21 Smooth-hounds in the Northeast Atlanticices.dk/sites/pub/Publication Reports/Expert Group...

544 | ICES WGEF REPORT 2016

21 Smooth-hounds in the Northeast Atlantic

21.1 Stock distribution

Three species of smooth-hound (Triakidae) occur in the ICES area. The most frequent

species in the northern part of the area is starry smooth-hound Mustelus asterias. Com-

mon smooth-hound Mustelus mustelus may also occur in northern European seas, alt-

hough no confirmed specimens have been found in recent years and historical records

may be unreliable. Separating these two species on the presence or absence of spots is

unreliable (Compagno et al., 2005; Farrell et al., 2009), and information and data from

northern Europe referring to M. mustelus likely refers to M. asterias.

A third species, black-spotted smooth-hound Mustelus punctulatus, occurs in the Med-

iterranean Sea (Quignard, 1972) and off NW Africa and may occur in the southernmost

part of ICES Division 9.a.

M. asterias is the dominant smooth-hound in northern European waters. The develop-

ment of a molecular genetic identification technique has allowed the reliable identifi-

cation and discrimination of Northeast Atlantic Mustelus species (Farrell et al., 2009).

Subsequent studies involving the collection of 231 Mustelus from the Irish Sea, Bristol

Channel, Celtic Sea and west of Ireland, identified all to be M. asterias (Farrell et al.,

2010a, b). A further study from the North Sea and English Channel (McCully and Ellis,

2015 WD) that sampled 504 Mustelus, also found no specimens of M. mustelus.

Given the problems in separating M. asterias and M. mustelus and that data for these

two species are confounded, data in this chapter are generally combined at genus level.

Whilst assessments conducted by WGEF are based on Mustelus asterias, management

advice should be applied at the genus level, so as to avoid potential identification prob-

lems associated with management and enforcement.

In the absence of dedicated scientific studies on stock units, WGEF considers there to

be a single management unit of Mustelus asterias in the continental shelf waters of the

ICES area, comprising ICES Subareas 4, 6–9. This stock may extend to the northern part

of the CECAF area and possibly the Mediterranean Sea.

Improved studies to better understand the stock unit(s) are required. There are several

programmes that tag and release M. asterias in the North Sea and Celtic Seas ecoregions

(e.g. Burt et al., 2013 WD). In the North Sea between 2011–2014, Sportvisserij Nederland

in conjunction with the fisheries research institute IMARES, Wageningen UR, initiated

a tagging programme with anglers in the Dutch Delta, whereby 2244 M. asterias were

tagged, with 80 recaptures reported (Brevé et al., 2016). Recapture positions showed a

circannual migration, with fish spending the summer in the southern North Sea and

overwintering in the English Channel and Bay of Biscay, suggesting a degree of philo-

patry (Brevé et al., 2016). Cooperative large-scale analyses of all available tagging data

are required. Additionally, tagging studies from the more southern parts of the distri-

bution range could be usefully undertaken.

21.2 The fishery

21.2.1 History of the fishery

Smooth-hounds are taken as a seasonal bycatch in trawl, gillnet and longline fisheries.

Though they are discarded in some fisheries, other fisheries land this bycatch, depend-

ing on market demands. Some may also be landed to supply bait for pot fisheries.

ICES WGEF REPORT 2016 | 545

Smooth-hounds are also a relatively important species for recreational sea anglers and

charter boat fishing in several areas, with anglers and angling clubs often having catch-

and-release protocols, particularly in the Celtic and North Seas.

21.2.2 The fishery in 2015

There were no major changes to the fishery noted in 2015. Information from the fishing

industry suggests that the increased landings of smooth-hounds, since 2009, are partly

to supply market demand for ‘dogfish’ given the current restrictions on spurdog.

21.2.3 ICES Advice applicable

ICES first provided advice for this stock in 2012 for 2013 and 2014 (which was reiterated

for 2015), stating that “Based on ICES approach to data-limited stocks, ICES advises that

catches should be reduced by 4%. Because the data for catches of smooth-hounds are not fully

documented and considered highly unreliable (due to the historical use of generic landings cat-

egories), ICES is not in a position to quantify the result”.

In 2015 ICES advised that “when the precautionary approach is applied, landings should be

no more than 3272 tonnes in each of the years 2016 and 2017”. This was based on a survey-

based (category 3.20) assessment, with the stock size indicator based on four surveys

indices.

21.2.4 Management applicable

There are no specific management measures for smooth-hounds.

EC Council Regulations 850/98 for the `conservation of fishery resources through tech-

nical measures for the protection of juveniles of marine organisms’ details the mini-

mum mesh sizes that can be used to target fish. Although other dogfish (Squalus

acanthias and Scyliorhinus spp.) could be targeted in fixed nets of 120–219 mm and

>220 mm mesh size (in regions 1 and 2), Mustelus spp. would be classed under ‘all other

marine organisms’, and so can only be targeted in fixed nets of >220 mm. This has been

queried by some fishermen.

21.3 Catch data

21.3.1 Landings

No accurate estimates of catch are available, as many nations that land smooth-hounds

report an unknown proportion of landings in aggregated landings categories (e.g. dog-

fish and hounds). Official reports of landings (Table 21.1; Figure 21.1) are increasing,

with landings exceeding 3000 t since 2012. However new ICES estimates indicate that

landings have been over 3000 t since 2005. The main nations exploiting smooth-hounds

are France (> 88% of landings in 2014) and the UK, with the English Channel and south-

ern North Sea are important fishing grounds.

21.3.2 Discards

Although some discards data are available from various nations, data are limited for

most nations and fisheries. Four countries reported preliminary estimates of discards,

which ranged from 28 to 950 t in 2014. Given the seasonality of catches in some areas,

and that M. asterias is often taken by inshore vessels where observer data can be more

sporadic, further studies to evaluate the most appropriate methods of raising data from

observer trips to fleet level are required if catches are to be estimated appropriately.


Earlier studies have indicated that juveniles are typically discarded (Figure 21.2), alt-

hough the survival of these discards has not yet been evaluated (Silva et al., 2013 WD).

Smooth-hounds taken by beam trawl and Nephrops trawl were composed primarily of

juveniles and subadults (<70 cm LT), and most these were nearly all discarded. Gillnet

catches were comprised primarily of fish 60–110 cm LT, with fish <55 cm LT usually

discarded. Otter trawl catches covered a broad length range, and smooth-hounds <50

cm LT were usually discarded. The absence of full retention at length in these gears

may be due to various factors (e.g. catch quality and local market value) influencing

the discarding behaviour of fishers.

