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Coleoptera of Dawlish Warren Nature Reserve including a comparison of dry pitfall trapping with shoreline sampling

Coleoptera of Dawlish Warren Nature Reserve including a comparison of dry pitfall trapping with... Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 BioscienceHorizons Volume 11 2018 10.1093/biohorizons/hzy017 ............................................................................................ ..................................................................... Research article Coleoptera of Dawlish Warren Nature Reserve including a comparison of dry pitfall trapping with shoreline sampling E. Armstrong and G.J. Holloway Centre for Wildlife Assessment and Conservation, School of Biological Sciences, Harborne Building, The University of Reading, Berkshire RG6 6AS, UK *Corresponding author: Centre for Wildlife Assessment and Conservation, School of Biological Sciences, Harborne Building, The University of Reading, Berkshire RG6 6AS, UK. Email: ellis.armstrong@pgr.reading.ac.uk Supervisor: G.J. Holloway, Centre for Wildlife Assessment and Conservation, School of Biological Sciences, Harborne Building, The University of Reading, Berkshire RG6 6AS, UK. Email: g.j.holloway@reading.ac.uk ............................................................................................ ..................................................................... An examination of community structure and associative niche habitats on the Coleoptera of Dawlish Warren were carried out. The aim of this study was to better understand coleopteran communities frequently ignored in coastal habitats that are at significant risk of becoming endangered due to conflicting coastal human activities and rising sea levels resulting in a coastal squeeze. Here, we hypothesise that the shoreline Coleoptera are an extension of neighbouring coleopteran commu- nities and so will find little difference in community structure. Two techniques were used: pitfall trapping and shoreline sam- pling. A dry pitfall trap was designed to facilitate live trapping of insects and that guarded against the killing of vertebrates such as sand lizards. Shoreline sampling using a quadrat along a transect was carried out to establish whether this approach could be used as an alternative to pitfall trapping to assess the beetle fauna of the site. Beetles were captured from June until September 2017. A total of 56 species across 15 families were captured with 20 species a first recording for Dawlish Warren. Pitfall traps in the dune system produced 50 species while the shoreline sampling yielded just 13 species. The community structure was observably different between the shoreline and dune system in addition to the disparity in numbers. It was concluded that driftwood and seaweed were important attributes to shoreline coleopteran communities along with a lower pebble presence. Not all of the species in the dunes took to the air to disperse and in doing so exposed themselves to the possibility of becoming stranded in the sea. This point and the fact that a number of beetles captured were shoreline specia- lists meant that shoreline sampling could not serve as an alternative to pitfall trapping to survey for beetles on this site. Key words: Dawlish Warren, Coleoptera, pitfall trapping, shoreline sampling, management Submitted on 30 July 2018; editorial decision on 10 December 2018 ............................................................................................ ..................................................................... Union for Dune Conservation noted that 56 000 ha of sand Introduction dunes existed on the coastlines of the UK, of which 11 897 ha Coastal sand dunes are important to local communities for were on the coast of England. Sand dunes systems are fre- their socio-economic benefits. In the UK Biodiversity Action quently of high ecological value. Sandy coastlines often face Plan: Priority Habitat Descriptions (Biodiversity Reporting disturbance and subsequent modification through abiotic con- and Information Group, 2008), the Sand Dune Survey of ditions, such as strong winds (Wasson and Nanninga, 1986), Great Britain (1993–1995) in accordance with the European frequent change of composition due to tidal fluctuation ............................................................................................... .................................................................. © The Author(s) 2019. Published by Oxford University Press. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. (Curtiss, Osborne, and Horner-Devine, 2009) and deform- Spungis, 2002). Of course, this practice has its limitations; dis- ation from frequent human activities. As a result, sand dune proportionately more ground-dwelling taxa are captured in com- ecosystems are forever in flux but also offer a wide range of parison to those primarily occupying vegetation or living under habitat niches. This is associated with high levels of biodiver- the soil surface (Fanini and Lowry, 2016), while dry pitfall sam- sity. For example, 102 notable species of Coleoptera associated pling can result in species being predated upon by cohabitants or with sand dune habitat are listed in the UK Biodiversity Action escaping through flight. Pitfall sampling is simply unsuitable for Plan (Joint Nature Conservation Committee, 2018). Coleoptera the shoreline due to tidal fluctuations and exposure to disturb- play a key role in sand dunes as ecosystem engineers by feeding ance. Therefore, a less-frequently used quadrat-transect sampling upon carrion and vegetation. They are also vital as a food source method is more applicable. This approach allows for active for species of insectivorous dune-dwelling vertebrates, such as searching through debris over a limited time scale to standardise the sand lizard (Lacerta agilis L.). sampling effort along the shoreline as conducted by Hodge and Williams (2007). There are restrictions using this approach as the Disturbance occurs naturally in coastal regions with tidal weather conditions when sampling can influence the presence or fluctuation and strong wind resulting in frequent erosion and absence of invertebrates and it excludes species that are nocturnal the deposition of sediments. More recently, the rise of industrial- (Aloia et al., 1999). Each approach can create a sampling bias isation has altered climatic conditions (Rosenzweig et al., 2008; (Fanini and Lowry, 2016) and therefore skew which species are Vavrus et al., 2008). Since 1870, global sea level has risen captured and identified, potentially having a negative influence 195 mm (Church and White, 2006) while recent models predict on management technique success. a sea level rise of between 400 and 700 mm by 2100 at current rates (Horton et al.,2014). For coastal habitats, this means The current study focusses on the dune system of Dawlish more frequent extreme weather events (Mann et al.,2017)caus- Warren in south-western England. Dawlish Warren is a fine ing higher erosion rates and increased sea levels resulting in habi- example of a dune ecosystem in the UK and as a result is pro- tat migration. Human urbanisation interferes with the natural tected by several layers of legislation (Joint Nature Conservation migration of the coastal habitat (Doxa et al.,2017)asprotective Committee, 2017; Natural England, 2017). However, it is very measures such as sea walls cause a ‘coastal squeeze’ (Doody, important to know which species are on-site to inform manage- 2004; Pontee, 2013), the prevention of habitat migration inland ment procedures (Teignbridge District Council, 2010). This is in the face of elevated sea levels (Jackson and McIlvenny, 2011). the same for all designated wildlife areas. Legal protection As a result of this, many