Estuarine, Coastal and Shelf Science (1990) 30,131-140

Changes in the Heterogeneity of Mesoscale

Distribution Patterns of Larval Fish
Associated with a Shallow Coastal
Haline Front

A. Sabatts
Institute de Ci&cias de1 Mar, Paseo National s/n. 08003 Barcelona, Spain

Received 1.5 May 1989 and in revisedform 25 September 1989

Keywords: fish larvae; spatialheterogeneity;coastalhaline fronts; advection

process;WesternMediterranean
In June 1983there wasa largeoutflow of inland fresh-watersfrom the Rb6ne
River that coveredabroadareaoff the northern coastof Catalonia(Northwestern
Mediterranean),giving riseto a surfaceplumeof freshwaterwith adistinct haline
front at its southernedge.This hydrodynamic event brought about substantial
changesin the structure of the larval fish community in the area,which wasin
turn reflectedin an increasein spatialheterogeneity.Large numbersof species
and individualswerefound alongthe southernedgeof the plumein association
with the halinefront, whereas,with the exceptionof certain species,larval fish
concentrationsin the areacoveredby the plumetendedto beconsiderablylower
than in the rest of the area. Speciespredominantlylocatedin the surfacewater
layerswere probably transportedby the plume out to the plume’sedge.Other
species,which apparentlyenjoy a broadervertical distribution range,wereless
affectedby the surfaceadvectionprocesses.

Introduction
The distribution patterns of inshore larval fish communities naturally present a high level
of spatial and temporal heterogeneity as a result of the combined effects of a number
of often localized, sporadic factors reflected in mesoscale changes in zooplanktonic
communities in general (Denman &Powell, 1984). Factors on this scale,which, according
to a number of investigators, include such local hydrodynamic events asfronts (Le Fkre,
1986) and other advective processesgiving rise to convergence or divergence phenomena
(Haury et al., 1978), play a decisive role in determining the structure and variability of
zooplankton communities and hence the ichthyoplankton.
While mesoscalespatial heterogeneity may sometimesbe causedby biological factors,
local hydrodynamic factors would seemto furnish the most satisfactory explanations for
observed variations in the structure of larval communities (Flier1 & Wroblewski, 1985).
Mesoscale advective hydrodynamic processesmay transport larvae, thereby forming
aggregations in specific areas(Iles & Sinclair, 1982; Lobe1 & Robinson, 1986). However,
such aggregations do not appear to last more than a few days or weeks (Mackas et al.,
1985).
0272-7714/90/020131+ 10 $03.00/O @ 1990 Academic Press Limited
132 A. Sabatis

The general distribution pattern of fish larvae off the Catalan coast (Western
Mediterranean) basically follows a common pattern which, with certain seasonal vari-
ations, is repeated over the course of the year. Variability in the species composition of
the community is the most significant change, but on the whole the behaviour of the
community is rather conservative, particularly with respect to its spatial distribution. The
topography and hydrographic features of the area confer on the larval fish community a
general distribution pattern reflected in a gradual ‘ inshore-offshore ’ gradient of species
and individuals, with maximum concentrations of larvae occurring over the edge of the
continental shelf (Sabates, 1990). However, the presence of inland waters from the out-
flow of the RhBne River over the shelf in the Gulf of Lion, one of the most important
sources of inland water off the Catalan coast (Font, 1987), plays an important role in the
circulation pattern in the northern part of the region. The influx may, on occasion, rapidly
alter hydrodynamic conditions in the area, thereby affecting the structure of the epipelagic
larval fish community.
In June 1983, Castellon et al. (1985) reported the existence of a sizeable surface plume of
fresh water in the northern part of the Catalan coast which originated a strong haline front
at its southern edge. The authors pointed out that the influx of fresh water covered a much
larger area than was normal off the Catalan coast at the end of spring and attributed this
abnormal situation to an exceptionally high outflow of inland waters from the Rhone River
and other rivers in the area in spring and to the presence of a surface thermocline caused by
constant, heavy solar warming during a pronounced June calm, which combined to facili-
tate extension of the fresh water surface layer. This paper considers the effect of the above
mentioned surface plume of fresh water on the larval fish population off the Catalan coast.
The changes taking place in the spatial distribution pattern of larvae in response to this
phenomenon, and the increase in spatial heterogeneity with respect to the normal pattern
for larvae in the area, are discussed. The situation of the next month, July 1983, was
considered in order to compare the larval distribution pattern in the absence of the plume.
This situation can be considered as representative of the area and the species composition
was not very different between both months (Sabates, 1990).

