Cleaner fish biology and aquaculture applications (Ed. Jim Treasurer) - Chapter 6. Review of lumpfish biology.

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CHAPTER 6
Review of lumpfish biology
Adam Powell,1 Craig Pooley,2 Maria Scolamacchia2 and
CarlosGarciadeLeaniz2
1University of Gothenburg – Kristineberg, Department of Biological and
Environmental Sciences, Sweden
2Centre for Sustainable Aquatic Research, Department of Biosciences,
Swansea University, UK
6.1 Distribution
e lumpsh, Cyclopterus lumpus, occupies diering habitats depending on life stage. Adults are
generally solitary and spend much of the year far from land, and at up to several hundred metres
depth (Kudryavtseva and Karamushko, 2002; Parin et al., 2002). e following distribution
describes hatchlings, juveniles and adults during the breeding season, which takes place gener-
ally closer to shore, in shallow coastal water (after Davenport, 1985; Stein, 1986; Aquamaps,
2017).
Lumpsh may be regarded as abundant, potentially inhabiting c. 32,000 km of coast across both
sides of the Atlantic Ocean (see Davenport, 1985 for a summary of archive studies). Lumpsh
are distributed in the boreal region of the east and west North Atlantic coasts (Figure 6.1). For
the western Atlantic, the most northerly occurrence has been found on the island of Disko o
north-western Greenland; lumpsh are distributed from there southwards to Chesapeake Bay
(range: 70° – 37°N). is distribution incorporates most of eastern Canada, including Nunavut,
Hudson Bay, James Bay, Labrador, Newfoundland (Stephenson and Baird, 1988), Gulf of St
Lawrence, New Brunswick and Nova Scotia.
On the western side of the Atlantic Ocean, the species has been recorded over a wider range
of latitudes than the European coast (see Davenport, 1985). However, in Europe, recent records
of the species have been recently reported from lower latitudes o Spain, southern Portugal
(36°N) and a probable vagrant in the Mediterranean Sea (Banon et al., 2008; Vasconcelos et
al., 2004; Dulčić and Golani, 2006). e most northerly occurrences in Europe include Jan
Mayen (north of Iceland), Svalbard, and the White and Barents Seas including Novaya Zemlya
(Kudryavtseva and Karamushko, 2002), with populations found easterly into the Baltic Sea.
e species is commonly found o Iceland, the southerly tip of Greenland and the Faroes,
Norway (Holst, 1993) and countries bordering the North Sea, particularly France, the UK and
Ireland.
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Review of lumpsh biology ♦ 99
6.2 Description of the appearance of sh and taxonomy
e lumpsh or lumpsucker was formally named and initially characterised by Linnaeus (1758)
as Cyclopterus lumpus (‘round n’). Other common names in current usage across Europe (in
direct translation to English) include stone biter, sea chicken and fatsh, with names occasionally
diering for each sex due to pronounced sexual dimorphism (Davenport, 1985). e species is
a bony sh (class: Osteichthyes, infraclass: Teleostei) belonging to the Order Scorpaeniformes,
family Cyclopteridae. C. lumpus is unique and morphologically distinct, and is the only species
of the genus Cyclopterus. Related taxa include the less common Eumicrotremus spinosus (Atlantic
spiny lumpsh, found in the western Atlantic Ocean) and Aptocyclus ventricosus (Smooth lump-
sh, northern Pacic Ocean; Froese and Pauly, 2017). ese other species have no recorded
commercial value and have not been investigated for use as cleaner sh to our knowledge.
Davenport (1985) and Bigelow and Schroeder (2002) provide a detailed generic and specic
description of adult C. lumpus. Briey, in prole the body is about twice as long as it is deep, and
it is compressed anteriorly and posteriorly. e rst dorsal n forms a high crest with large com-
pressed tubercles, which increase in size with age, although these may be reduced in specimens
inhabiting particularly cold or low salinity habitats, such as the Baltic Sea. ere are three longi-
tudinal ridges along the length of the body marked by a line of pointed tubercles, with the most
obvious as a dorsal crest. e head is short with lateral moderate-sized eyes, while the opercula
have slit-like openings. e snout is blunt with a terminal, slightly upturned mouth containing
small teeth. In cross section, the body is vaguely triangular with a attened ventral surface (with
the exception of gravid females, when distended with roe), with a round, broad, muscular suck-
ing disc (Davenport and orsteinsson, 1991) that gives the species its generic name. e disc
constitutes approximately 20% of the body length and is a specialised organ that descends from
the pectoral ns. e vivid skin colouring in adults and associated sexually dimorphic character-
istics (see section 6.12) and skin texture (rubbery, tough and scaleless) are also diagnostic features
(Figure 6.2).
Fig. 6.1 Map showing coastal
distribution Cyclopterus lumpus
(shaded areas showing probable
extent of adult spawning grounds
and habitat of hatchlings and
substrate associated juveniles).
Credit: Redrawn by the authors
(after Davenport, 1985)
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100 ♦ Biology and rearing of wrasse and lumpsh
6.3 The lumpsh shery: vulnerability to shing and sources of
broodstock
Historically, the lumpsh shery has entered three distinct phases. Prior to the 1940s, there
was a small-scale artisanal shery for males that were sold smoked, salted or dried. is was
then largely replaced by a commercial shery emerging in the late 1940s, focusing on roe from
females (as a cheap alternative to caviar from sturgeon; see Davenport, 1985). is has continued
to the present day at scales of many thousand tonnes per year (Figure 6.3; Johannesson, 2006).
