Tuatara: Volume 30, Issue 1, December 1988
A Comparison of the Prey of Three Common Web-Building Spiders of Open Country, Bush Fringe and Urban Areas
A Comparison of the Prey of Three Common Web-Building Spiders of Open Country, Bush Fringe and Urban Areas
Abstract
The prey is examined of three species of web-building spiders which are commonly found in the open country, bush fringe and urban areas of Wellington, New Zealand. The species are: Achaearanea veruculata, F. Theridiidae (the small cobweb spider): Artaneus pustulosus, F. Araneidae (the garden orb web spider); and Ixeuticus martius. F. Dictynidae (the black house spider). Prey is compared numerically and from a biomass point-of-view. Web examinations revealed that A. veruculata captures a wider range of prey than either of the other two species. A significant amount of its food comes from predation on other spiders. A. pustulosus feeds mainly on Diptera (62% of its prey) while I. martius also feeds mainly on Diptera (57% of its prey). These two species are regarded as competing strongly for food, with a lesser degree of competition occurring between either of the two and A. veruculata. Young spiders of all three species fed mainly on prey which was smaller than that captured by the adult spiders, but it was concluded that some food niche shift occurs in the life cycle of the three species. A comparison of clutch sizes and clutch production showed that A. pustulosus has a far higher natality rate than either of the other species. A. pustulosus also suffers from many more sources of mortality than the other two species.
Keywords:Achaearanea veruculata, Araneus pustulosus, Araneidae, competition, Dictynidae, food selection. Ixeuticus martius, mortality, spiderling.
Introduction
This study was conducted to see if any marked overlap in prey occurred between three species of web-building spiders which commonly occupy the same part of the habitat in urban Wellington, and the fringe of nearby bush areas.
The three species are:
Achaearanea veruculata (Theridiidae) - the small cobweb spider. (Fig. 1). This species, by far the smallest of the three, reaches to about 5mm body length. It is fawn to brown in colour and builds a cobweb of untidy threads in crevices of rocks or trees, also on dwellings, shrubs and fences. Its captured prey is wrapped up with a technique of throwing threads around the animals. This technique is shared with other theridiids such as Steatoda and Latrodectus (the katipo).
Araneus pustulosus (Araneidae) - the garden orb web spider. (Fig. 2). This species grows to 15mm body length, and shows a wide colour variation from cream through brown and green to near-black. It builds an orb web of radiating threads with a spiral thread laid on to the radials. The web may be found on bushes, dwellings, fences and on rock faces where projecting surfaces are available. Prey are wrapped up by rolling with the legs while silk is dispensed from the spinnerets.
Ixeuticus martius (Dictynidae) - the black house spider. (Fig. 3). This species grows to 15mm body length and while brownish when young, darkens with age until parts of the body are black. Other parts carry whitish hairs giving this species a greyish appearance. The web is a series of struts on which a zig-zag pattern is woven, but as the web ages it becomes disordered and the strut and zig-zag pattern is often hard to see. Prey are grabbed and dragged to the spider's retreat to be dealt with there.
Ixeuticus is the only one of these three species to construct a definite retreat of silken thread. A. pustulosus lies exposed on a surface while A. veruculata hides in a crevice when possible.
page 24
Fig. 1. Achaearanea veruculata shown in a typical posture. The dorsal surface faces to the ground.
Fig. 2. Araneus pustulosus shown at the centre of its orb web. The spider normally faces head towards the ground. An insect prey can be seen above the spider, well wrapped in silk.
Fig 3. Ixeuticus martius in a typical posture at night, waiting for prey to contact the web.