coastal communities are now under ser- makes surveys that require sampling a sensitive issue particularly ious threat (Nicholls et al., 1999; Doxa et al.,2017). if high-profile species are considered to be at risk through the sampling. The aims of the current study were twofold: Humans are attracted to coastal sites, in particular sand dunes, for recreational purposes. Trampling associated with � To compare the beetle community collected along the human recreation threatens many species occupying sand dune strandline with the beetle community collected from the habitats (Avgın and Luff, 2010; Bessa et al.,2013; Schlacher dunes to establish whether the two communities are similar. et al.,2016). This interference is affecting trophic levels in dif- � To sample the site to contribute to our knowledge of spe- ferent ways. While most designated sites and conservation cies and associated shoreline debris utilising Dawlish as an strategies focus on megafauna and habitats, less is known aid to future management strategies. about the effects of disturbance on lower trophic levels. Invertebrates usually occupy the second trophic level as pri- mary consumers, although a small number can be classified as Methodology secondary consumers. They prop up the food web as sources of nutrition for many birds, reptiles and mammals. Dirzo et al. Shoreline transect (2014) observed a mean decrease in invertebrate abundance of Two 500-m stretches of shoreline were marked out (Fig. 1) 45% over a 35-year period when the human population with one facing the open ocean (outer) and one facing the doubled. Further analysis shows a decline of 30–60% of com- mouth of the river Exe (inner). The start of the outer shoreline mon insects in the UK over the last 40 years. However, elusive transect was marked by the final groyne along the 2.5 km species were not included in this data set, which casts doubt beach, then at every 50 m a 1 m quadrant was marked whilst over the accuracy of this study. The loss of invertebrates, maintaining a distance of 3 m perpendicular to the waveline including beetles, is of great concern and it is likely that effects (so the position of the quadrat on the beach varied depending at higher trophic levels are likely to be delayed. on the height of the tide). Ten sampling points were estab- Constant fluctuation of habitat type and structure in sand lished along each of the transects. Within each quadrat, the dune systems can be an issue when sampling. Pitfall traps provide percentage cover of four constituents was recorded: bare a standard method often used in arthropod sampling due to their sand, seaweed, driftwood and pebbles. Within each quadrat, simplicity, efficacy and low cost (Southwood, 1978). In coastal the debris was hand searched for beetles for 10 min, mirroring dune systems, proportionately more ground-dwelling taxa are the method used by Hodge and Williams (2007). The tip of nocturnal due to the heat stored in the sand during the day, mak- the dune system (Fig. 1B and C) was excluded from the two ing pitfall trapping an applicable method (Aloia et al.,1999; transects to separate the outer and inner communities. Data ............................................................................................... .................................................................. 2 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Bioscience Horizons � Volume 11 2018 Research article ............................................................................................... .................................................................. Figure 1. Aerial image of Dawlish Warren illustrating the outer (A and B) and inner (C and D) shoreline transects. The markers on the dune site indicate where 15 pitfall traps were randomly positioned. were gathered once a week over a period of 14 weeks from early June to mid-September. Fluctuation in tide and attempt- ing to gather data distanced relative to the waveline meant a quadrat-transect was the favoured sampling method. Pitfall traps Fifteen pitfall traps were randomly allocated to 15 locations Figure 2. The pitfall trap was designed to catch and collect the target (Fig. 1) across 0.05 km within the sand dune ecosystem on the species alive. An outer cup was dug carefully into the ground ensuring spit of Dawlish Warren Nature Reserve. Excel was used to ran- that the rim of the cup was exactly at the level of the soil. An inner cup domly generate a four-digit number that split into two two-digit was placed inside the outer cup to facilitate retrieval of captured numbers which marked a 10 m by 10 m location on the sand insects without disturbing the outer cup. A hole in the bottom of the dune system. A pitfall trap was placed in the centre of each ran- outer cup allowed drainage during wet weather. The inner cup had domly located square using GPS coordinates. If a pitfall trap five small holes in the bottom to drain rainwater to ensure that captured insects did not drown. A small amount of debris was placed was deemed to be too close to a distinct pathway the trap was in the base of the inner cup for beetles to shelter within. The funnel moved at least 1 metre away from the path. Pitfall sampling in was the top of a plastic drinks bottle cut-off, turned upside down and the dune system was preferred as this would capture coleopteran wedged into the top of the outer cup. This prevented vertebrates from species which were nocturnal (Aloia et al.,1999) while also pro- being caught and stopped captured insects from escaping. If any small viding longer exposure to capture species. Furthermore, disturb- vertebrates, e.g. juvenile lizards, did fall into the trap they were unlikely ance in this location would be minimal. to die. A large, flat stone propped upon three twigs was placed over the trap to reduce rainfall from entering the system. Due to site sensitivities and the presence of sand lizards (L. agilis), a priority species, the standard pitfall trap design (Greenslade, 1964)requiredmodification for use. A funnel in a dilute vinegar solution [40% malt vinegar and 60% water design was used to trap invertebrates but prevent the trapping of (Aristophanous, 2010)] and returned to the laboratory for iden- non-target vertebrates (Fig. 2). This consisted of an outer cup tification at a later date. Specimens were identified to species dug carefully into the ground ensuring the rim was level with the using a stero microscope in conjunction with Chinery (1993), soil surface. An inner cup was placed inside the outer cup to Duff (2012; 2016)and Telfer’s (2017) online resources. facilitate retrieval of captured insects without disturbing the out- Sampling effort and methods differed between the two er cup. A hole in the base of the outer cup allowed drainage dur- sampled locations, sand dunes and the shoreline. Therefore, ana- ing wet weather, whilst the inner cup had five small punctures in lysis was conducted using a Simple Correspondence Analysis, the base to facilitate drainage ensuring captured specimens did looking at the presence of relatively abundant species (>5 not drown. A small amount of debris was placed at the base of records) relative to their preferred location which spans across the inner cup to provide shelter for these captured species. The three categories; sand dunes, the outer shoreline and the inner funnel design used upturned cut-off tops of plastic drinks bottles shoreline. This helped visualise communities in each of these and wedged into the top of the outer cup. This prevented verte- three locations and determine the difference between the shore- brates being caught and stopped captured insects escaping. A lines and the dune system. Linear Regression Analysis was used flat stone propped upon three twigs was placed over the trap to to determine any associative relationship between the type of reduce the amount of rainfall entering the system. debris present within a shoreline transect, determined by mean percentage cover per week, and the presence with relative abun- Data recording and statistical analysis dance of Coleopteran families. It was uncertain whether individ- Where possible, beetles were identified on-site, recorded and ual species would appear in enough abundance to allow for released. If a specimen could not be identified it was preserved individual species analysis; Therefore the data were combined ............................................................................................... .................................................................. 