, 1993) was performed. Although size is a continuous
thickness, while in adult fish the outer rings decreased in width,
variable which changes continuously throughout ontogeny,
becoming irregular and equally wide.
size-related diet variations were investigated by dividing the
Analysis of 498 otoliths permitted an estimate of a maxi-
collected sample into four size-classes: Class I, 40–63 cm
mum age of 21 years, but specimens aged more than 14 years
( age 0); Class II, 64–90 cm ( age 1); Class III, 91–137 cm
were uncommon in this study area. The majority of samples
( age 2–3); Class IV, > 137 cm (age > 4). With respect to
belonged to the first age classes; in particular, 237 specimens
the factor ÔdepthÕ, four levels were considered: depth 1 (100–
(47.6%) were included in age class 0 (average
200 m), depth 2 (201–300 m), depth 3 (301–400 m) and depth 4
TL = 6.3 ± 0.7 cm), 57 were aged as 1 year old (average
(> 401 m). To reach a sufficient number of samples for the
TL = 9.0 ± 1.2 cm) and age class 2 was represented by 38
statistical analyses the last depth level was obtained by
H. d. dactylopterus (average TL = 11.6 ± 1.1 cm).
combining the specimens caught in the two bathymetric
ranges: 401–500 and 501–600 m.
Prior to the analysis, prey items were clustered into Growth
taxonomic categories at the order level. A binomial test, on
The relationship between weight and length estimated for H.
the frequency of occurrence values of each one, was applied to
d. dactylopterus in the study area is expressed by the equation:
select prevailing orders in the diet of bluemouth: a taxon was
W = 0.016 TL2.99, R2 = 0.99 (Fig. 3).
considered as rare when its frequency of occurrence was
Weight increased isometrically with size, since the value of
significantly < 5%, for P < 0.05; the rare orders and
b = 2.99 did not have a significant difference from the value
unidentified taxa were pooled into broad taxonomic groups.
3.0, as confirmed by the StudentÕs t-test (t496 = )0.1324;
The prey categories considered were: Euphausiacea, Decapoda
P < 0.05).
reptantia, Decapoda natantia, Amphipoda, Isopoda, Mysida-
The parameters of the von Bertalanffy growth curve fitted to
cea, Other Crustaceans, Cephalopoda, Polychaeta, Osteichth-
total lengths at age were estimated as Linf = 26.1 cm,
yes, Thaliacea, and Others. The data matrix based on the
k = 0.14 year)1, t0 = )1.92 years (Fig. 4). The theoretical
weight of these 12 prey categories was then standardized to the
total weight sample, transformed to square root and analysed
on the basis of Gower distance. 350
y = 0.016x2.99
300
Results R2 = 0.99

Overall, 524 specimens of bluemouth ranging from 3.3 to 250

27.0 cm total length (mean 11.1 ± 5.8 cm) and from 0.58 to
200
W (g)

288.6 g total weight (mean 40.8 ± 57.8 g) were examined.

Figure 2 shows the length (2 cm classes) frequency distribution 150
of the bluemouth specimens; 38.9% of samples belong to the
6 cm TL class (5.0–6.9 cm). 100

50

Age
0
Sagittal otoliths of H. d. dactylopterus appear elliptic, longer 0 5 10 15 20 25 30
than wide, with a rostrum moderately elongated. On a total of TL (cm)
524 sagittae only 5% (equally distributed in all size classes) Fig. 3. Helicolenus d. dactylopterus weight-length relationship
were unreadable, since they were broken or difficult to observed in the southern Tyrrhenian Sea. N = 498
 14390426, 2010, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1439-0426.2010.01467.x by Cochrane Portugal, Wiley Online Library on [24/10/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
586 P. Consoli et al.

Mean ±0.95 Conf. interval Table 1

Y = 26.1 (1–e–0.14 (X–(–1.92))) N = 498 Diet composition of Helicolenus d. dactylopterus with relative food
30 indexes: weight % (%W), number % (%N), frequency % (%F) and
Index of Relative Abundance % (%IRI). Unid. = unidentified
25
Prey types %W %N %F %IRI