Localities Studied
1. |
The fringe of Otari Reserve in Wilton, where A veruculata and A. pustulosus were present in large numbers, with I. martius less common. This locality is exposed to both of Wellington's main winds - north-westerlies and southerlies. Insects are carried from farmland to the north and west, and from within the bush, which is mixed broadleaf with some podocarps. From the south insects will be carried from the slopes of Tinakori Hills and the urban areas. |
2. |
The south-eastern slopes of the Tinakori Hills adjacent to Wadestown. Here A. veruculata and A. pustulosus were present on rock faces and trees, with some I. martius as well. This location is adjacent to macrocarpa, pine and page 25 smaller native trees such as mahoe, rangiora and various species of coprosma. Insects are carried here from farmland and the bush reserve at Otari. |
3. |
A crib-wall garden on a path running between Hill Street and Hawkestone Street in Thorndon. This faces north west and insects probably arrive here from Tinakori Hills, and further north from Otari and the farmland beyond. The spider webs here are mainly found on the ornamental plants of the crib-wall. These are the prostrate rosemary (Rosemarinus officinalis var. prostratus) and a yellow flowering succulent, Sedum sp. This urban area is rich in all three species of spider. |
4. |
A garden and roadside reserve at the author's home in suburban Northland. Here the exposure is to the north so insects could be carried from the farmland and bush remnants around Johnston Hill and further out. All three species were present here with webs on flowers, shrubs, house, fences, and rocky bank at the roadside. |
Method
A preliminary study provided the outline for a web-collecting programme in which 100 prey-containing webs were examined for each species in each of the four seasons. Thus a total of 400 webs were found for each species, accumulated at the rate of 10 per week spread over the four locations.
The webs of the species studied do not always contain recognisable prey remains. It was found in the preliminary survey that for every usable web, between five and 10 webs need to be examined. This gave a figure of between 6,000 and 12,000 web examinations over the year in order to find the 1200 prey-containing webs required for the three species comparison. Only prey which were actively being fed on, or had recently been fed on by the spiders were counted in the study. Familiarisation with the collection sites and positions of webs ensured that double counting of remains was avoided. This is an important point because some prey (e.g. beetles) can persist in webs for many weeks after the spider has fed on them.
The prey remains were identified to family level where possible and these data used to draw up results (Tables 1-7).
Spider Size and Prey Captured
Where possible a measurement was made to the body length of spiders found with prey in order to relate the prey captured to the maturity of the spider.
Spiders were placed into one of the folowing groups for this intraspecific comparison:
1. |
Spiderlings - those spiders in the first 1-2 months of independent life. |
2. |
Immature spiders - those which had been in their own webs for at least two months but which had not yet reached sexual maturity. |
3. |
Mature spiders - those which had reached sexual maturity. Body length measurement of each species in relation to maturity is as follows: |
Species | Body length in mm | ||
Spiderlings | immature spiders | mature spiders | |
A. veruculata | <1.5 | 1.5-3.5 | >3.5 |
A. pustulosus | <4.0 | 4.0-8.0 | >8.0 |
I. martius | <4.0 | 4.0-8.0 | >8.0 |
The dividing line between spiderlings and immature spiders is somewhat arbitrary, but that between immatures and matures was based on the body length of females found with egg sacs. The results of the survey are presented in Tables 1-7.
Prey Comparison Between Species
A. veruculata
In this species, Diptera (35.4%) were numerically the largest prey group, but they were not nearly as significant as they were to the other two species. The Hymenoptera (18.2%) were the next most significant prey for A. veruculata. Here ants (N = 74) were the single largest family. The web of this species seems well suited to the capture of walking insects and this would explain partly the high capture level of ants. Another feature of A. veruculata's feeding is its predation on spiders, particularly on I. martius. Close observations of these two species on the side of the author's house showed that the captures of I. martius were probably related to members of this species changing their web sites. The captured spiders in many instances having moved 30cm or more from their known web site. A. veruculata quickly and readily wraps up other spiders with the silk-throwing technique mentioned earlier. The other unusual feature of this species prey is its feeding on slaters (Isopoda), millipedes (Diplopoda) and harvestmen (Opiliones), all prey groups which are either absent or found in minimal numbers in the webs of A. pustulosus and I. martius.
Table 2 shows some niche differentiation between the spiderlings and the mature spiders, with smaller prey such as psyllids and phorids being important to spiderlings, while mature spiders utilise other spiders as the most numerous prey item. There is thus a shift towards predation on other spiders as A. veruculata matures. The importance of very large prey such as cicadas is discussed below. Seasonal prey differences were not especially marked in this species. Ants are most significant in summer (N = 28); craneflies (Tipulidae) in autumn (N = 21); and through the winter fungus gnats (Mycetophylidae) (N = 30) are an important food source.
A. pustulosus
Diptera (62%) were again by far the most numerous prey group, with Mycetophilidae (N = 351) being the most common item. Of the rest of the prey groups, only the Hemiptera (16%) and the Hymenoptera (14%) exceeded double figures as percent of prey caught. The orb web. which is strung in the flight path of insects, appears well suited to capturing Diptera. Because of its location in the habitat, it is less well suited for the capture of other prey groups which spend either all or most of their time walking. This was well illustrated by the capture of ants which were all winged forms in the orb-webs of A. pustulosus but walking ants in the cob-web traps of A. veruculata. A. pustulosus is definitely more restricted in its diet than A. veruculata, as shown in the relative paucity of items in the “other” prey category.
