Estimating consumable biomass from body length and order in insects and spiders

1. Current models used to estimate insect prey biomass for diet studies use whole weight. However, a large proportion of an arthropod's body is taken up by an indigestible exoskeleton, leading to erroneous estimation of the food intake of insectivorous animals. 2. Linear mixed effect models were used to obtain equations to predict consumable biomass from body length for a variety of Neotropical insects and spiders. These data were obtained by feeding taxa of various orders to groups of 100 social spiders and comparing pre‐ and post‐consumption weights using size‐matched controls. 3. Significant linear relationships were found relating body size to consumed biomass for all orders, with slopes ranging from 1.276 to 4.011 and R² values from 0.476 to 0.929. For orders other than spiders and Orthoptera, the increase in weight with size exhibited negative allometric scaling, suggesting a decrease in tissue density, or an increase in internal air space, with size. 4. Although there were significant differences across taxonomic orders in the proportion of biomass consumed, within most orders the proportion consumed did not differ significantly with body size. The estimated regression coefficients may be used by other workers to estimate consumable biomass of arthropod prey for studies requiring large sample sizes or non‐lethal sampling of rare or endangered species.     

Relationships between body size and biomass of aquatic insects

SUMMARY. Predictive equations were developed to estimate dry weight from body length measurements for forty-three taxa of aquatic insects. The inter-relationships between dry weight, body length and head capsule width of individuals grouped according to the eight major orders of aquatic insects were also examined. Regression analysis indicated that the relationship between biomass and body size was best expressed by a power equation, Y =_aX^b, rather than by linear or exponential equations. Changes in body length versus head capsule width were best expressed by linear equations, with three distinct relationships being observed. Body length estimated biomass better than head capsule width. Populations often species of insects collected from two different rivers generally did not differ significantly in their dry weight to body length relationships.     

Population structure, growth, mortality and estimated stock size of the introduced tench, Tinca tinca (L.), population in Lake Beyşehir, Turkey

Population structure, growth, length–weight relationship, mortality and stock size of tench, Tinca tinca (L.), was studied in Lake Beyşehir, Turkey in 2005. Totals of 3360 tench (1865 males; 1795 females) were captured with gill- and trammel-nets of various mesh sizes. Male to female ratio was 1.04 : 1. The study covered length year classes. Fork lengths and total weights ranged from 9 to 37 cm and 13 to 815 g. For all individuals, the von Bertalanffy growth equation and length–weight relationship were L _t = 54.2[1−exp(−0.1350( t  + 1.0281)] and W  = 0.0151  L ^2.9993, respectively. Growth performance index and mean condition factor of the tench population were 2.598 and 1.513, respectively. Mortality rates were Z  = 1.97 year⁻¹, M  = 0.29 year⁻¹ and F  = 1.68 year⁻¹ for total, natural, and fishing mortality, respectively. The exploitation rate was E  = 0.85, and the percentage of surviving fish was 13.9%. Tench stock was assessed as about 6–7 million individuals and 1450–1500 tonnes in biomass. It was determined that maximum sustainable yield could be obtained with an 80% level of the current fishing effort.     

The size–grain hypothesis: do macroarthropods see a fractal world?

Abstract.  1. In the size–grain hypothesis (a) long legs allow walking organisms to step over gaps and pores in substrate but prohibit them from entering those gaps; (b) the world is more rugose for small organisms; and (c) the relative cost of long legs increases as organisms grow smaller. The hypothesis predicts a positive allometry of leg length ( = mass ^b where b > 0.33 of isometry), a pattern that robustly holds for ants.
2. Toward testing for leg length allometries in other taxa, arthropods were extracted from the Panama leaf litter and measured. Three common taxa (spiders, diplopods, Coleoptera) yielded b s that exceeded 0.33 while three others (Acarina, Pseudoscorpiones, and Collembola) did not. The exponent b tended to increase ( P = 0.06, n = 7) with an arthropod taxon's average body mass.
3. Since leg length in cursorial organisms tends toward isometry in very small and very large taxa (i.e. mammals) this suggests that the size–grain hypothesis may best apply at a transition zone of intermediate body mass: the macroarthropods.
4. Body length was a robust predictor of mass in all groups despite variation in shape.     