3 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. melanocephalus (L.) with 107 recordings while Phylan gibbus (Fabricius), a specialist halobiontic species, was recorded 59 times. The families Carabidae, Staphylinidae and Curculionidae were particularly well represented in the pitfall traps with 14, 10, and 8 species identified in each family, respectively. Forty- three species were solely found on the sand dune system. Community structure and comparison Fig. 3 shows a simple correspondence analysis of community structure. All of the species identified were plotted against the three sample locations: sand dunes, outer shoreline and inner shoreline. When a species (blue dot) is close to the centre point (0,0), it is relatively evenly distributed between the shoreline community and the sand dune community. Species found only in one community occupy the same position in Fig. 3, which has been represented by enlarged blue dots and a corre- sponding number signalling the number of species plotted in the same location. If a species is close to a community centre (red square), then it is unevenly distributed between the com- Figure 3. Simple correspondence analysis illustrating the different munities. Few species were found present in both the sand beetle community structures (red squares) gathered from the sand dune and shoreline habitats, with only two species occurring dunes, the inner shoreline and the outer shoreline. Seventy-seven multiple times in each habitat. Therefore, the shoreline cole- percentage of species (43/56) identified were solely found in the sand opteran community can be defined as a separate community dune habitat while only 12.5% (7/56) were found on both the shoreline to the sand dune community rather than an extension to the and sand dune habitats. Positions of individual species in the two dune community. The inner and outer shoreline species show dimensional space are represented by small blue dots. Enlarged blue subtle differences but had proportionately more species that dots have a corresponding number relative to the number of species within the blue dot. The arrows represent the strength of distribution were present in both communities. towards each community where the centre point (0,0) represents an Table 1 shows results of regression analyses of beetle abun- even distribution between the shoreline’s community and sand dune community. Few species were found present in both the sand dune dance on amounts of different types of debris along the shore- and shoreline habitats, with only two species occurring multiple times line. On the outer shoreline, beetles from all families showed in each habitat. Therefore, the shoreline coleopteran community can a significant positive relationship with the number of pebbles be defined as a separate community to the sand dune community in a quadrat. Carabidae were also more likely to be found rather than an extension to the dune community. The inner and outer when more driftwood or seaweed was present. Staphylinidae shoreline species show subtle differences but had proportionately were associated with seaweed. The lowest beetle counts were more species that overlapped communities. found from bare sand. Fewer associations were found between beetle number and debris on the inner shoreline into families to identify how different families associate with although Carabidae were still closely associated with the debris. Life-history traits of the shoreline species identified were quantity of driftwood present. Beyond that the only associ- reviewed to determine the effectiveness of shoreline sampling. ation was a weak relationship between Tenebrionidae and All data were analysed using Mintab version 18.1. amount of seaweed. Discussion Results It has been widely acknowledged that coastal habitats face a Species summary significant threat of being lost as a result of rising sea levels A total of 56 species spread across 15 families were identified and a squeeze against coastal communities (Doody, 2004; (Table A1). Thirteen species were found along the shoreline; Doxa et al., 2017). One of the aims here was to distinguish seven species were found more than once and only four of these the relationship shoreline communities have with their neigh- were recorded more than five times: Broscus cephalotes (L.), bouring habitat, in this case, sand dunes. Fig. 3 shows these Calathus cinctus (Motschulsky), Phaleria cadaverina (Fabricius) neighbouring local habitats had little overlap of species and and Cafius xantholoma (Gravenhorst). Of the 50 species identi- therefore could be defined as separate communities. Looking fied in the sand dune system, 14 were recorded only once, while at the species identified from the shoreline, many are known only eight of these species were recorded more than 10 times. for being strandline specialists. Therefore, these shoreline Twenty species were a first recording for Dawlish Warren. coleopteran communities are facing a significant threat of The most common sand dune species was Calathus local extinctions where species migration perpendicular to the ............................................................................................... .................................................................. 4 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Bioscience Horizons � Volume 11 2018 Research article ............................................................................................... .................................................................. Table 1. Results of linear regression analyses [F, R-squared (%) and p values] of beetle abundance on the amount of debris along the shoreline over time (df = 1,12, significant values in bold) Bare sand Pebbles Driftwood Seaweed O I OI O I OI Carabidae F-stat 0 0.05 6.43 1.35 9.57 32.5 12.5 0.75 R-sq 0 0.38 34.9 10.1 44.4 73.0 51.1 5.88 p value 0.99 0.83 0.03 0.27 0.01 0.001 0.004 0.40 Tenebrionidae F-stat 0 0.63 8.95 3.47 0 1.74 4.22 5.16 R-sq 0 4.99 42.7 22.5 0.01 12.7 26.0 30.1 p value 0.99 0.44 0.01 0.09 0.97 0.21 0.06 0.04 Staphylinidae F-stat 0.03 0.19 14.7 0.05 0.12 1.23 8.17 0.75 R-sq 0.21 1.59 55.0 0.37 1.00 9.28 42.1 5.86 p value 0.87 0.67 0.002 0.83 0.73 0.29 0.01 0.40 Only relatively abundant families are considered (five or more recordings). O, outer shoreline; I, inner shoreline. coast is prevented as a result of a ‘coastal squeeze’. Furthermore, Twenty species new to Dawlish Warren were found, this study shows little presence of these shoreline specialists in a illustrating the importance of finding ways to carry out on- similar neighbouring habitat meaning the shoreline communities site surveys in protected areas. Dawlish Warren is a very could not be used to ‘seed’ populations on the shoreline. special site so it comes as no surprise that it holds a wide range of species. This study was limited both in terms of It is well known that shoreline walking can reveal extensive time and spatial scale and therefore further, more extensive numbers of beetles. One of the aims here was to identify how surveys will certainly reveal the presence of more