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Distribution pattern of decapod larvae off the north-western Iberian Peninsula

coast (NE Atlantic)

Article in Journal of Plankton Research · January 1991

DOI: 10.1093/plankt/13.1.217

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Journal of Plankton Research Vol 13 no 1 pp.217-228, 1991

Distribution pattern of decapod larvae off the north-western Iberian

Peninsula coast (NE Atlantic)

Xavier Fust6 and Josep-Maria Gili

Institut de Ciincias del Mar, Passeig National sin, 08003 Barcelona, Spain

Abstract. In October 1977 the model of general circulation of the water masses off the coast of
Galicia, and the presence of a coastal upwelling, led to a high primary productivity. This high
productivity in rum favoured the development of a rich population of decapod larvae. The

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abundance and distribution pattern of these organisms were closely linked (I) to the abundant
presence of the corresponding adult species in the area, (li) with the spatial distribution of
phytoplanlctonic populations concurrently studied by Estrada [/. Plankton Res., 6, 417-434 (1984)]
and (iu) with the hydrodynamic pattern in the area. Fifty-two decapod larvae taxa were identified
and Solenocera membranacea, Pisidia longicornis, Pdumnus hirtellus and Goneplax rhomboidcs
were the most representative species It was observed that the greatest concentrations of larvae (3387
larvae 10~2 m~3) were to be found near the mouth of the Rfas Baixas (situated in the south-west of
the coastal area) and in some zones further out to sea (863 larvae 10"2 m"3) (due to a process of
hydrodynamic transport)

Introduction
The north-western coast of the Iberian Peninsula is situated at the northern limit
of the areas of upwelling associated with the anticyclonic gyre in the North
Atlantic. The studies by Fraga (1981) and Fraga et al. (1982) reveal the existence
of a very heterogeneous hydrographic pattern, resulting from the upwelling
regime, the distribution and dynamics of the water masses and the continental
apports from the Rfas Baixas (flooded tectonic valleys). The model of general
circulation defines two differentiated areas (to the west and to the north of the
coast of Galicia), on the basis of the nature of the water masses that converge at
Cape Finisterre, giving rise to an area favourable to coastal upwelling. Along the
west coast, the subsurface North Atlantic Central Water (NACW) flows in a
northerly direction, while the surface water flows in the opposite direction
(towards the south). Along the north coast, the subsurface Central Water of the
Gulf of Biscay (CWGB) flows in a south-westerly direction, as does the surface
water in the area. Furthermore, on the west coast great quantities of fresh water
flow into the coastal waters from the Rfas Baixas, especially during the rainy
season (autumn); on the other hand, when upwelling is at its height (summer),
shelf water penetrates into the RJas.
In the coastal areas there is a spatial and temporal connection between
hydrodynamic and biological processes (Denman and Powell, 1984). This is due
essentially to the fact that physical processes become advective processes, which
in turn give rise to local enriching (areas of upwelling) or local accumulation
(areas of convergence or divergence) (Mackas et al., 1985). On the basis of these
considerations, it is to be expected that the hydrographical heterogeneity along
the coast of Galicia would give rise to changes in the overall distribution pattern
of planktonic organisms. This increase in the heterogeneity of the plankton
might be reflected in specific areas of high concentration and production, as

© Oxford University Press 217

X.Fust* and J.-M.G01

opposed to other poorer areas that are nevertheless not very far away. This
hypothesis is corroborated by the results obtained by Estrada (1984), who made
a concurrent study of the phytoplankton distribution in the area.
Of the organisms that form the neritic zooplankton, decapod larvae constitute
an important part of the meroplankton (e.g. Kurian, 1956; Bourdillon-
Casanova, 1960; Roff et al, 1984; Fustf, 1987; Paula, 1987; Locke and Corey,
1988). Local hydrographical conditions, especially the regime of currents, are
vital if the viability of larvae is to guarantee the recruitment and continuity of the
populations of benthic adults (e.g. Makarov, 1969; Scarrat, 1973; White et al.,
1988). Nevertheless, despite the fact that the larval phase forms part of the

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development process of the great majority of crustaceans in temperate areas,
there is a singular lack of studies describing the spatial distribution of larval
populations (e.g. Sastry, 1983).
Decapod crustacean fauna is very abundant off the coast of Galicia, and has
been widely studied by Gonzalez-Gurriaran (1982, 1986, 1987). However,
studies carried out on decapod larvae are limited to general observations
concerning the zooplankton in certain Rias (Corral and Alvarez-Ossorio, 1978).
This present work sets out to establish the distribution pattern of decapod
crustacean larvae in the area, and to determine its relationship with local
hydrodynamic and biological processes.