Silva et al. (2013 WD) also noted that a greater proportion of smooth-hounds were re-

tained since landing opportunities for spurdog had become restrictive. Over the time

period 2002–2005, the retention of Mustelus spp. ≥70 cm LT was 1% and 39% in gillnet

and otter trawl fisheries, respectively. In the period 2006–2011, however, retention in-

creased to 73% (gillnets) and 49% (otter trawl).

21.3.3 Quality of catch data

Landings data have historically been of poor quality, as much of the landings data have

been reported under generic landings categories. Most nations have made efforts to

improve the recording of species in recent years. Some northern European nations re-

port more M. mustelus than M. asterias in official statistics, but WGEF combine these

data.

Smooth-hounds are often taken in inshore fisheries, and landings data for vessels <10

m may not be complete.

The availability of landings data from outside the ICES area (e.g. Mediterranean Sea)

is limited, and the quality uncertain. In 2010, the European Commission collated land-

ings data as an average across 2008–2010 and three species of Mustelus were repre-

sented in these data; M. punctulatus (269 t from Italy), M. mustelus (14 t combined from

Italy, Spain, Malta and Slovenia) and M. asterias (1 t from Malta) (ICES, 2012). Catch

data are absent from off NW Africa.

Better estimates of discarding are required, with information on discard survival also

needed, as a proportion of discarded smooth-hounds may survive.

21.3.4 Discard survival

Survival appears to be quite variable across this family (Ellis et al., 2014 WD). Whilst

quantitative data are limited in European waters, Fennessy (1994) reported at-vessel

mortality of 29% for Arabian smooth-hound Mustelus mosis taken in a prawn trawl

fishery. Mortality ranged from 57–93% for three triakid sharks taken in an Australian

gillnet fishery, despite the soak times being <24 hours (Braccini et al., 2012). High sur-

vival of triakids has been reported in longline fisheries (Frick et al., 2010a; Coelho et al.,

2012).

21.4 Commercial catch composition

Studies to better understand the composition by size and sex (and species where there

is spatial overlap) are required. Given the potential for sexual and sex-based segrega-

tion of smooth-hounds, appropriate levels of monitoring would be required to fully

understand catch composition over appropriate spatial and temporal scales.


21.4.1 Length Composition of landings

In a UK study, 504 starry smooth-hound samples (266 female, 238 male, Figure 21.3)

were examined (McCully Phillips and Ellis, 2015), of which 286 (with a length range of

52–124 cm LT) were landed by commercial vessels.

21.4.2 Length composition of discards

Silva et al. (2013 WD) analysed the discard and retention patterns of Mustelus spp. taken

as bycatch in UK commercial fisheries. Beam trawlers caught proportionally more ju-

veniles (most records were for fish of about 35–70 cm LT), consequently, discarding was

quite high (95–99%). High rates of discarding (of smaller fish, <65 cm LT) were also

apparent in otter trawls, where about 75–80% of the total catches were discarded in the

Celtic Seas and North Sea, respectively. Gillnets were more selective for larger fish

(with the majority of fish 60–100 cm LT), where typically only the larger fish (>70 cm

LT) were retained.

21.4.3 Sex ratio of landings

Of 286 commercially landed samples from the southern North Sea and eastern English

Channel in May–November, 155 were female and 131 were male (McCully Phillips un-

published). Due to smooth-hounds aggregating by sex and size, the sex ratio (and

length–frequency) may vary over the year and area.

21.4.4 Quality of data

Mustelus length measurements may be collected as part of the concurrent sampling of

the DCF. These data should be made available for future analysis.

21.5 Commercial catch and effort data

There are no data available.

21.6 Fishery-independent information

21.6.1 Availability of survey data

Several fishery-independent surveys operate in the stock area. Analyses of survey data

need to be undertaken with care, as smooth-hounds are relatively large-bodied species

(maximum size of M. asterias is about 140 cm LT) and adults are strong swimmers.

Hence, larger individuals may not be sampled effectively in IBTS surveys. Given their

aggregating nature, some surveys may have a large number of zero hauls and a few

hauls with relatively large numbers.

They are often caught in GOV trawl and other otter trawl surveys in the area. For fur-

ther details of trawl surveys in the stock area, see Section 15 (North Sea ecoregion),

Section 18 (Celtic Seas) and Section 19 (Biscay-Iberia). Summary details from IBTS 2011

are shown in Figure 21.4.

Larger individuals are not sampled effectively in beam-trawl surveys (because of low

gear selectivity). For example, the UK western English Channel beam-trawl survey

only occasionally records fish >100 cm LT (Silva et al., 2014 WD; Figure 21.5).

Although two species of smooth-hound have previously been reported in most sur-

veys, the discrimination of these species was usually been based on the presence or

absence of spots, which is not a reliable characteristic. WGEF consider that survey data


for these two species should be combined in any analyses, and that starry smooth-

hound Mustelus asterias is likely to be the only species in the Celtic Seas and North Sea

ecoregions.

21.6.2 Survey trends

Trends in most of the fisheries-independent surveys have been increasing in recent

years. Over the longer term time-series of each survey, all show an increasing cpue,

which has been substantial in recent years. Of the six surveys providing an index, four

reported a mean annual cpue for 2013–2014 ranging from 40–56% above that seen in

the preceding five years (2008–2012).

The UK (England and Wales) beam-trawl survey of the Irish Sea (Division 7.f) and

Bristol Channel (Division 7.f) catches reasonable numbers of smooth-hounds. The

trend in abundance is derived from the catch rates from fixed stations (97 stations

fished at least 19 years out of the 22 year time-series Figure 21.6; Ellis 2015 WD), and is

currently at its highest level (since 1993) of 3.23 ind.h–1 (2013–2014). This was 40.5%

above the mean average cpue for the preceding five years (2008–2012; 2.30 ind.h–1).