There is a clear niche shift in prey taken at different stages of development of A. pustulosus. The spiderlings take a variety of small prey such as psyllids, aphids, page 27 ants and psocopterans. The mature spiders capture much larger prey with bees featuring prominently along with larger types of dipterans such as blowflies (Calliphoridae) and cranefiles. The bees were nearly all honey bees (Apis mellifera) with an occasional bumble bee (Bombus sp.) being found. Bees are a problem prey item for A. pustulosus; they yield a large amount of food for the spider and so are well worth capturing, but the hazards of being stung are always present and a number of adult spiders were observed misjudging the wrapping-up of either honey bees or bumble bees - and were seen to be stung. In all cases the spider died as a result (see Table 12).
There were some seasonal differences apparent in this species, but these tended to be obscured by the predominance of mycetophylids in the prey count. In winter this group assumed even greater significance. 169 captures being found. In summer psyllids were numerically important but would be significant only for the small spiders. In autumn aphids and ants (N = 34 and 47 respectively) are important but mainly for smaller spiders.
I. martius
Like A. pustulosus, I. martius relies heavily on Diptera for its food intake. This order made up 57% of the total prey caught. The difference between these two spiders is the wider spread among the families within the Diptera by I. martius. The Mycetophylidae are not the dominant item as in A. pustulosus. Here Muscidae. Acalyptrata and Tipulidae are all very important with Calliphoridae and Stratiomyiidae being significant too. As in A. pustulosus, flying ants were caught and formed a useful addition to the diet in summer and autumn. Cicadas form an important item in summer (N = 15) and because of their large size must be even more important than their numbers alone suggest. In winter prey are scarce and flies such as craneflies, fungus gnats, muscids and blowflies are particularly important for the continued growth and maturation of I. martius. As the spider grows, smaller prey, like psyllids and ants which are present in significant numbers in the food of the spiderlings, become unimportant for mature spiders. The larger flies such as tipulids and stratiomyiids along with the cicadas are very important in the diet of the mature spiders.
Prey Size
The numerical data given in Tables 1-7 give no indication of the volume of food the prey contains, or its biomass. It is clear that a small fungus gnat is not as valuable a catch as a honey bee or a cicada. An attempt was made to relate the prey groups to one another in terms of food volume. The standard used was a very common prey item: a mycetophylid fly of 5mm body length. It was estimated that with a body diameter of 1.5mm, such a fly would contain approximately 8.8mm3 volume of food. The food volumes of other prey were estimated in the same manner and were then converted to “mycetophilid equivalents”. For example, a honey bee with a volume of 150 mm3 is equal to 15 mycetophylid units. A list of mycetophylid equivalents follows in Table 8.
Comparisons between the 12 major prey items for each spider species were made by multiplying the number of prey captured by the “mycetophilid equivalent”, (e.g. cicadas for A. veruculata: 60 × 13 = 780). These figures are presented in Table 9. Here the following prey items become much more significant than they were on a numberical count:
A. veruculata - cicadas, bees, tipulids and slaters
A. pustulosus - blowflies, bees and tipulids
I. martius - cicadas, bees and soldier flies page 28 This approximate biomass measurement clearly shows how important large-sized prey items are in the diet of these three spiders.
Cicadas could be particularly important for females of A. veruculata and I. martius. This is because egg sac production requires a substantial food input and without the addition of cicadas to the diet in summer and autumn it is probable that this activity would be curtailed in these two species.
Interspecific Predation
This study revealed high levels of interspecific predation by the small A. veruculata on I. martius and to a lesser degree on A. pustulosus (Table 10). Predation on other spiders by I. martius and A. pustulosus was less evident. The captures occurred in two ways. Firstly when the webs of two spiders were closely adjacent to one another and contact with the web of one was made by the other.
Secondly when spiders wandered from an older web site. I. martius was particularly prone to capture by A. veruculata in the second of these circumstances.