species. shoreline coleopteran species are associated with coastal debris Some of these species might be scarce or restricted to and to consider whether shoreline Coleoptera could be used to sur- Dawlish and help to inform appropriate management in the vey the wider site removing the need to set pitfall traps. During the field (Teignbridge District Council, 2010). Without this study, a much wider range of species were found in the dune sys- knowledge it is possible that current management techni- tem than along the shoreline. This suggests that not all of the beetle ques could be disadvantageous to one or more potential species in the dunes take to the air readily and expose themselves speciesofinterest. to the risk of being stranded along the shoreline. Some dune spe- cies certainly do fly and get trapped on the shoreline as shown by Reticence among wardens of nature reserves to allow sur- the presence of two dune-dwelling species of Curculionidae, veys using lethal pitfall trapping to sample invertebrate com- Polydrusus cervinus (L.) and Coelositona cambricus (Stephens). In munities is frequently encountered. This degree of caution is addition, although the shoreline represents very harsh and danger- usually a result of one or more scarce target species on the ous conditions, some beetle species are specialists in this type of reserve that might be susceptible to being caught in the pitfall environment (Topp and Ring, 1988; Brown, 1996), such as traps. In this case, the species of concern was sand lizard, Cafius xantholoma (Staphylinidae) (Moore and Legner, 1976). A L. agilis. To counter this concern a modified pitfall trap was number of beetle species found along the shoreline appeared to be designed that facilitated the trapping of invertebrates live to seeking out certain features, such as pebbles and driftwood either release on-site if the species was known or to selectively (Table 1). Presumably, these structures were offering refugia or return to the laboratory for subsequent examination. The traps foraging opportunities. The outcome is that collecting along the appeared to work well although we do not know about per- shoreline cannot serve as an appropriate alternative to on-site sam- formance relative to lethal traps as the latter could not be set. pling, in this case using pitfall traps. No sand lizards or other vertebrates were caught in the traps. ............................................................................................... .................................................................. 5 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. Doxa, A., Albert, C. H., Leriche, A. et al. (2017) Prioritizing conservation Acknowledgements areas for coastal plant diversity under increasing urbanization, Journal of Environmental Management, 201, 425–434. We would like to thank Devon Wildlife Trust and Teignbridge District Council for allowing access to Dawlish Warren National Duff, A. (2012) Beetles of Britain and Ireland. Volume1: Sphaeriusidae to Nature Reserve and providing permission to conduct this research. Silphidae, A.G.Duff, Norfolk. We would also like to extend our thanks to Philip Chambers and Duff, A. (2016) Beetles of Britain and Ireland. Volume 4: Cerambycidae to the team for their help during the research on the site. Curculionidae, A.G.Duff, Norfolk. Fanini, L. and Lowry, J. (2016) Comparing methods used in estimating Author biography biodiversity on sandy beaches: pitfall vs. quadrat sampling, Ecological Indicators, 60, 358–366. Ellis Armstrong studied for a degree in Ecology and Wildlife Greenslade, P. J. M. (1964) Pitfall trapping as a method for studying Conservationgraduationin2018withaparticularinterestin populations of Carabidae (Coleoptera), The Journal of Animal Coleoptera. During the academic year 2018/2019, he will be Ecology, 33, 301–310. studying towards a Masters by Research in Entomology. 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Observations on rove beetles (Staphylinidae) from san- Press, New York. dy beaches, Canadian Journal of Zoology, 66, 2464–2468. Vavrus, S., Ruddiman, W. F. and Kutzbach, J. E. (2008) Climate model tests Spungis, V. (2002) Invertebrates of the sandy coastal habitats in Latvia, of the anthropogenic influence on greenhouse-induced climate Latvijas Entomologs, 39, 10–19. change: the role of early human agriculture, industrialization, and Teignbridge District Council. (2010) Dawlish Warren National Nature vegetation feedbacks, Quaternary Science Reviews, 27, 1410–1425. Reserve: Management Plan 2010–2020. Wasson, R. J. and Nanninga, P. M. (1986) Estimating wind transport of Telfer, M. (2017) Identifying Beetles, accessed at: http://www. sand on vegetated surfaces, Earth Surface Processes and Landforms, markgtelfer.co.uk/beetles/ (8 December 2017). 11, 505–514. ............................................................................................... .................................................................. 7 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. Appendix Table A1. Summary of the 56 Coleoptera species found at Dawlish Warren and their abundance in each of the habitats sampled Family Genus Species Sand dunes Outer shoreline Inner shoreline Carabidae Broscus cephalotes 48 4 Carabidae Syntomus foveatus 20 0 0 Carabidae Amara tibialis 22 0 0 Carabidae Dyschirius thoracicus 10 0 Carabidae Calathus melanocephalus 107 0 4 Carabidae Harpalus affinis 10 0 Carabidae Nebria salina 10 0 Carabidae Bembidion quadrimaculatum 10 0 Carabidae Calathus cinctus 11 12 Carabidae Pterostichus madidus 11 0 Carabidae Calathus fuscipes 20 0 Carabidae Trechus obtusus 20 0 Carabidae Trechus fulvus 10 0 Carabidae Pterostichus longicollis 40 0 Leiodidae Catops grandicollis 60 0 Leiodidae Agathidium Seminulum 10 0 Leiodidae Leiodes furva 20 0 Leiodidae Leiodes sp. 1 0 0 Tenebrionidae Phaleria cadaverina 029 37 Tenebrionidae Phylan gibbus 59 0 0 Hydrophilidae Anacaena globulus 20 0 Hydrophilidae Megasternum concinnum 60 0 Hydrophilidae Cercyon littoralis 01 0 Curculionidae Otiorhynchus atroapterus 13 0 0 Curculionidae Otiorhynchus rugifrons 40 0 Curculionidae Rhinonchus castor 50 0 Curculionidae Coelositona cambricus 30 1 Curculionidae Philopedon plagiatum 20 0 Curculionidae Orthochaetes insignis 10 0 Continued ............................................................................................... .................................................................. 8 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Bioscience Horizons � Volume 11 2018 Research article ............................................................................................... .................................................................. Table A1. Continued Family Genus Species Sand dunes Outer shoreline Inner shoreline Curculionidae Pselactus spadix 10 0 Curculionidae Polydrusus cervinus 01 0 Chrysomelidae Longitarsus succineus 10 0 Chrysomelidae Longitarsus flavicornis 30 0 Chrysomelidae Cryptocephalus pusillus 11 0 0 Elateridae Agrypnus murinus 90 0 Apionidae Apion haematodes 90 0 Apionidae Perapion hydrolapathi 20 0 Histeridae Hypocaccus dimidiatus 12 1 Histeridae Saprinus aeneus 10 0 Silphidae Silpha tristis 20 0 Latridiidae Corticaria sp. 1 0 0 Oedemeridae Nacerdes melanura 02 1 Coccinellidae Scymnus frontalis 10 0 Coccinellidae Coccinella septempunctata 60 0 Coccinellidae Coccinella undecimpunctata 10 0 Aphodiidae Aphodius foetens 40 0 Staphylinidae Ocypus olens 18 0 0 Staphylinidae Cafius xantholoma 012 3 Staphylinidae Cafius fucicola 01 0 Staphylinidae Ocypus brunnipes 30 0 Staphylinidae Platydracus stercorarius 10 0 Staphylinidae Stenus clavicornis 10 0 Staphylinidae Lordithon thoracicus 10 0 Staphylinidae Mycetoporus sp. 1 0 0 Staphylinidae Gabrius sp. 2 0 0 Staphylinidae Aleocharinae sp. 10 0 1 ............................................................................................... .................................................................. http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png BioScience Horizons Oxford University Press

