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Species Identification | Stock Definitions | Distribution | Age and Growth | Fecundity | Nutrition and Feeding Habits | Reproduction and Recruitment | Physiology and Role in Ecology | References

The horse mackerel (Trachurus trachurus and T. trachurus capensis) is a member of the large carangid family, which includes many important commercial species world wide. The name of the horse mackerel is misleading, as the true mackerel-like fishes such as tunas or bonitos belong to the scombrid family. Horse mackerel is a schooling species, caught mainly with pelagic nets, but close to the sea floor. T. trachurus is commonly found from the waters off West Africa/Cape Verde Islands to the Norwegian Sea, including Iceland, as well as in the Mediterranean and Black Sea. It is the most northerly representative of the trachurid sub-family, which is widely distributed in the world’s seas and often supports important fisheries.

Three species of the Trachurus genus, T. trachurus, T. mediterraneus and T. picturatus, are found together in the northeast Atlantic waters and are commercially exploited in parts of the ICES Sub-area VIII and Division IXa. Studies on genetic differentiation showed three clear groups corresponding to each species of Trachurus with no intermediate principal component scores, excluding the possibility of hybrids between the species (ICES, 1998).

The most obvious feature to distinguish these three species is the length of the accessory (dorsal) lateral line (Nümann, 1959). In T. trachurus, the accessory lateral line extends to below the soft dorsal rays 23-31, in T. mediterraneus it ends below the 8th dorsal spine to the 3rd soft ray (see Figure 1 below). T. picturatus shows an intermediate length of the accessory lateral line, ending below soft dorsal rays 5-10. Other features used for species discrimination are the shape, number and diameter of the scales of the curved lateral line, and the number of gill rakers.

Fig. 1: Characteristic features of the different horse mackerel species in European waters. After Nümann, 1959, modified.

It is assumed that there are three distinct spawning populations of T. trachurus in the northeast Atlantic (section 4.3 of ICES, 1998):

• the Southern horse mackerel around the Iberian peninsula;
• the Western horse mackerel in the Norwegian Sea, northern North Sea, western part of Skagerrak, west and south off the British Isles, western Channel and west off France;
• the North Sea horse mackerel, mainly restricted to the central and southern North Sea, eastern part of Skagerrak, Kattegat and eastern English Channel.

Earlier stock discrimination studies have not provided firm evidence for the existence of true horse mackerel stocks (Polonsky & Baydalinov, 1964, Nazarov, 1976 cit. in ICES, 1992). However, one genetic study provided a significant separation between a southern and a northern component (Nevedov et al., 1978). The southern stock is commonly mixed with T. mediterraneus (Polonsky & Baydalinov, 1964). From circumstantial evidence it is concluded by ICES (1992) that there is little exchange between the southern and the western units and these could therefore be regarded as stocks. This separation is also based on the observed egg distributions and the spatial and temporal distribution of the fishery. The 1995 (ICES, 1996) and 1998 egg surveys covered both the southern area and western area. Spawning areas have been defined for the North Sea by a series of egg surveys from 1988-1991 (Eltink, 1990, 1991 and 1992). However, there are transition zones with low egg abundances between the main spawning areas of the western and the southern as well as the western and North Sea area (ICES, 1996), indicating no clear separation of stocks.

Attempts to separate stocks have included discriminative measurements of the length of otoliths at age 1 (L1) (Marecos, 1986). A tagging experiment carried out off Spain in 1997 did not result in any returns. As a result of this, and a lack of any newer conclusive evidence to indicate that the Western and Southern stocks are independent units, a degree of uncertainty exists concerning the true identity of these stocks (ICES, 1992).

Kerstan (1991) found significant morphological differences between North Sea and Western stock members, at least when using age stratified samples. However, horse mackerel is a migratory species and the distribution of the Western and North Sea component overlap extensively during over-wintering in the western Channel. The overlap during summer feeding in the Skagerrak is relatively small (ICES, 1998).

Shelf attachment is a predominant distributional pattern of horse mackerel. For the North Sea stock, a distribution close to the southern and eastern coast was already given by Olsen (1883). Western and Southern stock components are closely connected to the shelf contour, as demonstrated on a number of occasions (e.g. Dornheim, 1987; Macer, 1977; Dornheim & Kerstan, 1985; Eaton, 1989; Dornheim, 1993; Porteiro et al., 1993) (see Figures 2 and 3 below).