The UK (England and Wales) beam-trawl survey of the southern North Sea (Division

4.c) and eastern English Channel (Division 7.d) catches lower numbers. The trend in

abundance of smooth-hounds (derived from the catch rates from 76 fixed stations

fished at least 18 years out of the 22 year time-series; Figure 21.6) has increased over

the time-series, and they were also being observed in an increasing proportion of hauls

until 2011 (ICES, 2011). Mean annual cpue for 2013–2014 (0.85 ind.h–1) was just below

the mean average cpue for the preceding five years (2008–2012; 0.88 ind.–1). This slight

(ca. 3% less) decrease should be viewed in the context of the longer term increase in

cpue during the overall survey series, and the mean cpue observed in the last two years

were both above the long-term mean.

The IBTS surveys of the North Sea, undertaken in Q1 and Q3 by seven and six countries

respectively, catch relatively low numbers (which may relate to smooth-hounds being

more abundant in only the southern parts of the area sampled by the IBTS). Neverthe-

less, the long-term trend in abundance of smooth-hounds has increased over both of

the 24- (Q3) and 25-year (Q1) time-series. In the NSIBTS-Q1, the mean annual cpue for

2013–2014 (1.41 ind.h–1) was far greater than the mean average cpue for the preceding

five years (2008–2012; 0.92 ind.h–1); this is a 52.7% increase, and the cpue is currently at

its highest level of 1.97 ind.h–1 (2014). A similar trend is also seen in the NSIBTS-Q3,

with the mean annual cpue for 2013–2014 (1.32 ind.h–1) far greater than the mean aver-

age cpue for the preceding five years (2008–2012; 0.88 ind.h–1); this is a 51.1% increase,

and the cpue is currently at its highest level (1.38 ind.h–1 in 2014).

The increasing long-term trend is mirrored (albeit with lower catch rates) in the Irish

Groundfish Survey with the mean annual cpue for 2013–2014 (0.08 ind.h–1) exceeding

the mean average cpue for the preceding five years (2008–2012; 0.05 ind.h–1). Once

again, cpue is currently at its highest level (0.11 ind.h–1 in 2014).

The EVHOE survey of ICES Divisions 7.g-k and 8.a.b.d has an 18-year time-series of

data, however for three years of this survey, data were either available, or with zero

catch; the lack of these records needs verification. Unfortunately, two of these years

were in 2009 and 2010, thus when calculating the mean average cpue for the preceding

five years, this was based on just three years’ data, which consisted of high catch rates,

based on a few large catches. The actual proportion of hauls where smooth-hounds

were caught is at its highest level (ca. 16%; similar to that seen in 2012) of the time-


series. The catch rates fluctuated highly across this survey series, from zero to more

than 25 ind.h–1.

The UK (Northern Ireland) western IBTS Q4 survey of the Irish Sea also indicated an

increase in mean catch rates in previous year’s analyses, but recent data were not avail-

able to WGEF.

A further UK (England and Wales) beam-trawl survey of the western English Channel

also encounters smooth-hounds in good numbers. Across the survey time-series (2006–

2014), a total of 658 have been caught, accounting for 7.6% of the elasmobranch catch

by numbers; the observed length range was 31–115 cm LT (Silva et al., 2014 WD). Stand-

ardised indices of relative abundance have not yet been developed for this survey.

Although smooth-hounds are not subject to routine biological sampling in any of the

surveys, all UK (England and Wales) surveys tag and release starry smooth-hounds,

and the individual weights and sex (all fish) and maturity (male fish only) are recorded

prior to release (See Section 21.7.5).

21.7 Life-history information

Biological data are not collected under the Data Collection Regulations, however some

ad hoc data are collected on fishery-independent surveys and there are some published

studies resulting from biological investigations of Mustelus spp. in European seas, in-

cluding from the NE Atlantic and Mediterranean Sea.

21.7.1 Habitat

The distribution of Mustelus spp. around the British Isles has been described, with more

detailed studies on the habitat utilization only examined in the English Channel (Mar-

tin et al., 2010; 2012).

21.7.2 Spawning, parturition and nursery grounds

Pups of M. mustelus are born at a size of 34–42 cm in the Mediterranean (Saidi et al.,

2008) and 36 to 45 cm off Senegal (Capapé et al., 2006). Pups are taken in trawl surveys,

and such data might be able to assist in the preliminary identification of general pup-

ping and/or nursery areas. Most of the records for M. asterias pups recorded in UK

beam-trawl surveys are from the southern North Sea, English Channel and Bristol

Channel (Ellis et al., 2005).

Recent biological studies have indicated that full-term pups range in size from 205–329

mm LT and pup size was positively correlated with maternal length (McCully Phillips

and Ellis, 2015; Figure 21.7). The smallest free-swimming neonate reported in this

study was 24 cm LT.

Parturition of M. asterias occurred in February in the western English Channel and

June–July in the eastern English Channel and southern North Sea (Figure 21.8), indi-

cating either protracted spawning or asynchronous parturition for the stock as a whole

(McCully Phillips and Ellis, 2015).

Studies on other species of smooth-hound have shown high site fidelity of immature

individuals on nursery grounds (Espinoza et al., 2011).

21.7.3 Age and growth

Farrell et al. (2010a) studied the age and growth of M. asterias in the Celtic Seas ecore-

gion. Growth parameters for males (n = 106) were L∞ = 103.7 cm LT, L0 = 38.1 cm, k =


0.195 year–1). Growth parameters for females (n = 114) were (L∞= 123.5 cm LT, L0= 34.9

cm, k = 0.146 year–1). Estimates of longevity were 13 years (males) and 18.3 years (fe-

males).

Age and growth of M. mustelus has been studied in South African waters, with males

and females estimated to mature at 6–9 and 12–15 years, respectively (Goosen and

Smale, 1997). The maximum age reported in this study was 24 years.

The length–weight relationship of Mustelus spp. caught during the Cefas tagging pro-

gramme, 2000–2010 is illustrated in Figure 21.9.

21.7.4 Reproductive biology

Studies in the Celtic Seas ecoregion had indicated that the total length (and age) at 50%

maturity for male and female M. asterias are 78 cm LT (4–5 years) and 87 cm LT (six

years), respectively (Farrell et al., 2010b). Studies of M. asterias primarily from the

southern North Sea and English Channel estimated 50% maturity for male and females

at ca. 70 cm LT and 82 cm LT respectively (McCully Phillips and Ellis, 2015; Figure

21.10).

