Lipid-rich copepods form an essential link between primary producers and higher trophic levels in high-latitude oceans. These zooplankton can take advantage of ephemeral phytoplankton blooms to fuel development and reproduction. However, we have limited understanding of how the physiological condition of these animals varies in relation to environmental factors such as food availability. Due to high advection, it is likely that physiological plasticity, rather than local adaptation, is primarily responsible for physiological differences within a region. We use transcriptomics and other physiological metrics to understand how two species of copepods (Calanoides acutus and Calanus propinquus) vary across environmental gradients along the West Antarctic Peninsula. For the primarily herbivorous C. acutus, physiological separation between sampling locations appears to be driven by feeding status, and gene expression differences indicate differential expression of genes regulating lipid metabolism, reproduction, aerobic metabolism, and protein translation. For the more omnivorous C. propinquus, physiology and gene expression did not segregate as clearly by location, showed minimal signs of food deprivation at any location, and had a weaker relationship with chlorophyll compared to C. acutus. By comparing these results with concurrent starvation experiments, we find that spatial variation in gene expression reflects short-term differences in food availability (particularly for C. acutus), and we identify genes whose expression indicates recent feeding status. Further examination of the relationships between food availability, copepod physiology, and population dynamics will ultimately improve our capacity to predict how copepod populations will respond to rapidly changing environmental conditions in the West Antarctic Peninsula ecosystem.

Respiration of lipids by copepods during diapause (overwintering dormancy) contributes to ocean carbon sequestration via the seasonal lipid pump (SLP). Parameterizing this flux in predictive models requires a mechanistic understanding of how life history adaptation in copepods shapes their timing of exit from diapause. We investigate the optimal phenology of Calanus finmarchicus in the Norwegian Sea using an individual-based model in which diapause exit is represented as a trait characterized by phenotypic mean and variance. Without interannual variability, optimal exit correlated with the onset of the spring phytoplankton bloom and phenotypic variance was of no benefit. In contrast, copepods endured reduced fitness and adopted bet-hedging strategies when exposed to interannual variability in bloom timing and predation: later exit from diapause and phenotypic variance maintained adult numbers in anomalous late-bloom years. Exit nevertheless remained well before the peak of the bloom which is a favorable strategy when low predation early in the year enhances survival of eggs and early developmental stages. Our work highlights the complex interactions between C. finmarchicus and its environment and the need for improved understanding of bet-hedging strategies and the cues of diapause exit to progress the representation of the SLP in global biogeochemical models.

Climate change may exacerbate the impact of invasive parasites from warmer climates through pre-existing temperature adaptations. We investigated temperature impacts on two closely related marine parasitic copepod species that share the blue mussel (Mytilus edulis) as host: Mytilicola orientalis has invaded the system from a warmer climate <20 years ago, whereas its established congener Mytilicola intestinalis has had >90 years to adapt. In laboratory experiments with temperatures 10-26°C, covering current and future temperatures as well as heat waves, the development of both life cycle stages of both species accelerated with increasing temperature. In the parasitic stages, the growth of the established invader increased evenly from 10°C to 22°C, whereas the recent invader barely grew at all at 10°C and grew faster already at 18°C. In contrast, temperature had little effect on the transition success between life cycle stages. However, the highest temperature (26°C) limited the egg development success of the established invader and the host entry success of both species, whereas the infection success of the established invader increased at 18°C and 22°C. In general, our experiments indicate that the main effect of temperature on both species is through development speed and not life cycle stage transition success. Based on regional long-term temperature data and predictions, the numbers of completed life cycles per year will increase for both parasites. The established invader seems better adapted for low current temperatures (around 10°C), whereas the more recent invader barely develops at these temperatures but can cope in high temperatures (around 26°C). Hence, pre-existing temperature adaptations of the recent invader may allow the species to better cope with heat waves.