Coleoptera of Dawlish Warren Nature Reserve including a comparison of dry pitfall trapping with shoreline sampling

BioScience Horizons , Volume 11 – Jan 1, 2018

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Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 BioscienceHorizons Volume 11 2018 10.1093/biohorizons/hzy017 ............................................................................................ ..................................................................... Research article Coleoptera of Dawlish Warren Nature Reserve including a comparison of dry pitfall trapping with shoreline sampling E. Armstrong and G.J. Holloway Centre for Wildlife Assessment and Conservation, School of Biological Sciences, Harborne Building, The University of Reading, Berkshire RG6 6AS, UK *Corresponding author: Centre for Wildlife Assessment and Conservation, School of Biological Sciences, Harborne Building, The University of Reading, Berkshire RG6 6AS, UK. Email: ellis.armstrong@pgr.reading.ac.uk Supervisor: G.J. Holloway, Centre for Wildlife Assessment and Conservation, School of Biological Sciences, Harborne Building, The University of Reading, Berkshire RG6 6AS, UK. Email: g.j.holloway@reading.ac.uk ............................................................................................ ..................................................................... An examination of community structure and associative niche habitats on the Coleoptera of Dawlish Warren were carried out. The aim of this study was to better understand coleopteran communities frequently ignored in coastal habitats that are at significant risk of becoming endangered due to conflicting coastal human activities and rising sea levels resulting in a coastal squeeze. Here, we hypothesise that the shoreline Coleoptera are an extension of neighbouring coleopteran commu- nities and so will find little difference in community structure. Two techniques were used: pitfall trapping and shoreline sam- pling. A dry pitfall trap was designed to facilitate live trapping of insects and that guarded against the killing of vertebrates such as sand lizards. Shoreline sampling using a quadrat along a transect was carried out to establish whether this approach could be used as an alternative to pitfall trapping to assess the beetle fauna of the site. Beetles were captured from June until September 2017. A total of 56 species across 15 families were captured with 20 species a first recording for Dawlish Warren. Pitfall traps in the dune system produced 50 species while the shoreline sampling yielded just 13 species. The community structure was observably different between the shoreline and dune system in addition to the disparity in numbers. It was concluded that driftwood and seaweed were important attributes to shoreline coleopteran communities along with a lower pebble presence. Not all of the species in the dunes took to the air to disperse and in doing so exposed themselves to the possibility of becoming stranded in the sea. This point and the fact that a number of beetles captured were shoreline specia- lists meant that shoreline sampling could not serve as an alternative to pitfall trapping to survey for beetles on this site. Key words: Dawlish Warren, Coleoptera, pitfall trapping, shoreline sampling, management Submitted on 30 July 2018; editorial decision on 10 December 2018 ............................................................................................ ..................................................................... Union for Dune Conservation noted that 56 000 ha of sand Introduction dunes existed on the coastlines of the UK, of which 11 897 ha Coastal sand dunes are important to local communities for were on the coast of England. Sand dunes systems are fre- their socio-economic benefits. In the UK Biodiversity Action quently of high ecological value. Sandy coastlines often face Plan: Priority Habitat Descriptions (Biodiversity Reporting disturbance and subsequent modification through abiotic con- and Information Group, 2008), the Sand Dune Survey of ditions, such as strong winds (Wasson and Nanninga, 1986), Great Britain (1993–1995) in accordance with the European frequent change of composition due to tidal fluctuation ............................................................................................... .................................................................. © The Author(s) 2019. Published by Oxford University Press. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. (Curtiss, Osborne, and Horner-Devine, 2009) and deform- Spungis, 2002). Of course, this practice has its limitations; dis- ation from frequent human activities. As a result, sand dune proportionately more ground-dwelling taxa are captured in com- ecosystems are forever in flux but also offer a wide range of parison to those primarily occupying vegetation or living under habitat niches. This is associated with high levels of biodiver- the soil surface (Fanini and Lowry, 2016), while dry pitfall sam- sity. For example, 102 notable species of Coleoptera associated pling can result in species being predated upon by cohabitants or with sand dune habitat are listed in the UK Biodiversity Action escaping through flight. Pitfall sampling is simply unsuitable for Plan (Joint Nature Conservation Committee, 2018). Coleoptera the shoreline due to tidal fluctuations and exposure to disturb- play a key role in sand dunes as ecosystem engineers by feeding ance. Therefore, a less-frequently used quadrat-transect sampling upon carrion and vegetation. They are also vital as a food source method is more applicable. This approach allows for active for species of insectivorous dune-dwelling vertebrates, such as searching through debris over a limited time scale to standardise the sand lizard (Lacerta agilis L.). sampling effort along the shoreline as conducted by Hodge and Williams (2007). There are restrictions using this approach as the Disturbance occurs naturally in coastal regions with tidal weather conditions when sampling can influence the presence or fluctuation and strong wind resulting in frequent erosion and absence of invertebrates and it excludes species that are nocturnal the deposition of sediments. More recently, the rise of industrial- (Aloia et al., 1999). Each approach can create a sampling bias isation has altered climatic conditions (Rosenzweig et al., 2008; (Fanini and Lowry, 2016) and therefore skew which species are Vavrus et al., 2008). Since 1870, global sea level has risen captured and identified, potentially having a negative influence 195 mm (Church and White, 2006) while recent models predict on management technique success. a sea level rise of between 400 and 700 mm by 2100 at current rates (Horton et al.,2014). For coastal habitats, this means The current study focusses on the dune system of Dawlish more frequent extreme weather events (Mann et al.,2017)caus- Warren in south-western England. Dawlish Warren is a fine ing higher erosion rates and increased sea levels resulting in habi- example of a dune ecosystem in the UK and as a result is pro- tat migration. Human urbanisation interferes with the natural tected by several layers of legislation (Joint Nature Conservation migration of the coastal habitat (Doxa et al.,2017)asprotective Committee, 2017; Natural England, 2017). However, it is very measures such as sea walls cause a ‘coastal squeeze’ (Doody, important to know which species are on-site to inform manage- 2004; Pontee, 2013), the prevention of habitat migration inland ment procedures (Teignbridge District