wildlife surveys chapter 5: pitfall results with specific

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1 RSPB/NE Countdown 2010: Bringing Reedbeds to Life Project Wildlife surveys CHAPTER 5: Pitfall results with specific reference to the Coleoptera C J Hardman, D B Harris With helpful comments on draft by Alan Stubbs (Buglife) Contents SUMMARY ............................................................................................................................................... 1 METHODS ................................................................................................................................................ 2 Pitfall survey field methods .................................................................................................................... 2 Analysis methods .................................................................................................................................... 4 RESULTS .................................................................................................................................................. 6 Composition of species caught ............................................................................................................... 6 Habitat associations with overall species diversity................................................................................. 7 Habitat associations with wetland specialists in pitfall traps ............................................................... 12 Habitat associations with pitfall conservation scores........................................................................... 15 Habitat associations with Coleoptera species diversity in pitfall traps ................................................ 18 Habitat associations with wetland specialist Coleoptera species in pitfall traps ................................. 23 Habitat associations with Coleoptera conservation scores in pitfall traps........................................... 26 Composite habitat variables ................................................................................................................. 29 References ............................................................................................................................................ 31 Appendix to invertebrate results .......................................................................................................... 31 SUMMARY Species diversity 24 pitfall were set at Stodmarsh, 22 at Hickling Broad and 12 at Ham Wall, all points were dry during surveys (August). At least 7 421 individual invertebrates were trapped in total: 41 % Coleoptera (beetles), 37% Araneae (spiders), 12% Hymenoptera (bees, ants, sawflies and wasps), 5% Stylommatophora (snails and slugs) and more. 288 species were identified including 26 Rare or Nationally Scarce. One Endangered rove beetle: Quedius balticus, a Vulnerable spider: Clubiona juvenis and a Data Deficient rove beetle Philhygra terminalis were trapped. 132 wetland specialists and 4 reedbed specialists were trapped. There was not enough variation in number of reedbed specialists per trap for analysis.

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1

RSPB/NE Countdown 2010: Bringing Reedbeds to Life Project

Wildlife surveys

CHAPTER 5: Pitfall results with specific reference to the Coleoptera

C J Hardman, D B Harris

With helpful comments on draft by Alan Stubbs (Buglife)

Contents

SUMMARY ............................................................................................................................................... 1

METHODS ................................................................................................................................................ 2

Pitfall survey field methods .................................................................................................................... 2

Analysis methods .................................................................................................................................... 4

RESULTS .................................................................................................................................................. 6

Composition of species caught ............................................................................................................... 6

Habitat associations with overall species diversity................................................................................. 7

Habitat associations with wetland specialists in pitfall traps ............................................................... 12

Habitat associations with pitfall conservation scores ........................................................................... 15

Habitat associations with Coleoptera species diversity in pitfall traps ................................................ 18

Habitat associations with wetland specialist Coleoptera species in pitfall traps ................................. 23

Habitat associations with Coleoptera conservation scores in pitfall traps ........................................... 26

Composite habitat variables ................................................................................................................. 29

References ............................................................................................................................................ 31

Appendix to invertebrate results .......................................................................................................... 31

SUMMARY

Species diversity

• 24 pitfall were set at Stodmarsh, 22 at Hickling Broad and 12 at Ham Wall, all points were dry during surveys (August).

• At least 7 421 individual invertebrates were trapped in total: 41 % Coleoptera (beetles), 37% Araneae (spiders), 12% Hymenoptera (bees, ants, sawflies and wasps), 5% Stylommatophora (snails and slugs) and more.

• 288 species were identified including 26 Rare or Nationally Scarce.

• One Endangered rove beetle: Quedius balticus, a Vulnerable spider: Clubiona juvenis and a Data Deficient rove beetle Philhygra terminalis were trapped.

• 132 wetland specialists and 4 reedbed specialists were trapped. There was not enough variation in number of reedbed specialists per trap for analysis.

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Relationship to habitat variables

Overall diversity in pitfall traps (measured by bootstrapped species richness) tended to be higher in traps that were surrounded by a greater diversity of plants.

Greater numbers of wetland specialists (both across all species and for Coleoptera alone) were trapped at points surrounded by taller, thicker and denser reed.

Relationships between conservation score across all species and habitat variables did not produce clear results. When conservation scores were calculated just for Coleoptera, trends became evident at Stodmarsh but not at the other sites. Traps with higher conservation scores at Stodmarsh tended to be surrounded by thicker and taller reed stems, and were closer to scrub.

Number of Coleoptera species per trap was higher at Stodmarsh than the other two sites. Site dominated the analysis and trends. However at Stodmarsh alone, points with taller, thicker reed stems, lower plant species richness and shallower litter tended to trap more Coleoptera species. Only three traps had litter that had been fully saturated in the months before surveys, but these points trapped a similar average number of Coleoptera species to dry points.

Since pitfall traps had to be placed in dry areas during the survey month of August, 29 of the points had a dry seasonal litter saturation category (May to August) and only 3 had litter that was totally flooded in this period. 11 had partially flooded litter in the months before surveys, but these were all at Hickling Broad. Seasonal water level information for this time period was not available for 9 of the points. Therefore overall conclusions about the relationship between ground-dwelling invertebrates and reedbed hydrology are limited.

Trends derived from algorithmic modelling were compared to trends from principle components analysis and similar trends were seen. Sampling points that caught reedbed specialist invertebrates did not cluster together based on their environmental attributes.

Pitfall results were more limited in their interpretation than water trap/moth results. Pitfall traps caught a more varied assemblage of invertebrates in terms of the number of orders of invertebrates within samples. There were fewer reedbed specialists than in water traps or moth traps so the analysis is very coarse scale, looking at a wide variety of generalist species. Attempts to narrow the analysis to wetland specialists, Coleoptera or species with a conservation status created clearer trends. Future work should focus on what representation of reed ecologies are needed to embrace all the scarce and special species in viable populations may not be possible for species with too few specimens to give a sound answer. The results were limited by only a few sites being surveyed, the sites having large differences in habitat variables and geography, and the range of habitat variables within each site being a narrow section of the entire spectrum within reedbeds (dry points sampled predominantly). However the result about wetland Coleoptera being associated with taller, thicker denser reed seems one of the firmest conclusions.

METHODS

Pitfall survey field methods

Pitfall traps were set at 24 locations at Stodmarsh, 22 locations at Hickling Broad and 12 locations at Ham Wall. At Hickling Broad, pitfalls were set between 13-14th August 2009 and 29-30 August 2009. At Ham Wall, pitfalls were put out on 11th August 2009 and collected on 27th August 2009. At

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Stodmarsh, pitfalls were put out on 8-9th August 2009 and collected on 24-25 August 2009. Each trap was left out for 16 days.