Method
Samples were collected during the Galicia IV cruise, carried out by the Marine
Sciences Institute of Vigo between October 7 and October 28, 1977. During the
cruise, hauls were made between 45°N 7°W and 41°N 11°W. For the study of the
composition and distribution of decapod larvae, 39 inclined hauls were carried
out (Figure 1). These hauls were made from 200 m below the surface, or from
the sea bottom to the surface in shallower stations. Bongo-type nets (60 cm
mouth diameter) were used with 505 u-m of mesh size. Samples were collected
over a 24 h cycle but mainly were made during the night. All the decapod larvae
thus caught were separated and classified in the laboratory. The density of the
larvae has been established at number of individuals per 100 m3, on the basis of
the theoretical volume filtered by the nets. The information related with
environmental factors (temperature and salinity) and chlorophyll, concurrently
studied during the cruise has been summarized in Figures 2 and 3. Additional
information concerning the hydrographical and biological parameters obtained
during the cruise can be found in Fraga et al. (1978) and Manriquez et al. (1978).
A Principal Component Analysis (PCA) was used to obtain a simplified and
objective representation of the information obtained in the larvae inventories. A
self-developed program package was used (LAWI, Dr J.Lleonart). The analysis
was based on the correlation matrix of abundance correlations between the 31
most common species (only those species constituting > 10% of the total hauls
were considered), after smoothing of the data by logarithmic transformation
(Legendre and Legendre, 1979). For the interpretation of factors, the sample
scores were plotted on the station map.

218
 Distribution pattern of decapod larvae

8° W

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Fig. 1. Cruise Galicia IV. Position of stations and geographical locatjon of study area

Results
Generally speaking the distribution of decapod larvae off the coasts of Galicia
responds to the overall model described by different authors for different oceans
(Rees, 1955; Kurian, 1956; Bourdillon-Casanova, 1960; Makarov, 1968; Sastry,
1983). Larvae are most abundant in neritic zones, while their concentrations
decrease offshore. Nevertheless, this general distribution pattern has particular
characteristics here, specifically a greater spatial heterogeneity as regards the
overall model: the highest concentrations of larvae were detected near the
south-west coast, at the mouths of certain Rfas (Figure 4). In other stations in
the area relatively large quantities of these organisms were found, the greatest
anomalies being the concentrations encountered in stations 2 and 3, at a
considerable distance from the coast. In the north-east zone relatively high
concentrations were found at station 23 (Figure 4) to the north of Finisterre. As
we hope to show later, these high levels of concentration are probably associated
with the overall circulation model, which determines a greater upwelling
influence in this area.
When the species are examined one by one, different distributions were
shown: some, e.g. Pisidia longicornis, are particularly abundant right in the
mouth of the Rfas Baixas (Figure 4), while others, e.g. Goneplax rhomboides,
219
X.Fustt and J.-M.GUi

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8° W

Fig. 2. Surface salinity and temperature, during the Galida IV cruise (October 1977) Redrawn from
Estrada (1984)

05

10° 8° W

Fig. 3. Weighted averages of chlorophyll a concentration in a layer from 0 to 50 m depth (or from
0 m to the bottom when it was shallower than 50 m). Redrawn from Estrada (1984).