20 Crustacea
Stomatopoda
TL (cm)

Pseudosquillidae 0.17 0.20 0.42 0.01

15 Euphausiacea
Euphausia krohni 0.01 0.20 0.42 0.00
10 Unid. Euphausiacea 0.17 0.98 1.27 0.07
Decapoda
Brachynotus sp. 0.01 0.20 0.42 0.00
5 Goneplax rhomboides 50.37 9.98 19.49 56.77
Monodaeus couchii 2.10 0.20 0.42 0.05
0
Unid. Brachiura 1.58 1.37 2.97 0.42
0 2 4 6 8 10 12 14 16 18 20 22 Unid. Reptantia 0.11 0.20 0.42 0.01
Age (years) Alpheus glaber 0.18 0.20 0.42 0.01
Philocheras echinulatus 1.25 0.59 1.27 0.11
Fig. 4. Average observed length (± 0.95 confidence interval) by age- Philocheras bispinosus 0.03 0.20 0.42 0.00
class and von Bertalanffy growth curves determined from annuli Pontocaris lacazei 0.91 0.20 0.42 0.02
interpretation in otoliths of Helicolenus d. dactylopterus, southern Unid. Crangonidae 1.19 0.59 0.85 0.07
Tyrrhenian Sea Pasiphaea sp. 0.27 0.20 0.42 0.01
Chlorotocus crassicornis 3.75 0.39 0.85 0.17
Pandalina sp. 0.23 0.20 0.42 0.01
asymptotic total length (Linf = 26.1 cm) gave results lower Processa sp. 3.71 0.78 1.69 0.37
than the largest specimen (TL = 27.0 cm) caught in this Unid. Natantia 2.57 1.17 2.12 0.38
study. The value of the growth performance index obtained for Unid. Decapoda 0.41 0.39 0.85 0.03
all individuals was F = 1.98. Total Decapoda 68.67 16.83 32.20 58.44
Amphipoda
Gammaridea
Arrhis mediterraneus 0.00 0.20 0.42 0.00
Feeding habits Cheirocratus assimilis 0.00 0.20 0.42 0.00
The gastric contents analysis was performed on a total of 455 Hippomedon bidentatus 0.01 0.39 0.85 0.02
individuals; the remaining samples were utilized for other Rhachotropis caeca 0.00 0.20 0.42 0.00
Rhachotropis grimaldii 0.04 0.98 2.12 0.10
purposes. Of the 455 stomachs analysed, 219 were empty Stegocephaloides 0.01 0.20 0.42 0.00
(%V = 52) whereas 236 had at least one food item christianiensis
(%V = 48). Specimens ranged from 3.3 to 24.9 cm TL (mean Unid. Gammaridea 0.14 1.96 3.39 0.34
8.4 ± 3.9 cm). Hyperiidea
Scina sp. 0.00 0.20 0.42 0.00
The cumulative prey type curve for the entire data set was a Vibilia sp. 0.00 0.20 0.42 0.00
better fit with a logistic curve (R2 = 0.953, F(1,234) = 4795.8, Unid. Hyperidea 0.02 2.54 1.27 0.16
P < 0.001) than with a linear relation (R2 = 0.826, Total Amphipoda 0.23 7.05 8.47 0.65
F(1,234) = 1114.1, P < 0.001); therefore, the sample size was Cumacea
considered sufficient to describe bluemouth feeding habits in Diastylis cornuta 0.03 0.20 0.42 0.00
Unid. Cumacea 0.00 0.59 0.85 0.02
this area. Isopoda
Stomach contents analysis led to the identification of 512 Flabellifera
prey for a total weight of 43.52 g (mean prey weight in each Cirolanidae 1.31 3.91 5.93 1.50
stomach = 0.64 ± 2.68 g) belonging to 67 taxa, 45 of which Unid. Flabellifera 0.96 3.72 5.93 1.34
Asellota
were represented by benthic or epibenthic crustaceans Joeropsidae 0.07 0.98 0.85 0.04
(Table 1). Gnathiidea
In fact, crustaceans were the most important taxon in terms Gnathiidae 0.38 6.65 2.12 0.72
of %W (80.3), %N (83.8), %F (86.0) and %IRI (92.8). Among Unid. Gnathiidea 0.07 0.98 0.85 0.04
them, decapods were largely dominant by weight (%W = 68.7) Unid. Isopoda 0.15 3.13 2.97 0.47
Total Isopoda 2.97 20.16 17.80 4.11
and the most representative species was the brachiura Goneplax Mysidacea
rhomboides (%IRI = 56.8). Mysids were the best-represented Paramysis sp. 0.02 0.20 0.42 0.00
taxon in terms of abundance (%N = 27.4), followed by Unid. Mysinae 0.01 0.20 0.42 0.00
isopods (%N = 20.2) and decapods (%N = 16.8). Siriella sp. 0.25 2.35 4.24 0.53
Lophogaster typicus 3.06 8.41 12.71 7.04
Molluscs, Anellids, Fishes, Tunicates and Others repre- Unid. Mysida 0.09 3.13 4.24 0.66
sented only a small fraction of the H. d. dactylopterus diet; Unid. Mysidacea 0.48 13.11 12.71 8.34
their contributions in terms of the relative importance index Total Mysidacea 3.90 27.40 33.90 16.58
were < 6%. Copepoda
LevinÕs standardized index, calculated for the evaluation of Copepoda 0.00 1.17 1.27 0.07
Unid. Crustacea 4.15 9.78 19.07 12.83
diet breadth, was 0.41 for the biomass of 67 prey items. Total Crustacea 80.28 83.76 86.02 92.78
The dietary pattern of H. d. dactylopterus is graphically Mollusca
represented in Fig. 5a where the prey-specific abundance (Pi) is Gasteropoda
plotted against the frequency of occurrence (%F). As food Opistobranchia 1.55 0.59 1.27 0.13
Cephalopoda
categories, the lowest taxonomic level of the different food Sepiolidae 0.11 0.20 0.42 0.01
items was considered. Information on prey importance and
 14390426, 2010, 4, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1439-0426.2010.01467.x by Cochrane Portugal, Wiley Online Library on [24/10/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Age, growth and feeding habits of the bluemouth rockfish 587