The trap points were randomly located within reedbed, at a minimum of 50m separation for sample independence, so as not to interfere with each others' catches (Telfer, M pers. comm.). Sample sizes were constrained by the availability of dry reedbed, which explains the lower sample size at Ham Wall where less dry reedbed was available. However, where possible we tried to distribute the pitfall trapping points between reedbed that was dry all year or periodically dry (and obviously dry at the time of trapping). The aim, as with the water trap survey plan, was to obtain a representative sample for analysis of seasonal water levels rather than a comparison between hydrological categories per se. Again, the points that were perceived as dry all year or periodically dry were based on available information at the time, from our water trapping work and from annual hydrology information from site managers, but more detailed water level information was collected during the survey.

The original stratification was:

Stodmarsh: 12 pitfalls in seasonally dry reedbed and 12 in permanently dry reedbed

Hickling Broad: 10 fringing broad and 10 on reserve reedbed (all seasonally dry reedbed)

Ham Wall: 6 traps in seasonally dry reedbed, 6 in adjacent dry, recently cut reed stubble

Pitfall trap locations were not randomly assigned but positioned according to our knowledge of water level fluctuations and accessibility. Disturbance to nesting birds was always a concern and we worked closely with site managers to avoid sensitive areas. If the mapped point fell close to a habitat feature which may bias the sample, the point was relocated by at least 6m to the south side of any tree/scrub or away from a pond/ditch. All pitfall trap points were placed at least 10m from edge habitat, paths, or open water. These measures should have limited the bias from specialist communities or edge effects. All pitfall trap points were also placed a minimum distance of 15m from previous access routes and survey locations to eliminate any bias caused by open, disturbed micro-habitat.

The traps were 400ml plastic flip top containers (Alana Ecology) that were dug in to the ground ensuring that the soil is compact and flush to the rim. The containers were 1/3 full with propylene glycol solution (50%), an antifreeze solution that is non-toxic to vertebrates and acts as a preservative until the traps were collected. A couple of drops of washing up liquid were added to reduce surface tension so that invertebrates didn’t float or escape. We added a plastic garden mesh cover and pinned it down with wire pegs to prevent larger creatures falling in and drowning. Unfortunately this still happened on several occasions (frogs, shrews and a harvest mouse were found).

Figure 5.1: Pitfall trap in situ (Anna Doeser)

On approaching the trap to collect the contents, the surveyor noted the presence of any vegetation protruding into the pitfall trap that may have aided escape of trapped invertebrates. The liquid level was also checked and non-target species capture was recorded. Samples were washed through filter mesh and stored in 60% Industrial Methylated Spirits. Surveys were designed by Donna Harris and carried out by Donna Harris and Anna Doeser.

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Sorting specimens to Order level was done by Donna Harris and John Baxter in autumn 2009. Any non-invertebrate material was removed. The remainder of the sample consisted mainly of Coleoptera (beetles) but also Hemiptera (true bugs). Coleoptera were stored in 60% IMS solution and others were stored in 70% IMS solution. Coleoptera were identified by Mark Telfer, who discarded larvae. Ian Dawson identified spiders. Aculeate hymenoptera (bees, wasps and ants) were identified by Mike Edwards and Symphata (sawflies) by David Sheppard. Parasitic Hymenoptera were identified where possible by Gavin Broad. Diptera were not identified as the water trap survey was thought sufficient for this group.

Habitat surveys

During pitfall trap set-up two 50 x 50 cm reed quadrats 2m north and 2m south from the pitfall were surveyed. In each quadrat, dead and live reed stems and stems with panicles were counted, four reed stem base diameters and reed height (dead and live) were measured using the same method as in water trap habitat surveys. 16 days later during trap collection, plant surveys were carried out. Eight 50 x 50cm quadrats at N, NE, E, SE, S, SW, W, NW, directions at 2m from the pitfall trap were surveyed and all plants within the quadrat recorded. Litter depth (surface to silt/root layer) was measured at 1m from the pitfall en route to every other plant quadrat (N, E, S, W).

Analysis methods

Invertebrate species data

Analysis methods were as described in water trap samples. The response variables available were number of species, bootstrapped number of species, wetland specialists, conservation scores, number of Coleoptera species, number of wetland Coleoptera species, Coleoptera conservation scores. Number of species was bootstrapped to 26 individuals, based on minimum sample sizes for pitfall traps. When Coleoptera and wetland specialists were analysed alone, these subsets were not bootstrapped due to sample sizes being too small. There were not enough reedbed specialists caught per trap to analyse as a response variable. Only 9 traps contained reedbed specialists and the number of reedbed specialists per trap varied between 0 and 2.

For wetland specialists, lists created from experts were used for consistency of methodology but it should be noted that unlike Diptera and Lepidoptera, the lists created from ISIS matched expert lists well for pitfall samples. 129 species trapped in pitfall samples were listed on ISIS under BAT categories W2 and W3. The lists provided by experts had 132 wetland specialists, 114 of which were also on the ISIS list. The habitat association results not changing depending on which list was used. The wetland specialist list from experts was composed of 81 % Coleoptera 11 % Araneae plus small numbers of Basommatophora, Hemiptera, Stylommatophora, Isopoda and Orthoptera species.

As for water trap analysis, a conservation score for each trap was calculated using:

Score of 10 for: Vulnerable, RDB2 (similar to vulnerable), RDB3 (similar to near threatened), RDB1 (similar to endangered), Rare

Score of 5 for: UK BAP, Scarce, Notable, Data deficient

Each species score was only counted once per trap. For pitfall traps, species contributing to conservation scores were predominantly Coleoptera (24 species) and Araneae (5 species).

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Habitat data

From the habitat variables recorded in the field, the following set of explanatory variables was derived. Averages were calculated for habitat variables that were measured at more than one point around the sampling location.

Table 5.1: Explanatory variables used in models relating habitat variables to invertebrate species diversity in pitfall traps

Water level data

Pitfall traps had to be placed in dry areas, so information on the wetness of sampling points is limited. A litter saturation variable was created using water levels measured between May and August 2009 as these had the maximum amount of data available. Litter levels measured in October 2009 (as these correspond to zero AOD) were compared to the water level predictions.

Variable Type Description

Litter saturation Explanatory categorical 3 categories; dry, partial and wet

Dry if predicted water level (May to Aug 2009) never exceeds zero. Partial if predicted water level greater than zero but less than litter depth. Wet if predicted water level above litter depth.

Litter depth Explanatory continuous Average of 4 litter depth measurements in 1 m radius of trap point.

Total stem density Explanatory continuous Average of 4 counts of number of stems in 50 x 50 cm quadrat, 1 m away from trap point.

Dead stems percentage Explanatory continuous percentage

Percentage of total stems that were dead

Mean reed height Explanatory continuous Mean height of live and dead reed, average of 4 measures in 1m radius of trap point.

Stem diameter Explanatory continuous Stem diameter at base, average of 4 measures in 1m radius of trap point.

Distance to scrub Explanatory continuous Distance to nearest patch of scrub/trees calculated from aerial photos by CDMU using an add-on tool in MapInfo 6.5

Plant species richness Explanatory continuous Total number of plants (excluding Phragmites australis) found over four 50 x50 cm quadrats in 1 m radius of trap point.