Horse mackerel show distinct areas for spawning, feeding and over-wintering, what is most evident in the case of the western stock (Borges et al., 1995). Migration might be mainly driven by water temperature. In autumn, at a temperature falling below ca. 10°C, T. trachurus retreat from the feeding areas in the southern Norwegian and the North Sea and migrate to the over-wintering areas further south. These are situated in the English Channel (Lockwood & Johnson, 1977, Macer, 1974 and 1977) and along the continental slope (Macer, 1977) in the Bay of Biscay and Celtic Sea (Eaton, 1983, Figure 2.3). In winter they form dense schools in deeper water. In spring the fish become far more dispersed (Polonsky, 1965) and migrate northward again with increasing water temperature (e.g. Chuksin & Nazarov, 1989). The North Sea component appears in April in the southern North Sea and reaches the western Jutland coast and southern Norway by August. Parts of the Western stock may reach Trondheim Fjord in July-August (ICES, 1998). Other parts of this component feed in areas west of Ireland or at the Bay of Biscay continental slopes. The Southern stock shows a large overlap between spawning and feeding areas. In fact, in the Cantabrian Sea and Galician waters, the horse mackerel population appears more stable and stationary than typically migratory although there are also variations of small magnitude (Villamor et al., 1997).

Apparently, the water temperature of 8°C is the lower limit for horse mackerel which it avoids during over-wintering (Polonsky, 1965). Laboratory investigations have shown that they stop feeding at water temperatures below about 10°C (Hermann, pers. comm.). Lozano Cabo (1952) gives optimal water temperatures of 19-23 °C, higher temperatures seem to be avoided.

Fig. 2: Schematic outline of assumed migration routes, spawning and feeding areas for the three Horse Mackerel stocks. Depth line drawn is the 200 m contour. For over-wintering areas see Fig. 3. From ICES 1992, redrawn.	Fig. 3: Schematic outline of over-wintering areas and assumed migration routes, Depth line drawn is the 200 m contour. For feeding and spawning areas, see Fig. 2. From Eaton 1983, redrawn.

Horse mackerel is a fairly long lived species. The cohort analysis of ICES (1998) shows that the 1982 year class has been extraordinarily strong and can be identified for 15 years in the annual age compositions (Figure 4 below). Ages of nearly 40 have been reported. However, for assessment purposes the higher ages are not differentiated and summarised as 15+ ages.

Borges (1991) described the growth of horse mackerel by length frequency analysis of commercial catch data, disaggregated on a monthly scale. She described two cohorts recruiting to the fishery each year. Both cohorts grew into each other in the course of the first two years and were later on not distinguishable any more. Depending on the method employed (Battacharya or Shepherd) L inf was in the region of 40 to 50 cm TL with K = 0.29 and 0.14, respectively. Considerably larger horse mackerel were however observed in the field (e.g. 59 cm TL in Portuguese waters, Murta et al., 1993).

The species T. picturatus seems to show a similar growth. Isidro (1990) found an L inf of 52.9 cm (fork length) for individuals from the Azores. For T. mediterraneus, Lucio (1996) provides a similar L inf (52.01 cm TL).

The shape (outline) of the otoliths changes considerably with increasing fish age (click here to see animation). Based on this fact, there are approaches to use otolith shape analysis as an ageing tool (see http://www.ImageScience.de/fabosa/welcome.htm).

Tab. 1: Length weight relationships (in cm and g)

Several authors have given v. Bertalanffy growth parameters for T. trachurus (Lucio, 1990; Kerstan, 1985; Fariña Pérez, 1983; Lourdes Marecos et al., 1978; Trouvery, 1977; Nazarov, 1978). As these are highly dependent on ageing method and accuracy, which are currently regarded as uncertain, they are not displayed here.

Fig. 4: Age composition of Western Horse Mackerel in international catches 1983-1999. from ICES (2001).

For the North Sea stock, after spawning in summertime the fat and energy content of horse mackerel is lowest (Sahrhage, 1970, Hermann pers. comm.). In August and September the horse mackerel energy content rises rapidly, apparently as a result of extensive feeding. As mentioned earlier, feeding ceases as soon as the water temperature drops below 10°C (Hermann, pers. comm.). At 8-9°C the fish stop feeding totally and leave the area for over-wintering. In spring, before spawning, the horse mackerel have only little fat reserves in the gut (Leloup and Gilis, 1964). In accordance to this, Polonsky (1965) found that the muscle fat content is lowest in March and highest in September.

According to these findings, one should expect the development of opaque zones in the otoliths during periods of extensive feeding, which enables fast growth and enhanced calcification. On the other hand, starving during winter, the energy consuming gonad development in winter/spring and the period of spawning will lead to a formation of translucent zone.

Several investigations indicate that T. trachurus is a filter feeder, swimming at low but constant speed (see physiology section), mainly ingesting zooplankton (e.g. Ben Salem, 1988). In the Eastern part of the North Sea (off Jutland) horse mackerel were found to forage predominantly on fish (Dahl and Kirkegaard, 1987), with 0-group whiting being the most important prey item, followed by other gadoids and herring. More to the south invertebrates constituted the bulk of ingested food items. Of these, surprisingly few Crangon crangon were taken and by far more decapods and other undetermined crustaceans. No clear shift of prey fraction with size could be shown. However, a clear diurnal feeding pattern was observed, with highest food intake during midday.

