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Ocean temperature
The independent bottom water temperature estimates all showed variable but increasing autumn temperatures from 1996 to 2010, followed by a decline through to 2018 in the mid-depth offshore stations and the survey, but less of a decline in the inshore or deep offshore stations (Fig. 1, 2a). Both deep offshore stations warmed by ~ 0.2 °C over the time series. The ~ 1 °C temperature range evident in the southwestern mid-depth offshore FX8 autumn hydrographic series was larger than that of the less variable northern mid-depth offshore SI7 hydrographic station, but neither of the mid-depth offshore stations showed any net warming over the 22-year period. The shallow inshore stations both warmed considerably (1.5–2 °C) over the time series. Although no net increase was apparent in either of the mid-depth offshore hydrographic station autumn time series, 0.25–0.40 °C increases were observed in both the winter and spring measurement time series at both mid-depth hydrographic stations.
Figure 1

Map of 245 autumn groundfish survey core stations (black dots) sampled around Iceland every October between 1996 and 2018 (except 2011). Bottom-temperature profiles were collected independently at hydrographic stations FX3, FX8 and FX9 (red triangles) and SI1, SI7 and SI8 (blue triangles). Depth contours are 200, 500 and 1000 m.

Full size image

The mean environmental temperature from the autumn survey (Tempe) lay midway between that of the two mid-depth offshore hydrographic stations, over a similar ~ 1 °C temperature range. Based on a fitted regression to the environmental temperature time series, there was a net warming of 0.33 °C across the survey area over the 22-year time period, although the increase was certainly not linear (Fig. 2c). There were large differences in both the temperature range and temperature-at-depth among some of the regions (Fig. 2b). Bottom water temperatures at 200 m depth were 3–4 °C warmer in the south (SW = 7.3 °C; SE = 8.2 °C) than in the north (NW = 5.4 °C; NE = 4.5 °C), and 4–5 °C warmer at a depth of 500 m (NW = 1.4 °C; NE = −0.5 °C; SW = 6.6 °C; SE = 3.9 °C). Although most stations deeper than 500 m in the NE were  3 °C.
Figure 2

Bottom-temperatures from hydrographic stations (A: left panel) and from the autumn groundfish survey (B: top right and C: bottom right panel). (A) Near-bottom temperatures at hydrographic stations FX3 and SI1 (~ 70 m), FX8 and SI7 (~ 400 m), and FX9 and SI8 (~ 1000 m) between 1996 and 2018 are shown for winter (January–March), spring (May–June) and autumn (October–December). A geometric smooth has been fitted to the annual means. Note different scales on the y-axes. (B) Mean depth-temperature profiles for each region based on bottom temperatures from the autumn survey. Fitted lines are loess regressions. (C) The mean annual environmental temperature time series estimated from a GLM of the autumn survey bottom temperature measurements (see text for details). The linear regression fitted to the annual means is intended only to show the overall rate of warming through the time series.

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Survey catches
A total of 5390 tows were fished over the 22-year time series. Station depths ranged from 26 to 1203 m (mean of 377 m) while bottom temperatures ranged between − 2 and 11 °C (mean of 4.85 °C). Of the 7,246,474 fish representing 200 species that were caught, 82 species were caught during at least 19 of the survey years, and thus are the primary subject of this analysis (Suppl. Table 1). The mean standardized annual abundance of these 82 species ranged between 1 and 27,513 fish, with an overall mean across species of 4096; 70% of the species had a mean standardized annual abundance of less than 100.
There was no clear identifier of species that were rare, and thus caught sporadically, versus those that were newly immigrated to the survey area in response to the warming environment. Nor was it possible to exclude the possibility that certain rare species were incorrectly identified in some survey years. However, there were 6 species that were never caught prior to 1998, yet appeared in increasing numbers in subsequent years (Table 1). All of these species were caught in at least 8 survey years, and as many as 19. However, three of the species were deepwater ( > 800 m), and thus unlikely to have been subject to temperature increases of > 0.2 °C. Of the remaining three species, Atlantic mackerel (Scomber scombrus), blackbelly rosefish (Helicolenus dactylopterus) and hollowsnout grenadier (Coelorinchus caelorhincus) were all warmwater (TB  > 2) and stenothermal (Steno  More

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