Shifts in reciprocal river‐riparian arthropod fluxes along an urban‐rural landscape gradient

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Biomass estimates of terrestrial arthropods based on body length

The relationship between body length and biomass (dry weight) was investigated for nineteen taxa of terres trail arthropods and for the combined sample that included all insect taxa. The specimens were collected from forests of Bicholim taluk of Goa. Four models were evaluated, a linear function, a logarithmic function, a power function and an exponential function. The linear function best describes dry weights in the order Isopoda of class Crustacea and dictyopteran, coleopteran larvae of class Insecta. The logarithmic function fits well for only one group namely Opiliones of class Arachnida. The power functions fits best for Scutigeromorpha and Scolopendromorpha of class Chilopoda, Araneae of class Arachnida, Collembola, Thysanura, Orthoptera, Hemiptera, Homoptera, coleopteran adults, Diptera, lepidopteran adults and Hymenoptera of class Insecta and the combined data on adult insects. An exponential function fits well for the dermapteran and lepidopteran larvae of class Insecta. The usefulness of these estimates of Arthropod biomass in community ecology is discussed.     

An evaluation of two biochemical methods of age determination in insects (pteridines and lipofuscins) using the ant Polyrhachis sexpinosa Latrielle (Hymenoptera: Formicidae)

Accurate information on the age of wild-caught animals is valuable for a variety of areas, but can be particularly difficult to obtain for small holometabolous insects, whose body size is fixed at the time of pupal eclosion. A variety of chemical groups, such as lipofuscins and pteridines accumulate in body tissues through time and can be used to predict age in a variety of arthropod taxa. Here we use spectrofluorometry to confirm the presence of extractable levels of lipofuscins and pteridines in individual social insects (using the ant Polyrhachis sexpinosa Latrielle, average body size 25 mg, as an example) and evaluate their ability to predict age. Pteridine levels were independent of age but lipofuscin levels increased with age in a predictable manner ( r ² = 72.8%). Lipofuscin levels therefore represent a new method of age determination for social insects that should be applicable to both individual laboratory and wild-caught animals.     

Arthropod responses to experimental fire regimes in an Australian tropical savannah: ordinal‐level analysis

Fire is widely used for conservation management in the savannah landscapes of northern Australia, yet there is considerable uncertainty over the ecological effects of different fire regimes. The responses of insects and other arthropods to fire are especially poorly known, despite their dominant roles in the functioning of savannah ecosystems. Fire often appears to have little long‐term effect on ordinal‐level abundance of arthropods in temperate woodlands and open forests of southern Australia, and this paper addresses the extent to which such ordinal‐level resilience also occurs in Australia’s tropical savannahs. The data are from a multidisciplinary, landscape‐scale fire experiment at Kapalga in Kakadu National Park. Arthropods were sampled in the two major savannah habitats (woodland and open forest) using pitfall traps and sweep nets, in 15–20 km² compartments subjected to one of three fire regimes, each with three replicates: ‘early’ (annual fires lit early in the dry season), ‘late’ (annual fires lit late in the dry season), and ‘unburnt’ (fires absent during the five‐year experimental period 1990–94). Floristic cover, richness and composition were also measured in each sampling plot, using point quadrats. There were substantial habitat differences in floristic composition, but fire had no measured effect on plant richness, overall composition, or cover of three of the four dominant species. Of the 11 ordinal arthropod taxa considered from pitfall traps, only four were significantly affected by fire according to repeated‐measures ANOVA . There was a marked reduction in ant abundance in the absence of fire, and declines in spiders, homopterans and silverfish under late fires. Similarly, the abundances of only four of the 10 ordinal taxa from sweep catches were affected by fire, with crickets and beetles declining in the absence of fire, and caterpillars declining under late fires. Therefore, most of the ordinal taxa from the ground and grass‐layer were unaffected by the fire treatments, despite the treatments representing the most extreme fire regimes possible in the region. This indicates that the considerable ordinal‐level resilience to fire of arthropod assemblages that has previously been demonstrated in temperate woodlands and open forests of southern Australia, also occurs in tropical savannah woodlands and open forests of northern Australia.     

Aquatic Terrestrial Linkages Along a Braided-River: Riparian Arthropods Feeding on Aquatic Insects

Rivers can provide important sources of energy for riparian biota. Stable isotope analysis (δ¹³C, δ¹⁵N) together with linear mixing models, were used to quantify the importance of aquatic insects as a food source for a riparian arthropod assemblage inhabiting the shore of the braided Tagliamento River (NE Italy). Proportional aquatic prey contributions to riparian arthropod diets differed considerable among taxa. Carabid beetles of the genus Bembidion and Nebria picicornis fed entirely on aquatic insects. Aquatic insects made up 80% of the diet of the dominant staphylinid beetle Paederidus rubrothoracicus. The diets of the dominant lycosid spiders Arctosa cinerea and Pardosa wagleri consisted of 56 and 48% aquatic insects, respectively. In contrast, the ant Manica rubida fed mainly on terrestrial sources. The proportion of aquatic insects in the diet of lycosid spiders changed seasonally, being related to the seasonal abundance of lycosid spiders along the stream edge. The degree of spatial and seasonal aggregation of riparian arthropods at the river edge coincided with their proportional use of aquatic subsidies. The results suggest that predation by riparian arthropods is a quantitatively important process in the transfer of aquatic secondary production to the riparian food web.     