Material and methods

Samples were taken over the entire area of the Catalan coast (Northwestern
Mediterranean) in June and July 1983. A total of 44 stations on 17 transects perpendicular
to the coast were visited. The number of stations on each transect depended upon shelf
width. The distance between the stations on a given transect, and between the transects,
was approximately 18 km (Figure 1).
Plankton was sampled by a Bongo net with a mouth diameter of 40 cm and a 300 l.trn
mesh size, equipped with a flow meter. Oblique tows were made from just above the
bottom to the surface or from 200m below the surface where the bottom depth was
greater. Samples were fixed in 50/Q formalin, buffered with borax immediately after
capture. Data were standardized to number of larvae 10 mP2.
Hydrographic conditions were recorded at all sampling stations by a conductivity-
temperature-depth probe (Neil Brown). Surface chlorophyll a was ascertained by con-
centrating samples in a Whatman GFjC filter, homogenizing the filtrate in acetone and
measuring the fluorescence of the extract (Yentsch & Menzel), 1963).
In order to obtain an objective description of the distribution pattern of the larval
population, a principal component analysis (PCA) was carried out for each month. The
 Larvalfish heterogeneity associated with a halinefront 133

E 10 20 30 40

42’ , _

00 50

41 0.

Western
Mediterranean

4c

Figure 1. Map of the study area and sampling stations.

PCA were performed on the speciescorrelation matrix after smoothing of the data by a
logarithmic transformation. To better appraise the spatial distribution of the patterns
observed, the samplescoresfor the first two factors obtained in each analyseswere plotted
on the station map.

Results

Temperature, salinity, and chlorophyll a

The surface distribution of temperature, salinity and chlorophyll a in July, followed the
general tendency observed in the area in the previous studies (Font, 1987; Font et al.,
1988; Estrada, 1981). In July, the temperature was slightly colder in the north than in the
southern part (Figure 2). In contrast, in June maximum values were recorded in the
northern third of the area, though values generally tended to be somewhat higher inshore
(Figure 2). In July, surface salinity values were similar between the north and the southern
part of the area, although they were slightly higher at the edge of the continental shelf
(Figure 2). In June, the main feature of note wasthe discontinuity between the Blanesand
Palambs submarine canyons. In this month, surface salinity ranged from 37% to below
30%0,reaching a minimum of 295%0in the centre of the plume of surface water. In the rest
of the area salinity wasslightly higher at the edgeof the continental shelf (Figure 2). Below
20 m salinity values were normal for the area, oscillating between 37.14 and 38.04%.
 134 A. Sabatds

(b)
4”
-A--
I

42

Temperature (“Cl
c

Salinity (%.I
-<

Chlorophyll 0 (/& [-‘I

Figure 2. Distribution of surface temperature (-C), salinity (%o) and chlorophyll a

(pg 1-l) values in (a) June and (b) July. The plot of surface salinity in June shows the
location of the haline front (after Castell6n et al., 1985).
 Larvalfish heterogeneity associated with a haline front 135

TABLE 1. Mean number of individuals per 10 m2 and frequency of occurrence of species

in the samples affected by the surface plume of fresh water (10 sampling stations) and in
the rest of the area (34 sampling stations)

Surface plume area Remaining area

n f(lO) n f (34)

1 Oblada melanura 25 30 113 79

2 Serranus cabrilla - 20 26
3 Sparidae Type 1 60 20 216 59
4 Trachurus trachurus 15 30 38 41
5 Mullus surmuletus - 17 18

6 Cepola macrophtalma 51 70 84 56
7 Buglossidium luteum 41 50 33 9
8 Engraulis encrasicholus 2658 100 1369 94
9 Arnoglossus thori 113 90 80 12
10 Callionymus risso 34 70 20 20
11 Gobiidae Type 1 140 70 41 68

12 Ceratoscopelus maderensis 15 20 21 38
13 Hygophum hygomii 11 10 30 21
14 Lampanyctus crocodilus 12 30 25 41
15 Maurolicus muelleri 9 10 11 18
16 Cyclothone braueri 6 10 47 18
17 Benthosema glaciale 18 10 26 21