More recently, the shery has diversied somewhat to include male and female broodstock for
the emerging cleaner sh industry. Aquaculture production of the species currently depends on
the capture of wild broodstock, which together with current sheries may impact on natural
populations.
e species has a trophic level of 3.9, typical of secondary consumers, and a low resilience to
shing pressure, with an estimated time for doubling of population size of 4.5 to 14 years. It
also has a moderate to high vulnerability to shing (47 of 100; Cheung et al., 2005; Friese and
Pauly, 2017). Although the conservation status of the species has not been assessed by the IUCN
(International Union for Conservation of Nature), the characteristics outlined above suggest a
species vulnerable to overexploitation. A signicant decrease in some spawning stocks has been
recorded in recent decades, suggesting that some stocks may already be overexploited (Pampoulie
et al., 2014). Global lumpsh production from sheries is seasonably variable (Johannesson,
2006; FAO, 2017), potentially inuenced by the combined impacts of exploitation, with emerg-
ing problems such as invasive species that feed on lumpsh eggs (Mikkelsen and Pedersen, 2012),
climate change (e.g. Perry et al., 2005) and emerging diseases (e.g. Freeman et al., 2013).
e reported catch of lumpsh has steadily increased from a few hundreds of tonnes per year
in the early 1950s to 20,365 tonnes in 2013 (FAO 2017, Figure 6.3). Mean catch since the year
2000 has been 15,997 tonnes (SE = 1,002) with unreported catches probably varying between
1,600 and 4,600 tonnes (i.e. 9–22% of the total). However, historical catch records are proba-
bly misleading because most of the former catch was unreported, mostly due to discards on the
Fig. 6.2 Caption: Cyclopterus lumpus adult male
and female a. Lateral view. b. Ventral view. Credit:
Photograph by Craig Pooley and Maria Scolamacchia,
CSAR, Swansea University
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Review of lumpsh biology ♦ 101
east coast of Canada. Data reconstruction indicates that unreported catches may have reached c.
124,000 tonnes (98% of total) by 1965, when the declared catch was only 2,400 tonnes (Pauly
and Zeller, 2015).
Nowadays, the most important lumpsh sheries operate in the Canadian eastern Atlantic–
west Greenland (c. 70% of catch) and Iceland (c. 23% of catch), which are exploited by the
Greenlandic and Icelandic eets. Data from 2010 indicate that these are mostly mixed artisanal
(57% of catch) and commercial sheries (43% of catch), with only 30 tonnes being classied as
subsistence (0.1% of catch; Pauly and Zeller, 2015).
e artisanal sheries consist of small boats, shing close to shore in up to ~30 m depth, and
which are typically manned by one to three people. Gears vary among locations (Johannesson,
2006). In Norway, nets with 252 mm mesh size are used, these typically being c. 47 m wide
with a 3.4 m drop (Bertelsen, 1994). Mesh size in Iceland varies between 267 and 286 mm
(orsteinsson, 1996). Monolament nets with a 256–281 mm mesh size are used in Canada
to catch lumpsh, nets being 100m long (Benson et al., 1998). eir round shape and short ns
means that lumpsh do not entangle in the nets to the same extent that other species do, and
this is the reason lumpsh nets have as much ‘bag’ as possible, as this is thought to increase catch
eciency. Typically, nets are set up on 45 m oat and foot lines, with 90–125 cm verticals to
maintain the bags and prevent lumpsh escape (Johannesson, 2006).
Traditionally, lumpsh have been harvested for their roe, which can be processed into an alter-
native to caviar (Johannesson, 2006). No reliable data are available for the number of females
removed from the wild for the caviar industry, but this probably amounts to several million sh
annually; eorts to harvest lumpsh eggs non-destructively have met with limited success (Grant,
2001). e number of adult lumpsh taken by the incipient cleaner sh industry (c. 300 tonnes
in the UK in 2014, personal observation) is currently small compared with the commercial and
artisanal sheries (20,365 tonnes in 2013, FAO 2017), but there is a growing concern from some
conservation bodies regarding the sustainability of the catch (Anon, 2013a). Maximum lumpsh
Fig. 6.3 Reported global catch for
lumpsh (tonnes), 1950–2013 (FAO
FishStat). Credit: FAO Fishstat
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102 ♦ Biology and rearing of wrasse and lumpsh
production is attained when males reach 20 cm and females reach 32 cm (equivalent to two and
three years of age, respectively (Hedeholm et al., 2014), and that means that removing brood-
stock older than two to three years of age may have a disproportionately high impact on wild
populations. Both Iceland and Greenland have adopted strict shing regulations (Stevenson and
Baird, 1988) and their lumpsh sheries have recently been awarded MSC certication (Anon,
2013b; Lassen et al., 2015).
While it is possible to use means other than gillnets to capture lumpsh, for example by
scuba diving (Killen et al., 2007a), these tend to be more labour intensive and much less e-
cient. Lumpsh lack a swim bladder, and adults hauled quickly to the surface may experience
barotrauma, even using static gear. is can be rectied by returning animals to depth in a cage,
followed by gradual decompression.
6.4 Commercial uses of lumpsh
Up until the 20th century lumpsh had little economic value. Small sheries existed on both
sides of the North Atlantic for local consumption, but sh caught as bycatch were often used
as animal feed or bait (Stevenson and Baird, 1988). A dedicated shery targeting lumpsh roe
started in the late 20th century. Ripe females yield 15–36% of roe by weight (Davenport, 1985,
Stevenson and Baird, 1988, Mitamura et al., 2007, Hedeholm et al., 2014) and were rapidly
targeted by the shery. us, in Newfoundland, roe production grew from 21 tonnes in 1970
to 3,000 tonnes by 1989 (Stevenson and Baird, 1988). Similarly in Norway, 100 tonnes of roe
were taken annually in the 1950s, compared with 500 tonnes of roe by the middle of the 1980s
(c. 2,500 tonnes of sh).
Lumpsh eggs are marketed in two ways: either as whole roe, which is then dried, salted, or
smoked, or as processed eggs, which are separated from the ovaries and then further elaborated
into lumpsh caviar. Annual production of lumpsh caviar is about 4 million kg and has a
market value of about $ 60 million, with Canada (35%), Iceland (31%) and Norway (15%) as
the main producers (Johannesson, 2006). e price of lumpsh caviar uctuates depending on
supply, but is currently sold in supermarkets at $36–72 per kg.
Excluding the sale of caviar, the shery has a low value (always below $5 million since 2000),
and the species is classed as a ‘low value’ sh in the prize category of FishBase (Sumaila et al.,
2007; FishBase, 2017). is is in stark contrast to the situation during 1950–1970, when the
value of the shery peaked at c. $200 million in 1961, mostly generated by the European eet
(Spain, Portugal, Germany) operating in eastern Canada. However, in some places such as
Iceland, the UK and Norway, this situation has changed in recent years, with the development
of the cleaner sh industry. Fishermen are currently paid US $87 (GBP £60) for ripe females in
some parts of the UK, and a kilogram of fertilised eggs for export to the cleaner sh industry may
cost several hundred US dollars in Iceland personal observation), which may put new pressures
on wild stocks.