Council, 2010). This is in the face of elevated sea levels (Jackson and McIlvenny, 2011). the same for all designated wildlife areas. Legal protection As a result of this, many coastal communities are now under ser- makes surveys that require sampling a sensitive issue particularly ious threat (Nicholls et al., 1999; Doxa et al.,2017). if high-profile species are considered to be at risk through the sampling. The aims of the current study were twofold: Humans are attracted to coastal sites, in particular sand dunes, for recreational purposes. Trampling associated with � To compare the beetle community collected along the human recreation threatens many species occupying sand dune strandline with the beetle community collected from the habitats (Avgın and Luff, 2010; Bessa et al.,2013; Schlacher dunes to establish whether the two communities are similar. et al.,2016). This interference is affecting trophic levels in dif- � To sample the site to contribute to our knowledge of spe- ferent ways. While most designated sites and conservation cies and associated shoreline debris utilising Dawlish as an strategies focus on megafauna and habitats, less is known aid to future management strategies. about the effects of disturbance on lower trophic levels. Invertebrates usually occupy the second trophic level as pri- mary consumers, although a small number can be classified as Methodology secondary consumers. They prop up the food web as sources of nutrition for many birds, reptiles and mammals. Dirzo et al. Shoreline transect (2014) observed a mean decrease in invertebrate abundance of Two 500-m stretches of shoreline were marked out (Fig. 1) 45% over a 35-year period when the human population with one facing the open ocean (outer) and one facing the doubled. Further analysis shows a decline of 30–60% of com- mouth of the river Exe (inner). The start of the outer shoreline mon insects in the UK over the last 40 years. However, elusive transect was marked by the final groyne along the 2.5 km species were not included in this data set, which casts doubt beach, then at every 50 m a 1 m quadrant was marked whilst over the accuracy of this study. The loss of invertebrates, maintaining a distance of 3 m perpendicular to the waveline including beetles, is of great concern and it is likely that effects (so the position of the quadrat on the beach varied depending at higher trophic levels are likely to be delayed. on the height of the tide). Ten sampling points were estab- Constant fluctuation of habitat type and structure in sand lished along each of the transects. Within each quadrat, the dune systems can be an issue when sampling. Pitfall traps provide percentage cover of four constituents was recorded: bare a standard method often used in arthropod sampling due to their sand, seaweed, driftwood and pebbles. Within each quadrat, simplicity, efficacy and low cost (Southwood, 1978). In coastal the debris was hand searched for beetles for 10 min, mirroring dune systems, proportionately more ground-dwelling taxa are the method used by Hodge and Williams (2007). The tip of nocturnal due to the heat stored in the sand during the day, mak- the dune system (Fig. 1B and C) was excluded from the two ing pitfall trapping an applicable method (Aloia et al.,1999; transects to separate the outer and inner communities. Data ............................................................................................... .................................................................. 2 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Bioscience Horizons � Volume 11 2018 Research article ............................................................................................... .................................................................. Figure 1. Aerial image of Dawlish Warren illustrating the outer (A and B) and inner (C and D) shoreline transects. The markers on the dune site indicate where 15 pitfall traps were randomly positioned. were gathered once a week over a period of 14 weeks from early June to mid-September. Fluctuation in tide and attempt- ing to gather data distanced relative to the waveline meant a quadrat-transect was the favoured sampling method. Pitfall traps Fifteen pitfall traps were randomly allocated to 15 locations Figure 2. The pitfall trap was designed to catch and collect the target (Fig. 1) across 0.05 km within the sand dune ecosystem on the species alive. An outer cup was dug carefully into the ground ensuring spit of Dawlish Warren Nature Reserve. Excel was used to ran- that the rim of the cup was exactly at the level of the soil. An inner cup domly generate a four-digit number that split into two two-digit was placed inside the outer cup to facilitate retrieval of captured numbers which marked a 10 m by 10 m location on the sand insects without disturbing the outer cup. A hole in the bottom of the dune system. A pitfall trap was placed in the centre of each ran- outer cup allowed drainage during wet weather. The inner cup had domly located square using GPS coordinates. If a pitfall trap five small holes in the bottom to drain rainwater to ensure that captured insects did not drown. A small amount of debris was placed was deemed to be too close to a distinct pathway the trap was in the base of the inner cup for beetles to shelter within. The funnel moved at least 1 metre away from the path. Pitfall sampling in was the top of a plastic drinks bottle cut-off, turned upside down and the dune system was preferred as this would capture coleopteran wedged into the top of the outer cup. This prevented vertebrates from species which were nocturnal (Aloia et al.,1999) while also pro- being caught and stopped captured insects from escaping. If any small viding longer exposure to capture species. Furthermore, disturb- vertebrates, e.g. juvenile lizards, did fall into the trap they were unlikely ance in this location would be minimal. to die. A large, flat stone propped upon three twigs was placed over the trap to reduce rainfall from entering the system. Due to site sensitivities and the presence of sand lizards (L. agilis), a priority species, the standard pitfall trap design (Greenslade, 1964)requiredmodification for use. A funnel in a dilute vinegar solution [40% malt vinegar and 60% water design was used to trap invertebrates but prevent the trapping of (Aristophanous, 2010)] and returned to the laboratory for iden- non-target vertebrates (Fig. 2). This consisted of an outer cup tification at a later date. Specimens were identified to species dug carefully into the ground ensuring the rim was level with the using a stero microscope in conjunction with Chinery (1993), soil surface. An inner cup was placed inside the outer cup to Duff (2012; 2016)and Telfer’s (2017) online resources. facilitate retrieval of captured insects without disturbing the out- Sampling effort and methods differed between the two er cup. A hole in the base of the outer cup allowed drainage dur- sampled locations, sand dunes and the shoreline. Therefore, ana- ing wet weather, whilst the inner cup had five small punctures in lysis was conducted using a Simple Correspondence Analysis, the base to facilitate drainage ensuring captured specimens did looking at the presence of relatively abundant species (>5 not drown. A small amount of debris was placed at the base of records) relative to their preferred location which spans across the inner cup to provide shelter for these captured species. The three categories; sand dunes, the outer shoreline and the inner funnel design used upturned cut-off tops of plastic drinks bottles shoreline. This helped visualise communities in each of these and wedged into the top of the outer cup. This prevented verte- three locations and determine the difference between the shore- brates being caught and stopped captured insects escaping. A lines and the dune system. Linear Regression Analysis was used flat stone propped upon three twigs was placed over the trap to to determine any associative relationship between the type of reduce the amount of rainfall entering the system. debris present within a shoreline transect, determined by mean percentage cover per week, and the presence with relative abun- Data recording and statistical analysis dance of Coleopteran families. It was uncertain whether individ- Where possible, beetles were identified on-site, recorded and ual species would appear in enough abundance to allow for released. If a specimen could not be identified it was preserved individual species analysis; Therefore the data were combined ............................................................................................... .................................................................. 