220
 Distribution pattern of decapod larvae

8° W

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Fig. 4. Spatial distribution of total number of larvae and of three species showing the three main
. distribution pattern explained in the text.

were captured mainly over the continental shelf as from the 100 m isobath, being
very scarce in the coastal area (Figure 4). On the other hand, larvae of species
such as the sergestid Solenocera membranacea were invariably captured far from
the coast (Figure 4).
The principal component analysis (PCA) of the 31 species selected (Table I)
offers a more concise representation of the larval population distribution in the
area. The first two components account for 51.58 and 13.4% respectively of
the total variance, and the representation of the sample scores for these
221
X.Fustt and J.-M.Glli

Table I. Mean (jt) in larvae 100 m'} and standard deviation (s) of the species, calculated considering
the number of samples (n) in which they occurred from a total of 39 samples

Speaes x s n

1 Solenocera membranacea (Risso, 1816) 17 44 28.95 25

2 Sergestes spp. 1 33 0.51 6
Palaemon elegans Rathke, 1837 3.25 2.85 3
Palaemon serratus (Pennant, 1777) 500 _ 1
3 Palaemon spp. 15.12 23 01 8
4 Alpheus glaber (Olivi, 1792) 306 1.86 15
5 Alpheus spp 3.33 3 42 9
Athanas nuescens (Leach, 1814) 200 0 2

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6 Hippolitidae undetermined 20.21 4166 14
7 Processa spp. 1.66 1 11 9
8 Plesionika spp 25 85 30.97 20
9 Pandahna brevirostns (Rathke, 1843) 266 308 9
Pandalidae undetermined 1.75 1.50 3
10 Crangon crangon (Linnaeus, 1758) 9.20 860 10
Phdocheras bispmosus (Hailstone, 1835) 200 — 1
Phdochcras sculptus (Bell, 1847) 35.00 - 1
11 Phdocheras spp. 32 58 38 09 17
12 Scyllarus arctus (Linnaeus, 1758) 1 46 1 30 15
13 Calhanassa subterranea (Montagu, 1808) 4 14 484 7
Jaxea nocturna (Nardo, 1847) 100 0 3
14 Upogebia sp 39.55 81.71 9
15 Diogenes pugilator (Roux, 1829) 31.25 57 08 8
16 Catapaguroides timidus (Roux, 1830) 7.91 10.39 12
Pagurus bernhardus (Linnaeus, 1758) 300 264 3
17 Pagurus cuancnsis Bell, 1846 2.20 1.64 3
Pagurus prideauxi Leach, 1814 3.00 - 1
18 Pagurus spp. 14 31 19.31 16
Anapagurus laevis (Bell, 1846) 2.50 3.00 3
19 Anapagurus spp. 14.85 18 74 14
Pagundae undetermined 100 0 2
20 Galathea intermedia Lilljeborg, 1851 3.57 4.96 7
Galathea nexa Embleton, 1834 00 0 2
Galathea disperse Bate, 1859 1.50 0.70 2
Galathea sp. ;»50 2 12 2
21 Pisidia longicomis (Linnaeus, 1757) 3<j.33 670.90 12
22 Latreilla elegans (Roux, 1830) 14 0 37 7
23 Ebalia spp I 77 1 09 9
Macropodia sp .00 - 1
24 Atelecyclus spp 6 47 9.35 17
Cancer pagurus (Linnaeus, 1758) 1.33 0 57 3
25 Corystes cassivelaunus (Pennant, 1777) 17.83 22.52 6
Pinmela denticulala (Montagu, 1808) 1.00 _ 1
26 Thia scutellata (Fabricius, 1793) 3 78 7.39 14
Caranus maenas (Linnaeus, 1758) 3.50 3 53 2
Portumnus latipes (Pennant, 1777) 1.66 1.15 3
27 Necora puber (Linnaeus, 1777) 320 2.94 3
28 Polybiinae undetermined 81.58 127.83 24
29 Pdumnus hirtellut (Linnaeus, 1761) 101.21 146.74 14
30 Goneplax rhomboides (Linnaeus, 1759) 20.95 24 31 24
31 Brachynotus sexdentatus (Risso, 1827) 6.25 10.13 8
Grapsidae undetermined 1.00 _ 1
Palicus corona (Roux, 1830) LOO 0 2

The numbers in the left margin correspond to the codes of the species in the representation of the
PCA

222
 Distribution pattern of decapod larvae

components can be seen in Figure 5. In this figure, the most positive values of
the first factor are to be found in the area of influence of the Rias, the zone
where the greatest concentration of larvae was found. To the south of the Rias
(stations 1 and 4), positive values for the sample scores were also observed, in an
area where larvae were also abundant. In this latter zone the chlorophyll a and
phytoplankton values (Estrada, 1984) were relatively high. The same can be said
of the area near La Coruna (stations 23, 24 and 27) and for the north-east zone
(stations 35 and 36) (Figure 5). This afirmation can be corraborated with the
high correlation between this first PCA factor and the hydrographic and
biological data (see Table II).