Currently, all lumpsh used as cleaner sh are culled at the end of the salmon production
cycle, when they reach about 500 g. Although this is done on biocontainment grounds, to pre-
vent transmission of diseases between salmon and cleaner sh, some have considered this prac-
tice as wasteful and have urged the industry to nd alternative uses for farmed lumpsh (Anon
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Review of lumpsh biology ♦ 103
2013a; Powell et al., 2017). is has prompted research into new markets and a consideration
of possibilities for nutrient recycling, either as animal feed or into more valuable products, par-
ticularly in the lucrative Asian markets, where the species could perhaps nd a market niche as a
delicacy (Nøstvold, 2017). Another alternative to culling would be to recondition large juveniles
in captivity and use them as broodstock for the production of eggs, either for caviar or for the
cleaner sh industry (Powell et al., 2017), although such animals would need to be certied free
from disease.
6.5 Abundance
Lumpsh occupy a wide distribution, and have been recorded in 24 countries in Europe and
North America. Animal distribution also varies according to life history stage and season, with
many historic reports unaware of the semi-pelagic adult phase outside the spawning season.
Davenport (1985) summarised this as a fundamental problem with assessments of the total area
occupied by the species. A recent study of stocks in the Barents Sea (Eriksen et al., 2014) sug-
gested a mean annual biomass (48 000–143 000 t) and mean annual abundance (53 –132 mil-
lion individuals) since 1980. Some data on stock assessment are available for the Icelandic and
Greenland sheries that were recently awarded MSC certication, but not for other areas. On
the other hand, reliance on catch data (Figure 6.3) to infer trends in abundance is fraught with
diculties, as the majority of lumpsh caught during 1950–1970 went unreported.
6.6 Food
e diet of lumpsh was summarised by Davenport (1985), who concluded that the overall
impression was of a species that subsisted mainly on large planktonic organisms living in surface/
mid waters, but which sometimes browsed upon benthic organisms, particularly those dwelling
upon weed. For adults, historic studies found a high proportion (c. 70–80%) of adults with
empty stomachs in sampled populations. A similarly high proportion of empty stomachs has also
been reported in juveniles (Ingólfsson and Kristjánsson, 2002). e intestine is long, being more
than twice the length of the body in adult sh, with many bends and numerous pyloric caecae
(Davenport, 1985) suggesting ecient digestion and absorption of food. Gut contents of adults
are variable across all studies, with small crustaceans (mysids, amphipods, euphausids, isopods,
decapod zoeae), ctenophores, polychaetes, seagrass, insects, small sh and sh eggs recorded
(Davenport, 1985; Davenport and Rees, 1993).
Juvenile lumpsh (c. 5–55 mm length) are apparently year-round seaweed specialists, inhabit-
ing and feeding on surface plankton after hatching, and weed-associated invertebrate fauna when
larger (Daborn and Gregory, 1983; Vandendriessche et al., 2007). Motile crustaceans larger than
0.5 mm dominate diet, with the natural abundance of these prey items on seaweed correlating
with their abundance in the gut of juvenile lumpsuckers (Ingólfsson and Kristjánsson 2002).
Furthermore, as hatchlings increased in size and yolk reserves became exhausted, the proportion
of larvae with full guts increased. Hatchlings tend to consume harpacticoid copepods or halac-
rid mites, with older juveniles switching to physically bigger and more active crustacean prey
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104 ♦ Biology and rearing of wrasse and lumpsh
such as amphipods or decapod larvae (Daborn and Gregory, 1983; Tully and Ó Ceidigh, 1989;
Ingólfsson and Kristjánsson 2002). Since cannibalism has been observed between individuals
of similar age and size, mouth gape is probably not a limiting factor to the size of prey that is
consumed. Rather, a change in behaviour corresponding to age and feed density may allow them
to enter a dierent trophic group (See section 6.15; Brown, 1986; Tully and Ó Ceidigh, 1989;
Killen 2007a, b).
More recently, interest has switched to the gut contents of lumpsuckers deployed in salmon
pens. Large lumpsh juveniles 54 g in weight deployed in sea pens are seemingly highly oppor-
tunistic, not limiting themselves to one food item (Imsland et al., 2015a). Over a 77-day trial,
the most common food items observed were salmon pellets, although crustaceans, hydrozoans,
mussels and, as the study progressed, sea lice, became an increasing part of the diet. Further
research suggests that food preference in pens may also depend on genetic provenance, and dier
between juveniles originating from distinct families (Imsland et al., 2016a), size and co-existence
with other species during deployment (Imsland et al., 2016b, 2016c).
6.7 Physiology
Lumpsh anatomy is unusual, with most studies focusing on the skeleton, skin and ventral
sucker, tissue composition and buoyancy, and reproduction.
Although the species has no swim bladder and contains large quantities of dense eggs, gravid
females have a body density very similar to seawater (Davenport and Kjörsvik, 1986). e low
density is achieved by extensive subcutaneous jelly, low osmolarity ovarian uid, and a dorsal
musculature, which is loose-bred and has high water content. Males do not present these char-
acteristics to such an extent, but have a higher lipid composition. is enables adult lumpsh to
reside in the open ocean for long durations. However, during the breeding season, they return to
shallow, rocky waters that experience strong wave surge. e low body density, large surface area
and limited swimming power demands an extremely powerful and ecient muscular, cartilagi-
nous sucker (or disc), which enables lumpsuckers to attach themselves to substrate, and to rest
without being swept away (Davenport and Kjorsvik, 1986; Davenport and orsteinsson, 1990).
Lumpsuckers can also colour-match their skin to local substrates within minutes (Davenport and
Bradshaw, 1995). Suitable resting places and substrates have also been recommended during
deployment in salmon pens, to promote sh welfare (Imsland et al., 2015b).
Unfertilised eggs in the large ovaries are bathed and protected in copious ovarian uid, which
has a low divalent ion concentration. As portions of ripened eggs are released during spawning
events, they harden and clump together in seawater due to the presence of divalent ions, particu-
larly calcium and magnesium. Copious uid and a strong sphincter muscle reduces the risk of
seawater reux into the oviduct, hardening remaining eggs inside the sh and blocking further
egg release. e kidney likely removes divalent ions, while female sh have large urinary blad-
ders that may assist with urine storage and water reabsorption (Davenport and Lönning, 1983;
Lönning et al., 1984).