Herbivorous arthropods on bracken (Pteridium aquilinum (L.) Khun) in Australia compared with elsewhere

Abstract The herbivorous arthropod fauna of bracken fern (Pteridium aquilinum (L.) Khun), at a site near Sydney, Australia, is described and compared with previously reported bracken faunas in other geographic regions. Monthly sampling over 18 months found 17 species of herbivorous arthropods (15 insect and two mite species) from five orders. At the ordinal level, the mixture differed substantially from the bracken faunas of sites in Britain and Papua New Guinea. Notable was the presence of Thysanoptera and Acari, and the absence of Coleoptera and Hymenoptera. The mixtures of orders/families represented in the site bracken faunas in Britain, and less so in Australia, resembled those in the pool of herbivorous arthropods in those regions. Further, the mixture of orders on bracken was more similar to the mixture of orders on other ferns than to the mixture of orders among herbivorous insects on all plants; such similarity was not evident at family level. Compared with sites in other regions, the Sydney site had an abundance of pinna-sucking species and a dearth of mining species. Differences between regions in feeding niches most occupied tended to correspond with the differences in orders represented. Not all features of the fauna of bracken near Sydney reflected differences in the general herbivorous arthropod fauna of Australia compared with other regions, or differences between the herbivore faunas of ferns and seed plants. Its composition must be attributed in part to stochastic aspects of the speciation of herbivorous arthropods onto host plants.     

Bayesian inferences of arthropod movements between hedgerows and orchards

Hedgerows are agro-ecological infrastructures that are assumed to enhance biodiversity in an agro-ecosystem and the control of crop pests. However, local movements of arthropod predators from hedgerows to crops remain poorly understood. In this research, these movements were analysed in eleven commercial apple orchards over two weeks in the spring of 2014. Predators were indirectly marked by spraying ovalbumin on the hedgerows. Canopy and ground predators were captured using beating and pitfall traps, respectively, in both the orchards and the hedgerows, and individuals marked by ovalbumin were detected using ELISAs. Approximately 20% of the 1272 captured predators were identified as marked. Movements between the orchards and the hedgerows of the most abundant predator taxa (Forficula auricularia, Chrysoperla sp., Philodromus spp., Cheiracanthium mildei, and Nebria brevicollis) and of four guilds (ground spiders, ground beetles, canopy spiders and canopy insects) captured across the eleven locations were estimated using a Bayesian model. On one hand, canopy insects and ground spiders were less likely to stay in the hedgerows than were the canopy spiders and ground beetles. On the other hand, the canopy spiders and ground beetles were less likely to stay in the orchard than were the canopy insects and ground spiders. However, there were exceptions within these groups: F. auricularia and N. brevicollis exhibited a high probability of staying in the hedgerow and in the orchard, respectively. Overall, these results demonstrate the frequent movements that occurred between the orchard and the adjacent hedgerow in a diverse range of predator taxa. The probabilities of movement were further affected by the characteristics of the orchard for most taxa. Therefore, agro-environmental measures that focus on hedgerow management require consideration of the local arthropod predator communities and their characteristics to enhance pest control in apple orchards.     

Length–weight relationships of Wallago attu and Sperata sarwari from the Indus River, southern Punjab, Pakistan

The present study describes the length–weight parameters of Wallago attu and Sperata sarwari from the Indus River, southern Punjab, Pakistan with W  = 0.001698  L ^3.27 for W. attu and W  = 0.001698  L ^3.28 for S. sarwari . The values of the slope b are significantly higher than b  = 3.0, which shows that the weight grows more rapidly as compared to the cube of the length. Thus it was concluded that body proportions changed as fish grew in size.     