Estimates of fecundity range from 8–27 (ovarian fecundity) and 6–18 (embryonic fe-

cundity), with a gestation period of about twelve months (Farrell et al., 2010b), and

there may also be a resting period of a year between pregnancies, giving a two year

reproductive period. However, within mature female fish sampled by McCully Phil-

lips and Ellis (2015), seventeen late gravid females with term pups (uterine fecundity

4–20) were also found to have numerous mature follicles (n = 6–22; follicle diameters

6–10 mm). This indicates that the reproductive cycle is likely to extend beyond one

year, and coupled with the extended parturition periods (Section 21.7.2) either pro-

tracted spawning or asynchronous parturition for the stock as a whole also occurs. Fur-

ther studies are required to confirm or reject this hypothesis, including more samples

of fish from winter and spring.

The smallest mature female that Farrell et al. (2010a) reported was 83 cm; a lot larger

than the smallest female (69 cm LT; summarised below) recorded by McCully Phillips

and Ellis (2015). This is interesting, as the two studies use slightly different maturity

keys, with Farrell et al. (2010a) assigning a female to be mature when oocytes were

present, yellow, and countable at >3 mm in diameter, whereas Cefas maturity keys

(Table II of McCully Phillips and Ellis, 2015), which are comparable to those keys de-

veloped within ICES, assign a female as mature when the oocytes are slightly larger

(>5 mm).

McCully Phillips and Ellis (2015) estimates the length at 50% maturity to be 81.9 cm for

females (smallest mature = 69 cm; largest immature = 87 cm) and 70.4 cm for males

(smallest mature = 65 cm; largest immature = 74 cm).

The number of mature follicles ranged from 0–28 in the mature females. These will not

all necessarily develop into embryos, however, and estimates of ovarian fecundity are

known to exceed estimates of uterine fecundity. The size spectra of the mature follicles

(within mature females) ranged from 4.1 mm (mid-term gravid female) to 20.7 mm

(mature female).

The uterine fecundity ranged from 4–20, which exceeds the maximum uterine fecun-

dity (18) found by Farrell et al. (2010a); however they stated that their values may be

underestimated due to females aborting pups on capture. The female identified with a

fecundity of 20, was found with full-term pups. Uterine fecundity increased with total


length (Figure 21.11). Furthermore, there were also positive linear relationships iden-

tified between maternal length and average pup length and weight (Figure 21.7).

In the Mediterranean Sea, Mustelus asterias reach maturity at about 75 cm (males) and

96 cm (females), with estimates of fecundity ranging from 10–45 (ovarian fecundity)

and 10–35 (uterine fecundity), with fecundity increasing with length (Capapé, 1983),

although it is possible the higher fecundity in this study may relate to data being con-

founded with other species of smooth-hound.

Studies on Mustelus mustelus in the Mediterranean have found that females matured at

107.5–123 cm LT (50% maturity at 117.2 cm) and that males matured at 88–112 cm LT

(50% maturity at 97.1 cm) (Saidi et al., 2008). This study also found that embryonic fe-

cundity ranged from 4–18 embryos, with fecundity increasing with length. Further

south, off Senegal, the lengths at first (and 100%) maturity for M. mustelus were found

to be 82 cm (95 cm), for males, and 95 cm (104 cm) for females (Capapé et al., 2006). This

study reported litters of 4–21.

21.7.5 Movements and migrations

Although the movements and migrations of smooth-hounds are not fully known, there

have been relatively high numbers of Mustelus spp. tagged and released during various

elasmobranch research programmes (e.g. Burt et al., 2013 WD; Figure 21.12). A recent

(2011–2014) study by Sportvisserij Nederland, in conjunction with the fisheries re-

search institute IMARES, Wageningen UR, involved a tagging programme with an-

glers in the Dutch Delta, whereby 2244 M. asterias were tagged, with 80 recaptures

reported (Figure 21.13; Brevé et al., 2016). Recapture positions showed a circannual mi-

gration between summertime grounds in the southern North Sea and overwintering in

the English Channel and Bay of Biscay, suggesting philopatry (Brevé et al., 2016).

21.7.6 Diet and role in ecosystem

Mustelus spp. are primarily carcinophagous, predating on various crustaceans, includ-

ing hermit crabs (Paguridae), stomatopods, brachyuran crabs, squat lobsters and

shrimps, with teleosts occasionally eaten by larger individuals (Ellis et al., 1996; Morte

et al., 1997; Jardas et al., 2007; Santic et al., 2007; Saidi et al., 2009; Lipej et al., 2011;

McCully and Ellis, 2014). They can be important predators of commercial crustaceans,

feeding on velvet swimming crab Necora puber and small edible crab Cancer pagurus.

21.7.7 Conversion factors

The relationship between total length and weight in the smooth-hounds sampled by

sex and maturity stage are summarised below and in Figures 21.14 and 21.15 (McCully

Phillips and Ellis, 2015). The relationship for males differs slightly to that of females,

largely driven by the larger maximum length of females and the weights of females

about to give birth. Of note is the 119 cm outlier, which related to a post-partum female

with a very low body mass. Samples of the smaller size classes were obtained from

scientific trawl surveys, while the larger individuals were from commercially landed

specimens. Smooth-hounds are traditionally landed for the market gutted, and so con-

version factors to length is also a useful parameter to augment data collected during

market sampling programmes.


Relationship

Y=axb Sex/Stage a b r2 n

Total weight to Total

length

All Females 0.0014 3.2 0.992 248

All Males 0.0020 3.1 0.995 237

Immature Female

(stage A/B)

0.0020 3.1245 0.994 170

Immature Male

(stage A/B)

0.0014 3.2159 0.991 113

Mature Female (incliding

early gravid)

(stage C/D)

0.0021 3.1396 0.913 54

Mature Male

(stage C/D)

0.0077 2.8084 0.938 123

Mid-/late-term gravid

females

(stage E/F)

0.0002 3.7072 0.935 21

Gutted weight to

Total Length

Sexes combined 0.0014 3.1580 0.995 484

Female 0.0016 3.1 0.994 249

Male 0.0014 3.2 0.996 235

21.8 Exploratory assessment models

21.8.1 Previous studies

No previous assessments of NE Atlantic smooth-hounds have been made. However,

there have been assessment methods developed for the Australian species Mustelus

antarcticus (e.g. Xiao and Walker, 2000; Pribac et al., 2005) which may be applied to

European species when relevant data are available.

21.8.2 Data exploration and preliminary assessments

Although no modelling or quantitative stock assessments have been undertaken,

trends in relative abundance have been used to inform on current status (see Section

21.6).