Sperm extracted from males was found to remain viable for several days after removal of the
sh, although Davenport (1983) suggested that stripping eggs or milt from adults was not pos-
sible and euthanasia was required to extract gametes for articial fertilisation. Powell et al. (2017)
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Review of lumpsh biology ♦ 105
found that it was possible to strip eggs from very ripe females, while recently caught males often
release milt upon handling (although this was never achieved a few days later, in aquaria) requir-
ing culling and manipulation of male gonadal tissue. However, with well-considered logistics, it
may be possible to store sperm from wild ‘running’ males and, using enhancers or cryopreserva-
tion techniques (Norðberg et al., 2015), to reduce the necessity for maintaining or euthanising
adults.
6.8 Environmental requirements and ecotoxicology
Overall, C. lumpus is regarded as ‘hardy’ (i.e. robust); with the southern and northern limits of
distributional range approximating to the 20°C and 0°C August surface water isotherms, indicat-
ing that the lumpsucker is eurythermal but capable of inhabiting very cold water (overview by
Davenport, 1985). Adults are generally not found in low salinities within their latitudinal range,
although populations exist in the Baltic Sea and Hudson Bay that have permanently low salini-
ties; these populations also show some dierences in skin, body shape and smaller comparative
size (Davenport, 1985), suggesting a discrete race or even subspecies inside a particular oceano-
graphic area that experiences low salinity.
Hatchlings are precocious with a functional sucker, allowing immediate attachment to rock
or weed substrate, or even momentarily to the tending male shortly after hatching (Davenport,
1985). Hatchlings and juveniles appear to be almost obligate inhabitants of seaweed, living and
feeding on and around weed beds (Vandendriessche et al., 2007), although young juveniles are
also commonly found in parts of the intertidal zone such as rockpools (Moring, 2001).
Few ecotoxicological studies have been made on the species, although very low concentra-
tions of mercury have been found in the esh of wild adult lumpsuckers (Freeman, 1974).
Juveniles proved relatively resilient to experimental oil exposures designed to simulate oil spills
(Frantzen et al., 2015). An antifoulant candidate (medetomidine) has been tested on lump-
sh larvae; sub-lethal eects (a reduction respiration rate and skin colour adaptation rate) were
found when exposed at nanomolar concentrations for 72–98 hours. However, larvae returned to
baseline levels after 48 hours in untreated seawater (Bellas et al., 2005). is nding may have
implications for deployment near antifoulant-treated structures.
6.9 Movements and migrations
e species is benthopelagic and can be found at depths <868 m (Parin et al., 2002), although
they usually live in waters between 50 and 150 m deep (Stein, 1986). e semi-pelagic status of
adult females has been conrmed with a recent tagging study (Kennedy et al., 2016). Lumpsh
may undertake extensive annual migrations between their feeding grounds found in deeper waters
in the winter and the shallower waters preferred for spawning in spring and summer (Blacker,
1983; Davenport, 1985; Kaspar et al., 2014). Tagging studies indicate that the species displays
homing behaviour and may return to breed in the same areas more than once (Davenport, 1985,
Stevenson and Baird, 1988, Kennedy et al., 2014). Tagged females were found to reside in a ord
for up to a week and then disappeared, possibly returning oshore after spawning (Mitamura
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106 ♦ Biology and rearing of wrasse and lumpsh
et al., 2012). Following spawning, females can travel up 49 km per day (Kennedy et al., 2014),
unlike males, which remain in the same location for several weeks to tend the eggs (Davenport,
1983).
6.10 Genetic diversity and genetic stock differentiation
Lumpsh have 25 haploid chromosomes (n) and 50:50 diploid chromosomes (2n; Li and
Clyburne, 1977; Klinkhardt et al., 1995), but little is known about its genetic diversity or extent
of population dierentiation. Twenty-two novel microsatellite DNA loci were characterised
recently for the species (Skirnisdottir et al., 2013) and these have revealed three distinct genetic
groups in the North Atlantic: Maine–Canada–Greenland; Iceland–Norway, and the Baltic Sea
(Pampoulie et al., 2014; Garcia-Mayoral et al., 2016) with little evidence of gene ow among
these zones. However, data for other parts of the range are currently lacking. Preliminary results
from populations in the English Channel suggest that lumpsh there have low to moderate levels
of genetic diversity and low genetic dierentiation (Expected Heterozygosity, He = 0.53–0.61;
Pooley et al., 2015). is initial data suggests indistinct genetic populations in this particular
geographic area. is demands further investigation to inform a sustainable shing strategy, and
potential end uses after deployment as cleaner sh.
6.11 Behaviour
Adult lumpsh are typically solitary in the wild, though hatchlings tend to aggregate in clumps
in tanks during captivity (personal observation). Adult spawning behaviour has been described
by Goulet et al. (1986), Goulet and Green (1988) and is summarised in section 6.13. At spawn-
ing, the male exhibits parental care and tends the eggs, which are kept oxygenated through
fanning and air pung.
Unlike most other Actinopterygians (ray-nned sh), lumpsh lack Mauthner neurons that
are involved in the startle response and escape behaviour (Hale et al., 2000). ey have a longer
startle response latency than other shes, but are nevertheless capable of performing a fast anti-
predatory response, albeit not as eective. ey appear to have a relatively small brain for their
size (encephalisation coecient = 0.07–0.21; Albert et al., 1999), though this may be due to
the unusual shape and large body mass of the species. e species is apparently an intelligent,
inquisitive sh that can be entrained to accept feed (personal observation) and can change colour
rapidly for camouage (Davenport, 1989; Davenport, 1995).
e ventral sucking disc enables lumpsh to forage dierently from most other shes. ey
can cling and feed passively, or forage actively in pursuit of prey. e larvae become more active
a few weeks post-hatch (Brown, 1986) and their foraging mode appears to depend on prey abun-
dance. us, they adopt a ‘passive cling’ foraging mode when food is abundant, and switch to a
more ‘active swim’ mode when food is more scarce (Killen et al., 2007a).