3 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. melanocephalus (L.) with 107 recordings while Phylan gibbus (Fabricius), a specialist halobiontic species, was recorded 59 times. The families Carabidae, Staphylinidae and Curculionidae were particularly well represented in the pitfall traps with 14, 10, and 8 species identified in each family, respectively. Forty- three species were solely found on the sand dune system. Community structure and comparison Fig. 3 shows a simple correspondence analysis of community structure. All of the species identified were plotted against the three sample locations: sand dunes, outer shoreline and inner shoreline. When a species (blue dot) is close to the centre point (0,0), it is relatively evenly distributed between the shoreline community and the sand dune community. Species found only in one community occupy the same position in Fig. 3, which has been represented by enlarged blue dots and a corre- sponding number signalling the number of species plotted in the same location. If a species is close to a community centre (red square), then it is unevenly distributed between the com- Figure 3. Simple correspondence analysis illustrating the different munities. Few species were found present in both the sand beetle community structures (red squares) gathered from the sand dune and shoreline habitats, with only two species occurring dunes, the inner shoreline and the outer shoreline. Seventy-seven multiple times in each habitat. Therefore, the shoreline cole- percentage of species (43/56) identified were solely found in the sand opteran community can be defined as a separate community dune habitat while only 12.5% (7/56) were found on both the shoreline to the sand dune community rather than an extension to the and sand dune habitats. Positions of individual species in the two dune community. The inner and outer shoreline species show dimensional space are represented by small blue dots. Enlarged blue subtle differences but had proportionately more species that dots have a corresponding number relative to the number of species within the blue dot. The arrows represent the strength of distribution were present in both communities. towards each community where the centre point (0,0) represents an Table 1 shows results of regression analyses of beetle abun- even distribution between the shoreline’s community and sand dune community. Few species were found present in both the sand dune dance on amounts of different types of debris along the shore- and shoreline habitats, with only two species occurring multiple times line. On the outer shoreline, beetles from all families showed in each habitat. Therefore, the shoreline coleopteran community can a significant positive relationship with the number of pebbles be defined as a separate community to the sand dune community in a quadrat. Carabidae were also more likely to be found rather than an extension to the dune community. The inner and outer when more driftwood or seaweed was present. Staphylinidae shoreline species show subtle differences but had proportionately were associated with seaweed. The lowest beetle counts were more species that overlapped communities. found from bare sand. Fewer associations were found between beetle number and debris on the inner shoreline into families to identify how different families associate with although Carabidae were still closely associated with the debris. Life-history traits of the shoreline species identified were quantity of driftwood present. Beyond that the only associ- reviewed to determine the effectiveness of shoreline sampling. ation was a weak relationship between Tenebrionidae and All data were analysed using Mintab version 18.1. amount of seaweed. Discussion Results It has been widely acknowledged that coastal habitats face a Species summary significant threat of being lost as a result of rising sea levels A total of 56 species spread across 15 families were identified and a squeeze against coastal communities (Doody, 2004; (Table A1). Thirteen species were found along the shoreline; Doxa et al., 2017). One of the aims here was to distinguish seven species were found more than once and only four of these the relationship shoreline communities have with their neigh- were recorded more than five times: Broscus cephalotes (L.), bouring habitat, in this case, sand dunes. Fig. 3 shows these Calathus cinctus (Motschulsky), Phaleria cadaverina (Fabricius) neighbouring local habitats had little overlap of species and and Cafius xantholoma (Gravenhorst). Of the 50 species identi- therefore could be defined as separate communities. Looking fied in the sand dune system, 14 were recorded only once, while at the species identified from the shoreline, many are known only eight of these species were recorded more than 10 times. for being strandline specialists. Therefore, these shoreline Twenty species were a first recording for Dawlish Warren. coleopteran communities are facing a significant threat of The most common sand dune species was Calathus local extinctions where species migration perpendicular to the ............................................................................................... .................................................................. 4 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Bioscience Horizons � Volume 11 2018 Research article ............................................................................................... .................................................................. Table 1. Results of linear regression analyses [F, R-squared (%) and p values] of beetle abundance on the amount of debris along the shoreline over time (df = 1,12, significant values in bold) Bare sand Pebbles Driftwood Seaweed O I OI O I OI Carabidae F-stat 0 0.05 6.43 1.35 9.57 32.5 12.5 0.75 R-sq 0 0.38 34.9 10.1 44.4 73.0 51.1 5.88 p value 0.99 0.83 0.03 0.27 0.01 0.001 0.004 0.40 Tenebrionidae F-stat 0 0.63 8.95 3.47 0 1.74 4.22 5.16 R-sq 0 4.99 42.7 22.5 0.01 12.7 26.0 30.1 p value 0.99 0.44 0.01 0.09 0.97 0.21 0.06 0.04 Staphylinidae F-stat 0.03 0.19 14.7 0.05 0.12 1.23 8.17 0.75 R-sq 0.21 1.59 55.0 0.37 1.00 9.28 42.1 5.86 p value 0.87 0.67 0.002 0.83 0.73 0.29 0.01 0.40 Only relatively abundant families are considered (five or more recordings). O, outer shoreline; I, inner shoreline. coast is prevented as a result of a ‘coastal squeeze’. Furthermore, Twenty species new to Dawlish Warren were found, this study shows little presence of these shoreline specialists in a illustrating the importance of finding ways to carry out on- similar neighbouring habitat meaning the shoreline communities site surveys in protected areas. Dawlish Warren is a very could not be used to ‘seed’ populations on the shoreline. special site so it comes as no surprise that it holds a wide range of species. This study was limited both in terms of It is well known that shoreline walking can reveal extensive