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Since dry points were chosen for sampling, there was little variation in litter saturation in this seasonal period. Hickling Broad had 1 wet, 10 dry and 11 partially saturated and points, Ham Wall points were all dry, and Stodmarsh had 2 wet points and the remainder dry. At Stodmarsh, lack of gauge board reading between May and August 2009 meant no litter saturation category was available for 9 points. Therefore when interpreting the results we must note that the sampling points were biased towards dry areas. Therefore the habitat surveyed around pitfall traps represents a narrow spectrum within the overall environmental gradients in reedbeds (diameters were thinner and heights shorter than average reedbed points in audit data). Six of the points at Ham Wall were in reed stubble so were not considered comparable enough to others for analysis.

RESULTS

Notable records from pitfall traps

Some notable Coleoptera records were obtained. In the pitfall traps at Hickling Broad, Boreophila eremite was a new record for Norfolk and the finding of Baeocrara variolosa made the fourth modern record of (the previous three of which were from Hickling).

Further information on some of the invertebrates with conservation statuses in pitfall traps

Quedius balticus (Staphylinidae) Order: Coleoptera

This rove beetle is endangered, classed as a reedbed specialist on the ISIS database and included on the Buglife list of notable species associated with fen habitats. In the analysis of the Broads Fen Invertebrate Survey 2007-09, Quedius balticus was noted as a species of conservation interest that could be affected by changes in site management. It was recorded once at each of ten pitfall points at Hickling Broad, both in the reed fringing the broad and in Hundred Acre reedbed (for grid references and coordinates see appendix).

Eight out of ten points where Quedius balticus was trapped had litter that was predicted to be dry in the four months before trapping (HBPDD6 was partially saturated and HPPDD8 was wet). Points where Quedius balticus was found had slightly higher density of live stems, slightly deeper litter and were slightly closer to scrub than the average of all sampling points at Hickling Broad. This was not tested statistically due to the uneven sample size between points with and without Quedius balticus.

Paradromius longiceps (Carabidae) Order: Coleoptera

This nationally scarce Carabid ground beetle is a reedbed specialist. One individual was recorded from dry reedbed at Hickling Broad where it was trapped at pitfall trap HBPDD3.

Composition of species caught

There were 7 421 individuals caught in pitfall traps, 41% of which were Coleoptera (beetles), 37% of which were Araneae (spiders and harvestmen), 12% of which were Hymenoptera (wasps and bees), 5 % of which were Stylommatophora (air-breathing snails and slugs) and the remaining 5 % of which were Basommatophora (air-breathing snails and slugs), Diptera (true flies), Hemiptera (true bugs), Isopoda ( e.g woodlice) and Orthoptera (grasshoppers, crickets and locusts). Note there were more Diptera than this but pitfall Diptera samples were not identified. A total of 289 species were caught in pitfall traps over the three sites. At least 7 421 individual invertebrates trapped in total: 41 % Coleoptera (beetles), 37% Araneae (spiders), 12% Hymenoptera (bees, ants, sawflies, wasps), 5% Stylommatophora (snails and slugs) and more. 26 Rare or Nationally Scarce species were recorded and several red listed species (one Endangered rove beetle: Quedius balticus, a Vulnerable spider:

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Clubiona juvenis , a Data Deficient rove beetle Philhygra terminalis). 132 wetland specialists and 4 reedbed specialists were trapped.

Differences in number of species between sites

Stodmarsh had the greatest number of species and abundance of the three sites. It also had the greatest variance of number of species between sampling points. When abundance was accounted for by bootstrapping, Hickling Broad and Stodmarsh had similar number of species. Ham Wall had the lowest bootstrapped number of species and average abundance but fewer traps were set at this site due to a lack of suitable dry habitat. Considering there were almost half the number of traps set out at Ham Wall, and because it is a much newer site, we would have expected even lower diversity than that recorded (table 5.2).

Table 5.2: Measures of number of species per trap per site

Hickling Broad Ham Wall Stodmarsh

Number of traps (n=22)

(n=12 but 6 used in analysis)

(n=24)

Total number of species 136 113 196 Total abundance of individuals

1563 1192 3280

Average of richness 22.05 26.17 33.08

Variance of richness 45.19 30.57 105.04

Average of abundance 71.05 117.17 136.67

Average of bootstrap 8.44 7.56 8.43

Habitat associations with overall species diversity

Number of species

Analysis relating number of species per trap to habitat variables in the vicinity were highly site dependent so general conclusions were limited. Many trends were evident mostly at Stodmarsh where there was a higher number of species trapped in pitfalls. Habitat associations for number of species have not been reported because they were skewed by a few points at Stodmarsh so were unrepresentative of wider trends.

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Figure 5.2: Relative importance of habitat variables in explaining variation in number of species in pitfall traps. These models explained 90 % of the variance.

Site

Stodmarsh trapped a higher number of species per trap on average than the other two sites.

Figure 5.3: Relationship between number of species per pitfall trap and site

Bootstrapped number of species

When number of species was bootstrapped to a set number of individuals per trap, the following relationships with habitat variables emerged.

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Figure 5.4: Variable importance for bootstrapped number of species in pitfall traps. These models explained 90 % of the variance.

Plant species richness

Points with a greater number of plants were associated with greater bootstrapped number of species. Plant species richness was important at Hickling Broad but not at Stodmarsh, where many points did not have any other plant species present except reed. There were not enough traps set at Ham Wall to test if it was important at this site alone.

Figure 5.5: Relationship between plant richness and bootstrapped number of species

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Stem density

The model showed stem densities below 250 stems per m2 to be associated with higher species diversity in pitfall traps. However general conclusions should not be drawn because the relationship was not clear cut and will be a result of many different relationships with different taxonomic groups.

Figure 5.6: Relationship between stem density and bootstrapped number of species in pitfall traps.

Mean reed height

The model showed a mid-range of reed heights to be associated with high diversity in pitfall traps.

The trend is not clear, and this is probably due to both the coarse scale of the species data (including many taxonomic groups) and the variation in reed height between sites.

Figure 5.7: Relationship between mean reed height and bootstrapped number of species

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Dead stems percent

The model showed points with a lower proportion of dead stems were related to a higher bootstrapped number of species. However, this includes trends with a whole range of species ecologies, both phytophagous on live reed and saprophagous. Perhaps this trend reflects a greater number of invertebrates being trapped that are dependent on live reed. Further work into the ecologies of the individual species would be needed to investigate this trend in more detail.

Figure 5.8: Relationship between dead stems (%) and bootstrapped number of species in pitfall traps

Site

The model showed there were not large differences between bootstrapped number of species per pitfall traps at the different sites. The raw data showed Ham Wall had lower species diversity per trap than the other two sites; this is likely to be affected by the smaller sample size at Ham Wall.

5.9: Relationship between site and bootstrapped number of species in pitfall traps.

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Litter depth

The very shallowest litter depths were associated with lower diversity of invertebrates in pitfall traps. This may be because litter above a certain depth provides an advantage of shelter (from weather, poor light resting places or perhaps avian predation). At Hickling Broad, points with deeper litter were associated with a greater diversity of species in pitfall traps.