Little is known about the welfare requirements for the species. Some information is avail-
able on the substrate and colour preferences following deployment in salmon pens (Imsland et
al., 2014a, 2015b), but studies are also needed during the juvenile phase. In the wild, lumpsh
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Review of lumpsh biology ♦ 107
match the colour of seaweed, suggesting that light intensity, photoperiod and tank colour may
also aect juvenile growth, since these factors have been observed to alter melanin concentrations
under experimental conditions (Davenport and Bradshaw, 1995). In salmon pens at night, they
prefer to aggregate together on smooth plastic and concrete substrates (thought to be similar to
seaweed), rather than on stones or car tyres (Imsland et al., 2015b). In comparison with Ballan
wrasse, Labrus bergylta (Helland et al., 2014), availability of suitable substrates appears to be
important for health and welfare.
e behaviour of lumpsh in cages has been studied via underwater cameras (Imsland et al.,
2015b,), and an ethogram has been constructed that identied a range of dierent behavioural
elements, which varied with the presence of Atlantic salmon (Imsland et al., 2014a). When
salmon were present, lumpsh were more active and spent less time resting, but little antagonis-
tic behaviour was observed between the two species. However, lumpsh did show some aggres-
sion to co-deployed goldsinny wrasse (Ctenolabrus rupestris), particularly across disparate size
grades between the two species (Imsland et al., 2016b).
Lumpsh are opportunistic omnivores in salmon cages, eating not only sea lice, but also
salmon pellets and organisms attached to the cages (Imsland et al., 2015a, 2016b. Signicant
variation appears to exist among families in the ecacy of sea lice grazing (Imsland et al., 2016a),
and this may provide the basis for the selective breeding of lumpsh with improved delousing
eciency.
6.12 Sexual dimorphism and sex ratio
Lumpsh display pronounced sexual dimorphism, allowing straightforward dierentiation of
sexually mature specimens. Males are typically smaller (30 ±10 cm length) and during the spawn-
ing season attain startling red, orange or purple coloration of the ns, eyes and ventral surfaces
(e.g. Davenport and orrsteinsson, 1989). Females are larger (42 ± 10 cm length), less colourful
and usually grey or blue-green, and are markedly rotund, likely due to increased subcutaneous
gelatinous tissue, urinary bladder and copious ovarian tissue, eggs and uid; they also have a
larger vent (Davenport and Lonning 1983; Davenport, 1985; Goulet et al., 1986; Figure 6.2).
Although the available data on sex ratio is scant, it is clear that the species deviates from the
Fisherian 1:1 ratio, the extent of which is likely to dier in space and time. e apparent itin-
erant, oceanic and solitary nature of sexually mature lumpsh outside spawning seasons, and
contrasting habitat favoured by juveniles (and diculty to assign gender) makes an accurate and
meaningful estimation of the sex ratio somewhat challenging. It is also likely that dierent arrival
and residence times between sexes during the spawning period (females arrive later and for only
a few days; Davenport, 1985; Mitamura et al., 2007) would further complicate the description
since sex ratio would change during the season. For instance, the reported sex ratio of 1:1.57
(F:M; n=54; Gregory and Daborn, 1982) is likely to be specic for the sample taken in shallow
water, in the Bay of Fundy, in early summer.
Furthermore, some have speculated that males outnumber females since they are apparently
capable of guarding more than one egg mass at a time; however, females may spawn their eggs
in a number of batches at dierent locations (Mochek, 1973; Davenport, 1985; Davenport and
Kjvosvik, 1986). In addition, it has yet to be conrmed if sexually mature females spawn annually,
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108 ♦ Biology and rearing of wrasse and lumpsh
at greater intervals or if they are eectively semelparous due to potentially low recovery rates
after spawning (which becomes progressively more expensive with increasing size; orsteinsson,
1983; Kaspar et al., 2014). An added complication is that males mature one to two years earlier
than females, however females appear to experience greater longevity (Davenport, 1985; Albert
et al., 2002; Hedeholm et al. 2014). erefore, it seems the operational sex ratio (i.e., the local
ratio of fertilisable females to sexually active males at any given time) for lumpsh is apparently
quite complex.
In areas of their range where a roe shery is signicant, there may be further bias toward
recorded numbers due to the dierence in size and behaviour between genders, and stronger
swimming ability of males. Males are likely to be underestimated in the gillnet shery since
they may be able to evade capture or are unlikely to move signicant distances while guarding
eggs (Davenport, 1985). Furthermore, the very presence of a shery that targets gravid female
sh will bias the sex ratio. Landing data is often expressed by biomass or tonnage rather than by
number (e.g. Marine Research Institute, 2015) although analysis of long-term survey data from
Newfoundland found that the sex ratio has decreased progressively from approximately 2.24:1
to 1.09:1 (F:M) between 1985–1994, despite an increase in Catch per Unit Eort (CPUE;
Stansbury, 1995; Hoenig and Hewitt, 2005). However, human activity may not be the sole
reason for female mortality. Emerging diseases such as Nucleospora cyclopteri were more prevalent
in females in a recent survey of the species o Iceland (Freeman et al., 2013).
6.13 Reproduction, spawning and geographical patterns
Early in the breeding season, males establish territories in shallow water on rocky and/or sea-
weed-covered substrate, prior to the arrival of females; females arrive asynchronously, allowing
more than one pairing and breeding event to occur per animal. is process occurs in spring and
early summer (Davenport, 1985) with a likely latitudinal variation. In Iceland, northern Norway
and Newfoundland, females start to move inshore during March and April, with evidence for
spawning occurring in early July and hatching until late August (Brown et al., 1992; Kennedy
et al., 2014; Mitamura et al., 2012) whereas in the English Channel gravid females have been
caught from early January until early May only (personal observation).
Davenport (1985) and Goulet et al. (1986) provide accounts of courtship, which appears to
be extended and of several hours’ duration. is includes showing anks to one another, pectoral
n brushing, quivering, and long periods of sucker attachment in close proximity; there was also
evidence for olfactory (perhaps pheromone) communication. In the wild, this included cleaning
a nest site (which may be a simple crevice or depression in bedrock, boulders and/or vegetation).
Mating appears to correlate with night (or darkness in aquaria) and potentially with high tide.
