Sea Lice Working Group Report

The collective term “sea lice” is colloquially used to refer to numerous species of copepod crustaceans of the family Caligidae that are externally parasitic on the skin of marine and anadromous fishes. The most intensively studied species ‐ Lepeophtheirus salmonis ‐ is, as its specific name implies, a specialist parasite of salmonid fishes. It is commonly associated with a total of 12 host salmonid fish species of the genera Salmo, Oncorhynchus and Salvelinus in the Pacific and Atlantic Oceans. Along the Pacific coasts of Alaska and British Columbia, L. salmonis as well as Caligus clemensi and Lepeophtheirus cuneifer (both host generalist lice species) have been recorded on wild and farmed salmonids. In British Columbia, Chile and Tasmania Atlantic salmon (Salmo salar) is the principal salmonid species in culture. While the Tasmanian industry apparently suffers no especial problems from caligid infestation, the Chilean industry has been heavily impacted by Caligus species, initially C. teres but more recently and significantly C.\nrogercresseyi (both of which are host generalists). In Japan, Caligus orientalis is the most pathogenic sea louse on cultured Pacific salmon, although L. salmonis also remains a problem. L. salmonis is associated with wild chum and pink salmon in Japan, but also infests cultured coho salmon and rainbow trout. C. orientalis – like C. elongatus in the North Atlantic – is a host generalist; C. orientalis occasionally impacts salmonids, and it is an especial problem to cultured rainbow trout.\nThe current scientific literature refers to Lepeophtheirus salmonis Krøyer as occurring on salmonids in both the North Pacific and North Atlantic Oceans. At first sight it might appear curious that the same species should occur in two separate and geographically distant oceans, but there is convincing geological, molecular and ecological evidence of past trans‐Arctic connectivity of the marine fauna of the North Pacific and North Atlantic basins — and specifically of Pacific species having tended to colonize the North Atlantic rather than vice versa — following the recent opening of the Bering Strait (~5 million years ago). The presently available molecular (DNA) results cannot provide conclusive evidence regarding the specific status of Pacific and Atlantic L. salmonis, but recent DNA sequence analyses do indicate clear\ngenetic distinction between the Atlantic and Pacific lineages. Similarly, there now is a body of mitochondrial DNA sequence and ecological (host association) data indicating that the host generalist parasite, Caligus elongatus, actually comprises taxonomically separable entities. It is, however, too early to affirm that there are definitely two (or more) species of either “C. elongatus” or “L. salmonis”; for clarity and consistency with the contemporary scientific literature we continue in the present report to refer to single species in both cases.\nAll female caligids undergo internal fertilization of the eggs prior to their extrusion into a pair of external egg sacs or “eggstrings”. The nauplius I is the hatching stage and at this point the eggstring disintegrates and the larvae are released to become planktonic. The nauplius I molts into a nauplius II and then again into the infective copepodid stage. None of the three planktonic stages feed; all the reserves the larvae require to complete development to the infective copepodid are provided by the parent female. Initial attachment for the copepodid typically occurs on the fins of the host fish (especially the dorsal, ventral and anal fins) or to the scales. Several chalimus stages follow, attached to the host by a sort of tether. Depending upon the species of caligid there then may be two, one, or no pre‐adult stages between chalimus IV and the mature adult. The pre‐adult and adult stages are all mobile, i.e., able to move about on the host fish’s body.\nWhen large numbers of farmed salmon are introduced to the marine environment in open net cage salmon farms, three things are virtually inevitable for these fish:\n they will become hosts to sea lice (Lepeophtheirus spp. and/or Caligus spp.) since these occur naturally on wild host species in the vicinity of most farms;\n they will become part of a dynamic host‐parasite system involving wild hosts, because they can produce large numbers of infective larvae in a restricted spatial area if gravid females are allowed to develop; and\n because they carry sea lice, and because some of these fish may escape from the farms, the dispersal of parasites is likely to be even more widespread on occasion. Given the above, it may be concluded that it is next to impossible to (1) avoid infection of farmed fish, all of which go into the pens as clean smolts, and (2) also subsequently avoid infection of wild fish that are found in the vicinity (“infective field”) of an open cage farm.