18 Myctophum punctatum 17 40 10 29
19 Coris julis 8 10 11 26
20 Callionymus maculatus 17 20 30 47
21 Sparus pagrus 6 10 11 29
22 Serranus hepatus 18 50 33 41
23 Solea Iascaris - 3 9

In July, the concentration of chlorophyll a in the surface layer showed slightly higher
values inshore all along the area (Figure 2). In June this tendency was observed in the
southern two-thirds of the area. Nevertheless, chlorophyll a values were generally higher
in the northern third of the area, in associationwith the plume of lesssaline water. At the
haline front the concentration was much higher, about three times the maximum value in
the rest of the area (Figure 2).

LarvalJish distribution
Table 1 setsout the mean concentration and frequency values for speciescollected in June
at stations in the northern part of the area, where salinity values were very low, aswell asin
the rest of the area. The 23 speciescollected in June were classified into four groupings
according to the PCA (Figure 3) representing the main distribution patterns of fish larvae
for the particular month. The first group consisted of five specieswhich were scarcein the
zone affected by the surface plume of fresh water but were present throughout the rest of
the area. These species,like Oblada melanura, Sparidae Type 1, and Trachurus trachurus
were present at high density close to the southern edge of the plume (Figure 4). The
secondgroup wasmade up of six speciesabundant in the zone covered by the surface water
plume (e.g. Buglossidium luteum and Callionymus risso) (Figure 4). Some of the species
included in this group, such asEngraulis encrasicholus, were widely distributed over the
136 A. Sabatis

entire area, and others (e.g. Cepola macrophtalma) showed high concentrations of indi-
viduals at the southern edge of the plume (Figure 4). The third group comprised six
speciesof mesopelagicfishes. The larvae of this specieswere located preferentially over
the shelf break all along the area, including somestations in the area affected by the plume.
The fourth group wasmade up of the few abundant speciesand others with poorly defined
spatial patterns.
Figure 3 alsoshowsthe larval distribution pattern in June and July obtained by plotting
the sample scoresfor the first two factors of both PCA on the stations map. The spatial
distribution of the sample scoresfor the first factor in July followed the general distri-
bution pattern of larval population in the area (Sabates, 1990) where, the positive values
were located along the edgeof the continental shelf and the negative values closer inshore.
Nevertheless, in June this general pattern wasonly observed in the southern two-thirds of
the area. In the north, a sharp discontinuity wasobserved in the region between the Blanes
and Palamossubmarine canyons, where the most positive values were aggregated, while
the most negative values were located in the remainder of the northern third of the area. In
both months, this first factor is closely related to the number of speciesand number of
larvae, since the most positive factor scoreswere clustered in the area where speciesand
individual abundance were highest. The spatial distribution of the samplescoresfor the
secondfactor in July, separatedthe zones where the continental shelf is broadest (positive
values) from the rest of the area. This tendency was one of the most important character-
istics of the spatial larval distribution in the Catalan coast,after the tendency explained for
the first factor (Sabates, 1990). Nevertheless, in June the tendency observed was com-
pletely different and the northern third of the area (negative values) was clearly differ-
entiated from the rest of the area. This second factor in June is closely related to the
abundance of somespeciesin the zone influenced by the surface plume of fresh water.

Discussion
The plot of the two first factors from the PCA of June and July pointed out the existence of
a well-differentiated zone in the northern part of the area during the month of June,
coinciding with the strong haline front. Larval fish concentrations tended to be lower in
the region covered by the surface plume of fresh water than in the rest of the study area.
This suggeststhat many of the speciesmay have been transported by the surface plume,
resulting in higher speciesand individual abundance levels in the region of the haline
front, owing either to passiveaggregation or to high productivity levels associatedwith the
front.
Speciespresent in high densities along the edge of the plume of fresh water but very
scarcein the region covered by the plume, displayed a preference for the uppermost layers
of the water column. Certain authors have referred to the distinctly neustonic habitat of
the larvae of someof these species,e.g. Serranus cabrilla (Fage, 1918), Oblada melanura
(Aboussouan, 1964) and other speciesof the samegenus as Mullus barbatus (Sabates &
Palomera, 1987), which probably occupies a similar habitat to that of Mullus surmuletus.
Trachurus trachurus hasa wider vertical distribution, although it may display a tendency
to concentrate in the upper layers (Russell, 1976; John, 1985). This may explain why they
were swept along by the plume to form concentrations at its edge, besidesoccurring in the
rest of the area. Species which were found in the region covered by the plume have a
broader distribution through the water column, e.g. Callionymus risso and Arnoglossus
spp. (Russell, 1976; Southward & Bary, 1980). The distribution pattern of these species
 Larvalfish heterogeneity associated with a haline front
137