Table 1 (a) 100

(Continued)

Prey types %W %N %F %IRI

Prey-specific abundance (%)

Teuthoidea 0.11 0.20 0.42 0.01 Lophogaster typicus
Unid. Cephalopoda 0.27 0.98 2.12 0.13
Total Mollusca 2.04 1.96 3.81 0.27 Goneplax rhomboides
Anellida Unid. Crustacea
Polychaeta 50
Alciopidae 0.69 0.20 0.42 0.02
Unid. Mysidacea
Fabriciinae 0.05 0.20 0.42 0.01
Nephtyidae 0.29 0.39 0.85 0.03
Polynoidae 0.08 0.59 1.27 0.04
Chloeia venusta 0.07 0.20 0.42 0.01
Unid. Polychaeta 0.03 0.20 0.42 0.00
Total Anellida 1.20 1.76 3.81 0.10
Teleostea 0
Stomiiformes
Argyropelecus hemigymnus 0.51 0.39 0.85 0.04 0.0 0.5 1.0
Perciformes Frequency of occurrence
Deltentosteus quadrimaculatus 0.46 0.20 0.42 0.01
Gadiformes
Macrouridae 0.54 0.20 0.42 0.02 (b)
Gadidae larvae 0.69 0.20 0.42 0.02
Myctophiformes
Myctophum punctatum 0.94 0.39 0.85 0.05
Hygophum benoiti 0.35 0.39 0.85 0.03
Lampriformes
Trachipteridae 0.70 0.20 0.42 0.02
Unid. Teleostea 7.17 4.50 9.32 5.25
Total Teleostea 11.37 6.46 13.14 5.44
Tunicata
Pyrosomida
Pyrosoma atlanticum 0.34 0.20 0.42 0.01
Salpida
Salpida 4.35 4.89 2.97 1.32
Total Tunicata 4.69 5.09 2.97 1.33 Fig. 5. Graphic representation of diet composition in Helicolenus
Others d. dactylopterus. (a) Diagram representing prey taxa. Only most
Siphonophora important species specified. (b) Explanatory diagram for interpretation
Calycophorae 0.04 0.39 0.85 0.02 of feeding strategy, prey importance and niche width contribution
Echinodermata (adapted from Amundsen et al., 1996)
Ophiuroidea 0.37 0.59 1.27 0.06

differences in diet observed between fish groups pooled in

feeding strategy of the predator can be obtained by examina- relation to depth and size (Table 2). These differences were
tion of the point distributions along the diagonals and axes of both statistically significant, attaining an R value of 0.11
the diagram (Fig. 5b). The percent abundance, increasing (P < 0.01) and 0.22 (P < 0.01), respectively, thus allowing
along the diagonal from the lower left to the upper right rejection of the null hypothesis (Table 3).
corner, provides a measure of prey importance, with dominant Pairwise comparisons showed that the dietary compositions
prey at the upper, and rare or unimportant prey at the lower were significantly different among depth groups except
end. The vertical axis represents the feeding strategy of the between depth 3 and 4 (Table 3). On the other hand, pairwise
predator in terms of specialization or generalization. tests confirmed significant differences in diet only between size
Feeding strategy indicates a specialization of individual fish groups 1 vs 3 and 1 vs 4.
(high between-phenotype contribution) on food items of high Applying the SIMPER routine, we obtained the relative
specific abundance and low occurrence; most of the species or contribution of each prey category to the dissimilarities
food categories are located on the left side of the diagram,
close to the vertical axis in a region of low importance. Table 2
Conversely, G. rhomboides located near the central area of the Number of samples for both variables (depth and size) considered in
graph, can be considered the most important prey at popula- statistical analyses of bluemouth diet
tion level, mainly due to the highest value of frequency of
Size 1 Size 2 Size 3 Size 4
occurrence (%F = 19.5). 4.0–6.3 cm 6.4–9.0 cm 9.1–13.7 cm > 13.7 cm Total