Having been successfully bred in semi-intensive and intensive aquaculture systems, oval squids of the Sepioteuthis lessoniana species complex are emerging as promising candidates for research and industry. Nevertheless, information about pathogens and diseases that may affect squid aquaculture remains sparse. In this study, we identify new parasitic copepod species that causes squid mortality and decreases squid hatching rates, and we also offer a solution to eliminate the pathogen during incubation of squid eggs. The newly discovered copepod Ikanecator primus gen. et sp. nov. was identified on oval squid eggs for the first time using both morphological and molecular diagnostic markers. In the genomes of the copepod and associated microbiome, we identified multiple genes for enzymes involved in cephalopod eggshell degradation in genomes of the copepod and associated microbiome. Furthermore, we conducted experiments to assess efficacy of peracetic acid in inhibiting the I. primus gen. et sp. nov. both in vitro and in vivo using immersion treatment. We established that a 2-min exposure to a concentration of 250 μl/L of peracetic acid containing product (PAA-product; 35 mg/L PAA and 15 mg/L H2O2) inhibited the development of nauplii in vitro. All parasites exposed to a concentration of 500 μl/L of PAA-product (70 mg/L PAA and 30 mg/L H2O2) were eliminated within two minutes. On top of this, the immersion treatment with 500 μl/L of PAA-product (70 mg/L PAA and 30 mg/L H2O2) improved survival of squid embryos and increased size of squid hatchlings compared with control and the immersion treatment with 125 μl/L of PAA-product (17.5 mg/L PAA and 7.5 mg/L H2O2) and the immersion treatment with 250 μl/L of PAA-product (35 mg/L PAA and 15 mg/L H2O2). These findings suggest that PAA holds a great potential as inhibitor and controller of parasitic copepod infections and for overall health management in cephalopod culture.

Fast-sinking zooplankton carcasses and fecal pellets appear to contribute significantly to the vertical transport of particulate organic carbon (POC), partly because of low temperature that decreases microbial degradation during the descent into the deep ocean. Increasing hydrostatic pressure could further reduce the degradation efficiency of sinking POC, but this effect remains unexplored. Here, the degradation of carcasses and fecal pellets of the abundant marine copepod Calanus finmarchicus was experimentally studied as a function of pressure (0.1-100 MPa). Samples were either exposed to elevated pressure in short 1-day incubations or a gradual pressure increase, simulating continuous particle sinking during a 20-day incubation. Both experiments revealed gradual inhibition of microbial respiration in the pressure range of 20-100 MPa, corresponding to 2-10-km depth. This suggests that hydrostatic pressure impedes carbon mineralization of fast-sinking carcasses and fecal pellets and enhances the deep-sea deposition rate of zooplankton-derived organic material.

The copepod family Shiinoidae Cressey, 1975 currently comprises nine species of teleost parasites with unusual morphology and a unique attachment mechanism. Female shiinoids possess greatly enlarged antennae that oppose a rostrum, an elongate outgrowth of cuticle that originates between the antennules. The antennae form a moveable clasp against the rostrum which they use to attach to their host. In this study, we use micro-computed tomography (microCT) to examine specimens of Shiinoa inauris Cressey, 1975 in situ attached to host tissue in order to characterize the functional morphology and specific muscles involved in this novel mode of attachment and to resolve uncertainty regarding the segmental composition of the regions of the body. We review the host and locality data for all reports of shiinoids, revise the generic diagnoses for both constituent genera Shiinoa Kabata, 1968 and Parashiinoa West, 1986, transfer Shiinoa rostrata Balaraman, Prabha & Pillai, 1984 to Parashiinoa as Parashiinoa rostrata (Balaraman, Prabha & Pillai, 1984) n. comb., and present keys to the females and males of both genera.

Climate change is resulting in increasing ocean temperatures and salinity variability, particularly in estuarine environments. Tolerance of temperature and salinity change interact and thus may impact organismal resilience. Populations can respond to multiple stressors in the short-term (i.e., plasticity) or over longer timescales (i.e., adaptation). However, little is known about the short- or long-term effects of elevated temperature on the tolerance of acute temperature and salinity changes. Here, we characterized the response of the near-shore and estuarine copepod, Acartia tonsa, to temperature and salinity stress. Copepods originated from one of two sets of replicated >40 generation-old temperature-adapted lines: ambient (AM, 18°C) and ocean warming (OW, 22°C). Copepods from these lines were subjected to one and three generations at the reciprocal temperature. Copepods from all treatments were then assessed for differences in acute temperature and salinity tolerance. Development (one generation), three generations, and >40 generations of warming increased thermal tolerance compared to Ambient conditions, with development in OW resulting in equal thermal tolerance to three and >40 generations of OW. Strikingly, developmental OW and >40 generations of OW had no effect on low salinity tolerance relative to ambient. By contrast, when environmental salinity was reduced first, copepods had lower thermal tolerances. These results highlight the critical role for plasticity in the copepod climate response and suggest that salinity variability may reduce copepod tolerance to subsequent warming.