Figure 5.10: Relationship between bootstrapped number of species and litter depth in pitfall traps

Habitat associations with wetland specialists in pitfall traps

Figure 5.11: Importance of habitat variables in describing variation in number of wetland specialist species. These models explained 89% of the variance.

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Plant species richness

In general, points with lower plant species richness were related to points with a greater number of wetland specialist species trapped in pitfalls. Stodmarsh points were often in areas where the only plant present was reed (plant richness score 0). Points at this site also had high numbers of wetland specialist species in pitfall traps so the trend may be exaggerated by this confounding factor. The trend was not evident when sites were analysed separately.

Figure 5.12: Relationship between plant richness and number of wetland specialists in pitfall traps

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Stodmarsh had a greater number of wetland specialist species trapped in pitfalls than the other two sites.

Figure 5.13: Relationship between site and number of wetland specialist species in pitfall traps

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Points with greater reed stem densities were associated with higher numbers of wetland specialist species. However this trend was not evident when sites were analysed separately. A relationship with higher stem densities could be expected because this would provide more shelter for invertebrates. Another explanation could be that dense stems tend to trap more invertebrates because more get blown in to this habitat where the air is more stable.

Figure 5.14: Relationship between stem density and number of wetland specialist species in pitfall traps.

Mean reed height

In general points where the reed was taller were associated with a greater number of wetland specialist species in pitfall traps. (This was true for Hickling Broad and Stodmarsh analysed separately).

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Figure 5.15: Relationship between mean reed height and number of wetland specialist species in pitfall traps

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Diameter

Traps surrounded by thicker reed stems tended to trap more wetland specialists in pitfall traps.

However this trend was not important when each site was analysed individually.

Figure 5.16: Relationship between reed stem diameter and number of wetland specialists in pitfall traps

Habitat associations with pitfall conservation scores

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Figure 5.17: Relative importance of habitat variables in explaining the total conservation score in each pitfall trap. These models explained 93 % of the variance.

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Diameter

Points at Stodmarsh and Hickling Broad where the reed stems were thicker tended to be associated with more species with higher conservation statuses.

Figure 5.18: Relationship between diameter and conservation score of Coleoptera in pitfall traps.

Plant species richness

The model indicates points with lower plant species richness are associated with higher conservation scores. However the raw data suggests this is mainly due to points at Stodmarsh having low plant richness and high conservation scores.

Figure 5.19: Relationship between plant species richness and conservation score of pitfall traps

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13

.51

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14

.5

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Plant richness

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Actual data

Plant richness

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Litter saturation

Points that were dry between May and August 2009 tended to be associated with high conservation scores in pitfall traps. However this is biased by the tendency for pitfall traps to be placed in areas that were dry in August.

Figure 5.20: Relationship between litter saturation category and conservation score in pitfall traps

Litter depth

The model suggests points with deeper litter tended to be associated with more species with higher conservation statuses in pitfall traps. But the scatter plot shows a wide range of conservation scores were found throughout the different litter depths so further work is recommended.

Figure 5.21: Relationship between litter depth and conservation score in pitfall traps

5 10 15 20

11

.01

1.5

12

.01

2.5

13

.01

3.5

Partial Plot

Litter depth

5 10 15 20

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10

15

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35

Actual data

Litter depth

Co

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Mean reed height

The model suggests that up to a point, taller reed was associated with greater conservation scores. This was most evident at Stodmarsh. However the trends relating to mean reed heights were very site specific, therefore general conclusions cannot be drawn.

Figure 5.22: Relationship between mean reed height and conservation score in pitfall traps

Habitat associations with Coleoptera species diversity in pitfall traps

Since Coleoptera made up 52% of the species caught and identified in pitfall traps, there was scope to analyse Coleoptera alone to make deductions about the habitats associated with greatest Coleoptera diversity and abundance at the sites. The number of species was not bootstrapped as the minimum number of species in any one trap was three, which was too small to make meaningful bootstrapped numbers.

Differences between sites

Stodmarsh had the highest number of species, abundance and variance of number of species. This was followed by Hickling Broad in total number of species and abundances, but Ham Wall was second in terms of average number of species and abundance.

Table 5.3: Coleoptera number of species and abundances at each site

Hickling Broad Ham Wall Stodmarsh

(n=22) (n=12 but 6 used in random forest models)

(n=24)

Total number of Coleoptera species 78 69 116

Total abundance of Coleoptera individuals 552 667 1802

1.0 1.5 2.0 2.5

12

.01

2.5

13

.01

3.5

Partial Plot

Mean reed height

1.0 1.5 2.0 2.5

05

10

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35

Actual data

Mean reed height

Co

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67

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standing.water.summer.2010

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19

Average of Coleoptera number of species 9.73 12.58 16.46

Variance of Coleoptera number of species 17.06 3.72 30.43 Average of abundance of Coleoptera individuals

25.09 55.58 75.08

Figure 5.23: Importance of habitat variables in explaining variation in number of Coleoptera species in pitfall traps. These models explained 90% of the variance.

20

Site

Of the three sites, Stodmarsh had the most Coleoptera species in total; however Ham Wall was not represented as well as the other two sites so it is not an entirely even comparison. The differences in numbers of Coleoptera per pitfall trap were best explained by which site the trap was located at. Stodmarsh had the greatest number of Coleoptera species per pitfall trap on average. Other habitat variables were of much lower importance in explaining variation Coleoptera diversity between traps.

Figure 5.24: Differences in number of Coleoptera species caught in pitfall traps and site

Mean reed height

There appears to be a tendency for points with taller reed to be associated with more Coleoptera species. This was most evident at Stodmarsh and further work is recommended to see if this trend is true at a wider range of sites.

21

Figure 5.25: Relationship between mean reed height and number of Coleoptera species in each

pitfall trap

Plant species diversity

At Stodmarsh, points with fewer plant species were associated with more Coleoptera species, however this trend was not seen at other sites.

Figure 5.26: Relationship between plant species richness and number of Coleoptera in pitfall traps

1.0 1.5 2.0 2.5

12

.21

2.4

12

.61

2.8

13

.01

3.2

Partial Plot

Mean reed height

1.0 1.5 2.0 2.5

51

01

52

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53

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Actual data

Mean reed height

No

. o

f C

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0 2 4 6 8 10 12

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Plant richness

0 2 4 6 8 10 12

51

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52

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Actual data

Plant richness

No

. o

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22

Litter depth

A tendency for points with shallower litter to be associated with more Coleoptera species was shown by the model and this was most evident at Stodmarsh.

Diameter

The model showed a tendency for points with thicker reed stems to be associated with greater numbers of Coleoptera in pitfall traps. This was most evident at Stodmarsh.