21.9 Stock assessment

No quantitative stock assessment has been undertaken. The stock is evaluated on the

basis of abundance trends in fishery-independent trawl surveys, as these are the long-

est time-series of standardised, species-specific data available for the stock (Figure

21.16).

The abundance trends of the long-term time-series of six different surveys covering a

large proportion of this species’ distribution range consistently show an increase,

which has been substantial in recent years.

Of the six surveys providing an individual survey index, four reported a mean annual

cpue for 2013–2014 ranging from 40–56% above that seen in the preceding five years

(2008–2012). These indices were standardised, and a mean index calculated using four

different surveys, providing a 22-year standardised time-series. This gave a mean an-

nual cpue for 2013–2014 of 2.3 ind.h–1, which was a 33% increase from the preceding

five years (2008–2012).


The Irish Groundfish Survey was excluded from the mean standardised survey index,

as it did not begin until 2003, and thus by excluding this, and utilising the other five

surveys, enabled a 22-year index to be calculated, thus informing on the interpretation

of longer-term trends in relative abundance. However, this survey provides important

supporting information and shows a similar increase in abundance for the north-west-

ern part of the stock area.

The EVHOE survey was also excluded from the mean standardised survey index, as

again, it would reduce the time-series down to 18-years and thus hamper interpreta-

tion of longer term trends in abundance. Furthermore, three years of this survey had

either missing or zero catch data, which need verification. However, this survey pro-

vides important supporting information, especially for the southern area (ICES Divi-

sions 8.a.b.d) of the stock distribution (Figure 21.17). Although catch rates in 2013 and

2014 were at a low level for the time-series, the distribution of this species remains

consistent and the proportion of hauls where smooth-hounds were present are at a

high level for the time-series.

21.10 Quality of the assessment

Commercial landings data are available for recent years, but may be compromised by

poor data quality. Whilst fishery-independent trawl surveys provide the best time-se-

ries information, such surveys may under-represent the largest size classes. It is unclear

as to how recent increases in cpue may relate to increased stock abundance and/or a

possible northward shift in distribution.

The positions of survey hauls containing smooth-hounds in the EVHOE survey were

plotted over the 18-year time-series (Figure 21.17). The number of stations catching

smooth-hounds increased over the survey, however, the distribution of the catches has

remained constant, occurring northwards of 46°N. There is no evidence from this sur-

vey to support the theory of a northward shift in the distribution, suggesting it is plau-

sible that the recent increase in catch rates are a result of population growth.

21.11 Reference points

No reference points have been proposed for this stock.

21.12 Conservation considerations

The most recent IUCN Red List Assessment for European marine fishes (Nieto et al.,

2015) upgraded all three Mustelus spp. identifying them as increasing conservational

importance. They are now listed as Near Threatened (M. asterias; previously Least Con-

cern), and Vulnerable (M. mustelus; previously Least Concern and M. punctulatus; pre-

viously Least Concern).

21.13 Management considerations

Smooth-hounds appear to be increasing in relative abundance in trawl surveys, and

also in commercial landings data. Given the potential expansion in fisheries for

smooth-hounds (which may reflect an increased abundance and that fishing opportu-

nities for S. acanthias are limited), further work to understand the dynamics of this stock

is required.

It should be noted that smooth-hounds taken by beam trawl and Nephrops trawl were

composed primarily of juveniles and subadults (<70 cm LT), and these were nearly all

discarded, as were smooth-hounds <50 cm LT in the otter trawl fishery (Figure 21.2).


Discard mortality is not known, and nor is the proportion of recruits that may survive

to maturity and marketable size. Discard survival within this family is variable (Ellis

et al., 2014 WD). Further studies on the mortality and survival rates of juveniles in

these fisheries are needed to evaluate impacts on recruitment.

Smooth-hounds are also an important target species in some areas for recreational fish-

eries; though there are insufficient data to examine the relative economic importance

of these fisheries, or the degree of mortality associated with recreational fisheries.

Other species of smooth-hound are targeted elsewhere in the world, including Aus-

tralia/New Zealand and South America. Although smooth-hounds are generally quite

productive stocks (relative to some other elasmobranchs), evidence from these fisheries

suggests that various management controls can be used for their appropriate manage-

ment.

21.14 References

Braccini, M., Van Rijn, J., and Frick, L. 2012. High post-capture survival for sharks, rays and

chimaeras discarded in the main shark fishery of Australia? PloS One, 7(2), e32547, 1–9.

Brevé, N.W.P., Winter, H.V., Van Overzee, H.M.J., Farrell, E.D. and Walker, P.A. 2016. Seasonal

migration of the starry smooth-hound shark Mustelus asterias as revealed from tag-recap-

ture data of an angler-led tagging programme. Journal of Fish Biology, in press

Burt, G. J., Silva, J. F., McCully, S. R., Bendall, V.A. and Ellis, J. R. 2013. Summary results from

opportunistic tagging programmes for smooth-hound Mustelus spp., greater-spotted dog-

fish Scyliorhinus stellaris and tope Galeorhinus galeus around the British Isles. Working Doc-

ument to the Working Group on Elasmobranch Fishes, Lisbon, Portugal, 17–21 June 2013.

12 pp.

Capapé, C. 1983. New data on the biological of the reproduction of Mustelus asterias Cloquet,

1821 (Pisces, Pleurotremata, Triakidae) from Tunisian coasts. Vie et Milieu, 33: 143–152.

Capapé, C., Diatta, Y., Diop, M., Vergne, Y. and Guelorget, O. 2006. Reproductive biology of the

smoothhound, Mustelus mustelus (Chondrichthyes: Triakidae) from the coast of Senegal

(eastern tropical Atlantic). Cybium, 30: 273–282.

Coelho, R., Fernandez-Carvalho, J., Lino, P.G., and Santos, M.N. 2012. An overview of the hook-

ing mortality of elasmobranchs caught in a swordfish pelagic longline fishery in the Atlantic

Ocean. Aquatic Living Resources, 25: 311–319.

Compagno, L., Dando, M. and Fowler, S. 2005. Sharks of the World. Collins Field Guide. Harper

Collins Publishers, London.

