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We carried out monitoring of ECM wintering at a roost in Gibraltar that consists of a series of caves alongside each other at sea level, primarily during the autumn–winter period of 2019-2020. The monitoring consisted of weekly counts of birds returning to roost, and of regular ringing and measuring sessions. The ringing data for 2019–2020 were augmented with data collected at the site between 2016 and 2018.The eight caves at the site are all micro-sites within a single ECM roost and three of them, which lie just above the current sea level and are the only ones accessible from land, were studied: Gorham’s Cave (36° 07′ 13.86″ N 5° 20′ 32.57″ W UTM 30 N 289190.5 3999856.5), Vanguard Cave (36° 07′ 18.89″ N, 005° 20′ 31.62″ W UTM 30 N 289218.0 4000011.0) and Cave F (36° 07′ 19.8″ N, 005° 20′ 30.088″ W UTM 30 N 289257.0 4000038.1) (Figs. 4 and 5).Figure 4The position of Gorham’s Cave (1), Vanguard Cave (2) and Cave F (3) at the Gorham’s Cave Complex UNESCO World Heritage Site.Full size imageFigure 5Plan map of Gorham’s Cave, Vanguard Cave and Cave F at the Gorham’s Cave Complex UNESCO World Heritage Site, indicating key roosting areas for ECM and the position of the mist nets at each cave.Full size imageTwenty-three weekly evening counts were conducted of birds returning to the roosting site during the period 4th October, 2019 to 12th March, 2020, from a fixed point overlooking the study site. We attempted to space these evenly in time, but adjustments were made due to unfavorable weather (mean number of days between counts = 6.96 ± 1.45 SE). The approach of the birds as they return to roost is described elsewhere12. It occurs along a fixed trajectory and our vantage point optimized the viewing of these movements. All ECM returned to the site along a common trajectory and birds only broke up and headed towards the different caves once at the site, so that in principle, every bird had equivalent opportunities to access each cave on arrival to the site.We used a combination of the results of the counts and the Schnabel Index for mark-release-recapture data from a series of dates33 to estimate the total roosting population size of ECM at the site during the 2019–2020 period. The latter was achieved by estimating the number of birds roosting at each cave and then combining these for a total population size, although we recognize that birds also use other micro-sites7; (pers. obs.). Due to differences in sample sizes of birds recaptured, 95% confidence limits for the estimated roosting population size at each cave were drawn from the t-distribution for Gorham’s Cave and the Poisson distribution for Vanguard Cave and Cave F33.Trapping and ringing were carried out at the three caves at least once a week. All licences required under the laws of Gibraltar were obtained and protocols were approved by the Ethics Committee of the University of Gibraltar. Ringing and handling of birds was carried out under the auspices of the Gibraltar Ornithological & Natural History Society (GONHS), which carries out its bird ringing under licence from the Ministry for the Environment, HM Government of Gibraltar, under the 1991 Nature Protection Act. Gibraltar-based ringers are licensed by the British Trust for Ornithology (BTO), and we adhered closely to the technical and ethical standards of the BTO for handling and ringing birds34. Routinely, birds are released without ringing when their condition is poor. One bird was captured in a condition that was too poor for it to be ringed. The reporting recommendations of the ARRIVE guidelines35 were followed.The majority of the data used in this study were collected between October 29th 2019 and March 4th 2020. In addition, trapping and ringing had taken place intermittently at Vanguard and Cave F during the winter since 2016, and trapping took place at the site throughout autumn 2020. We used the BTO A-sized rings, in accordance with guidelines for other European hirundines34. Due to the different dimensions of the caves, we used different mist net sizes at each one. A 6m-length net was used at Vanguard Cave, 12 m and 3 m nets at Cave F, and 3 × 6 m nets mounted vertically on triple high poles at Gorham’s Cave.The number of trapping sessions, and the range of dates of these at each cave during the 2019–2020 autumn-winter season, was: 10 Gorham’s Cave (29/10/2019–04/03/2020; mean number of days between sessions 14.11 ± 2.23 SE), 11 Vanguard Cave (13/11/2019–04/03/2020; mean number of days between sessions 11.20 ± 1.81 SE), 11 Cave F (13/11/2019–04/03/2020; mean number of days between sessions 11.20 ± 1.81 SE). Seven extra trapping sessions took place at Vanguard Cave and Cave F before the 2019-2020 autumn-winter season, on: 01/28/2016, 02/16/2016, 02/13/2018, 02/21/2018, 12/04/2018, 01/08/2019 and 02/21/2019. There were eight additional trapping sessions during the autumn of 2020, on: 10/29/2020, 11/02/2020, 11/12/2020, 11/15/2020, 11/19/2020, 11/24/2020, 12/02/2020 and 12/03/2020. 