Figure 5.28: Relationship between reed stem diameter and number of Coleoptera in pitfall traps

5 10 15 20

12

.61

2.8

13

.01

3.2

13

.4

Partial Plot

Litter depth

5 10 15 20

51

01

52

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Actual data

Litter depth

No

. o

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sp

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.01

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Partial Plot

Diameter

2 3 4 5

51

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52

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53

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Actual data

Diameter

No

. o

f C

ole

op

tera

sp

p

Figure 5.27: Relationship between litter depth and number of Coleoptera in pitfall traps

0 10 20 30 40

45

67

89

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11

standing.water.summer.2010

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Litter saturation

Points that were dry between May and August 2009 tended to trap more Coleoptera species in August. However the points were biased towards dry sites. Interestingly, wet points (n=3) had on average higher numbers of Coleoptera trapped than partially saturated points (n=11). However this comparison was not even so further work is needed on the variation in Coleoptera diversity along hydrological gradients in reedbeds.

Figure 5.29: Relationship between litter saturation categories and number of Coleoptera species in

pitfall traps

Habitat associations with wetland specialist Coleoptera species in pitfall traps

This analysis looks at the habitat variables related to the number of wetland specialist beetles in pitfall traps.

Figure 5.30: Importance of habitat variables in explaining variation in number of wetland specialist Coleoptera species in pitfall traps. These models explained 90% of the variance.

-4-202468

1012

Var

iab

le im

po

rtan

ce (

%In

cMSE

)

Habitat variables

Coleoptera wetland specialists

24

Plant richness

Points with lower plant species richness were associated with higher numbers of wetland specialist Coleoptera species. However this seems to be skewed by the fact that Stodmarsh points had high Coleoptera diversity and low plant species diversity.

Figure 5.31: Relationship between plant species richness and number of wetland Coleoptera species

Site

Stodmarsh had a higher number of wetland specialist Coleoptera, reflecting the higher overall Coleoptera species diversity at this site compared to Ham Wall and Hickling Broad.

Figure 5.32: Relationship between site and number of wetland specialist Coleoptera in pitfall traps

0 2 4 6 8 10 12

7.5

8.0

8.5

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.0

Partial Plot

Plant richness

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Actual data

Plant richness

No

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Mean reed height

Points with taller reed on average tended to trap more wetland specialists Coleoptera species in pitfall traps. This appeared to be true at Stodmarsh and Hickling analysed separately.

Figure 5.33: Relationship between mean reed height and number of wetland specialist Coleoptera species

Diameter

Points with greater reed stem diameters were associated with higher numbers of wetland specialist Coleoptera species. This relates to the trend with taller reed height. Again it could be skewed by Stodmarsh points tending to have thicker stems and higher Coleoptera diversity.

Figure 5.34: Relationship between reed stem diameter and number of wetland Coleoptera species

1.0 1.5 2.0 2.5

8.0

8.2

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8.6

8.8

9.0

9.2

Partial Plot

Mean Reed Height

1.0 1.5 2.0 2.5

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Actual data

Mean Reed Height

No

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Diameter

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Diameter

No

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Stem density

Points with higher stem densities were associated with higher numbers of wetland specialist Coleoptera species and this appeared to be the case at Stodmarsh and Hickling Broad alone.

Figure 5.35: Relationship between stem density and number of wetland Coleoptera specialist species

Litter saturation was of low importance and was biased towards dry points (only 3 were wet).

Habitat associations with Coleoptera conservation scores in pitfall traps

This analysis looks at the conservation scores of pitfall traps calculated just for Coleoptera. It shows which habitat variables were associated with traps with beetles with a conservation status.

Figure 5.36: Relative importance of habitat variables in explaining variation in Coleoptera conservation scores. These models explained 93% of the variance.

100 200 300 400 500

8.0

8.5

9.0

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.01

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.0

Partial Plot

Stems per square metre

100 200 300 400 500

05

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Actual data

Stems per square metre

No

. o

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0123456789

10

Var

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%In

cMSE

)

Habitat variables

Coleoptera conservation score

0 10 20 30 40

45

67

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standing.water.summer.2010

Ere

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Diameter

Points with thicker reed stems were associated with more Coleoptera species with conservation status at Stodmarsh and Hickling Broad.

Figure 5.37: Relationship between reed stem diameter and Coleoptera conservation score

Litter depth

All but the shallowest litter depths were capable of trapping high numbers of Coleoptera species with conservation statuses.

Figure 5.38: Relationship between litter depth and Coleoptera conservation score

2 3 4 5

10

.01

0.5

11

.01

1.5

12

.0

Partial Plot

Diameter

2 3 4 5

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Actual data

Diameter

Co

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5 10 15 20

9.5

10

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Partial Plot

Litter Depth

5 10 15 20

05

10

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Actual data

Litter Depth

Co

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Scrub distance

The relationship between scrub distance and number of Coleoptera species with a conservation status was unclear and needs further investigation.

Figure 5.39: Relationship between scrub distance and Coleoptera conservation score

Plant species richness

Points with low plant species diversity tended to trap more Coleoptera species with a conservation status, however these tended to be points at Stodmarsh so further work would be needed to see how general this trend is. It is interesting to note that Coleoptera species with a conservation score were trapped at points where the only plant was reed. Only two of the Coleoptera in pitfall traps with conservation scores were reedbed specialists.

Figure 5.40: Relationship between plant richness and Coleoptera conservation score

0 50 100 150 200 250 300

10

.51

1.0

11

.51

2.0

Partial Plot

Scrub distance

0 50 100 150 200 250 300

05

10

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35

Actual data

Scrub Distance

Co

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0 2 4 6 8 10 12

10

.01

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.01

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.0

Partial Plot

Plant richness

0 2 4 6 8 10 12

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35

Actual data

Plant richness

Co

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29

Litter saturation: Points that were dry tended to trap more Coleoptera species with a conservation score, however this is likely to be highly biased by the fact that pitfall traps were placed in dry areas.

Composite habitat variables

A number of the measured habitat variables are associated with each other, due to the nature of reedbed successional habitat. Although this inter-correlation was controlled for in random forest models, a separate principle component analysis (PCA) was carried out to see if similar results were found when composite habitat variables were used.

Here, the PCA has been done for each survey dataset individually, in order to relate gradients to the number of species caught. In a separate analysis, habitat variables across all surveys were combined to look at the main environmental gradients in reedbeds (Chapter 2). In addition the audit data was analysed to look at a wider range of sites. The principle components from these wider analyses could not be directly related to numbers of invertebrates in the traps, since they either combined different trapping techniques (pitfall and water trap) or were from points without trap data (audit points). Therefore to relate principle components to numbers of invertebrates in the traps, separate analysis on each survey dataset was carried out.

Summary of tests correlating principle components for pitfall traps with bootstrapped species richness in traps

The first four principle components explained 81% of the data. None of the principle components were significantly correlated with bootstrapped number of species in the pitfall traps. PC1 was closest to being significant, which was a gradient of increasing stem density, decreasing panicle percentage and decreasing plant species richness. PC1 was tending towards a negative correlation with bootstrapped species richness, which would have meant points with higher plant species richness, higher percentage of panicles and lower stem densities were associated with higher invertebrate diversity. Random forest models found high plant species richness to be important. Standing water was not included as a habitat variable since all points were dry. This means fewer associations between habitat variables and water levels exist as there is not the same gradient of wetness seen at the watertrap and moth points.