Fertilisation is external, with females releasing eggs freely into the water on to the surface of
a nest (or in the aquaria, near to any shallow depression or tank structure, personal observation)
with attendant males immediately fertilising the eggs, with the overall act lasting only ve to ten
seconds (Goulet, 1986). Females are thought to lay two to three batches of eggs over c. one to
two weeks (Davenport et al., 1985), and in one study actively moved in and out of a ord within
days of spawning (Mitamura et al., 2012). On contact with sea water, the eggs adhere to one
another to form large ovoid masses, which are fertilised by the male. Before the mass hardens, the
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Review of lumpsh biology ♦ 109
male may mould the eggs into the nest. Males also create funnel-like depressions that likely assist
gaseous exchange and removal of nitrogenous waste from the centre of the egg mass.
e eggs undergo paternal care throughout the incubation, with the females expending no
time in egg care after fertilisation. e most frequent behaviour includes water circulation by
fanning with the pectoral, but also dorsal and caudal ns (for a few seconds to hours in dura-
tion), and pung (expelling water from the mouth). is appears to occur more frequently for
a few hours after fertilisation, and also near the end of the incubation period when the embryos
are more developed, and to disseminate emergent larvae (Davenport, 1985; Goulet et al., 1986).
If eggs are exposed by low tides, males have been observed to spout water from the mouth to
maintain them (Davenport et al., 1984; Davenport, 1985). e male also becomes aggressive
and removes or defends the egg mass from conspecics and predators, including sea urchins,
periwinkles and even large predatory sh, although this is more challenging against schools of
ocean pout Zoarces americanus, cunners Tautogolabrus adspersus (Davenport, 1985 Goulet et
al., 1986) or large crabs such as the invasive red king crabParalithodes camtschaticus (Mikkelsen
and Pedersen, 2012). Males expend much eort in guarding egg masses, apparently not eating
during this period, potentially guarding more than one mass at a time or guarding successive
masses over the spawning season. e amount of time spent in parental care, or number of eggs
guarded, is independent of male size, while nest characteristics such as depth, distance from
shore, and topography do not correlate with hatching success (Davenport, 1985; Goulet et al.,
1986; Goulet and Green, 1988).
6.14 Egg stages and environmental preferences
Lumpsh eggs are relatively uniform in size (2.0–2.6 mm diameter) across their range (see
Davenport, 1985; Benfrey and Methven, 1986; Brown et al., 1992) and after extrusion undergo a
colour change from pink to a variety of colours that are homogenous within a particular egg mass.
Colour is lost with time as the pigments move from the yolk into the chromatophores of the
developing embryo. e hardening process occurs via a transparent adhesive material, secreted
by the ovary to coat the eggs. is becomes viscous and elastic on exposure to seawater and con-
denses within an hour at 5°C to form a dense layer around the eggs. Within 48 hours the eggs
themselves harden and become dense (Zhitenev, 1970; Lonning et al., 1984; Davenport, 1985).
Embryonic development has been pictorially described for the related smooth lumpsucker
Aptocyclus ventricosus (Kyûshin, 1975) and for C. lumpus in this chapter (Figure 6.4). With the
naked eye, the initial egg colour fades three to four days after fertilisation, and eggs ‘eye-up’ after
10–12 days, with full embryo pigmentation and a darker egg mass existing from about day 16
onwards. Infertile or undeveloped eggs remain unpigmented and opaque. Initial development
can be slow and variable in the egg mass (Davenport, 1983). In our studies, the following stages
were observed post-fertilisation at 10°C: morula (one to two days); blastula and blastodisc (two
to four days); gastrulation; initial somites visible (ve to six days); otic capsule visible, continued
segmentation (seven days); eye pigmentation and heart beat (eight to ten days); yolk vascularisa-
tion and head pigmentation (11–12 days); development of eyes, head and extensive vascularisa-
tion (15 days); uniform pigmentation, regular heart beats, mouth opening, movement (16–19
days).
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110 ♦ Biology and rearing of wrasse and lumpsh
Early observations showed that eggs develop more rapidly as temperature rises (Davenport,
1985), with studies showing a temperature of greater than 3.8°C is required for development,
with hatching after 40, 31 and 25 days at 5°C, 6.4°C and 9.8°C respectively (Collins, 1978;
Davenport, 1983) with hatching occurring mainly at night (Benfrey and Methven, 1986).
Studies at Swansea University have found a time to hatch of 20–30 days at 10°C (i.e. 200–300
degree days C) with hatching occurring over a seven- to ten-day period. Egg masses manipulated
into attened layers 5–10 mm thick immediately after fertilisation (but prior to hardening)
have generally led to better hatching success and more even development rate, perhaps due to
improved water circulation. is is in agreement with Brown et al. (1992), who had better suc-
cess with ‘monolayers’ rather than fragmented egg masses, and which developed at 28.5 days at
10°C. Lumpsh embryos at early developmental stages are capable of extracting oxygen at low
oxygen tensions (c. 40% of air saturation), or even withstand 30 minutes of anoxia, although sen-
sitivity is increased in more developed embryos nearing hatching (Davenport, 1983). Correct egg
development also demands salinities between c. 20–34 ppt (Kjorsovik et al., 1984). Little appears
to be known regarding optimal photoperiod or pH. Further details on lumpsh broodstock,
hatchery and deployment are reviewed by Powell et al. (2017) and are considered in Chapters
7, 8 and 9.
6.15 Juveniles
Juvenile lumpsh hatchlings are c. 5–6 mm length and 2–3 mg weight (Benfey and Methven,
1986; Brown et al., 1992). e sucker is present at hatching and allows immediate attachment to
surfaces, although in other respects newly hatched lumpsh do not resemble adults. A posterior,
protocercal n is present dorsally and ventrally that terminates at the tail. e median n breaks
up into separate ns by the length of about 8–9 mm. e rst dorsal n is gradually overgrown
Fig. 6.4 Overview of embryogenesis of C.
lumpus at 10°C. A. Morula stage, c. two days
post-fertilisation. B. Blastula stage, c. four days
post-fertilisation. C. Eye pigmentation, c. ten
days post-fertilisation. D. Full pigmentation and
development at hatch. Scale bar = 1mm. E. Part
of egg mass, corresponding to Blastula stage; eye
pigmentation has not developed but mass has
lost initial colour. F. Part of egg mass, just before
hatch. Note dark colour of eggs and occasional
light eggs denoting undeveloped embryos. Scale
bar = 10mm. Credit: Description and images
courtesy of Maria Scolomacchia
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Review of lumpsh biology ♦ 111
by the characteristic dorsal ‘hump’, and a 32 mm specimen is essentially a miniature of the adult
sh (Figure 6.5; Davenport, 1985). Feminisation of lumpsucker hatchlings has been demon-
strated, either by immersion in oestradiol or via enrichment in live feeds, which could promote
production of monosex populations (Martin-Robichaud et al.,1994).