Clutch Size and Clutch Production (Natality)
Observations on egg sac production were made during the prey study. These showed that A. pustulosus was by far the most prolific species, in some individuals producing up to five sacs per year with more than 800 eggs per sac being counted. A. veruculata and I martius had a lower natality rate, producing considerably fewer eggs per sac than A. pustulosus (see Table 11). In I. martius the spiderlings appear 6 weeks after the early clutches are produced in August. Dispersal takes place any time from early spring in this species. A similar pattern is seen in A. veruculata but A. pustulosus egg sacs and spiderlings are not seen for almost a month later than those of the other two species in Wellington.
Mortality
The prey study provides an opportunity to observe and record causes of mortality in known populations of the three spider species (Table 12). Although not a quantitative investigation, these data suggest that A. pustulosus appeared to suffer the highest mortality. Perhaps this is not surprising in view of its exposed web, without a protective retreat, and its much higher natality than the other two species. The spider has a bulky shape which makes it an easy target for bird and wasp attack. It also suffers more storm damage than the other spiders. I. martius is by far the best protected of these three spiders, with its woven silk retreat, but is also the most difficult to observe.
Conclusions
These three spider species often live in the same habitat and could thus compete for web sites. Inter-specific predation, particularly on I. martius, may well be an indication of this. The study has revealed considerable overlap between the spiders in their food items. In particular between Araneus pustulosus and Ixeuticus martius, both of which depend so heavily on the same types of Diptera. Achaearanea veruculata feeds on the widest range of prey diversity of the three species, partly due to its choice of web site and partly to its prey-capturing behaviour. A. pustulosus had the distinction of capturing prey at about double the rate of the other two species. Although no attempt was made to determine growth rates, it appears from the natality and mortality data gathered in this study that this species has a page 29 higher population turnover rate than either of the other spiders. Perhaps this is largely due to its exposed orb web which lacks a protective retreat.
Acknowledgements
My thanks to Mr R. Ordish, entomologist. National Museum, Wellington, for his help in identifying the prey items found in this study, and to Dr R.R. Forster. Otago Museum, for checking the identification of the three spider species.
References
Bristowe, W.S., 1958: The Spider's World. Collins, London.
Forster, R.R. and Forster, L.M. 1973: New Zealand Spiders - an Introduction. Collins. Auckland.
Prey taxon | Summer | Autumn | Winter | Spring | Full year | % of total |
Diptera | ||||||
Tipulidae | 6 | 21 | 2 | 10 | 39 | |