Table 5.4: Principle components of habitat variables measured at pitfall trap points

PC1 PC2 PC3 PC4

Litter depth (cm) -0.079 0.502 -0.498 0.093

Stem density (stems/m2) 0.531 0.045 -0.034 -0.269

Dead stems (%) 0.031 -0.233 -0.789 -0.239

Panicles (%) -0.490 0.001 -0.022 0.269

Mean reed height (m) 0.181 -0.529 -0.304 0.418

Diameter (mm) 0.445 -0.243 0.135 0.464

Distance to scrub (m) -0.111 -0.468 0.131 -0.619

Plant species richness -0.478 -0.366 -0.004 0.128

30

-3 -2 -1 0 1 2 3

-3-2

-10

12

x[, 1]

x[, 2

]

1

2

3

4

Standard deviation 1.538 1.355 1.110 1.048

Proportion of variance 0.296 0.230 0.154 0.137

Cumulative variance 0.296 0.525 0.679 0.816

Table 5.5: Results of Pearson’s Correlation Tests on bootstrapped number of species against principle components for pitfall traps.

Component Pearsons df t p

PC1 -0.213 50 -1.55 0.129 .

PC2 0.091 50 0.643 0.523

PC3 0.101 50 0.717 0.477

PC4 -0.088 50 -0.625 0.535

Reedbed specialists

To see if the traps where reedbed specialist invertebrates were caught were associated with a particular combination of habitat variables, these traps were highlighted on the ordination plot. Distinct clusters did not emerge so reedbed specialists are fairly widely distributed across the environmental gradients.

Figure 5.41: Ordination showing pitfall traps containing reedbed specialists with the number of reedbed specialists given different colours.

As the number of reedbed specialists in pitfall traps increased, the points are more central in PC1. Points with more than one reedbed specialist tended to have lower PC2 values, representing shallower litter depths and taller reed heights. However this is a vague visual trend and the data does not support a firm conclusion on the habitat preferences of reedbed specialists.

-3 -2 -1 0 1 2 3

-3-2

-10

12

x[, 1]

x[, 2

]

31

References

UK BAP http://www.ukbap.org.uk

Appendix to invertebrate results

Table 5A: List of reedbed specialists as defined by experts and the category “reedfen and pool” in ISIS for pitfall trap samples

Higher Taxon Family

Experts reedbed specialists from pitfalls Higher Taxon Family

ISIS reedbed and fen from pitfalls

Araneae Salticidae Marpissa radiata Coleoptera Carabidae Demetrias imperialis

Coleoptera Carabidae Demetrias imperialis Coleoptera Carabidae Odacantha melanura

Coleoptera Carabidae Paradromius longiceps Coleoptera Carabidae Oodes helopioides

Hemiptera Lygaeidae Ischnodemus sabuleti Coleoptera Carabidae Paradromius longiceps

Coleoptera Malachiidae Anthocomus rufus

Coleoptera Malachiidae Cerapheles terminatus

Coleoptera Staphylinidae Ocyusa picina

Coleoptera Staphylinidae Pachnida nigella

Coleoptera Staphylinidae Paederus riparius

Coleoptera Staphylinidae Quedius balticus

Coleoptera Staphylinidae Stenus carbonarius

Table 5B: Wetland specialists (ISIS and experts lists compared) from pitfall samples