1511 different birds were processed between 2016–2020, of which 156 were captured at least twice. 796 individuals were processed during the 2019–2020 autumn-winter season, the period for which most of our analyses are based: 369 at Gorham’s Cave, 221 at Vanguard Cave and 206 at Cave F. Of the birds recaptured that had been ringed at the site during previous seasons, eighteen were from the 2019–2020 season (ten ringed at Gorham’s Cave, two at Vanguard Cave, seven at Cave F), fifteen were from the 2018–2019 season (eight at Vanguard Cave, six at Cave F), four were from the 2017–2018 season (three at Vanguard Cave, one at Cave F), and one was from the 2015–2016 season (from either Vanguard Cave or Cave F; unspecified and excluded from the analysis). A bird was recaptured that had been ringed elsewhere in Gibraltar (the GONHS Jews’ Gate Field Centre) on the 14/01/2014, 2233 days before it was captured again on the 25/02/2020.Biometric measurement of all birds was carried out by a single person (CP) in order to maximize consistency. We followed the standard processing procedure of the BTO34, which includes recording the weight of birds in grams (g) to 0.1 g and length of wing in millimeters (mm) to 0.5 mm. Birds were aged whenever this was possible but ageing of ECM became increasingly difficult towards the end of the winter period, increasing the possibility of confusion with adults9. For this reason, age was excluded from most of the analyses. Birds could not be sexed because sexes are similar in appearance, including size4,36. We captured birds only during the evening, to ensure that condition of birds was not a factor of weight-loss whilst roosting, since ECM at the site are known to weigh less during mornings than the evenings13. Birds captured were roosted in boxes and released at the site the following morning.Although Elkins & Etheridge12 assumed that movement of birds between different parts of the roost at Gibraltar is considerable, this was never tested. The proximity of different parts of the roost from each other means that all micro-sites are potentially equally accessible to ECM using the site. It is expected that they should be able to use micro-sites interchangeably, given especially their approach during evenings along a fixed narrow route. Any fidelity to micro-sites must thus be explained by factors other than distance between individual micro-sites. The multiple cavities at the roosting site, and the ease with which we were able to access these, allowed a unique opportunity to test whether individual birds repeatedly used the same micro-sites within the roost, both within and between winters. We used the data gathered to test the following hypotheses: (1) that a degree of fidelity to different spaces within the roost (‘micro-sites’) exists among ECM, with individuals more likely to be recaptured at the same cave than in a different cave, (2) that any fidelity observed will translate to a difference in quality of roosting sites, as indicated by differences in condition of birds according to micro-site, and (3) that the incidence of recapture should be highest at the cave at which birds are in the best condition.Statistical analyses followed Sokal & Rohlf37 and were carried out on SPSS statistical software (IBM). We used a binomial Z test to analyze whether recaptured birds that were initially ringed during the 2019–2020 season were returning to the cave where they were first trapped/ringed, (1) within the 2019–2020 season and (2) between this and separate seasons. We also used a 3 × 2 Fisher’s exact test to test for differences, between caves, in the frequency with which birds ringed at one cave were captured at another. Multiple recaptures of birds were excluded from all of these analyses on fidelity in order to avoid bias.We explored the relationship between wing length and weight using linear regression analysis, to control for the possible effect of body size on weight—on the basis that wing length provides a good measure of body size in passerines38—using only data collected during the 2019–2020 season. For individual birds that were trapped more than once, we used wing length and weight on the date of first capture. We then grouped, by cave, the residuals of the regression and used a one-way ANOVA to explore differences in mean condition of birds between caves, with condition expressed as the relationship between wing length and weight. We also used linear regression to explore the relationship between daily recapture rate at all caves and the number of days from the first day of trapping at each cave, with the latter as the explanatory factor. Again, we segregated the residuals of the regression by cave and used a one-way ANOVA to explore differences in recapture rates between caves. We used Pearson’s chi-squared test with Yates’s correction for small sample sizes39 to explore differences in the likelihood of recapture of birds on more than one occasion at each cave. Because differences in weight and wing length have been recorded between adult and juvenile ECM in Gibraltar13, we used Pearson’s chi-squared test to explore the relationship between age and use of the different micro-sites for all the birds that we were able to age (n = 395 of 796 birds processed), to see whether this was consistent with our other findings. More