Wild juveniles feed on plankton, and substrate-associated small invertebrates, and have been
oered Artemia and/or small dry feed pellets in hatcheries (e.g. Benfey and Methven, 1986;
Brown, 1986; Nytro et al., 2014, Powell et al., 2017). Juvenile lumpsh show limited aerobic
scope compared with other sh species (Killen, 2007a, b) and thus feed and behave dierently
from most other cultured sh due to the ventral sucker. is enables hatchlings to choose a
‘passive cling’ mode or a more costly ‘active swim’, with the latter change in behaviour induced
at low prey densities. Higher prey densities allow scope for other physiologically demanding
processes such as growth and digestion (Killen 2007a, b). However, lumpsh larvae grew faster
when food was administered in short pulses than when it was administered continuously (Brown
et al., 1997).
A few weeks post-hatch, hatchlings become more active (Brown, 1986), and also have redu-
ced yolk stores (Ingólfsson and Kristjánsson, 2002) allowing the start of weaning. Immobile dry
feed pellets <800 µm have been oered after 21 days, and hatchlings are maintained in shallow
raceways (Nytro et al., 2014), or circular tanks (Powell et al., 2017). e use of live feeds early
in the ontogeny of lumpsh larvae seems crucial, although the use of preserved or dry feeds in
older hatchlings switched on genes responsible for lipid metabolism and in some cases yielded
improved growth and survival (Belova, 2015). A strategy for species-specic improvement of
live feed enrichments or dry feeds is to match their composition with that of early life stages of
the species. Indeed, vitellogenin amino acid composition has been described for lumpsh eggs
(Yao and Crim, 1996) and proximate carcass analysis for juveniles (Sayer et al., 2000). Further
research eort could yield a range of specic feeds for lumpsh hatcheries, particularly as the
sector increases in scale.
Fig. 6.5 C. lumpus. Larval hatchlings and juveniles (not to
scale). Credit: Davenport, 1985 after Cox, 1920
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112 ♦ Biology and rearing of wrasse and lumpsh
6.16 Adults
Davenport (1985) provides an overview on the length–weight relationship of lumpsh (Fig. 6.6)
and age of sexual maturity. Historic studies suggest that spawning lumpsh are at least four years
of age but most common at ve to seven years up to nine to ten years, while males in the North
Sea may attain sexual maturity one or two years earlier. More recent studies (using updated oto-
lith readings, age estimates and length frequencies) suggest that males may actually spawn for the
rst time at age two to three, and females at age three to four in the North Sea, Norway, Iceland
and Greenland (Albert et al., 2002; Hedeholm et al., 2014).
O Greenland, the maximum total production rate (somatic and gonadal tissue) for males
and females was 0.47 and 0.92 kg wet weight per year, attained at 20 and 32 cm total length
respectively. For both sexes, somatic production declined steeply after the onset of matura-
tion (Hedeholm et al., 2014; Kasper et al., 2014). e anatomy of the gonads is described by
Davenport and Lönning (1983). In mature, ripe females, up to two-thirds of the visceral cavity
contains pink roe. e ovaries and oviducts are fused to form a single sac that is strongly bifur-
cated anteriorly, although the left horn is reduced to allow space for other organs. e dorsal
and dorso–lateral portions of the ovary-oviduct are lined by a thick, viscous layer that contains
whiteish opaque, unripe eggs in most females. Ripe eggs, which are clear and rose pink, fall from
this layer into the lumen of the structure, which is lled with 200–500 ml protective ovarian
uid. Egg numbers (100,000 to 400,000 per female) vary with the overall size of the sh, and
geographic population. e gonads of the male lumpsucker are simple and unremarkable white
structures.
Mortality appears to be size dependent. In Icelandic populations orsteinsson (1983) showed
that mean length increased with age until a certain length interval (c. 42–44 cm in Icelandic
females). ereafter, mean length decreased with age, indicating death of larger sh. Other than
shing pressure, adults are predated upon by dierent animals depending on their location.
During pelagic stages there is evidence of predation by sharks, seals and sperm whales. In shal-
low water during spawning season, males (which have a longer residency), are taken by gulls, sh
eagles and otters (see Davenport, 1985 for an overview). orsteinsson (1983) also suggested
that reproduction becomes progressively more expensive with increasing size, and that eventually
Fig. 6.6 C. lumpus. Length–weíght relationship of adult
lumpsuckers from Newfoundland waters. Credit: Davenport,
1985 after Cox, 1920
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Review of lumpsh biology ♦ 113
large sh do not recover from spawning (i.e. gonad production, spawning migration and the
associated period of prolonged starvation). Davenport and Kjörsvik (1986) suggested that spent
female sh might be positively buoyant (because the eggs are much denser than seawater) and
more vulnerable to predation. is is probably more likely for larger females following spawning.
6.17 Growth rate in the wild and hatcheries
e itinerant nature of adult lumpsh outside the breeding season makes growth assessments
challenging, with tagging experiments highlighting some limitations of this approach in the
species (e.g. Kaspar et al., 2014). Archive data on tagging, summarised by Davenport (1985)
suggests that large adults show a very small linear growth increment of a few centimetres after
about one year between tagging and recapture. Juveniles, however, are thought to possess rapid
allometric growth prior to leaving surface waters and follow a LW relationship of W = 8.7L3.36
x 10-6, (Newfoundland) and W = 2.309L3.053 (Norway; L in mm) with 55 mm length juveniles
thought to be about one year old (Myrseth, 1971; Daborn and Gregory, 1983). A long-term
study of wild juveniles found in summer rockpools by Moring (2001) reported that average
lengths increased by 23–43% and wet weight by 280–342% per month. Studies of both wild
and cultured larval and juvenile lumpsh show a rapid increase in growth rate from mid-July
to August, before decreasing in August–September (Benfey and Methven, 1986; Moring, 2001;
Ingolfsson and Kristjansson, 2002).