Higher taxon Family Experts wetland Higher taxon Family ISIS W2 and W3

Araneae Clubionidae Clubiona juvenis Araneae Araneidae Larinioides cornutus

Araneae Clubionidae Clubiona phragmitis Araneae Clubionidae

Clubiona juvenis

Araneae Linyphiidae Allomengea vidua Araneae Clubionidae Clubiona phragmitis

Araneae Linyphiidae Donacochara speciosa Araneae Clubionidae

Clubiona subtilis

Araneae Linyphiidae Gnathonarium dentatum Araneae Hahniidae

Antistea elegans

Araneae Linyphiidae Gongylidiellum murcidum Araneae Linyphiidae

Allomengea vidua

Araneae Linyphiidae Hypomma fulvum Araneae Linyphiidae Bathyphantes approximatus

Araneae Linyphiidae Microlinyphia impigra Araneae Linyphiidae

Donacochara speciosa

Araneae Linyphiidae Oedothorax gibbosus Araneae Linyphiidae

Gnathonarium dentatum

Araneae Linyphiidae Silometopus

Araneae Linyphiidae Gongylidiellum

32

elegans murcidum

Araneae Linyphiidae Walckenaeria vigilax Araneae Linyphiidae

Hypomma fulvum

Araneae Lycosidae Arctosa leopardus Araneae Linyphiidae Leptorhoptrum robustum

Araneae Lycosidae Pirata latitans Araneae Linyphiidae Lophomma punctatum

Araneae Lycosidae Pirata piscatorius Araneae Linyphiidae Microlinyphia impigra

Basommatophora Planorbidae Anisus leucostoma Araneae Linyphiidae

Oedothorax gibbosus

Coleoptera Cantharidae Cantharis thoracica Araneae Linyphiidae

Silometopus elegans

Coleoptera Carabidae Acupalpus dubius Araneae Lycosidae Arctosa leopardus

Coleoptera Carabidae Agonum emarginatum Araneae Lycosidae

Pardosa amentata

Coleoptera Carabidae Agonum fuliginosum Araneae Lycosidae

Pardosa proxima

Coleoptera Carabidae Agonum thoreyi Araneae Lycosidae Pirata hygrophilus

Coleoptera Carabidae Anisodactylus binotatus Araneae Lycosidae Pirata latitans

Coleoptera Carabidae Badister dilatatus Araneae Lycosidae Pirata piraticus

Coleoptera Carabidae Bembidion assimile Araneae Lycosidae

Pirata piscatorius

Coleoptera Carabidae Bembidion biguttatum Araneae Salticidae

Marpissa radiata

Coleoptera Carabidae Bembidion fumigatum Araneae Tetragnathidae

Pachygnatha clercki

Coleoptera Carabidae Carabus granulatus Basommatophora Ellobiidae

Carychium minimum

Coleoptera Carabidae Demetrias imperialis Basommatophora Planorbidae

Anisus leucostoma

Coleoptera Carabidae Elaphrus cupreus Coleoptera Carabidae Acupalpus dubius

Coleoptera Carabidae Leistus terminatus Coleoptera Carabidae Agonum emarginatum

Coleoptera Carabidae Odacantha melanura Coleoptera Carabidae

Agonum fuliginosum

Coleoptera Carabidae Oodes helopioides Coleoptera Carabidae Agonum thoreyi

Coleoptera Carabidae Oxypselaphus obscurus Coleoptera Carabidae

Anisodactylus binotatus

Coleoptera Carabidae Paradromius longiceps Coleoptera Carabidae

Badister dilatatus

Coleoptera Carabidae Paranchus albipes Coleoptera Carabidae Bembidion assimile

33

Coleoptera Carabidae Pterostichus anthracinus Coleoptera Carabidae

Bembidion biguttatum

Coleoptera Carabidae Pterostichus diligens Coleoptera Carabidae

Bembidion fumigatum

Coleoptera Carabidae Pterostichus gracilis Coleoptera Carabidae Blemus discus

Coleoptera Carabidae Pterostichus minor Coleoptera Carabidae

Carabus granulatus

Coleoptera Carabidae Pterostichus nigrita Coleoptera Carabidae

Demetrias imperialis

Coleoptera Chrysomelidae Aphthona nonstriata Coleoptera Carabidae

Elaphrus cupreus

Coleoptera Clambidae Clambus armadillo Coleoptera Carabidae Odacantha melanura

Coleoptera Coccinellidae Coccidula rufa Coleoptera Carabidae Oodes helopioides

Coleoptera Dytiscidae Hydroporus angustatus Coleoptera Carabidae

Oxypselaphus obscurus

Coleoptera Dytiscidae Hyphydrus ovatus Coleoptera Carabidae Paradromius longiceps

Coleoptera Dytiscidae Ilybius quadriguttatus Coleoptera Carabidae

Pterostichus anthracinus

Coleoptera Haliplidae Haliplus flavicollis Coleoptera Carabidae Pterostichus diligens

Coleoptera Helophoridae Helophorus brevipalpis Coleoptera Carabidae

Pterostichus gracilis

Coleoptera Helophoridae Helophorus grandis Coleoptera Carabidae

Pterostichus minor

Coleoptera Helophoridae Helophorus strigifrons Coleoptera Carabidae

Pterostichus nigrita

Coleoptera Heteroceridae Heterocerus fenestratus Coleoptera Chrysomelidae

Aphthona nonstriata

Coleoptera Hydraenidae Hydraena testacea Coleoptera Coccinellidae Coccidula rufa

Coleoptera Hydraenidae Ochthebius dilatatus Coleoptera Dytiscidae

Hydroporus angustatus

Coleoptera Hydraenidae Ochthebius minimus Coleoptera Dytiscidae

Hyphydrus ovatus

Coleoptera Hydrophilidae Anacaena globulus Coleoptera Dytiscidae

Ilybius quadriguttatus

Coleoptera Hydrophilidae Anacaena limbata Coleoptera Haliplidae Haliplus flavicollis

Coleoptera Hydrophilidae Cercyon convexiusculus Coleoptera Helophoridae

Helophorus brevipalpis

Coleoptera Hydrophilidae Cercyon marinus Coleoptera Helophoridae Helophorus grandis

Coleoptera Hydrophilidae Cercyon sternalis Coleoptera Helophoridae Helophorus strigifrons

34

Coleoptera Hydrophilidae Cercyon tristis Coleoptera Heteroceridae Heterocerus fenestratus

Coleoptera Hydrophilidae Cercyon ustulatus Coleoptera Hydraenidae Hydraena testacea

Coleoptera Hydrophilidae Coelostoma orbiculare Coleoptera Hydraenidae

Ochthebius dilatatus

Coleoptera Hydrophilidae Hydrobius fuscipes Coleoptera Hydraenidae

Ochthebius minimus

Coleoptera Malachiidae Anthocomus rufus Coleoptera Hydrophilidae Anacaena globulus

Coleoptera Malachiidae Cerapheles terminatus Coleoptera Hydrophilidae

Anacaena limbata

Coleoptera Phalacridae Stilbus oblongus Coleoptera Hydrophilidae Cercyon convexiusculus

Coleoptera Ptiliidae Baeocrara variolosa Coleoptera Hydrophilidae

Cercyon marinus

Coleoptera Scirtidae Cyphon hilaris Coleoptera Hydrophilidae Cercyon sternalis

Coleoptera Scirtidae Microcara testacea Coleoptera Hydrophilidae Cercyon tristis

Coleoptera Scydmaenidae Euconnus hirticollis Coleoptera Hydrophilidae

Cercyon ustulatus

Coleoptera Scydmaenidae Stenichnus collaris Coleoptera Hydrophilidae Coelostoma orbiculare

Coleoptera Silvanidae Psammoecus bipunctatus Coleoptera Hydrophilidae

Hydrobius fuscipes

Coleoptera Staphylinidae Anotylus rugosus Coleoptera Malachiidae Anthocomus rufus

Coleoptera Staphylinidae Atheta graminicola Coleoptera Malachiidae

Cerapheles terminatus

Coleoptera Staphylinidae Boreophilia eremita Coleoptera Scirtidae Cyphon hilaris

Coleoptera Staphylinidae Brachygluta fossulata Coleoptera Scirtidae

Microcara testacea

Coleoptera Staphylinidae Brachygluta helferi Coleoptera Silvanidae

Psammoecus bipunctatus

Coleoptera Staphylinidae Bryaxis bulbifer Coleoptera Staphylinidae Anotylus rugosus

Coleoptera Staphylinidae Carpelimus corticinus Coleoptera Staphylinidae

Atheta graminicola

Coleoptera Staphylinidae Carpelimus lindrothi Coleoptera Staphylinidae

Bryaxis bulbifer

Coleoptera Staphylinidae Carpelimus zealandicus Coleoptera Staphylinidae

Carpelimus corticinus

Coleoptera Staphylinidae Dacrila fallax Coleoptera Staphylinidae Carpelimus lindrothi

Coleoptera Staphylinidae Deinopsis erosa Coleoptera Staphylinidae Dacrila fallax

Coleoptera Staphylinidae Dilacra luteipes Coleoptera Staphylinidae Deinopsis

35

erosa

Coleoptera Staphylinidae Erichsonius cinerascens Coleoptera Staphylinidae

Dilacra luteipes

Coleoptera Staphylinidae Euaesthetus ruficapillus Coleoptera Staphylinidae

Erichsonius cinerascens

Coleoptera Staphylinidae Gabrius bishopi Coleoptera Staphylinidae Euaesthetus ruficapillus

Coleoptera Staphylinidae Lathrobium brunnipes Coleoptera Staphylinidae

Lathrobium elongatum

Coleoptera Staphylinidae Lathrobium elongatum Coleoptera Staphylinidae

Lathrobium impressum

Coleoptera Staphylinidae Lathrobium impressum Coleoptera Staphylinidae

Lathrobium quadratum

Coleoptera Staphylinidae Lathrobium quadratum Coleoptera Staphylinidae

Lathrobium terminatum

Coleoptera Staphylinidae Lathrobium terminatum Coleoptera Staphylinidae Lesteva sicula

Coleoptera Staphylinidae Lesteva sicula Coleoptera Staphylinidae Myllaena dubia

Coleoptera Staphylinidae Mycetota laticollis Coleoptera Staphylinidae Myllaena minuta

Coleoptera Staphylinidae Myllaena dubia Coleoptera Staphylinidae Ochthephilum fracticorne