Mycetophilidac | - | 8 | 30 | 21 | 59 | |
Cecidiemyidae | - | 7 | 1 | - | 8 | |
Chironomidae | - | 2 | 2 | 1 | 5 | |
Simuliidae | 1 | - | - | - | 1 | |
Psychodidae | 1 | - | 6 | 2 | 9 | |
Stratiomyidae | 3 | 3 | - | 6 | 12 | |
Phoridae | - | 3 | - | 1 | 4 | |
Muscidae | 2 | - | - | - | 2 | |
Calliphoridae | - | - | 1 | 2 | 3 | |
Anthomyiidae | - | 12 | 4 | 1 | 17 | |
Tachinidae | - | - | - | 1 | 1 | |
Acalyptrata | 1 | 3 | 2 | 3 | 9 | |
Unidentified | 1 | 4 | 3 | 2 | 10 | |
Totals | 15 | 63 | 51 | 50 | 179 | 35.4 |
Lepidoptera | ||||||
Hepialidae | 1 | - | - | 2 | 3 | |
Tineidae | 1 | 3 | 1 | - | 5 | |
Tortricidae | 4 | 4 | 3 | - | 11 | |
Crambidae | 2 | 2 | 1 | 1 | 6 | |
Noctuidae | 1 | - | - | 1 | 2 | |
Geometridae | - | 1 | - | - | 1 | |
Unidentified | - | 2 | 5 | 1 | 8 | |
Totals | 9 | 12 | 10 | 5 | 36 | 7.1 |
Hemiptera | ||||||
Cixiidae | 4 | 1 | - | - | 5 | |
Flatidae | 3 | 2 | 3 | - | 8 | |
Cicadidae | 8 | - | - | 5 | 13 | |
Cicadellidae | 5 | 2 | 1 | - | 8 | |
Psyllidae | 5 | - | 3 | - | 8 | |
Aphididae | - | 1 | 2 | 1 | 4 | |
Totals | 25 | 6 | 9 | 6 | 46 | 9.0 |
Coleoptera | ||||||
Carabidae | - | - | 1 | - | 1 | |
Cincindelidae | - | 1 | - | - | 1 | |
Coccinellidae | - | - | 1 | - | 1 | |
Anobiidae | 2 | - | - | - | 2 | |
Elateridae | 2 | - | - | 2 | 4 | page 30 |
Tenebrionidae | - | 4 | - | 3 | 7 | |
Scarabaeidae | 1 | 1 | - | 5 | 7 | |
Cerambycidae | 3 | 2 | 3 | 3 | 11 | |
Curculionidae | - | 2 | 3 | 6 | 11 | |
Unidentified | 2 | 2 | 1 | 2 | 7 | |
Totals | 10 | 12 | 9 | 21 | 52 | 10.3 |
Hymenoptera | ||||||
Ichnemuonidae | - | 2 | - | - | 2 | |
Formicidae | 28 | 19 | 12 | 15 | 74 | |
Pompilidae | - | 1 | - | - | 1 | |
Apidae | - | - | 1 | 1 | 2 | |
Unidentified | 5 | 6 | - | 2 | 13 | |
Totals | 33 | 28 | 13 | 13 | 42 | 18.2 |
Other groups | ||||||
Dermaptera | 1 | - | 1 | - | 2 | |
Dictyoptera | ||||||
Blattidae | 4 | 3 | - | 3 | 10 | |
Opilones | 7 | - | 1 | 2 | 10 | |
Diplopoda | 5 | 3 | 4 | 1 | 13 | |
Isopoda | 3 | 4 | 8 | 7 | 22 | |
Amphipoda | 5 | 1 | 1 | - | 7 | |
Gastropoda | 1 | - | - | - | 1 | |
Araneae | 8 | 14 | 5 | 9 | 36 | |
Total, Other group | 34 | 25 | 20 | 22 | 101 | |
Total prey taken by Achaeranea veruculata: | 506 |
spider body length (mm) | |||||
<1.5 | 1.5-3.5 | >3.5 | |||
Mycetophilidae | 13 | Formicidae | 49 | Spiders | 18 |
Anthomyiidae | 6 | Mycetophilidae | 26 | Mycetophilidae | 14 |
Psyllidac | 2 | Spiders | 15 | Tipulidae | 10 |
Pheridae | 2 | Tipulidae | 10 | Formicidae | 8 |
Acalyptrata | 2 | Anthomyiidae | 8 | Blattidae | 7 |
Prey taxon | Summer | Autumn | Winter | Spring | Full year | % |
Diptera | ||||||
Tipulidae | 4 | 11 | 6 | 3 | 24 | |
Mycetophilidae | 44 | 76 | 169 | 62 | 351 | |
Cecidemyidae | 7 | 55 | 29 | 21 | 112 | |
Chirenomidae | 1 | - | - | - | 1 | |
Simuliidae | 2 | 1 | - | - | 3 | |
Bibionidae | - | - | - | 1 | 1 | |
Psychodidae | 13 | 3 | 12 | 3 | 31 | |
Stratiomyiidae | - | - | -3 | 3 | ||
Empididae | 5 | - | - | - | 5 | |
Dolichopodidae | 3 | - | - | - | 3 | |
Phoridae | - | 4 | 1 | 1 | 6 | |