Depth 1
Diet variations with predator size and sampling depth 100–200 m 33 16 4 3 56
Depth 2
Twelve prey categories were selected by means of the binomial 201–300 m 10 19 10 6 45
test: Euphausiacea, Decapoda reptantia, Decapoda natantia, Depth 3
Amphipoda, Isopoda, Mysidacea, Other Crustaceans, Ceph- 301–400 m 25 52 16 8 101
alopoda, Polychaeta, Osteichthyes, Thaliacea, and Others. Depth 4
> 401 m 6 12 3 13 34
Two-way crossed ANOSIM (based on the data matrix of
Total 74 99 33 30 236
these 12 prey categories) allowed us to test the significance of
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588 P. Consoli et al.

Table 3 Sea. Results showed that the maximum total length of

Two-way crossed ANOSIM testing for differences in bluemouth diet H. d. dactylopterus in the study area was 27 cm, corresponding
among depths and sizes. Depth 1 (100–200 m), depth 2 (201–300 m),
depth 3 (301–400 m) and depth 4 (> 401 m); Size 1: 4.0–6.3 cm (age
to an estimated age by otolith readings of 21 years, even if only
0), Size 2: 6.4–9.0 cm (age 1), Size 3 9.1–13.7 cm (age 2–3), Size 4 a few samples were over 14 years old. Our maximum TL
> 13.7 cm (age > 4). *P < 0.05; **P < 0.01; ns, not significant recorded is, in most cases, lower than those reported for other
Mediterranean areas (Table 5); in particular, specimens of
R-value P R-value P 36 cm were found in the Alboran Sea by Massutı́ et al. (2000).
Among depths 0.112 ** Among sizes 0.221 * Observed differences in bluemouth length, as shown in
Pairwise tests R-value P Pairwise tests R-value P Table 5, could be due to the different fishing pressure in the
study areas, to the maximum sampling depth, and also to the
Depth 1 vs depth 2 0.243 ** Size 1 vs size 2 0.051 ns fishing gear employed. In fact, according to Massutı́ et al.
Depth 1 vs depth 3 0.220 ** Size 1 vs size 3 0.109 **
(2001), smaller sized individuals are concentrated at shallow
Depth 1 vs depth 4 0.183 ** Size 1 vs size 4 0.194 **
Depth 2 vs depth 3 0.194 ** Size 2 vs size 3 0.031 ns depths while larger ones prefer deeper areas, showing a clear
Depth 2 vs depth 4 0.352 ** Size 2 vs size 4 0.039 ns preference for rocky bottoms, which are not very accessible to
Depth 3 vs depth 4 0.066 ns Size 3 vs size 4 0.084 ns trawling, and thus they can escape or avoid the net.
Although we used two types of fishing gear (trawling and
bottom longlines to catch H. d. dactylopterus specimens, it is
among sizes and among depths at capture (Table 4). Mysids, possible that undersampling with bottom longlines introduced
isopods, walking decapods and other crustaceans were main a bias towards small fishes whereby the size-frequency distri-
contributors to the dissimilarity between size groups. In bution obtained could not reflect that of the population. In
particular, small fishes (size 1) fed mainly on mysids, whereas fact, a high percentage of the sample studied ( 59%)
walking decapods became an important food item for larger consisted of fish between 4 and 12 cm TL classes, correspond-
fishes (size 4). Isopods were consumed mainly by size 3 ing to juveniles from 0 to 1 years of age.
specimens. Regarding diet changes in relation to the bathym- It is clear that comparisons of the estimated bluemouth ages
etry, mysids represented the food item contributing most to with other authors may be done only when considering similar
the dissimilarity among depth 1 and the others, except size ranges. Nevertheless, maximum age obtained in our study
between depths 1 and 2; in particular they were more seems higher than that reported in other Mediterranean areas
abundant on specimens caught on the bottom at 100–200 m. (Ligurian Sea, Ionian Sea, lower Adriatic), as shown in
Walking decapods were the dominant prey of bluemouth Table 5.
specimens belonging to depth 2 (201–300 m) and contributed On the other hand, other authors (White et al., 1998; Kelly
the most to the diet dissimilarity between this bathymetric et al., 1999; Allain and Lorance, 2000; Abecasis et al., 2006)
range and the others. estimated a higher H. d. dactylopterus longevity in the Atlantic
Ocean. These differences could be due to the different age
reading methodologies as those authors used thin-sectioning
Discussion whereas we used whole otoliths. According to Allain and
The present study provides new information on biology and Lorance (2000) and Abecasis et al. (2006), when comparing the
ecology of H. d. dactylopterus from the southern Tyrrhenian two methods the sliced otoliths readings appear to be more