Coleoptera Staphylinidae Myllaena minuta Coleoptera Staphylinidae Ocyusa maura

Coleoptera Staphylinidae Ochthephilum fracticorne Coleoptera Staphylinidae Ocyusa picina

Coleoptera Staphylinidae Ocyusa maura Coleoptera Staphylinidae Olophrum fuscum

Coleoptera Staphylinidae Ocyusa picina Coleoptera Staphylinidae Oxypoda elongatula

Coleoptera Staphylinidae Olophrum fuscum Coleoptera Staphylinidae Oxypoda procerula

Coleoptera Staphylinidae Oxypoda elongatula Coleoptera Staphylinidae

Pachnida nigella

Coleoptera Staphylinidae Oxypoda procerula Coleoptera Staphylinidae

Paederus riparius

Coleoptera Staphylinidae Pachnida nigella Coleoptera Staphylinidae Philhygra gyllenhalii

Coleoptera Staphylinidae Paederus riparius Coleoptera Staphylinidae Philhygra malleus

Coleoptera Staphylinidae Philhygra gyllenhalii Coleoptera Staphylinidae

Philhygra terminalis

Coleoptera Staphylinidae Philhygra malleus Coleoptera Staphylinidae Philhygra volans

Coleoptera Staphylinidae Philhygra terminalis Coleoptera Staphylinidae

Philonthus fumarius

Coleoptera Staphylinidae Philhygra volans Coleoptera Staphylinidae Philonthus quisquiliarius

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Coleoptera Staphylinidae Philonthus fumarius Coleoptera Staphylinidae

Quedius balticus

Coleoptera Staphylinidae Philonthus quisquiliarius Coleoptera Staphylinidae

Quedius boopoides

Coleoptera Staphylinidae Quedius balticus Coleoptera Staphylinidae Quedius fuliginosus

Coleoptera Staphylinidae Quedius boopoides Coleoptera Staphylinidae

Rybaxis longicornis

Coleoptera Staphylinidae Rybaxis longicornis Coleoptera Staphylinidae

Stenus bimaculatus

Coleoptera Staphylinidae Stenus bimaculatus Coleoptera Staphylinidae

Stenus canaliculatus

Coleoptera Staphylinidae Stenus canaliculatus Coleoptera Staphylinidae

Stenus carbonarius

Coleoptera Staphylinidae Stenus carbonarius Coleoptera Staphylinidae Stenus juno

Coleoptera Staphylinidae Stenus juno Coleoptera Staphylinidae Stenus latifrons

Coleoptera Staphylinidae Stenus latifrons Coleoptera Staphylinidae Stenus melanopus

Coleoptera Staphylinidae Stenus melanopus Coleoptera Staphylinidae Stenus nitens

Coleoptera Staphylinidae Stenus nitens Coleoptera Staphylinidae Stenus pallipes

Coleoptera Staphylinidae Stenus pallipes Coleoptera Staphylinidae Stenus palustris

Coleoptera Staphylinidae Stenus palustris Coleoptera Staphylinidae Tachyporus transversalis

Coleoptera Staphylinidae Tachyporus transversalis Hemiptera Lygaeidae

Ischnodemus sabuleti

Hemiptera Lygaeidae Ischnodemus sabuleti Hemiptera Nepidae Nepa cinerea

Hemiptera Nepidae Nepa cinerea Hemiptera Saldidae Chartoscirta cincta

Hemiptera Saldidae Chartoscirta cincta Hemiptera Veliidae

Microvelia reticulata

Hemiptera Veliidae Microvelia reticulata Orthoptera Tetrigidae Tetrix subulata

Isopoda Ligiidae Ligidium hypnorum Stylommatophora Agriolimacidae

Deroceras laeve

Isopoda Trachelipidae Trachelipus rathkei Stylommatophora Vertiginidae

Vertigo antivertigo

Orthoptera Tetrigidae Tetrix subulata Stylommatophora Zonitidae Zonitoides nitidus

Stylommatophora Agriolimacidae Deroceras laeve

Stylommatophora Vertiginidae

Vertigo antivertigo

Stylommatophora Zonitidae Zonitoides nitidus

Quedius balticus

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Table 5C: List of grid references of points where Quedius balticus was found

Trap Code

Total Number of specimens

Grid reference

HBA1 4 642458 321925 HBA2 4 642490 321954 HBA4 1 642468 322116 HBA5 5 642423 322087 HBA6 1 642408 322037 HBA8 10 642359 322092

HBA9 1 642434 322139

HBPDD3 1 643420 320840

HBPDD6 2 643613 320704 HBPDD8 2 643691 320591

Maps of the locations where Quedius balticus was trapped

Figure 5A: Map of locations at Hickling Broad and hydrological units.

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Figure 5B: Map of locations at Ham Wall and hydrological units.

Table 5D: GPS coordinates of pitfall traps

Site Trap code Easting Northing

HB HBA1 642458 321925

HB HBA10 642499 322153

HB HBA2 642490 321954

HB HBA3 642477 322023

HB HBA4 642468 322116

HB HBA5 642423 322087

HB HBA6 642408 322037

HB HBA7 642337 322040

HB HBA8 642359 322092

HB HBA9 642434 322139

HB HBPDD1 643090 320900

HB HBPDD10 643652 320967

HB HBPDD11 643819 321019

HB HBPDD12 643800 321078

HB HBPDD2 643047 320851

HB HBPDD3 643420 320840

HB HBPDD4 643460 320880

HB HBPDD5 643622 320657

HB HBPDD6 643613 320704

HB HBPDD7 643640 320610

HB HBPDD8 643691 320591

HB HBPDD9 643631 320906

HW HWPDD1 346002 139957

HW HWPDD13 346038 139997

HW HWPT1 346411 139993

HW HWPT2 346392 139954

HW HWPT3 346360 139911

HW HWPT4 346352 139945

HW HWPUCD1 346491 139992

HW HWPUCD2 346472 139925

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HW HWPUCD3 346456 139882

HW HWPUCD4 346410 139879

HW HWPUCD5 346412 139923

HW HWPUCD6 346452 139957

SM SMPDD1 622730 161813

SM SMPDD10 623361 162428

SM SMPDD11 623530 162320

SM SMPDD12 623680 162360

SM SMPDD2 622790 161830

SM SMPDD3 623490 162786

SM SMPDD4 623579 162750

SM SMPDD5 623671 162635

SM SMPDD6 623708 162650

SM SMPDD7 623655 162694

SM SMPDD8 623297 162464

SM SMPDD9 623340 162370

SM SMPWD1 622233 161127

SM SMPWD10 621588 161213

SM SMPWD11 621590 161150

SM SMPWD12 623183 162560

SM SMPWD2 622260 161170

SM SMPWD3 622264 161240

SM SMPWD4 622223 161271

SM SMPWD5 621580 161580

SM SMPWD6 621610 161532

SM SMPWD7 621560 161521

SM SMPWD8 621629 161323

SM SMPWD9 621670 161280