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Those species with high correlations with the first component (Figure 6) are
characterized by being highly abundant near the mouth of the Rias (e.g.
P.longicornis), or else by having been captured only in the zones of influence of
10°

Fig. 5. Representation of the curves calculated with the values of the factor scores in all the samples
of decapod larvae for the first axis (left) and the second axis (right) in the PCA

Table n . Sperman Rank Correlation between the two first factors of the PCA /scores for decapod
larvae species and the surface temperature, surface salinity, chlorophyll a (average of 0-50 m),
number of individuals and species of larvae collected

1 axis 2 axis Temp Sal. Chlo. Indiv. Species

PCA 1 axis 1
PCA 2 axis 0 49* 1
Temperature -0.69** - 0 42* 1
Salinity -0.78'* -0.31 0 83" 1
Chlorophyll a 0 74" 0 24 -0.52* -0.61* 1
Number of individuals 0.96" 0.52* - 0 69" -0 75" 0 69" 1
Number of species 0.97" 0 57* -0 67" - 0 77" 0.70" 0.97** 1

Only the significant correlations are shown, and probability is indicated by * * / > & 0 9 9 ; or
" / » ; = 0.95.

223
X.Fustl and J.-M.GUi

PC II

J
'.30

,22
,2
.9
.12 .23 .28

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24* .11
3
.16
PCI .29
.5
17
'•3 .26 ' l 0
.6

•15

Fig. 6. Representation in the space of the first two axes, of the 31 most common species See
explanation in the text (for the code, see Table I)

the Rfas and in the most coastal areas (Crangon crangon, Corystes cassivelaunus,
PUumnus hirtellus, Callianassa subterranea). The area of dispersion of the larvae
of these species is very similar to the area of distribution of their corresponding
adults, which mostly have coastal habitats (Zariquiey Alvarez, 1968; Gonzalez-
Gurriaran, 1986). On the other hand, Philocheras spp. and Plesionika spp. have
a wider distribution over the platform than the aforementioned species. These
larvae are more abundant in the areas influenced by the remains of upwelling, in
the north-eastern part of the coast. The negative values of the sample scores for
this first factor correspond to the dispersion area of the species captured far out
from the shelf (Solenocera membranacea, Latreilla elegans and Sergestes spp.).
The second PCA factor (Figure 5) makes a clear differentiation between the
zones where larvae of not strictly coastal species dominate (more positive sample
score values) and the areas characterized by the abundance of larvae preferring
estuarine or littoral habitats (more negative values). The temperature follows
the same spatial pattern indicated by the larvae and shows a good correlation
with this second factor (Table II).
The species with high positive correlations with the second component, e.g.
Solenocera membranacea, Alpheus glaber and G.rhomboides, are characterized
224
 Distribution pattern of decapod larvae

by their relatively high rate of capture in hauls a long way out from the coast
(S.membranacea), or else by their positive abundance gradient towards the open
sea. The most negative values correspond to species so abundant in the area of
the Rfas that their distribution is practically restricted to these zones (P.longi-
cornis, C.cassivelanus, C.subterranea, Upogebia spp.).