time and spatial scale and therefore further, more extensive numbers of beetles. One of the aims here was to identify how surveys will certainly reveal the presence of more species. shoreline coleopteran species are associated with coastal debris Some of these species might be scarce or restricted to and to consider whether shoreline Coleoptera could be used to sur- Dawlish and help to inform appropriate management in the vey the wider site removing the need to set pitfall traps. During the field (Teignbridge District Council, 2010). Without this study, a much wider range of species were found in the dune sys- knowledge it is possible that current management techni- tem than along the shoreline. This suggests that not all of the beetle ques could be disadvantageous to one or more potential species in the dunes take to the air readily and expose themselves speciesofinterest. to the risk of being stranded along the shoreline. Some dune spe- cies certainly do fly and get trapped on the shoreline as shown by Reticence among wardens of nature reserves to allow sur- the presence of two dune-dwelling species of Curculionidae, veys using lethal pitfall trapping to sample invertebrate com- Polydrusus cervinus (L.) and Coelositona cambricus (Stephens). In munities is frequently encountered. This degree of caution is addition, although the shoreline represents very harsh and danger- usually a result of one or more scarce target species on the ous conditions, some beetle species are specialists in this type of reserve that might be susceptible to being caught in the pitfall environment (Topp and Ring, 1988; Brown, 1996), such as traps. In this case, the species of concern was sand lizard, Cafius xantholoma (Staphylinidae) (Moore and Legner, 1976). A L. agilis. To counter this concern a modified pitfall trap was number of beetle species found along the shoreline appeared to be designed that facilitated the trapping of invertebrates live to seeking out certain features, such as pebbles and driftwood either release on-site if the species was known or to selectively (Table 1). Presumably, these structures were offering refugia or return to the laboratory for subsequent examination. The traps foraging opportunities. The outcome is that collecting along the appeared to work well although we do not know about per- shoreline cannot serve as an appropriate alternative to on-site sam- formance relative to lethal traps as the latter could not be set. pling, in this case using pitfall traps. No sand lizards or other vertebrates were caught in the traps. ............................................................................................... .................................................................. 5 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. Doxa, A., Albert, C. H., Leriche, A. et al. (2017) Prioritizing conservation Acknowledgements areas for coastal plant diversity under increasing urbanization, Journal of Environmental Management, 201, 425–434. We would like to thank Devon Wildlife Trust and Teignbridge District Council for allowing access to Dawlish Warren National Duff, A. (2012) Beetles of Britain and Ireland. Volume1: Sphaeriusidae to Nature Reserve and providing permission to conduct this research. Silphidae, A.G.Duff, Norfolk. We would also like to extend our thanks to Philip Chambers and Duff, A. (2016) Beetles of Britain and Ireland. Volume 4: Cerambycidae to the team for their help during the research on the site. Curculionidae, A.G.Duff, Norfolk. Fanini, L. and Lowry, J. (2016) Comparing methods used in estimating Author biography biodiversity on sandy beaches: pitfall vs. quadrat sampling, Ecological Indicators, 60, 358–366. Ellis Armstrong studied for a degree in Ecology and Wildlife Greenslade, P. J. M. (1964) Pitfall trapping as a method for studying Conservationgraduationin2018withaparticularinterestin populations of Carabidae (Coleoptera), The Journal of Animal Coleoptera. During the academic year 2018/2019, he will be Ecology, 33, 301–310. studying towards a Masters by Research in Entomology. 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(2017) Identifying Beetles, accessed at: http://www. sand on vegetated surfaces, Earth Surface Processes and Landforms, markgtelfer.co.uk/beetles/ (8 December 2017). 11, 505–514. ............................................................................................... .................................................................. 7 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Research article Bioscience Horizons � Volume 11 2018 ............................................................................................... .................................................................. Appendix Table A1. Summary of the 56 Coleoptera species found at Dawlish Warren and their abundance in each of the habitats sampled Family Genus Species Sand dunes Outer shoreline Inner shoreline Carabidae Broscus cephalotes 48 4 Carabidae Syntomus foveatus 20 0 0 Carabidae Amara tibialis 22 0 0 Carabidae Dyschirius thoracicus 10 0 Carabidae Calathus melanocephalus 107 0 4 Carabidae Harpalus affinis 10 0 Carabidae Nebria salina 10 0 Carabidae Bembidion quadrimaculatum 10 0 Carabidae Calathus cinctus 11 12 Carabidae Pterostichus madidus 11 0 Carabidae Calathus fuscipes 20 0 Carabidae Trechus obtusus 20 0 Carabidae Trechus fulvus 10 0 Carabidae Pterostichus longicollis 40 0 Leiodidae Catops grandicollis 60 0 Leiodidae Agathidium Seminulum 10 0 Leiodidae Leiodes furva 20 0 Leiodidae Leiodes sp. 1 0 0 Tenebrionidae Phaleria cadaverina 029 37 Tenebrionidae Phylan gibbus 59 0 0 Hydrophilidae Anacaena globulus 20 0 Hydrophilidae Megasternum concinnum 60 0 Hydrophilidae Cercyon littoralis 01 0 Curculionidae Otiorhynchus atroapterus 13 0 0 Curculionidae Otiorhynchus rugifrons 40 0 Curculionidae Rhinonchus castor 50 0 Curculionidae Coelositona cambricus 30 1 Curculionidae Philopedon plagiatum 20 0 Curculionidae Orthochaetes insignis 10 0 Continued ............................................................................................... .................................................................. 8 Downloaded from https://academic.oup.com/biohorizons/article/doi/10.1093/biohorizons/hzy017/5304540 by DeepDyve user on 18 August 2022 Bioscience Horizons � Volume 11 2018 Research article ............................................................................................... .................................................................. Table A1. Continued Family Genus Species Sand dunes Outer shoreline Inner shoreline Curculionidae Pselactus spadix 10 0 Curculionidae Polydrusus cervinus 01 0 Chrysomelidae Longitarsus succineus 10 0 Chrysomelidae Longitarsus flavicornis 30 0 Chrysomelidae Cryptocephalus pusillus 11 0 0 Elateridae Agrypnus murinus 90 0 Apionidae Apion haematodes 90 0 Apionidae Perapion hydrolapathi 20 0 Histeridae Hypocaccus dimidiatus 12 1 Histeridae Saprinus aeneus 10 0 Silphidae Silpha tristis 20 0 Latridiidae Corticaria sp. 1 0 0 Oedemeridae Nacerdes melanura 02 1 Coccinellidae Scymnus frontalis 10 0 Coccinellidae Coccinella septempunctata 60 0 Coccinellidae Coccinella undecimpunctata 10 0 Aphodiidae Aphodius foetens 40 0 Staphylinidae Ocypus olens 18 0 0 Staphylinidae Cafius xantholoma 012 3 Staphylinidae Cafius fucicola 01 0 Staphylinidae Ocypus brunnipes 30 0 Staphylinidae Platydracus stercorarius 10 0 Staphylinidae Stenus clavicornis 10 0 Staphylinidae Lordithon thoracicus 10 0 Staphylinidae Mycetoporus sp. 1 0 0 Staphylinidae Gabrius sp. 2 0 0 Staphylinidae Aleocharinae sp. 10 0 1 ............................................................................................... ..................................................................

Journal

BioScience HorizonsOxford University Press

Published: Jan 1, 2018

References