Salmon louse (Lepeophtheirus salmonis) is a skin- and blood-feeding ectoparasite, infesting salmonids. While feeding, labial gland proteins from the salmon louse may be deposited on the Atlantic salmon (Salmo salar) skin. Previously characterized labial gland proteins are involved in anti-coagulation and may contribute to inhibiting Atlantic salmon from mounting a sufficient immune response against the ectoparasite. As labial gland proteins seem to be important in the host-parasite interaction, we have, therefore, identified and characterized ten enzymes localized to the labial gland. They are a large group of astacins named L. salmonis labial gland astacin 1-8 (LsLGA 1-8), one serine protease named L. salmonis labial gland serine protease 1 (LsLGSP1), and one apyrase named L. salmonis labial gland apyrase 1 (LsLGAp1). Protein domain predictions showed that LsLGA proteins all have N-terminal ShK domains, which may bind to potassium channels targeting the astacins to its substrate. LsLGA1 and -4 are, in addition, expressed in another gland type, whose secrete also meets the host-parasite interface. This suggests that LsLGA proteins may have an anti-microbial function and may prevent secondary infections in the wounds. LsLGAp1 is predicted to hydrolyze ATP or AMP and is, thereby, suggested to have an immune dampening function. In a knockdown study targeting LsLGSP1, a significant increase in IL-8 and MMP13 at the skin infestation site was seen under LsLGSP1 knockdown salmon louse compared to the control, suggesting that LsLGSP1 may have an anti-inflammatory effect. Moreover, most of the identified labial gland proteins are expressed in mature copepodids prior to host settlement, are not regulated by starvation, and are expressed at similar or higher levels in lice infesting the salmon louse-resistant pink salmon (Oncorhynchus gorbuscha). This study, thereby, emphasizes the importance of labial gland proteins for host settlement and their immune dampening function. This work can further contribute to anti-salmon louse treatment such as vaccine development, functional feed, or gene-edited salmon louse-resistant Atlantic salmon.

Organismal thermal limits affect a wide range of biogeographical and ecological processes. Copepods are some of the most abundant animals on the planet and play key roles in aquatic habitats. Despite their abundance and ecological importance, there is limited data on the factors that affect copepod thermal limits, impeding our ability to predict how aquatic ecosystems will be affected by anthropogenic climate change. In a warming ocean, one factor that may have particularly important effects on thermal limits is the availability of food. A recently proposed feedback loop known as \"metabolic meltdown\" suggests that starvation and exposure to high temperatures interact to drastically reduce organismal thermal limits, increasing vulnerability to warming. To investigate one component of this feedback loop, we examined how starvation affects thermal limits (critical thermal maxima: CTmax) of Acartia tonsa, a widespread estuarine copepod. We found that there was no effect of short-duration exposure to starvation (up to 2 days). However, after 3 days, there was a significant decrease in the CTmax of starved copepods relative to the fed controls. Our results provide empirical evidence that extended periods of starvation reduce thermal limits, potentially initiating \"metabolic meltdown\" in this key species of coastal copepod. This suggests that changes in food availability may increase the vulnerability of copepods to increasing temperatures, amplifying the effects of climate change on coastal systems.

The clade Pancrustacea, comprising crustaceans and hexapods, is the most diverse group of animals on earth, containing over 80% of animal species and half of animal biomass. It has been the subject of several recent phylogenomic analyses, yet relationships within Pancrustacea show a notable lack of stability. Here, the phylogeny is estimated with expanded taxon sampling, particularly of malacostracans. We show small changes in taxon sampling have large impacts on phylogenetic estimation. By analyzing identical orthologs between two slightly different taxon sets, we show that the differences in the resulting topologies are due primarily to the effects of taxon sampling on the phylogenetic reconstruction method. We compare trees resulting from our phylogenomic analyses with those from the literature to explore the large tree space of pancrustacean phylogenetic hypotheses and find that statistical topology tests reject the previously published trees in favor of the maximum likelihood trees produced here. Our results reject several clades including Caridoida, Eucarida, Multicrustacea, Vericrustacea, and Syncarida. Notably, we find Copepoda nested within Allotriocarida with high support and recover a novel relationship between decapods, euphausiids, and syncarids that we refer to as the Syneucarida. With denser taxon sampling, we find Stomatopoda sister to this latter clade, which we collectively name Stomatocarida, dividing Malacostraca into three clades: Leptostraca, Peracarida, and Stomatocarida. A new Bayesian divergence time estimation is conducted using 13 vetted fossils. We review our results in the context of other pancrustacean phylogenetic hypotheses and highlight 15 key taxa to sample in future studies.