Active ingredients detected in bee collected pollenAll 188 pollen samples had at least 12 active ingredients detected in each sample, with a maximum of 31 AIs and an average of 22.0 ± 0.3 per sample. Over both years, 80 of the 259 screened pesticide active ingredients were detected in the pollen. These included 28 fungicides, 26 insecticides, 21 herbicides, two miticides, and one rodenticide. We also detected one synthetic antioxidant and one pesticide synergist (Table S1). We detected approximately twice as many AIs in pollen collected by honey bees (68 AIs) in 2019 than in pollen collected by bumble bees (32). All AIs detected in pollen from bumble bees were also collected by honey bees in either 2018 or 2019. Honey bee collected pollen also had significantly more AIs on average detected at each site (35.0 ± 0.9 S.E. AIs per site) compared to bumble bees (18.6 ± 0.6) in 2019 (R2m = 0.73; X2 = 68.2, df = 1, p  More

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The location, diameter, height and circumference of the coral were measured (Table 1, Fig. 2). The Porites was brown to cream in colour and hemispherical in shape (Fig. 2). It was identified as either Porites lutea (Hump or Pore coral) or P. lobata (Lobe coral)14.The primary habitat on the Porites was live coral (70%), followed by sponge, live coral rock and a small amount of macroalgae (Table 2). No recently dead coral, coral rubble or sand was recorded (Table 2). We observed competition between the Porites and other species of coral and invertebrate including encrusting sponge, plating and branching Acropora spp., Montipora, Chlorodesmis, soft coral and zoanthids (Table 2, Figs. 3, 4).Table 2 Reef Health Impact Survey (RHIS) of habitat and species categories on Porites sp.Full size tableFigure 3Detail of the sub-habitats and competitive interactions Porites sp. and boring sponge Cliona viridis (left) and live coral Porites sp. and Montipora sp. (right) along interspecific contact zones.Full size imageFigure 4Detail of Reef Health Impact Survey (RHIS) of Porites.Full size imageThe boring sponge, Cliona viridis, is abundant on the Great Barrier Reef15. It is a common bioeroding species advancing laterally at around 1 cm and to a depth of 1.2 cm per annum15. Abundance of Cliona viridis is often correlated to substrate availability and water energy with the greatest abundance often on the windward side of bommies15. This correlates to our observations as the large proportion of the substrate estimated to cover the bommie (15%) was on the windward side. The sponge’s advances will likely continue to compromise the colony size and health.We recorded marine debris at the base of the Porites. The debris was 2–3 m of rope that appeared to have been wrapped around the base of an adjacent coral. Adjacent to the bommie were three concrete blocks.How big is the Porites coral at Goolboodi compared to other big corals in the GBR, and the world? Potts et al.6 reported a very large, rounded Porites colony, 6.9 m in diameter which is 3.1 m smaller than this study. Lough et al.16 reported coral cores from colonies between 1.6–8.0 m in height with the largest corals of 6.0 m at Havannah, North Molle and Masthead Islands, 7.5 m at Abraham Reef and 8.0 m at Sanctuary Reef. Recognising the limitations of published data, the Porites coral at Goolboodi is the largest diameter coral that has been measured, and the 6th tallest in the GBR. It is unknown if the other corals are still alive or dead.Other comparatively large massive Porites have previously been located throughout the Pacific. These have included multiple bommies measuring more than 10 m4 and one exceptionally large colony observed measuring 17 m × 12 m in American Samoa17. Additionally, large Porites sp. bommies have been observed at Green Island, 30 km east of Taiwan18 as well as an 11 m diameter Porites australiensis at Sesoko Island, Okinawa, Japan19.How old is this massive Porites? In discussions with the Australian Institute of Marine Science (AIMS), there is a robust, linear relationship ( > 80% variance explained) between Porites average linear extension rate and average annual