\nA parasitic infection becomes a disease when host behavior and physiology (and ultimately host health, survivorship and fitness) are altered or compromised to an exceptional extent. For example, the increased metabolic demand exerted by the parasite may cause slower host growth, making the wild fish more likely to be captured by predators, or conversely causing them to take greater risks to feed, with the same end result. Reduced host condition also can affect swimming ability, with several negative ecological consequences ranging from reduced competitive ability to slower migration. Slower migration rates through coastal waters might elevate the risk of infestation by sea lice copepodids. Skin damage caused by the feeding behavior of sea lice can increase the physiological cost of osmotic regulation, or provide sites for secondary bacterial or fungal infection. Contrary to typical perceptions that it is not “in the interest” of parasites to kill their host, it is worth noting that sufficiently high sea lice loads will kill individual wild fish, but the definition of “high” will depend on sea louse stage, fish size and developmental stage. While not exhaustive, this list illustrates some of the direct and more subtle indirect ways that sea lice may cause disease, as defined above.\nSea lice abundance on farmed salmon only rarely attains levels where the health or welfare of these fish is negatively affected. When this occurs there are legal and regulatory requirements in many countries that demand treatment, but it is clearly also in the economic interests of the farmer to treat the infection promptly and effectively. For wild fish, disease is likely to be an issue whenever sea lice intensity on individual hosts is sufficiently high as to cause significant stress, or to increase their vulnerability to secondary pathological infection or other mortality agents, as noted above. For example, newly‐migrated smolts exposed to the challenge of osmoregulating in saline waters will be physiologically stressed by that environmental challenge and will be more vulnerable than post‐smolts that are older and fully adapted to seawater.\nSea lice disease of wild salmonids is potentially problematic in areas with intensive Atlantic salmon aquaculture in British Columbia, Canada, on the west coasts of Ireland and Scotland, and throughout Norway (Chile lacks endemic species of anadromous salmonids). In British Columbia, the focus of attention has been on the much studied and discussed Broughton Archipelago region, where there is particular concern regarding the impact of Lepeophtheirus salmonis on wild stocks, particularly juvenile pink (Oncorhynchus gorbuscha) and chum (O. keta) salmon.\nUnlike Atlantic salmon, sea trout (the anadromous form of the brown trout, Salmo trutta) spend extended periods of time in nearshore or coastal waters, and this feature may render them particularly vulnerable to sea lice infestation. As has been the case in British Columbia, analyses concerning the potential interaction between farmed and wild salmonids in Irish bays, Scottish sea lochs and Norwegian fjords subject to intensive aquaculture have not been without controversy. The circumstantial evidence of farm‐produced larval sea lice contributing to parasite loadings on wild sea trout in Ireland is considerable. Correlations have been drawn between abundances of sea lice on wild sea trout and on Irish farms up to 30 km distant. As is the case for sea trout, Arctic charr (Salvelinus alpinus) are effectively confined to coastal waters (often in narrow fjord systems) and these areas commonly are home to a high density of captive farmed salmon. Sea trout and Atlantic salmon are also the species of most concern with regard to detrimental effects of sea lice in Norway, although Arctic charr also are impacted by these parasites there.\nIt has been demonstrated that salmon within a given farm site can be self‐reinfesting (because hatched nauplii drift back into the net pens having completed their development to the infective copepodid stage); it is also intuitive that nauplii exported from one farm site will infect salmon being grown in neighbouring farms or free‐ranging wild fish in the vicinity. Similarly, wild fish may well infect other wild fish or, if they are resident in coastal waters, adjacent farmed fish. The absolute abundances of farm and wild fish, the absolute abundances of sea lice on those fish and the relative strengths of farm‐farm and farm‐wild interactions (and any seasonal or annual variation thereof) will determine the overall infestation pressure on individual fish in a given locality.\nIt is far easier to monitor and assess the outcome of interventory treatment for sea lice infestations for farmed fish than it is for wild fish, and there are potentially many more strategies available to control sea lice on farmed fish. For example, in addition to medicinal treatments, a variety of management (and even informed environmental) decisions can be made which can have impacts on the control of sea lice on farms. The challenges in man

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