Seabirds and seals as drivers of plant succession on Surtsey

Plant colonization and succession on Surtsey have been monitored since 1965. In 2019, the 75th species of vascular plants was detected on the island, 62 species were present and about 40 species had established viable populations. Over the last decade colonization has slowed down and the number of present species not increased. The rising number of seagulls breeding on the island after 1985 had a great impact on plant colonization and vegetation development. While most parts of the island remained barren, a grassland area (13 ha in 2018) developed in the main seagull breeding colony on the southern part of the island. This development is attributed to transfer of nutrients from sea to land by the seagulls. In recent years a dense patch of vegetation, 2 ha in 2018, has also developed on the low, northern spit of the island, where a few pairs of seagulls breed in the spring and grey seals haul out and breed in the fall in considerable numbers. In a survey conducted during the grey seal pupping period in 2019, the seal abundance and spatial distribution was mapped accurately for the first time. The results show that the dense vegetation of the spit and seal distribution are clearly overlapping. The continuous shrinking of the island and its spit has led to an increasing concentration of the seals in their breeding area. Based on a literature survey we estimated the nitrogen (N) input from sea to land by the grey seals as 9-13 kg N/ha in 2019. This compares to an estimated input of 5-30 kg N/ha/yr by the seagulls breeding in the same area during 2015-2019. Within the grey seal and seagull breeding area on the spit of the island, a distinct community of shore plants has developed. Measurements of plant cover and biomass in permanent plots on Surtsey in 2018 and 2019 show that development on part of the spit is reaching a similar state as in the old gull colony on the southern part of the island. This suggests that the grey seals along with the seagulls are important drivers of plant succession on the northern spit. Further research on the effects of the seals on nutrient transfer from sea to land and ecosystem development on Surtsey is recommended.

Petrograpic and material observations of basaltic lapilli tuff, 1979 and 2017 Surtsey drill cores, Iceland

Petrographic studies of thin sections from the 1979 and 2017 Surtsey drill cores provide new insights into microstructural features in basaltic lapilli tuff sampled from the principal structural and hydrothermal zones of the volcano. These describe narrow rims of fine ash on altered glass pyroclasts in thin sections of the 2017 cores, characteristics of granular and microtubular structures in the original thin sections of the 1979 core, and glass alteration in diverse environments. The narrow ash rims follow the outlines of glass pyroclasts in the subaerial tuff cone and in submarine and sub-seafloor deposits; they suggest complex eruptive and depositional processes. The tubular microstructures resemble endolithic microborings in older oceanic basalt; they suggest possible microbial activity. Tubule lengths indicate rapid growth rates, up to 30 µm in ~15 years. Comparisons of glass alteration in thin sections prepared immediately after drilling in 1979 and 2017 indicate differential time-lapse alteration processes in the structural and hydrothermal zones of the volcano. In contrast, thin sections of the 1979 core prepared after 38 years in the repository reveal labile glass alteration during archival storage. The oven-dry density of the sub-seafloor lapilli tuff decreases in 2017 samples with high porosity and water absorption and increases in 2017 samples with a compact ash matrix and lower water absorption. The petrographic descriptions and material measurements provide a foundational reference for further investigations of explosive eruption and deposition of basaltic tephra at Surtsey and the subsequent alteration of these deposits in the volcanic environment and, potentially, the curatorial environment.

Sea sandwort from Surtsey: chromosomal evidence of active evolution via wide-hybridization

Sea sandwort (Honckenya peploides) was among the first species of vascular plants colonizing Surtsey. It is a member of the carnation family, Caryophyllaceae, a coastal plant with circumpolar distribution. The species is dioecious comprising separate female and hermaphrodite (male) plants. Our previous study of this plant revealed high molecular polymorphism, indicating rapid expansion and multiple origins, but low genetic differentiation, suggesting gene flow on Surtsey. The maintenance and/or expansion of populations with high gene diversity on the island are most likely fostered by several factors, one of them being the polyploid nature of the study species providing fixed heterozygosity. We therefore investigated chromosome number diversity of H. peploides from Surtsey, in comparison with accessions from Heimaey and other locations within and outside Iceland. Seeds were germinated with and without cold stratification. Chromosomes were isolated from root tips using the cellulase-pectinase enzymatic squash method. DAPIstained chromosomes were counted from microscopic images that were taken at 1000x magnification. The results show that the most common 2n somatic chromosome number of this species is 68 (2n=4x=68), but a tetraploid cytotype with 66 chromosomes also exists. The karyotype analysis shows that the species is an autotetraploid, most likely originating via chromosome doubling (whole genome duplication) in a diploid ancestor. Numerous other 2n numbers were discovered, from the diploid number 2n=2x=34 in Heimaey to several different numbers between 40 and 64. The triploid hybrid numbers 2n=3x=51 (50-52) were discovered from both Surtsey and Heimaey, as well as from other regions. Triploid hybrids serve as a bridge promoting gene flow within populations, promoting heterozygosity in the tetraploid species. All other numbers are aneuploids, most likely deriving from back-crossing of triploid hybrids and the euploid parents. The presence of aneuploids across the species distribution range is due to its ability to propagate asexually by clonal expansion. The presence of the lower ploidy levels within species, together with the extensive aneuploidy, may be an evolutionary characteristic of a pioneering plant, with great dispersal ability and genetic diversity, such as sea sandwort.