sea surface temperature (SST)20,21 that provides an estimate of colony age from its height. Using average annual SST at 18.5S, 146.5E of 26.12C (from HadiSST data set), the estimated linear extension rate is determined by (2.97 × 26.12) − 65.46 = 1.21 cm/year. Given the colony height of 5.1–5.3 m, this gives an estimated age of 421–438 years. This is well before European exploration and settlement of Australia. AIMS has investigated over 328 colonies of massive Porites corals from 69 reefs along the GBR and has aged them as being from 10–436 years21. AIMS has not investigated this coral (pers. comm Neal Cantin). Based on limitations of published data, the Porites coral at Goolboodi is one of the oldest corals on the GBR.Why is the Porites partially dead on top and living on the side? The proportion of live coral tissue on a colony reflects the cumulative, integrated effect of both beneficial and adverse environmental factors. Substantial portions of coral tissue can die from exposure to sun at low tides or warm water without lethal consequences to the colony as a whole10. Partial mortality of large bommies provides available real estate for opportunistic, fast growing sessile organisms. In this instance, multiple species of tabulate and branching Acropora sp., encrusting Montipora sp. and encrusting sponges are among the benthic organisms to have colonised 30% of the coral bommies’ surface area. Intraspecific competition is also evident from the skeletal barriers created along contact zones22 (Fig. 3). There was no observation of disease or coral bleaching.The Porites is located in a relatively remote, rarely visited and highly protected Marine National Park (green) zone. Its location had not been previously reported and there is no existing database for significant corals in Australia or globally. Cataloguing the location of massive and long-lived corals can have multiple benefits. Scientific benefits include geochemical and isotopic analyses in coral skeletal cores which can help understand century-scale changes in oceanographic events and can be used to verify climate models. Social and economic benefits can include diving tourism, citizen science23 culture and stewardship. Perhaps the Significant Trees Register, which was designed by the National Trust24 to protect and celebrate Australia’s heritage could be considered as a model. There are risks of cataloguing the location of massive corals. It could be damaged by direct and indirect human uses including anchoring, research and pollution.Indigenous languages are an integral part of Indigenous culture, spirituality, and connection to country. We consulted Manbarra Traditional Owners about protocol and an appropriate cultural name for the Porites and they considered: Big (Muga), Home (Wanga), Coral reef (Muugar), Coral (Dhambi), Old (Anki, Gurgu), Old man (Gulula) and Old person (Gurgurbu)25. The recommendation by Manbarra Traditional Owners is that the Porites is named as Muga dhambi (Big coral). The feedback from the process of consultation was very positive with acknowledgement of the respect that the scientists have demonstrated to acknowledge Traditional Owners in this way.The large Porites coral at Goolboodi (Orpheus) Island is unusually rare and resilient. It has survived coral bleaching, invasive species, cyclones, severely low tides and human activities for almost 500 years. In an attempt to contextualise the resilience of these individual Porites we have reviewed major historic disturbances such as coral bleaching which has occurred since at least 1575 and potentially 99 bleaching events in the GBR over the past 400 plus years26. Other indicators such as high-density ‘stress bands’ were recorded from 1877 and are significantly more frequent in the late twentieth and early twenty-first centuries in accordance with rising temperatures from anthropogenic global warming27. In addition there have been an average of 1–2 tropical cyclones per decade (40–80 in total) that have potentially impacted the coral adjacent to Goolboodi Island28,29; 46 tropical cyclones impacted the area between Ingham and Townsville from 1858 to 200830. The cumulative impact of almost 100 bleaching events and up to 80 major cyclones over a period of four centuries, plus declining nearshore water quality contextualise the high resilience of this Porites coral. Looking to the future there is real concern for corals in the GBR due to many impacts including climate change, declining water quality, overfishing and coastal development31,32. This field note provides important geospatial, environmental, and cultural information of a rare coral that can be monitored, appreciated, potentially restored and hopefully inspire future generations to care more for our reefs and culture. More

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