For adults, the largest reported lumpsh was an individual of body mass 9.5 kg, (maximum
total length 70 cm, maximum age of 14 years). However, the bulk of spawning females caught
in commercial gillnets are between 35 and 50 cm in length (roughly 2–5 kg) with slow growth
after maturity (Bagge, 1964; orsteinsson, 1983; Davenport, 1985).
Rearing studies started in the mid-1980s and attempted to culture larvae from the egg, oer-
ing dry (Benfey and Methven, 1986) and live (Brown, 1986) Artemia spp. to hatchlings. After
experimental periods of approximately one month, larvae reached a standard length of c. 12
mm on live feed, compared to 7 mm on dry feed. Brown et al. (1992) grew lumpsh from eggs
and reported a growth of 29 g after one year, and 510 g after two years at ambient temperatures
(between -1.8 and 14°C), although survival had declined to 0.5% after the rst year. Lumpsh
cultured in the laboratory became sexually mature at the end of their second year.
Sayer et al. (2000) attempted on-growing of wild caught juveniles, most success being achieved
by use of a low lipid diet formulated specically for the species (higher lipid diets allowed rapid
growth, but fat accumulated in brain and liver, causing early mortality). For sh under 350 g in
weight, a SGR of up to 3.8% per day was recorded, although the temperature varied between
6.1°C and 15.4°C during the on-growing period. e most recent studies of juvenile growth
have been derived from juveniles originating from eggs reared in-house and weaned on to appro-
priate dry feeds (Nytro, 2013; Nytro et al., 2014). e highest growth rates were comparable to
those reported by Sayer et al. (2000), and were observed for juveniles under 120g in warmer 13
and 16°C treatments, which resulted in an SGR of 3.65 and 3.60% per day. However, the eect
of rearing temperature on juvenile growth was reduced in size grades over 120 g. For lumpsh
deployed in sh pens at sea, a recent study has shown dierences in growth rate between juveniles
originating from dierent families (c. 1.2–2.1%/day; Imsland et al., 2016a).
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114 ♦ Biology and rearing of wrasse and lumpsh
Although there is likely genetic variation between families (Imsland et al., 2016a), given opti-
mal temperature, there is a high growth potential for juveniles (i.e. potentially increasing their
weight by an order of magnitude in under 80 days), thus allowing animals to reach deployable
size (10–15 g) within months, and achieving some annual synchrony with commercial salmon
farms.
6.18 Diseases and health implications
Maintaining high health status of all life stages of lumpsh is key during hatchery operations
to promote welfare and to attain numbers of deployable juveniles, and is particularly important
given the goal of closing the life cycle in captivity. Furthermore, understanding the susceptibility
of lumpsh to notiable diseases (or the potential to act as ‘carriers’) and thus onward transfer
to salmonid species during deployment, is also of clear importance to understand and manage
potential risks (Murray, 2016).
Lumpsh are a relatively novel candidate for aquaculture, with diseases and parasite records
found in academic literature (summarised in Powell et al., 2017), but also increasingly from ‘grey’
literature (such as veterinary reports) as cultured stock present with symptoms. e knowledge
of lumpsh diseases has increased in recent years, alongside research into the species immune
system that may foster vaccine development (e.g. Rønneseth et al., 2015). e topic will be con-
sidered in further detail in Chapter 13.
Briey, general (and treatable or preventable) pathogenic and non-pathogenic diseases in
hatcheries include ciliates (Trichodinia sp.), ukes (Gyrodactylus sp.), fungi (Exophiala sp), cata-
racts and post-capture stress (physical handling, transport, barotrauma, fatty deposits around
organs with high lipid diets) have been recorded in adults and reared juveniles (Scyborska, 1948;
Dawit, 2000; Sayer et al., 2000; personal observation).
Whilst parasitic worms (Rolbiecki and Rokicki, 2008) and myxosporeans (Cavin et al., 2012;
Kristmundsson and Freeman, 2014) have been recorded in wild lumpsh, endemic diseases are
likely more damaging. e fungal microsporidian Nucleospora cyclopteri (Mullins et al., 1994)
has been found at high (c. 25%) prevalence in wild adults (Freeman et al., 2013) although there
is currently no conrmed route of transmission or eective treatment. Lumpsh also appear to
suer from a high prevalence (61–100%) of sea lice such as Caligus elongatus (Boxshall, 1974;
Heuch et al., 2007). More recently, amoebic gill disease (Perry and Treasurer, 2015), pasteurel-
losis and other bacterial infections (e.g. Alarcon et al., 2015; Smage et al., 2016), and viral dis-
eases (Towers, 2015) have been detected.
6.19 Summing up: gaps in knowledge and research needs
Knowledge of the biology of lumpsh has advanced greatly over the last few years in response to
the needs of the new cleaner sh industry, but information on many critical areas is still missing.
A detailed gap analysis is provided in Powell et al. (2017) but in brief, the ultimate objective is to
produce disease-free juveniles that adapt well in captivity, do not pose a risk to salmon or other
shes, and are ecient at delousing.
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Review of lumpsh biology ♦ 115
To work toward this goal, the production of lumpsh needs to be closed in captivity, without
dependence on wild broodstock. is will require better knowledge of articial reproduction,
particularly with respect to control of maturation, gamete collection and storage. Commercial
production will need to be derived entirely from farmed strains, and this will require the devel-
opment of a genetic breeding programme, one that can produce elite lines with superior perfor-
mance and desirable traits, including disease resistance and delousing eciency. Targeting slow
growing lumpsh strains may be advantageous in order to prolong the time lumpsh can graze
on sea lice. Larval production also needs to be optimised, along with improved diets to help
reduce high post-weaning mortality. Vaccines, as well as more eective therapeutants, are also
required to combat emerging infectious diseases. Finally, reuse of deployed lumpsh needs to be
considered, perhaps through their use as broodstock, in animal feeds, or for human consumption.
Acknowledgements
e authors would like to thank an anonymous reviewer for constructive comments during the
preparation of this chapter.
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