While the species concept and a single unequivocal definition of species remains a topic of discussion, most regard reproductive incompatibility between two groups of individuals as irrefutable evidence of separate species. Thus, reproductive incompatibility between L. salmonis from the Pacific and Atlantic would support the case for separating them into separate species as has been previously suggested based upon their COI genetic diversities . In the present study, we successfully produced F1 and F2 hybrid L. salmonis strains using maternal lines from both the Pacific and the Atlantic. Previous studies of separate conspecific populations of copepods have shown that although defined as single species, the populations may or may not be reproductively compatible [36–38]. In studies of the intertidal copepod Tigriopus californicus it has been shown that “Dobzhansky-Muller incompatibilities” between strains, i.e. deleterious introductions of genetic variants into new genetic backgrounds, may result in reduced fitness in the F2 hybrids [39, 40]. Although the present study was primarily designed to qualitatively investigate the reproductive compatibility of Atlantic and Pacific L. salmonis, egg string hatchability and survival between infection and the pre-adult stage was quantified for the two hybrid strains in the F2 generation. The observed values (Table 1) fall within the range of values for these parameters in this salmon louse rearing facility [30, 41–43]. The infectivity and survival of Pacific L. salmonis reared in our facilities was not accurately quantified but did not appear to deviate from levels observed for Atlantic strains. Therefore, there are no data suggesting loss of fitness due to hybridization between Pacific and Atlantic L. salmonis up to and including the F2 hybrid generation. The genetic mtCOI divergence between T. californicus populations varies from 0.2% - 23% and the level of fitness reduction is correlated with the genetic divergence . The genetic divergence between Atlantic and Pacific L. salmonis varies between 4.8 and 7.1%  and is thus limited when compared to the differences reported for T. californicus. This may suggest that outbreeding depression in hybrids between L. salmonis from the Pacific and Atlantic oceans may be expected to be limited if present, and could have gone undetected in the present study. It is therefore concluded that Pacific-Atlantic hybrid fitness should be accurately quantified in future studies, but that L. salmonis from the Pacific and Atlantic are reproductively compatible at least until the F2 hybrid generation. Therefore, our results do not support separating Atlantic and Pacific L. salmonis into separate species.
The COI gene has been sequenced for a large number of species and is currently being used for species identifications via DNA barcoding [45, 46]. Looking at congeneric species (species within the same genus) across a wide range of taxa, 98% of species-pairs display COI divergence of 2% or more, and the average divergence across all taxa is 11.3% . Within Crustacea, the average diversity among congeneric species has been estimated to 15.4% . Looking specifically at copepods, congeneric COI sequence diversity was estimated between 13-22% . The calculated divergence for COI between L. salmonis from the Pacific and Atlantic (4.8-7.7%) [16, 17] is lower than what is typical for congeneric copepod species, but at the upper boundary of the reported variation of 1.3-7.9% within a crustacean species (i.e. between individuals of the same species) . However, the congeneric calanoids Calanus glacialis and C. finmarchicus, with a reported COI sequence divergence of 20% , produce hybrids capable of successful reproduction . Therefore, reproductive compatibility among Pacific and Atlantic L. salmonis may not be unexpected. Taken together, the reported sequence divergence of L. salmonis in the Pacific and the Atlantic oceans, which is well below the variations reported between congeneric copepods, and the evidence of reproductive compatibility with no apparent loss of fitness presented here, suggests that L. salmonis from the Pacific and Atlantic do not represent separate species.
While the genetic diversity within COI and 16S between L. salmonis from the Pacific and Atlantic is below that expected between species of copepods, it remains highly significant as illustrated by the phylogenetic analysis. This demonstrates minimal or non-existent genetic exchange between these allopatric L. salmonis components on a contemporary time-scale. This is in contrast to the very high level of gene-flow and lack of population differentiation reported among L. salmonis sampled from geographically distinct regions within each ocean [8, 10, 11, 17]. Stable barcoding gene sequence divergences between morphologically and geographically defined subspecies, below the divergence level expected between congeneric species, has previously been used to confirm subspecies validity [50, 51]. Lepeophtheirus salmonis from the Pacific and Atlantic exhibit morphological [20, 21] and apparent biological differences [24, 25, 27] in addition to considerable genetic sequence divergence . Despite these differences, Atlantic and Pacific L. salmonis are reproductively compatible and exhibit sequence divergence below the level typically observed between congeneric copepods and at the extreme upper boundary of the range found among conspecific crustaceans . Therefore, we suggest that Atlantic and Pacific L. salmonis should be regarded as two subspecies: Lepeophtheirus salmonis salmonis and Lepeophtheirus salmonis oncorhynchi subsp. nov. urn:lsid:zoobank.org:pub:8CABA147-FB3D-4DFA-AF37-7FFB74AFB454.
Due to the interaction of salmon lice with farmed and wild salmonids in both the Pacific and Atlantic, L. salmonis represents both an ecologically and economically significant parasite. In accordance with this is the growing volume of scientific studies investigating this parasite. The differences between L. salmonis in the two oceans make it crucial to correctly assign scientific results to the correct component of the species, which is presently taxonomically impossible. The proposed division is therefore well reasoned and will facilitate appropriate taxonomic indication of origin in the future.
Taxonomic summary of Lepeophtheirus salmonis oncorhynchisubsp. nov
Species: Lepeophtheirus salmonis oncorhynchi subsp. nov.
Etymology: ‘oncorhynchi’ reflecting the evolutionary association with the salmonid genus Oncorhynchus in the Pacific Ocean
Descriptions: Johnson and Albright [20, 22]
Holotype sequences: Lepeophtheirus salmonis oncorhynchi COI and 16S [GenBank:KF278676]
Museum specimens: Adult female ZMUB91335 (holotype, ethanol), adult male ZMUB91339 (allotype, karnovsky), paratype series, 3 specimens (ZMUB91336 – ZMUB91338, ethanol).