Lithofacies from the 1963-1967 Surtsey eruption in SUSTAIN drill cores SE-2a, SE-2b and SE-03

Surtsey was drilled in 2017 in the context of the Surtsey Underwater volcanic System for Thermophiles, Alteration processes and INnovative Concretes (SUSTAIN) project. Vertical drill holes, SE-02a and SE02b (drilled to 191.64 m), and angled drill SE-03 (drilled to 354.05 m), intersected armoured lapilli tuff and lapilli tuff generated mainly by explosive eruptions at Surtur from November 1963 to January 1964. The top ~20 m of lapilli tuff was erupted from Surtungur. Intervals of coherent basalt in SE-02b (15.7 to 17 m and <15 cm at the end) and in SE-03 (<1 m at ~60 m and ~238 m, and 10 m near the base) are probably intrusions that may have fed the small lavas erupted at Surtur ~2.5 years later. Although collared only a few m from the 1979 drill hole, neither SE-02a nor SE-02b intersected the 13-m-thick interval of basalt found in the 1979 drill hole. The 2017 drill cores are entirely lithified and variably altered, reflecting the effects of hydrothermal alteration and cement deposition on the originally fresh, unconsolidated ash and lapilli. Drill hole SE-03 was drilled on an azimuth of 264o and at 55o from horizontal, obliquely crossing the crater- and conduit-fill of Surtur. Although the exact trajectory of SE-03 is unknown (the drill hole was not surveyed), the drill hole ended at a vertical depth of ~100 m below the pre-eruption sea floor, however, sedimentary facies known to underlie the sea floor nearby were not intersected. Surtur eruptions therefore excavated the pre-eruption sea floor to a depth of several tens of m.

Observations on the structure of Surtsey

Comparison of investigations of the 1979 and 2017 cored boreholes coupled with continued observations of the dynamic surface of Surtsey has modified our concepts of the subsurface structure of the volcano. A geometrical analysis of the 2017 vertical and inclined cores indicates that near-surface layering dips westerly, indicating that the boreholes are located inside the Surtur crater. In subaerial deposits, as well as in deep deposits below sea level and below the pre-Surtsey seafloor, there are zones of porous tuff that contain abundant pyroclasts with narrow rims of fine ash. These features, typical of near-surface deposits, could have been carried down the vent by downslumping during fluctuating explosive activity. They support the hypothesis that a broad diatreme underlies the Surtur vent. No major intrusions were encountered in the 2017 drilling except for coherent basalt in deep sub-seafloor deposits below the center of Surtur crater. The 2017 borehole temperature measurements indicate that the peak temperature in the vertical boreholes was 124 °C at 105 meters below the surface (m.b.s.) and that in the inclined hole it was 127 °C at 115 m.b.s. immediately after drilling. These peak temperatures are 72 meters apart horizontally yet closely resemble each other in shape and magnitude, suggesting a broad heat source. In addition, measurements in the inclined hole from 200 to 290 m.b.s. indicate a temperature of 60±2 °C. This is apparently residual heat from the volcanic action that created the diatreme. These facts cast doubt on the previous concept that the heat anomaly in the 1979 borehole was due to a nearby intrusion. Instead they suggest that heat would have been conducted down from the 85-meter-thick hot lava shield within the Surtur crater into a warm diatreme substrate containing original volcanic heat. As the conducted heat moved down into the water-saturated substrate it would have elevated the temperature above the boiling point curve, baked out water, and created a vapor-dominated system below sea level. Eventually loss of heat by boiling and rise of steam caused the vapor-dominated system to retreat upward. The resulting steam rose and warmed the tephra adjacent to the lava shields where it produced broad areas of palagonitized tuff.