Dahl and Kirkegaard (1986) also found a clear diurnal feeding rhythm in the eastern part of the North Sea, but with highest food intake during early morning and lowest ingestion rates during the night. In this work, a shift in prey preference with age was proven: Smaller individuals (<20-24 cm) preyed mostly on crustaceans, gobies and haddock, while larger specimens shifted towards herring. Smaller fish did not forage on herring at all.

In the English Channel apparently adult horse mackerel were found to forage to nearly 70% on crustaceans and only to 17% on fish, with monthly varying proportions (Macer, 1977).

For the Bay of Biscay, Letaconnoux (1951) provided a description of the horse mackerel diet, noting possible seasonal differences. For specimens from the northwest of Spain, Lozano Cabo (1952) suggested that young specimens are planktophagous while adults are mainly ichthyophagous. These findings were supported by a recent work of Olaso et al. (in press) on the diet composition in the southern Bay of Biscay. They found seasonal differences: preying on crustaceans dominated during spring, while in autumn T. trachurus > 30 cm began to prey on fishes (blue whiting, gobiids, anchovy), which represented 45% of the food volume in this size-range. They also described a diurnal feeding mode, with feeding maximum around noon in spring (for fish > 30 cm), and at sunrise in autumn.

In Portuguese waters (ICES Division IXa) horse mackerel fed mainly on zooplankton, especially euphausids and copepods. Only at greater sizes (>19 cm TL) they also fed on fish and cephalopods (Murta et al., 1993).

The sex ratio at least in the southern stock is 1:1 if surveyed over a wider area. Deviations from this relationship may occur if investigations are spatially and temporally not wide enough (Abaunza et al., 1995). For the southern stock the length at first maturity is about 21 cm for males and approximately 22 cm for females (Borges & Gordo, 1991, Abaunza et al., 1995). Horse mackerel is a batch spawner (e.g. Borges et al., 1993). A potential fecundity of 1557 eggs per gramm female horse mackerel was determined for the conversion of egg production into biomass of western horse mackerel (Eltink and Vingerhoed, 1993; ICES., 1996).

The maturity cycle of horse mackerel of the southern stock (off the Portuguese coast) begins in December, attains a maximum in February with highest gonadosomatic index between February and April, and lowest values in July to October (Arruda, 1983).

Surveys on the egg production of horse mackerel were carried out repeatedly in the waters off Portugal up to the areas north of Shetlands for spawning stock biomass estimation, including the North Sea (ICES, 1996; Iversen et al., 1989; Eltink, 1992). The spatial and temporal distribution of the spawning process is given e.g. by Lockwood and Johnson (1977; see also Figure 2). However, more recent egg surveys have shown that the size of the spawning area also extends to the west of Ireland and Scotland later in summer (ICES, 1997).

Recruitment in the southern area is monitored by means of young fish surveys (e.g. Borges, 1983, 1984, 1986, Sánchez et al., 1991). As discussed earlier, the analysis of the recruitment of the western component showed that the 1982-year class has been extraordinarily strong. It is remarkable that this year class was apparently not able to produce any larger recruitment. This accounts for the hypothesis that the stock recruitment for this stock is independent of parental stock size. Recruitment success could be primarily determined by hydrographic factors or feeding conditions of larvae and young fish. By contrast, it can also be argued that the low recruitment observed throughout the recent 15 years is the normal level of reproduction and the 1982 year class represents a freak reproduction success, which should not be the measure of successful recruitment.

Horse mackerel is a batch spawner, which implies that horse mackerel spawn their eggs in several batches during the spawning season. There are two types of batch spawners: determinate and indeterminate spawners. For determinate spawners the fecundity is determined prior to spawning, which implies that in an individual fish the development of vitellogenic oocytes stops prior to spawning. In such case after starting a continuous increase in fecundity there might be a short period of a constant fecundity prior to the onset of spawning. This would be the right period for fecundity estimation and furthermore it would provide an indication that this species is a determinate spawner. For indeterminate spawners the fecundity is not determined prior to spawning, because in an individual fish the development of vitellogenic oocytes even continues after the onset of spawning in which case the potential fecundity can not be estimated. Fecundity estimations both prior to spawning and during spawning would underestimate the fecundity. If fecundity is estimated prior to spawning the fecundity will be underestimated because the eggs from de novo vitellogenesis are not taken into account. If fecundity is estimated at a time that no more vitellogenic oocytes develop then fecundity will be underestimated because of the loss of eggs by spawning.

