Representatives of the brachyuran family Varunidae occur in tropical to temperate regions throughout the world and currently include 150 identified species (Ng 2006). Varunid crabs are characterized by one of the most diverse habitat ranges, from freshwater to seawater and from shallow waters to the deep sea (Ng 2006; Ng et al. 2008). Some species occur in specific environments, such as anchialine caves and hydrothermal vents. Many species of Varunidae are known from only a few specimens due to their highly specialized habitats and often very cryptic habits (Ng 2006; Ng et al. 2008). According to Bos et al. (2016), only three species of Varunidae have been known so far from Dutch waters, namely
Samples were collected from 187 locations in different areas of the Dutch Exclusive Economic Zone – from Brown Bank, Dogger Bank, Frisian Front, Oyster Grounds and north of the Dutch Wadden Islands – in February and March 2019. Sampling was carried out by NIOZ, Eurofins AquaSense and eCOAST, commissioned by Rijkswaterstaat, using the vessel “Arca”.
All sites were sampled with a Triple-D dredge of NIOZ, which quantitatively samples larger, often more sparsely distributed and longer-lived in- and epifauna (Witbaard et al. 2013). At each sampling site, a strip of seabed was excavated to a depth of 20 cm (+/− 5 cm) below the sediment surface. The strip was 20 cm wide and about 100 m long (range within 90–110 m). The sediment was then washed through 5 × 5 mm meshes of a dredge case and a 7 mm stretched mesh net astern the dredge while towing the dredge through the sediment. If necessary, the contents of the dredge net were flushed over a 1 mm sieve on board to remove remaining mud and subsequently sorted on a table. Living organisms were sorted by species, counted and weighed. All organisms were subsequently discarded, except for three voucher specimens of each species at each of the sampling sites mentioned above and specimens unidentifiable on board, which were preserved in ethanol 70%. All dredge contents were examined by at least two analysts. Local field guides were used for identification, especially issues of the “Synopsis of the British Fauna”. Specimens that could not be identified on board were taken to the laboratory for identification using other literature sources. The specimen of
The study area, the areas of high ecological value and the sampling locations are shown in Figure 1. Soft sediments, sand and mud, dominate throughout the sampling area, while gravel, stones or boulders are extremely rare.
Brown Bank was visited as part of the sampling campaign. It is an area characterized by relatively high current velocities and sand waves in the north-south direction (van der Reijden et al. 2019). Brown Bank is located at the center of the southern North Sea (Fig. 1) and has an average depth of 32 m, with the shallowest part at 19 m. The ridges rise to approximately 20 m above the surrounding seabed. The sediment of Brown Bank consists of medium sand (Wentworth scale), with a median grain size varying between 250 and 300 μm, as well as clay and boulders between the sand ridges. Brown Bank is an area frequently visited by beam trawl vessels. The percentage of organic matter in the sediment is generally low and the oxidized sediment layer is roughly 20 cm thick (García et al. 2019). Large-scale sandbank structures, perpendicular sand waves and small sand waves are present on Brown Bank. Slopes of sand waves consist of finer sediments and sand wave valleys consist of coarser and more mixed sediments (van der Reijden et al. 2019).
At one sampling site on Brown Bank (site code 006_ BruineBank; dredge haul from 52°21′58.0788″N and 3°27′47.664″E to 52°21′55.5408″N and 3°27′51.3756″E; EPSG4258; average dredge haul depth of 32 m), one specimen of a crab species hitherto unknown from Dutch waters was collected on 24 March 2019 (Fig. 2). The sediment at this location was described during sampling as ‘sand with shell grit’. In the laboratory, a crab with a carapace width of 7.5 mm was identified as
The main diagnostic characters of this small crab are as follows (described by Bocquet 1963): (1) irregularly hexagonal carapace (about 1.6 to 1.8 times wider than long); (2) relatively small chelipeds; (3) the second and especially the fifth pereopods strongly reduced in size; (4) merus, carpus and propodus of the third and fourth pereopods clearly enlarged (while quite thin in juveniles); (5) in live specimens, the second to fifth pereopods are held between parallel tangents, running along the anterior and posterior boundaries of the carapace; and (6) the second to fourth pereopods with a distinct wine-red colored band in the distal half of the propodus. In our specimen, this colored band is present throughout the propodus and on the second and third pereopods also on the distal half of the carpus. This difference may be related to its smaller size, as Bocquet (1963) found that coloration changes across developmental stages.
Although not fully grown – the carapaces of the largest female and male found by Bocquet (1963) were respectively 14.5 mm and 12.9 mm wide, this crab was clearly a male specimen, as indicated by the narrow abdomen (Fig. 3). The propodus of the cheliped is enlarged, and the dactylus and finger of the propodus are relatively short as described for the male by Bocquet (1963).
Given the irregularly hexagonal carapace shape and the Atlantic distribution area of this small crab, we propose ‘Atlantisch zeshoekkrabbetje’ as the Dutch vernacular name. The single specimen of
The genus
Morphological differences between the three species of
Chronological distribution data and ecological data from the literature and our sampling
region | period | depth (m) | sediment | number | associates | source |
---|---|---|---|---|---|---|
Gold Coast (= Ghana) to Roscoff, including W Mediterranean Sea | 1932–1977 | 0–200 | mud, muddy sand, sandy mud | 1–40 | Glémarec & Hily (1979), partly cited from earlier sources | |
Bay of the Seine | 2008–2011 | 10–25 | mud, muddy sand, sand | 30 | Jourde et al. (2012) | |
Dieppe–Le Tréport | 2015–2018 | 12–25 | including coarse sand | Pezy & Dauvin (2016) | ||
southern North Sea | 2019 | 32 | sand with shell grit | 1 | this paper |
In general, water temperatures in shallow seas further from the ocean are more similar to air temperatures than those in deeper waters closer to the ocean. For example, the shallow southern North Sea exhibits more extreme temperature fluctuations than more stable waters of the central and northwest North Sea (MacKenzie & Schiedeck 2007). The recent (1983 to 2012) increase in annual mean sea surface temperature in the North-East Atlantic Ocean is even more pronounced in the eastern English Channel and the southern North Sea (Dye et al. 2013). Perhaps the most convincing evidence of increased average temperature for Brown Bank and the surrounding area is provided by Hughes et al. (2017; Fig. 10). They demonstrated an average increase in sea surface temperature in the southern North Sea of 0.5 °C per decade over the years 1984–2014. This is higher than for any other region in the NE Atlantic (Hughes et al. 2017; Fig. 10). According to Pezy & Dauvin (2016), the impact of climate change is indeed greater in the eastern English Channel than in the western English Channel, which is one of the main reasons why they hold climate change accountable for the range expansion of
Data on the depth range of
In our sample, we found the crab together with parts of thalassinid decapods