Ellis, J. R. 2015. Starry smooth-hound in the English beam trawl surveys of the eastern English

Channel and southern North Sea (VIId–IVc), and Irish Sea and Bristol Channel (Divisions

VIIa,f). Working Document to the ICES Working Group on Elasmobranch Fishes, 17–23

June, Lisbon, Portugal. 6 pp.

Ellis, J. R., Cruz-Martinez, A., Rackham, B. D. and Rogers, S. I. 2005. The distribution of chon-

drichthyan fishes around the British Isles and implications for conservation. Journal of North-

west Atlantic Fishery Science, 35: 195–213.

Ellis, J. R.., McCully, S. R. and Poisson, F. 2014. A global review of elasmobranch discard survival

studies and implications in relation to the EU ‘discard ban’. Working Document to the ICES

Working Group on Elasmobranch Fishes, Lisbon, Portugal. June 2014. 48 pp.

Ellis, J. R., Pawson, M. G. and Shackley, S. E. 1996. The comparative feeding ecology of six species

of shark and four species of ray (Elasmobranchii) in the North-East Atlantic. Journal of the

Marine Biological Association of the United Kingdom, 76: 89–106.


Espinoza, M., Farrugia, T. J. and Lowe, C. G. 2011. Habitat use, movements and site fidelity of

the gray smooth-hound shark (Mustelus californicus Gill, 1863) in a newly restored southern

California estuary. Journal of Experimental Marine Biology and Ecology, 401: 63–74.

Farrell, E. D., Clarke, M. W. and Mariani, S. 2009. A simple genetic identification method for

Northeast Atlantic smoothhound sharks (Mustelus spp.). ICES Journal of Marine Science, 66:

561–565.

Farrell, E. D., Mariani, S. and Clarke, M. W. 2010a. Age and growth estimates for the starry

smoothhound (Mustelus asterias) in the Northeast Atlantic Ocean. ICES Journal of Marine Sci-

ence, 67: 931–939.

Farrell, E. D., Mariani, S. and Clarke, M. W. 2010b. Reproductive biology of the starry

smoothhound shark Mustelus asterias: Geographic variation and implications for sustainable

exploitation. Journal of Fish Biology, 77: 1505–1525.

Fennessy, S. T. 1994. Incidental capture of elasmobranchs by commercial prawn trawlers in the

Tugela Bank, Natal, South Africa. South African Journal of Marine Science, 14: 287–296

Frick, L. H., Reina, R. D. and Walker, T. I. 2010. Stress related changes and post-release survival

of Port Jackson sharks (Heterodontus portusjacksoni) and gummy sharks (Mustelus anarcticus)

following gill-net and longline capture in captivity. Journal of Experimental Marine Biology

and Ecology, 385: 29–37.

Goosen, A. and Smale, M. J. 1997. A preliminary study of age and growth of the smoothhound

shark Mustelus mustelus (Triakidae). South African Journal of Marine Science, 18: 85–91.

ICES. 2012. Report of the 6th Meeting of the Mediterranean Planning Group for Methodological

Development (PGMED). 30 January to 5 February, 2012, Rome.

Jardas, I., Santic, M., Nerlovic, V. and Pallaoro, A. 2007. Diet of the smooth-hound, Mustelus

mustelus (chondrichthyes: Triakidae), in the eastern Adriatic Sea. Cybium, 31: 459–464.

Lipej, L., Mavric, B., Resek, S., Chérif, M. and Capapé, C. 2011. Food and feeding habits of the

blackspotted smooth-hound, Mustelus punctulatus (Elasmobranchii: Carcharhiniformes: Tri-

akidae), from the northern Adriatic. Acta Ichthyologica et Piscatoria, 41: 171–177.

Martin C. S., Vaz, S., Ellis, J. R., Coppin, F., Le Roy, D. and Carpentier, A. 2010. Spatio-temporal

patterns of demersal elasmobranchs in trawl surveys in the eastern English Channel (1988–

2008). Marine Ecology Progress Series, 417: 211–228.

Martin, C. S., Vaz, S., Ellis, J. R., Lauria, V., Coppin, F. and Carpentier, A. 2012. Modelled distri-

butions of ten demersal elasmobranchs of the eastern English Channel in relation to the en-

vironment. Journal of Experimental Marine Biology and Ecology, 418/419: 91–103.

McCully, S and Ellis, J. R. 2014. Biological studies to inform management of smooth-hounds

(Mustelus spp.) in the North-east Atlantic. Working Document to ICES Working Group on

Elasmobranch Fishes. Lisbon, Portugal. 16 pp.

McCully Phillips, S.R. and Ellis, J.R., 2015. Reproductive characteristics and life-history relation-

ships of starry smooth-hound Mustelus asterias in British waters. Journal of Fish Biology, 87:

1411–1433.

Morte, S., Redon, M. J. and Sanz-Brau, A. 1997. Feeding habits of juvenile Mustelus mustelus (Car-

charhiniformes, Triakidae) in the western Mediterranean. Cahiers de Biologie Marine, 38: 103–

107.

Nieto, A., Ralph, G.M., Comeros-Raynal, M.T., Kemp, J., Garcia Criado, M., Allen, D.J., Dulvy,

N.K., Walls, R.H.L., Russell, B., Pollard, D., Garcia, S., Craig, M., Collette, B.B., Pollom, R.,

Biscoito, M., Labbish Chao, N., Abella, A., Afonso, P., Alvarez, H., Carpenter, K.E., Clo, S.,

Cook, R., Costa, M.J., Delgado, J., Dureuil, M., Ellis, J.R., Farrell, E.D., Fernandes, P., Florin,

A-B., Fordham, S., Fowler, S., Gil de Sola, L., Gil Herrera, J., Goodpaster, A., Harvey, M.,

Heessen, H., Herler, J., Jung, A., Karmovskaya, E., Keskin, C., Knudsen, S.W., Kobyliansky,

S., Kovačić, M., Lawson, J.M., Lorance, P., McCully Phillips, S., Munroe, T., Nedreaas, K.,

Nielsen, J., Papaconstantinou, C., Polidoro, B., Pollock, C.M., Rijnsdorp, A.D., Sayer, C.,


Scott, J., Serena, F., Smith-Vaniz, W.F., Soldo, A., Stump, E. and Williams, J.T. 2015. European

Red List of marine fishes. Luxembourg: Publications Office of the European Union.