Up to now horse mackerel has been assumed to be a determinate spawner.

In 1998 the horse mackerel fecundity was estimated much lower compared to earlier years (ICES, 1999/G:5). This was expected be due to exceptional early spawning in 1998 and it was assumed that spawning fish had been used for the fecundity estimation. An important fact is that horse mackerel spawning can not easily be recognised in histological slides of the ovaries as having spawned in the current season. This is caused by the long time interval between two batches of spawning. It is that long that the post-ovulatory-follicles (POF’s) can have disappeared before other stages of spawning activity (migrating nucleus stage, hyaline oocyte stage) appear. Therefore, fecundity sampling should be carried out before any spawning takes place, because as soon as spawning starts individual fish can not be identified any more as not having spawned yet. If fish have spawned, fecundity will be underestimated, which then will cause spawning stock biomass to be overestimated.

In 2000 a small scale test sampling for fecundity was carried out as a test case for the sampling in 2001, which is the year in which the extensive international egg surveys will be carried out. The aim was to sample 25 horse mackerel for fecundity every two weeks from January to April 2000 to allow an investigation of the changes in fecundity over time until the start of spawning season and to determine the appropriate time for fecundity sampling.
The sampling for fecundity over the period January to April 2000 shows that the fecundity increases continuously over the whole period of sampling, but also after spawning started in March. Ovaries, which showed signs of spawning, had still a low fecundity. This is an indication that horse mackerel might be an indeterminate spawner.

The aim of this small-scale fecundity sampling in 2000 was to estimate the most appropriate time for the estimation of the maximum level fecundity before the onset of spawning, but this appears to be impossible with this early spawning of horse mackerel.

The oocyte development rate was estimated to be approximately 10 vitellogenic oocytes/g female/day. The historic estimate of the potential fecundity is 1557 eggs/gramme female, which has been used for the biomass calculation from all egg surveys up to 1998 (ICES, 2000/ACFM:5). If a development rate of 10 vitellogenic oocytes per gramme female per day is applied to this fecundity, it would require just over 5 months of development (5.2 * 30 * 10 = 1560). This would imply that the development of vitellogenic oocytes would stop around the middle of May assuming that the onset of vitelllogenic oocytes development starts in the middle of December. It should be taken into account that the production rate of vitellogenic oocytes might increase with increasing temperatures. Based on this development rate the historic estimate of 1557 eggs per gramme female does not seem to be a serious underestimate of the potential fecundity. However, in 2001 a lot more effort has to be put in to validate this historic fecundity estimate.

For the egg survey in 2001 fecundity information should be collected in such way that an extrapolated potential fecundity possibly can be calculated. This might be obtained from information on the production rate of vitellogenic oocytes and the duration of the period of vitellogenic oocytes development (oocyte diameter frequency distributions might help in determining at what time there is evidence that vitellogenesis stops). Recommendations concerning the fecundity sampling in 2001 are given in Eltink (WD 2000). A last possibility to discuss the fecundity problems and sampling in 2001 will be in December 2000 at a meeting on egg stageing and fecundity / atresia at CEFAS, Lowestoft, UK.

Horse mackerel is a predatory species, which is believed to consume considerable amounts of herring larvae and juveniles in the North Sea. It is regarded as one of the important predatory species in the North Sea ecosystem and the annual consumption is modeled by the Multispecies Virtual Population Analysis (MSVPA, Helgason & Gislason, 1979, Pope, 1979). The consumption is an input parameter for the MSVPA to determine natural mortality. To measure the consumption rate of horse mackerel, experiments are at present carried out at the University of Hamburg in the frame of an EU-project (Consumption rates of predatory fish relevant for multispecies assessment in the North Sea and the Atlantic off Spain and Portugal, CORMA). Two different methods are applied for the measurements and modeling of the consumption of horse mackerel, firstly by measuring the rate of gastric evacuation, and secondly by modeling bioenergetics. For the latter some physiological parameters are measured, such as the standard oxygen consumption for a weight range of 1.4 – 390 g, yielding a relationship of VO2 = 0.228 * WM^0.725 (at 13°C) (Enders, 1998). Further analyses confirmed that the horse mackerel is a good swimmer. Only moderate increases of oxygen consumptions were recorded at lower swimming speeds. This accounts for the hypothesis that for horse mackerel routine swimming is energetically not very costly, and that this species is adapted to swimming at a low but very constant speed.

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This text is mainly based on a "synopsis of the biology of horse mackerel in European waters", prepared by C. Hammer and C. Zimmermann, Hamburg, for the ICES/EU Workshop on Horse Mackerel Otolith Reading (WKHMO), held in Lowestoft, Jan. 1999. Information presented here might therefore not reflect latest scientific knowledge.

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