Discussion
The distribution pattern of the larval population observed during October 1977
mainly reflects the hydrographic structure, which originates from the interaction

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of upwelling produced by the action of the two hydrographic regimes, described
in the Introduction, and the important out-flows coming from the Rfas. In the
areas where the Rfas Baixas reach the sea, the interaction between the remains
of upwelling, which persist between Ria Arosa and Finisterre Cap, and the
relative stability of the water inside the Rfas, favoured the development of
phytoplanktonic populations (Estrada, 1984). This hydrographic structure
(interaction between the upwelling and the Rfas), which is permanent between
March and September and persists until the end of October, is favourable for
many benthic and planktonic species whose life cycle takes place totally or
partially in these estuaries (Corral and Alvarez-Ossorio, 1978; Tenore et al.,
1982). Many species of decapods, essentially Brachyura (Gonzalez-Gurriaran,
1986; Perez and Roman, 1979), carry out their breeding processes in this habitat.
Consequently the abundance of the larvae of some species of decapods,
observed in the neritic zones near the Rfas during the Galicia IV cruise, to the
presence of the corresponding adults in the area and the existence of
environmental conditions favouring their development. In this sense, the highly
negative salinity gradients, the result of the discharge of fresh water from the
Rfas (Estrada, 1984), do not appear to affect the adult and larval populations of
brachyural decapods, capable of breeding and developing in areas of high
environmental instability (Warner, 1977).
The faunistic composition and distribution pattern of decapod larvae reflect
mainly the influence of the Rfas, but also seem to be linked to the distribution of
water masses. In this sense, the relatively large quantities of these organisms
detected in the south-west sector might be explained by the model of circulation
of surface and subsurface waters (Fraga et al., 1982) and its corresponding
influence on fertilization processes (Estrada, 1984).
In zone A of Figure 7 (figure e in Fraga, 1981), the subsurface water formed
by NACW between 75 and 400 m moves in the opposite direction to the surface
water that flows along the coast of Galicia from the north-east to the south. This
opposition in the movement of the two bodies of water contributes to a growth in
fertility in the south-west region where, in the stations situated further south,
Estrada (1984) found relatively high values of chlorophyll a. This system of
circulation may influence the larval population, which might be transported
southwards by the surface water. Specifically, it has been observed that
numerous larvae of some coastal species, e.g. P.longicornis, P.hirtellus and
Palaemon spp., the first two very abundant in the coastal stations of the Rfas,
225
X.Fuste and J.-M.GU1

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6°W

Fig. 7. Scheme of circulation of the Central Water between 100 and 400 m depth. The shaded area is
the zone of lateral contact between NACW and the CWGB. Redrawn from Fraga el at. (1982)

have been captured in stations 1 and 4 (Figure 1); the surface water that
descends along the coast seems partially to scatter the coastal larval populations,
carrying them towards the south. While by no means an unusual phenomenon,
the passive transport of larvae of some coastal species offshore might, probably,
have a negative effect on recruitment. On the other hand, hydrodynamic
mechanisms similar to the one observed above may return the larvae to the
adults' area of development (Smith, 1979; Shanks, 1983). Thus, although it
would seem that larval loss takes place in this area, the combination between
processes of larval development and those of hydrodynamic transport linked to
production processes may in fact increase the larvae's survival.
The relative abundance of larvae in stations 2 and 3, at a considerable distance
from the coast, is due to the presence of larvae of non-coastal species
(S.membranacea), or those of widely distributed species {G.rhomboides), in an
area relatively rich in chlorophyll a and phytoplankton cells (Estrada, 1984).
In the zone G (north-eastern), the CWGB subsurface waters between 75 and
400 m and the surface waters between 0 and 75 m flow in the same direction.
Therefore the planktonic organisms are most probably carried towards
Finisterre Cap (Fraga, 1981). Decapod larvae are less common in this area than
in the south-western zone, and only in the upwelling area situated slightly to the
north of Finisterre has a relatively high concentration of larvae been observed.
This phenomenon might be the result of the lesser density of the adult
population in this area in relation to that of the Rias (Gonzalez-Gurriaran, 1986;
Perez and Roman, 1979).
In general, the spatial heterogeneity observed in the distribution of larvae is
determined both by the presence and density of adults and by the local
226
 Distribution pattern of decapod larvae

hydrographical pattern. Littoral flow and the effect of upwelling enhance

primary productivity and, in consequence, favour the accumulation of larvae in
the most productive areas of the coast of Galicia.

Acknowledgements
We wish to thank Dr Marta Estrada for her encouragement and help during the
writing of the manuscript and to all colleagues of the Galicia IV cruise.

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Received on October 28, 1990; accepted on October 6, 1990

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