Erosion and sedimentation in Surtsey island quantified from new DEMs

We present data from a photogrammetric study on Surtsey island that generated three new DEMs and orthoimages, two from scanned aerial images from 1967 and 1974 and one from high-resolution closerange images from a survey in 2019. DEM differencing allowed for quantification of the erosion and the sedimentation in the island since 1967. Of the subaerial volcanics, about 45% of the lava fields have eroded away but only about 16% of the tuff cones. The prevailing SW coastal wave erosion is evident from the erosive pattern in Surtsey, and the cumulative loss of the coastal margins amounts to 28±0.9x106 m3 since 1967, with the current average erosion rate of 0.4±0.02x106 m3 /yr. Wind deflation and runoff erode the tuff cones and the sediments at the flanks of the cones, with the total volume loss amounting to 1.6±0.2x106 m3 and the current erosion rate of 0.03±0.004x106 m3 /yr. A rapid decline in erosion rates characterized the first years post-eruption, and the coastal erosion rate during the winter of 1967–68 was about 5–6 times higher than the current erosion rate due to the thinner and less cohesive nature of the lava apron at the edge of the shelf. The cones eroded at a rate about 2–3 times higher during the first years due to the uncompacted and unconsolidated nature of the cones at that time. The 2019 area of 1.2 km2 and an extrapolation of the current erosion rate fits well with the projected erosion curve of Jakobsson et al. (2000) with the island becoming a tuff crag after approximately 100 years.

Availability of plant nutrients and pollutants in the young soils of Surtsey compared to the older Heimaey and Elliðaey volcanic islands

Surtsey and the older islands in the Vestmannaeyjar archipelago offer a unique possibility to study how sub-Arctic ecosystems develop from unvegetated mineral substrate that lacks soil cover to grasslands with thick Brown Andosol soils. The present study was carried out on Surtsey, Heimaey and Elliðaey in 2013 and involved an incubation of resin membranes in the 0-10 cm topsoil layer in different ecosystems, which were either inside or outside seabird colonies. We compared the effects of seabird presence on soil nutrient availability as well as the importance of time for soil development (at least ca.1600 years vs. 50 years). Further we looked for build-up of Cd and Pb within the seabird colonies. Seabird presence enhanced the availability of most nutrients (N, P, K, Mg, Ca, S, Fe, Mn and Zn) except B and Cu, irrespective of the age of the islands. Soil age was also a significant factor for nutrient availability for all macro- and micronutrients except B. Nutrient ratios indicated that N was the most limiting nutrient in all ecosystems, except in the thicker tephra soils on Surtsey where low P availability may lead to co-limitation. The role of P in ecosystem function on Surtesy warrants a futher study. No accumulation of Cd and Pb was found within the seabird colonies.

Hydrozoan colonization and succession in the tidal and subtidal zones in Surtsey during the period 1967 to 1984

This article reports on results of investigations of hydrozoans collected in Surtsey, Iceland in the period 1967 – 1984. Samples were collected in the intertidal zone and by divers in the subtidal zone down to 40 m. A list and illustrations of hydrozoan species found in the intertidal and subtidal rocky bottom in Surtsey are presented. Species numbers increased steadily during the study period and in 1984 a total of 37 species were recorded in Surtsey making hydrozoans one of the most diverse marine invertebrate groups in Surtsey. Among hydrozoans found during the study are 8 species not previously recorded in Iceland. Apart from dispersal by planktonic medusa, rafting of polyps on floating objects drifting to Surtsey is thought to be important for colonisation of hydrozoan fauna in Surtsey. At the end of the investigations period, 20 years after formation of rocky shores on the island, number of species seemed to be continually increasing

Effects of sea birds and soil development on plant and soil nutritional parameters after 50 years of succession on Surtsey

Surtsey, the island that rose from the sea in a submarine eruption during 1963 to 1967, has been the subject of many studies on primary succession. These studies have intensified after the establishment of a seagull colony on the island in 1986. This paper reports on the results of a short sampling expedition in 2013 that intended to characterize the interactive effects of the seagull colony and of soil development on soil nutritional characteristics in the tephra sands that cover the underlying lava, as well as in plants growing inside and outside the seagull colony. Feces and pellets of the gulls were extremely rich in both nitrogen (N) and phosphorus (P) and δ15N analyses showed that N was transferred from pellets and feces to the tephra soils and subsequently taken up by the plants. The tephra soils not affected by the birds showed a high concentration of P compared to N. The concentration of both nutrients was much lower than in the soils of the bird colony. In general, variation in tephra soil depth had little effect on nutritional characteristics, except for the very low N concentration in deep soils. Thus, our results confirm the overriding effect of the seagull colony on Surtsey on nutritional characteristics of the developing soils and vegetation. Due to the very high P availability of the volcanic soils in combination with the high P input by the birds, vegetation productivity is N limited, despite the extremely high N input of 47 kg N ha-1 yr-1 that the birds add to the system. Our findings emphasize the extreme importance of bird colonies on the nutritional ecology of young, N-poor ecosystems.