Syrphidae | - | - | - | 2 | 2 | |
Muscidae | - | 2 | - | 3 | 5 | |
Calliphoridae | - | 3 | 5 | 15 | 23page 31 | |
Anthomyiidae | - | 1 | 2 | 6 | 9 | |
Tachinidae | - | - | - | 2 | 2 | |
Acalyptrata | 20 | 4 | 5 | 8 | 37 | |
Unidentified | 5 | 4 | 2 | 3 | 14 | |
Totals | 104 | 164 | 231 | 133 | 632 | 62 |
Lepidoptera | ||||||
Tineidae | 2 | 1 | - | 1 | 4 | |
Tertricidae | - | 1 | - | - | 1 | |
Crambidae | - | - | - | 2 | 2 | |
Noctuidae | 1 | 1 | - | 6 | 8 | |
Unidentified | - | 1 | 1 | 4 | 6 | |
Totals | 3 | 4 | 1 | 13 | 21 | 2 |
Hemiptera | ||||||
Cixiidae | 6 | - | - | - | 6 | |
Flatidae | 2 | - | 1 | - | 3 | |
Cicadidae | 4 | 3 | - | - | 7 | |
Cicadellidae | 4 | 2 | - | 1 | 7 | |
Psyllidae | 28 | 17 | 7 | 17 | 69 | |
Aphididae | 11 | 34 | 4 | 9 | 58 | |
Unidentified | 4 | - | 1 | - | 5 | |
Hetereptera | 1 | 3 | - | 2 | 6 | |
Totals | 60 | 59 | 13 | 29 | 161 | 16 |
Coleoptera | ||||||
Anabiidae | 2 | - | - | - | 2 | |
Staphylinidae | 5 | - | - | - | 5 | |
Tenebrionidae | - | - | 1 | - | 1 | |
Scarabaeidae | - | - | - | 3 | 3 | |
Cerambycidae | - | - | 1 | - | 1 | |
Curculionidae | 1 | - | - | - | 7 | |
Unidentified | 5 | 1 | 1 | - | 7 | |
Totals | 13 | 1 | 3 | 3 | 20 | 2 |
Hymenoptera | ||||||
Ichneumonidae | 1 | 2 | 3 | 3 | 9 | |
Formicidae | 18 | 47 | - | 1 | 66 | |
Pompilidae | - | - | - | 1 | 1 | |
Vespidae | 1 | 1 | - | - | 2 | |
Apidae | 1 | 11 | 7 | 16 | 35 | |
Unidentified | 5 | 16 | 4 | 4 | 29 | |
Totals | 26 | 77 | 14 | 25 | 142 | 14 |
Other groups | ||||||
Dermaptera | ||||||
Blattidae | 1 | - | - | - | 1 | |
Psocoptera | - | 17 | 3 | 1 | 21 | |
Thysanoptera | 7 | - | - | - | 7 | |
Collembola | - | - | 3 | 1 | 4 | |
Amphipoda | - | - | - | 2 | 2 | |
Araneae | 3 | - | - | 3 | 6 | |
Totals | 11 | 17 | 6 | 7 | 41 | |
Total prey taken by Araneus pustulosus: | 1017 |
spider body length (mm) | |||||
<4 | 4-8 | >8 | |||
Psyllidae | 42 | Mycetephilidae | 212 | Mycetophilidae | 45 |
Mycetophilidae | 32 | Cecidomyiidae | 93 | Apidae | 25 |
Aphididae | 30 | Formicidae | 34 | Acalyptrata | 18 |
Formicidae | 26 | Psyllidae | 24 | Calliphoridae | 15 |
Psocoptera | 12 | Aphididae | 19 | Tipulidae | 12 |
Prey taxon | Summer | Autumn | Winter | Spring | Full year | % | |
Diptera | |||||||
Tipulidae | 12 | 7 | 11 | 9 | 39 | ||
Mycetophilidae | 13 | 6 | 15 | 19 | 53 | ||
Cecidomyiidae | 4 | 1 | - | 5 | 10 | ||
Chironomidae | 3 | - | 1 | - | 4 | ||
Simuliidae | 7 | - | - | - | 7 | ||
Psychodidae | 1 | - | - | - | 1 | ||
Stratiomyiidae | 7 | 6 | 1 | 15 | 29 | ||
Asilidae | 1 | - | - | - | 1 | ||
Empididae | 1 | - | - | - | 1 | ||
Pheridae | - | 3 | - | 1 | 4 | ||
Delichopodidae | 5 | - | - | 1 | 6 | ||
Syrphidae | 4 | 1 | - | 1 | 6 | ||
Muscidae | 10 | 21 | 20 | 3 | 54 | ||
Calliphoridae | 2 | 2 | 19 | 4 | 27 | ||
Anthemyiidae | 5 | 8 | - | 2 | 15 | ||
Tachinidae | - | 2 | - | - | 2 | ||
Acalyptrata | 27 | 3 | 1 | 4 | 35 | ||
Unidentified | 7 | 9 | 4 | 4 | 24 | ||
Totals | 109 | 69 | 72 | 68 | 318 | 57 | |
Lepidoptera | |||||||