Pribac, F., Punt, A. E. Taylor, B. L. and Walker, T. I. 2005. Using length, age and tagging data in

a stock assessment of a length selective fishery for gummy shark (Mustelus antarcticus). Jour-

nal of Northwest Atlantic Fishery Science, 35: 267–290.

Quignard, J. P. 1972. Note sur les especes mediterraneenes du genre Mustelus (Selachii, Gale-

oidea, Triakidae). Rev. Trav. Inst. Peches Marit., Nantes, 36(1): 15–29.

Saidi, B., Bradai, M. N. and Bouain, A. 2008. Reproductive biology of the smooth-hound shark

Mustelus mustelus (L.) in the Gulf of Gabes (south-central Mediterranean Sea). Journal of Fish

Biology, 72: 1343–1354.

Saidi, B., Enajjar, S., Bradai, M. N. and Bouain, A. 2009. Diet composition of smooth-hound shark,

Mustelus mustelus (Linnaeus, 1758), in the Gulf of Gabes, southern Tunisia. Journal of Applied

Ichthyology, 25: 113–118.

Santic, M., Nerlovic, V. and Pallaoro, A. 2007. Diet composition of blackspotted smooth-hound,

Mustelus punctulatus (Risso, 1826), in the eastern Adriatic Sea. Journal of Applied ichthyology,

23: 279–281.

Silva, J. F., Ellis, J. R., Catchpole, T. L. and Righton, D. 2013. Bycatch and discarding patterns of

dogfish and sharks taken in commercial fisheries around the British Isles. Working Docu-

ment to the Working Group on Elasmobranch Fishes, Lisbon, Portugal. 17–21 June 2013.

31 pp.

Silva, J. F., McCully, S. R., Ellis, J. R. and Kupshus, S. 2014. Demersal elasmobranchs in the Eng-

lish Channel. Working Document to the ICES Working Group on Elasmobranch Fishes. 28

pp.

Winter, E., and Brevé, N. W. P. 2014. Starry smooth-hounds tagged in the Netherlands on the

move. Proceedings of the European Elasmobranch Association Annual Scientific Confer-

ence. 7th to 9th November 2014, Leeuwarden, The Netherlands.

Xiao, Y. and Walker, T. I. 2000. Demographic analysis of gummy shark (Mustelus antarcticus) and

school shark (Galeorhinus galeus) off southern Australia by applying a generalized Lotka

equation and its dual equation. Canadian Journal of Fisheries and Aquatic Sciences, 57: 214–222.


Table 21.1. Smooth-hounds in the Northeast Atlantic. Reported species-specific landings (tonnes) for the period 1973–2014. These data are considered underestimates as some smooth-

hounds are landed under generic landings categories. Species-specific landings data are not available for the Mediterranean Sea and are limited for the northwestern African waters.

1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986

Belgium . . . . . . . . . . . . .

France 0 0 0 0 0 0 32 0 0 222 218 66 143 167

Netherlands . . . . . . . . . . . . . -

Portugal . . . . . . . . . . . . -

UK -E, W & NI 0 0 0 0 0 0 0 0 0 0 0 0 0 0

UK - Scotland 0 0 0 0 0 0 0 0 0 0 0 0 0 0

0 0 0 0 0 0 32 0 0 222 218 66 143 167

1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999

Belgium

France 119 64 117 126 93 90 102 138 145 228 187 197 0

Netherlands . . . . . . . . . . . .

Portugal

UK -E, W & NI 0 0 0 0 0 0 0 0 0 0 0 0 0

UK - Scotland 0 0 0 0 0 0 0 0 0 0 0 0 0

119 64 117 126 93 90 102 138 145 228 187 197 0


Table 21.1. (continued). Smooth-hounds in the Northeast Atlantic. Reported species-specific landings (tonnes) for the period 1973–2014. These data are considered underestimates as

some smooth-hounds are landed under generic landings categories. Species-specific landings data are not available for the Mediterranean Sea and are limited for the northwestern

African waters.

2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014

Belgium . . . . . . . . . . . . 8 10 1

France 306 377 585 589 682 767 714 908 522 926 969 706 2695 2955 2825

Netherlands . . . . . . . . . . 8 3 11 20 15

Portugal . . . . . . . . . . . 35 42 41 187

Spain . . . . . . . . . . 34 48 9 83 14

UK -E, W & NI 14 0 0 0 0 0 0 0 115 132 161 919 337 323 647

UK - Scotland 0 0 0 0 0 0 0 0 0 1 0 - - - -

320 377 585 589 682 767 714 908 637 1059 1172 1712 3101 3433 3690

Table 21.2 Smooth-hounds in the Northeast Atlantic. ICES estimated landings (tonnes; 2005–2015), based on data provided in the ICES Data Call.

2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015

Belgium - - - - - - - - 1 1 1

Spain 112 134 138 200 297 129 106 120 80 70 44

France 2685 2722 2958 3403 3082 3204 3241 2821 2942 2836 2946

UK 171 130 155 171 199 275 315 339 325 331 303

Ireland - - 0 1 0 0 0 - - 0 0

Netherlands - - - 4 9 3 23 26 24 24

Portugal 44 57 57 41 45 38 43 42 41 17 15

Total 3013 3043 3308 3816 3628 3655 3709 3345 3415 3280 3333


Figure 21.1. Smooth-hounds in the Northeast Atlantic. Working Group estimates of overall Mus-

telus spp. landings (top) and by country (Middle), 2000–2014. New ICES estimates (2005–2015) are

at the bottom. Data are considered underestimates.

a b

0

500

1000

1500

2000

2500

3000

3500

4000

2000 2002 2004 2006 2008 2010 2012 2014

Ton

nes

LandingsLandings

0

500

1000

1500

2000

2500

3000

3500

4000

2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014

Ton

nes

UK - Scotland

UK -E, W &NI

Spain

Portugal


C D

E F

Figure 21.2. Smooth-hounds in the Northeast Atlantic. Length–frequency of discarded (pale grey)

and retained (dark grey) smooth-hounds Mustelus spp. by (a) otter trawl (2002–2005), (b) otter trawl

(2006–2011), (c) gillnet (2002–2005), (d) gillnet (2006–2011), (e) beam trawl (2002–2011) and (f)

Nephrops trawl (2002–2011), as recorded in the Cefas observer programme. Data aggregated across

ecoregions (Source: Silva et al., 2013 WD).