Table 4
Results of SIMPERanalysis showing average abundances (Av Ab) of fish taxa most contributing to diet dissimilarities (Diss %), according to the
factor size and depth. Values given only for groups with significant differences in accordance with ANOSIM results. Size 1: 4.0–6.3 cm, Size 2 6.4–
9.0 cm, Size 3 9.1–13.7 cm, Size 4 > 13.7 cm, depth 1 (100–200 m), depth 2 (201–300 m), depth 3 (301–400 m) and depth 4 (> 401 m)

Av Ab Av Ab Av Ab Av Ab Av Ab Av Ab

Taxa Size 1 Size 3 Diss (%) Taxa Size 1 Size 4 Diss (%) Taxa Depth 1 Depth 2 Diss (%)

Mysidacea 3.31 1.48 18.26 Other 2.23 1.53 18.77 Decapoda 1.22 5.34 35.75
Crustaceans reptantia
Isopoda 0.6 3.01 14.84 Decapoda 1.87 3.75 17.82 Mysidacea 5.21 0.68 31.25
reptantia
Other 2.23 1.11 14.64 Mysidacea 3.31 0.48 14.87 Other 2.01 0.51 13.23
Crustaceans Crustaceans
Depth 1 Depth 3 Depth 1 Depth 4 Depth 2 Depth 3

Mysidacea 5.21 3.04 31.75 Mysidacea 5.21 0.31 29.73 Decapoda 5.34 1.65 32.21
reptantia
Other 2.01 1.91 20.07 Other 2.01 2.3 21.68 Mysidacea 0.68 3.04 18.91
Crustaceans Crustaceans
Decapoda 1.22 1.65 12.19 Isopoda 0.06 2.65 20.89 Other 0.51 1.91 11.38
reptantia Crustaceans
Depth 2 Depth 4

Decapoda 5.34 2.06 34.42

reptantia
Isopoda 0.88 2.65 22.85
Other 0.51 2.3 18.5
Crustaceans
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Age, growth and feeding habits of the bluemouth rockfish 589

Table 5
Age-classes, von Bertalanffy growth parameters (Linf = asymptotic length, cm; k = growth coefficient, year)1; t0 = theoretical age at zero
length, years) and growth performance indexes (F) for Helicolenus d. dactylopterus, reported in different areas of Mediterranean Sea. W = whole
population, F = females, M = males, N = number of specimens

Depth TL Age
Source Area range (m) Methods Sex N range (cm) classes Linf k t0 F

Peirano and Tunesi (1986) Ligurian Sea 0–700 Whole otoliths W 300 4–28 0–9 70.7 0.05 )0.41 2.35
Ragonese and Reale (1992) Sicilian Channel 100–750 Whole otoliths W 585 – – 39.2 0.13 )1.46 2.29
DÕOnghia et al. (1992) NE Ionian Sea 160–550 Length-frequency W 1370 4–29 – 30.0 0.20 )1.41 2.24
analysis
DÕOnghia et al. (1996) NE Ionian Sea – Whole otoliths W 410 3–24 1–7 30.7 0.16 )0.90 2.17
Ungaro and Marano (1995) S Adriatic – Whole otoliths W – – 29.9 0.19 )0.85 2.23
Romanelli et al. (1997) S Adriatic – Whole otoliths W 434 5–34 1–10 – – – –
Massutı́ et al. (2000) Alboran Sea 64–786 Whole otoliths W 1455 3–36 0–30 30.0 0.10 )2.86 1.95
Massutı́ et al. (2000) Alboran Sea 64–786 Whole otoliths F 561 8–32 1–26 27.1 0.12 )2.65 1.95
Massutı́ et al. (2000) Alboran Sea 64–786 Whole otoliths M 575 8–36 1–30 32.3 0.09 )3.31 1.97
Massutı́ et al. (2000) Balearic Sea 25–1713 Whole otoliths W 938 2–30 0–22 29.9 0.13 )1.75 2.06
Massutı́ et al. (2000) Balearic Sea 25–1713 Whole otoliths F 178 7–28 1–22 27.0 0.16 )1.62 2.07
Massutı́ et al. (2000) Balearic Sea 25–1713 Whole otoliths M 198 8–30 1–21 32.5 0.10 )2.62 2.02
Present study S Tyrrhenian 100–600 Whole otoliths W 498 3–27 0–21 26.1 0.14 )1.92 1.98
Sea