Hepialidae | - | - | 4 | - | 4 | ||
Tineidae | 1 | 2 | 1 | - | 4 | ||
Tortricidae | 3 | 1 | 4 | 1 | 9 | ||
Crambidae | - | 2 | 4 | 2 | 8 | ||
Noctuidae | 1 | 2 | 2 | - | 5 | ||
Unidentified | 1 | 2 | 5 | 1 | 9 | ||
Totals | 6 | 9 | 20 | 4 | 39 | 7 | |
Hemiptera | |||||||
Cixiidae | 3 | - | - | - | 3 | ||
Flatidae | 5 | 4 | - | - | 9 | ||
Cicadidae | 15 | 1 | - | - | 16 | ||
Cicadellidae | 6 | 2 | - | 1 | 9 | ||
Psyllidae | 9 | 1 | 1 | 6 | 17 | ||
Aphididae | 3 | - | - | 3 | 6 | ||
Unidentified | 2 | - | 2 | - | 4 | ||
Homoptera | |||||||
Heteroptera | 2 | 3 | 1 | - | 6 | ||
Totals | 45 | 11 | 4 | 10 | 70 | 12.5page 33 | |
Coleoptera | |||||||
Cicindelidae | 1 | - | - | - | 1 | ||
Coccinellidae | - | - | 3 | - | 3 | ||
Anobiidae | 2 | - | - | - | 2 | ||
Staphylinidae | 1 | 1 | - | - | 2 | ||
Elateridae | 2 | - | - | - | 2 | ||
Scarabaeidae | 1 | - | - | 3 | 4 | ||
Cerambycidae | 2 | - | - | 2 | 4 | ||
Curculionidae | - | 2 | - | 1 | 3 | ||
Unidentified | 3 | - | 2 | - | 5 | ||
Totals | 12 | 3 | 5 | 6 | 26 | 4.6 | |
Hymenoptera | |||||||
Ichneumonidae | 3 | 7 | 2 | 6 | 18 | ||
Formicidae | 15 | 7 | - | - | 22 | ||
Pompilidae | 1 | - | - | - | 1 | ||
Vespidae | 3 | 3 | - | - | 6 | ||
Apidae | 4 | 1 | 4 | 4 | 13 | ||
Unidentified | 9 | 6 | 1 | 2 | 18 | ||
Totals | 35 | 24 | 7 | 12 | 78 | 14 | |
Other groups | |||||||
Dermaptera | 2 | - | - | - | 2 | ||
Dictyoptera | |||||||
Blattidae | 5 | 5 | 2 | - | 12 | ||
Opiliones | - | 1 | - | 1 | 2 | ||
Diplopoda | - | - | - | 1 | 1 | ||
Isopoda | 1 | 1 | - | - | 2 | ||
Thysanoptera | 1 | - | - | - | 1 | ||
Araneae | 3 | 1 | 2 | 1 | 7 | ||
Totals | 12 | 8 | 4 | 3 | 27 | 4.9 | |
Total prey taken by Ixeuticus martius: | 558 |
spider body length (mm) | |||||
<4 | 4-8 | >8 | |||
Acalyptrata | 17 | Muscidae | 43 | Tipulidae | 16 |
Mycetophilidae | 15 | Mycetophilidae | 23 | Stratiomyiidae | 15 |
Psyllidae | 10 | Calliphoridae | 18 | Cicadidae | 13 |
Formicidae | 4 | Tipulidae | 18 | Mycetophilidae | 12 |
Tipulidae | 4 | Ichneumonidae | 15 | Blattidae | 8 |
Achaearanea veruculate | Araneus pustulosus | Ixeuticus martius | ||||
Prey Group | number | % | number | % | number | % |
Diptera | 179 | 35.4 | 632 | 62 | 318 | 57 |
Lepidoptera | 36 | 7.1 | 21 | 2 | 39 | 7 |
Hemiptera | 46 | 9.0 | 161 | 16 | 70 | 12.5 |
Coleoptera | 52 | 10.3 | 20 | 2 | 26 | 4.6 |
Hymenoptera | 92 | 18.2 | 142 | 14 | 78 | 14 |
Psocoptera | 21 | 2 | ||||
Isoptera | 22 | 4.34 | ||||
Araneae | 36 | 7.20 | ||||
Other prey | 43 | 8.46 | 20 | 27 | 4.9 | |
Totals | 506 | 1017 | 558 |
cicada | 60 | tenebrionid beetle | 5 |
calliphorid fly | 30 | harvestman | 3 |
noctuid moth | 26 | ichneumon wasp | 2.75 |
scarab beetle | 22 | anthomyiid fly | 2.5 |
click beetle | 16 | flattid bug | 2 |
syrphid fly | 16 | cicadellid bug | 2 |
vespid wasp | 16 | millipede | 2 |
honey bee | 15 | cixiid bug | 2 |
cerambycid beetle | 14 | weevil | 1.7 |
pompilid wasp | 12 | acalypterate fly | 1 |
staphylinid fly | 8 | ant | .68 |
tipulid fly | 6.5 | psyllid | .25 |