Figure 21.3. Smooth-hounds in the Northeast Atlantic. Number of starry smooth-hounds (n=504)

biologically sampled by length and sex. Source: McCully Phillips and Ellis (2015).


Figure 21.4a. Smooth-hounds in the Northeast Atlantic. Captures of Mustelus asterias as reported

in the 2011 IBTS. The catchability of the different gears used in the NE Atlantic surveys is not

constant; therefore the map does not reflect proportional abundance in all the areas but within each

survey. Source: ICES (2012).

35

40

45

50

55

60

15 10 5 0 5 10

15 10 5 0 5 10

35

40

45

50

55

60

LEGEND

6

18

30

60

> 70

SURVEYS:

NS-IBTS

SCOGFS

IGFS

NIGFS_Q4

EN-CEFAS_A

SP_Porc

EN-CEFAS_B

FR-EVHOE

FR-CGFS

SP_North

PT-GFS

SP_GC

Mustelus asterias

3 outliers between 226 and 406


Figure 21.4b. Smooth-hounds in the Northeast Atlantic. Captures of Mustelus mustelus as reported

in the 2011 IBTS. The catchability of the different gears used in the NE Atlantic surveys is not

constant; therefore the map does not reflect proportional abundance in all the areas but within each

survey. Source: ICES (2012).

35

40

45

50

55

60

15 10 5 0 5 10

15 10 5 0 5 10

35

40

45

50

55

60

LEGEND

5

15

25

50

SURVEYS:

NS-IBTS

SCOGFS

IGFS

NIGFS_Q4

EN-CEFAS_A

SP_Porc

EN-CEFAS_B

FR-EVHOE

FR-CGFS

SP_North

PT-GFS

SP_GC

Mustelus mustelus


Figure 21.5. Smooth-hounds in the Northeast Atlantic. Length–frequency by sex of smooth-hounds

Mustelus spp. From the UK Western Channel Q1 Beam-trawl survey. Source: Silva et al. (2014 WD).

Figure 21.6. Survey grid of the Irish Sea and Bristol Channel (left, 97 stations) and eastern English

Channel and southern North Sea (right, 76 stations) showing locations of fixed stations sampled

most consistently (1993–2014). Source: Ellis (2015 WD).

-7 -6 -5 -4 -3 -2

50.5

51.0

51.5

52.0

52.5

53.0

53.5

54.0

54.5

55.0

-2 -1 0 1 2

49.0

49.5

50.0

50.5

51.0

51.5

52.0

52.5

53.0


Figure 21.7. Smooth-hounds in the Northeast Atlantic. Relationship between maternal total length

and average length and weight of term pups. Source: McCully Phillips and Ellis (2015).

Figure 21.8. Smooth-hounds in the Northeast Atlantic. Percentage of mature females at each devel-

opmental stage (D: early gravid; E: mid-gravid; F: late gravid; G: post-partum) by month. Source:

McCully Phillips and Ellis (2015).


Figure 21.9. Smooth-hounds in the Northeast Atlantic. Length–frequency distributions of Mustelus

spp. (n = 715), and the length–weight relationships for (Mustelus spp. (n = 508) tagged during the

Cefas programme. Source: Burt et al. (2013 WD).

Figure 21.10. Smooth-hounds in the Northeast Atlantic. Maturity ogive for male (n= 237; L50 = 70.4

cm LT) and female (n= 248; L50 = 81.9 cm LT) M. asterias. Source: McCully Phillips and Ellis (2015).


Figure 21.11. Smooth-hounds in the Northeast Atlantic. Relationship between maternal total

length and number of term pups produced. Source: McCully Phillips and Ellis (2015).

Figure 21.12. Smooth-hounds in the Northeast Atlantic. Locations of smooth-hound, Mustelus spp.

(i) released and (ii) release and recapture positions for recaptured fish (2000–2013). Source: Burt et

al. (2013 WD).


Figure 21.13. Smooth-hounds in the Northeast Atlantic. The main map shows the more detailed

distribution of recaptures in the English Channel and southern North Sea. From three fish markets

(indicated with anchors), eight tagged M. asterias were reported (numbers next to the anchors rep-

resent the number of sharks from each fish market) with unknown recapture location. Inset (a)

shows the locations of recaptured Mustelus asterias (n = 80) reported by quarter for the years 2011–

2014. Their distribution pattern indicates a circannual migration between the Dutch Delta (sum-

mer), the English Channel and Bay of Biscay (winter). Inset (b) shows the tag and release location

with the main places fished indicated with open circles. Symbols: f = female; m = male; recaptures

per quarter are shown for January to March ( ),April to June ( ), July to September ( ) and October

to December ( ). Source: Brevé et al. (2016).


Figure 21.14. Smooth-hounds in the Northeast Atlantic. Length–weight relationship for female (n

= 248) and male (n = 237) M. asterias by maturity stage (shaded region showing 95% confidence

intervals). Source: McCully Phillips and Ellis (2015).

Figure 21.15. Smooth-hounds in the Northeast Atlantic.: Total length to gutted weight relationship

for female (n = 249) and male (n = 235) M. asterias (shaded region showing 95% confidence inter-

vals). Source: McCully Phillips and Ellis (2015).


Figure 21.16. Smooth-hounds in the Northeast Atlantic. Individual standardised survey indices

(top) and resultant overall stock size indicator of individuals per hour, using the mean standardised

indices from four surveys (Q1 NSIBTS, Q3 NSIBTS, UK-7d-BTS and UK-7af-BTS; bottom).

0,0

0,5

1,0

1,5

2,0

2,5

3,0

3,5

19

90

19

91

19

92

19

93

19

94

19

95

19

96

19

97

19

98

19

99

20

00

20

01

20

02

20

03

20

04

20

05

20

06

20

07

20

08

20

09

20

10

20

11

20

12

20

13

20

14

Ind

ivid

ual

s p

er

ho

ur

Standardised Survey CPUE, NEA Atlantic Smooth-hounds

IBTS Q1 EVHOE IGFS

0

0,5

1

1,5

2

2,5

19…

19…

19…

19…

19…

19…

19…

20…

20…

20…

20…

20…

20…

20…

20…

20…

20…

20…

20…

20…

20…

20…

Stock size indicator


Figure 21.17. Smooth-hounds in the Northeast Atlantic. Distribution of Mustelus spp. in catches

(green points vs. blue points for all sampling stations) in the Evhoe survey (1997–2014).

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