realistic and more accurate for ageing older fish while also aster typicus) show quite high frequencies of occurrence that,
being more time consuming and costly. Whole otolith readings combined with their high prey-specific abundances and %W,
are cheaper and less time consuming, but should not be used suggest a slight population specialization towards these two
for older fish since they significantly underestimate age because species (which again lead to a quite narrow population niche
the otoliths are more opaque and the rings closer together in width). The role of G. rhomboides as a preferential food in the
older fish. Nevertheless, as suggested by Beamish (1979) and bluemouth diet was also reported by Froglia (1976) in the same
Bennett et al. (1982), divergence of ages in other scorpionfish- area (southern Tyrrhenian Sea) where this burrowing crab is
es, estimated by whole or sliced otoliths, usually occurs after abundant (Castriota et al., 2004).
20–25 years of age. As our estimated maximum age for In contrast with other studies (Collignon and Alonclè, 1960;
H. d. dactylopterus is 21 years, the possibility for error may be Madurell and Labropoulou, 2000), fishes were only occasion-
considered as negligible. ally found in the bluemouth gastric contents. However, the
Analysis of growth rates indicates that this species is a slow- occurrence of myctophid fishes in the diet, together with other
growing fish with k = 0.14 year)1, as confirmed in the pelagic prey performing vertical migrations, indicates that
Mediterranean Sea by other authors (Table 5). The theoretical H. d. dactylopterus could play a role in the energy transfer
asymptotic total length (Linf = 26.1 cm) was lower than the from pelagic to benthic domains.
observed maximum (27 cm). This could depend on the Statistical analysis showed a shift between feeding habits of
sampling effort, which was perhaps too small for larger fishes belonging to size 1 (4.0–6.3 cm TL) and larger fishes. In
individuals. In fact, in our sample only two individuals particular, as reported also by other authors (Madurell and
outnumbered the Linf value and only exceptional samples Labropoulou, 2000; Sartor et al., 2003), small fishes feed
reached ages beyond 15 years. The growth performance index mainly on mysids with a preference for L. typicus whereas
(F) from our study was somewhat lower than those obtained adults are feeders of reptantian decapods (mostly G. rhombo-
in other areas (Table 5). However, it is clear that there could ides). This ontogenetic shift toward bigger prey could be
be some differences between growth characteristics from one related to a size increase in the mouth gape of adult fishes.
area to another for reasons of food availability, hydrog- Moreover, the contribution of prey types appeared to be depth
raphical and climatic conditions, and fishing mortality rates. related, maybe because of the different composition of the
The length-weight relationship describes an isometric growth macrobenthonic communities along the bathymetric gradient.
for H. d. dactylopterus in the study area, confirming the results This study provides new information on the biology and
reported for the Balearic Sea by Massutı́ et al. (2000). ecology of the bluemouth, contributing to the knowledge
The feeding study provides an accurate description of the required by resource management. Nevertheless, more research
diet composition with an identification of a higher number of effort should be finalized to standardize age reading methods in
prey taxa (67), an improvement on previous studies (Froglia, order to better compare results from different study areas.
1976; Wurtz, 1977; Sartor et al., 1993; Madurell and Labro-
poulou, 2000).
Results put in evidence the habits of the bluemouth rockfish Acknowledgements
as a benthic feeder. Preferential prey categories are crusta- We are grateful to D. Bianca for practical help in the
ceans, as also reported in studies carried out along the laboratory and P. Rinelli for the collection of the biological
Moroccan coasts, western Mediterranean, and the Ligurian, samples.
Ionian and Tyrrhenian seas (Collignon and Alonclè, 1960;
Froglia, 1976; Wurtz, 1977; Macpherson, 1981; Sartor et al.,
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