cockroach | 6 | aphid | .25 |
stratiomyid fly | 6 | psocopteran | .25 |
muscid fly | 5 | psychodid fly | .05 |
gall midge | .04 |
Prey | Food intake in “mycetophilid-equivalent” units | Food intake as biomass percentage of 12 listed prey groups |
Achaearanea veruculata | ||
Cicadidae | 780 | 38.5 |
Araneae | 360 | 17.7 |
Tipulidae | 234 | 11.5 |
Isopoda | 220 | 10.8 |
Cerambycidae | 154 | 7.6 |
Stratiomyiidae | 72 | 3.5 |
Formicidae | 63 | 3.1 |
Mycetophilidae | 59 | 2.9 |
Tortricidae | 55 | 2.7 |
Anthomyiidae | 40 | 2.0 |
Diplopoda | 26 | 1.3 |
Curculionidae | 16 | 0.8 |
Araneus pustulosus | ||
Calliphoridae | 690 | 37 |
Apidae | 525 | 28.1 |
Mycetophilidae | 351 | 18.8 |
Tipulidae | 156 | 8.37 |
Formicidae | 44.8 | 2.4 |
Acalyptrata | 37 | 2.0 |
Psyllidae | 17.25 | 0.92 |
Aphididae | 14.5 | 0.77 |
Small wasps | 14.5 | 0.77 |
Psocoptera | 5.25 | 0.28 |
Cecidomyiidae | 4.5 | 0.24 |
Pschodidae | 1.5 | 0.08 |
Ixeuticus martius | ||
Cicadidae | 960 | 63.67 |
Apidae | 195 | 12.93 |
Stratiomyiidae | 90 | 5.97 |
Blattidae | 72 | 4.77 |
Tipulidae | 58.5 | 3.88 |
Ichneumonidae | 49.5 | 3.28 |
Mycetophilidae | 19 | 1.26 |
Cicadellidae | 18 | 1.20 |
Flatidae | 18 | 1.20 |
Formicidae | 15 | 1.00 |
Small wasps | 9 | 0.60 |
Psyllidae | 4.25 | 0.28 |
Prey species | |||
Predator species | A. veruculata | A. pustulosus | I. martius |
A. veruculata | 5 | 9 | 17 |
A. pustulosus | 0 | 1 | 4 |
I. martius | 3 | 2 | 1 |
date egg sacs found | No. sacs/spider | mean No. eggs/sae | Range | No. egg sacs opened | |
A. veruculata | Aug.-Apr. | up to 5 | 72 | 25-151 | 18 |
A. pustulosus | Sep.-Jun. | up to 5 | 607 | 422-813 | 15 |
I. martius | Aug.-Mar. | up to 3 | 114 | 30-179 | 15 |
species | source of mortality | No. of deaths | effect on population |
A. veruculata | Predation by sphecid wasp, Pison morosum | up to 25% | heavy mortality in late spring to early autumn |
Intraspecific predation | 5 | moderate | |
Predation by I. martius | 3 | minor | |
Predation by salticid spiders | 3 | minor | |
Rain, wind, storms which remove spiders from web sites | 21 | moderate to heavy | |
A. pustulosus | Predation by Pison spinolae (mason wasp) | 4 | few of this species active in study area |
Predation by A. veruculata | 9 | moderate | |
Predation by I. martius | 2 | slight | |
Stung by Apis mellifera (honey bee) caught in web | 8 | moderate | |
Stung by Bombus spp (bumble bees) caught in web | 11 | moderate | |
Stung by Vespula spp (European wasps) caught in web | 5 | slight | |
Egg sacs parasitised by Ichneumonid wasp | up to 30% | heavy effect in some years | |
Predation by Sphictostethus nitidus (hunting wasp) | 2 | slighly | |
Predation by Zosterops lateralis (silvereye) | 4 | slight | |
Rain, wind, storms which remove spiders from web site | 34 | heavy | |
Summer-autumn drought which dehydrates egg sacs and kills eggs | 13 | moderate | |
I. martius | Predation by A. veruculata | 17 | heavy |
Intraspecific predation | 1 | slight | |
stung by Vespula spp. caught in web | 7 | moderate |