(0)
E I0 0 0 40
IU
(I 1 -0.5
/,

FIRST FACTOR

II
_’
,‘.I3 l 12 3

SECOND
FACTOR -. ‘-_. -L&I
,’
I

(b)

FIRST FACTOR SECOND FACTOR

Figure 3. (a) Spatial representation of the sample scores values for the first two factors
obtained in the PCA of June (i), and representation in the space of the first two axes, of
the species collected in June (ii) (For the code, see Table 1). (b) Spatial representation of
the sample scores values for the first two factors obtained in the PCA of July.
 138 A. Sabatis

I0 20 3” 4”
h cc

42’

41c

40’
0. melonuro
L
$Jc ; /’ S’ridae /

-rJ

Figure 4. Spatial distribution of six species representing different distribution patterns

observed in the area during June 1983 under the effect of the surface haline front.
 Larvalfish heterogeneity associated with a hafine front 139

made it unlikely that they would be affected by the plume; they were mostly located in
deeper layers, thereby avoiding its effects. Other species included in this group may
display a tendency to concentrate in the upper layers, e.g. Cepola macrophtalma. Thus,
probably only the larvae of those speciesoccupying the uppermost layer were transported
out to the edgeof the plume, while the larvae in the deeper layers were beyond the range of
the plume’s effects and remained just below the plume throughout the area. Engraulis
encrasicholus, the most abundant specieswith a broad spatial distribution in the area,
showed high densities in the zone affected by the plume (Palomera & Sabates, 1988). This
could be explained by the well-known ability of this speciesto spawn over a broad range of
salinities (Demir, 1965).
Mesopelagic fish larvae showed a clear distribution pattern; they were located over
the shelf break all along the areaand occurred in low densities within the zone covered by
the plume. Such speciesexhibit a relatively deep vertical distribution, with maximum
abundance levels occurring at depths below 25 m (Dekhnik & Sinyukova, 1966; Maso &
Palomera, 1984). The deep distribution of mesopelagicfish larvae implies that they were
located below the surface plume in the north of the area.
Transport mechanisms, particularly in the coastal zone, exert a considerable effect on
biological processes. Such mechanisms generate areas of hydrographic convergence
which result in a simultaneous aggregation of organismsof all sizes,giving rise to areasof
high productivity. These concentrations of individuals, brought about by the surface
advection processes,do not necessarily imply mobility of the organismsinvolved (Mackas
er al., 1985). Furthermore, the small size of most of the larvae collected at the edge of the
plume ( < 6 mm) indicates their reduced capacity of movement. It is, thus, more likely that
most of the larvae were transported into the frontal zone at the edge of the water plume.
Hydrographic phenomena, such asthose considered here, may benefit the larval popu-
lation by leading to the concomitant concentration of organisms that make up some of
their prey (Cote et al., 1986). This was true of the phytoplankton, which was present in
high concentrations in the frontal zone (Figure 2). Although larvae are most directly
related trophically with the microzooplankton (Jenkins, 1988), phytoplankton levels
like those observed probably occur in close association with similar microzooplankton
concentrations in the surface layers (Mackas et al., 1980).
In conclusion, the substantial and unpredictable increase in larval concentration
observed in a specific zone off the Catalan coast, appeared to be associatedwith local
hydrodynamic events. The perturbation took place only in the surface layers, and the
various specieswere affected differently, probably depending on their range of vertical
distribution.

Acknowledgements

This study was partly carried out with the financial support of the Fundacion Ramon
Areces. I wish to thank Mrs M. Mad for her useful comments and Mr R. Sacks for his
help with the English version. Also thanks to Dr J. M. Gili for helpful suggestionsand for
critical reading of the original manuscript.

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