Age, growth and food composition of Scorpaena porcus (Linnaeus, 1758) in the southeastern Black Sea

Samples of 525 Scorpaena porcus (Linnaeus, 1758) were collected from the southeastern Black Sea between January 2002 and May 2003 for information on age, growth, length–weight relationships and stomach contents. Total length of sampled fish ranged from 4.6 to 22.9 cm and total weight from 1.3 to 220 g. The sex ratio (1 : 1.61) was biased toward females (P < 0.05). Positive allometric growth was determined in the collected samples. The length–weight relationship for all individuals was described by the parameters a  = 0.0149 and b  = 3.09, with r ² = 0.99. The sample was composed of five age-classes (0–7 years). Three growth models were used to identify the growth characteristics: von Bertalanffy,     ; logistic,     ; and Schnute and Richards,     . Among these, the Schnute and Richards model was best fitted to the data as     The diet was composed of Carcinus mediterraneus (18.2%), Crangon crangon (12.1%), unidentified crab species (10.6%), Gobius sp. (4.6%), Palaemon sp. (4.5%), Macropipus sp. (4.5%), Engraulis encrasicholus (1.5%), Gastropoda (1.5%), and unidentified remains (34.9%).     

Lake‐derived midges increase abundance of shoreline terrestrial arthropods via multiple trophic pathways

Aquatic insects link adjacent ecosystems by transporting nutrients, energy, and material as they move from bodies of water into terrestrial habitats. Insects emerging from streams and rivers are known to benefit arthropod predators such as spiders, but their influence may extend to other arthropod feeding groups as well. We conducted a terrestrial arthropod survey at a series of lakes spanning a strong gradient of midge (Chironomidae, Diptera) emergence. These small, short‐lived insects reach high densities in some areas such that their carcasses litter the ground, and serve as a potential resource for non‐predatory arthropods. Our study revealed that arthropod assemblages in areas of high midge density were significantly different from those with few midges, the result of an increase of all taxa rather than changes in taxonomic composition. Eight of nine terrestrial arthropod taxa sampled showed a strong positive response to the presence of midges including detritivores and herbivores in addition to predators. Taxa that could consume living or dead midges directly responded especially strongly to midge gradients. Our results strongly suggest that midges enter the terrestrial arthropod food web through multiple pathways, increasing numbers of a wide range of arthropods. Furthermore, they emphasize the importance of lakes as sources of aquatic insects that significantly alter processes in the neighboring terrestrial environment.     

Phylogeny of the holometabolous insect orders: molecular evidence

Phylogenetic relationships among the holometabolous insect orders were reconstructed using 18S ribosomal DNA data drawn from a sample of 182 taxa representing all holometabolous insect orders and multiple outgroups. Parsimony analysis supports the monophyly of all holometabolous insect orders except for Coleoptera and Mecoptera. Mecoptera is paraphyletic with respect to Siphonaptera, which is nested within Mecoptera. Coleoptera is scattered as a paraphyletic assemblage across the tree topology. These data support a monophyletic Halteria (Strepsiptera + Diptera), Amphiesmenoptera (Trichoptera + Lepidoptera), Neuropterida (Neuroptera + (Megaloptera + Raphidioptera)), but Antliophora (Halteria + Mecoptera + Siphonaptera) and Mecopterida (Antliophora + Amphiesmenoptera) are paraphyletic. The limitations of using 18S ribosomal DNA as the sole phylogenetic marker for reconstructing insect ordinal relationships are discussed.     

复合荔枝园节肢动物群落动态的研究

对单一荔枝和复合荔枝园节肢动物群落进行了调查、分析,结果表明：复合园的捕食性、寄生性昆虫以及蜘蛛类群的数量和丰富度明显高于单一园,而植食性类群的数量和主要害虫发生数量明显少于单一园。复合园各种类型的多样性指数季节变化,其总趋势比单一园大,波动程度比单一园低。复合园的均匀度全年维持在０．８以上,而单一园的则波动很大。     

Relating body size to the role of aquatic subsidies for the riparian spider <Emphasis Type="Italic">Nephila clavata</Emphasis>

We examined the relationship between body size of the riparian spider Nephila clavata and the contribution of allochthonous (aquatic insects) and autochthonous (terrestrial insects) sources to its diet using stable isotope analysis. During the study period from July to September, the body size of the females increased remarkably (about 60-fold) but that of males remained small. The biomass of both aquatic and terrestrial insects trapped on the spider webs increased with spider size, with the biomass of the former ranging between 30 and 70% of that of the terrestrial insects. The average relative contribution of aquatic insects to the diet of the spiders, calculated from δ¹³C values, was 40–50% in spiders in the early juvenile and juvenile stages, 35% in adult males and 4% in adult females. There was a significant negative relationship between the relative contribution of aquatic insects and body size of the female spiders. We conclude that aquatic insects might be an important seasonal dietary subsidy for small spiders and that these allochthonous subsidies may facilitate the growth of riparian spiders, which may in turn enable the spiders to feed on larger prey.     

Arthropod community structure along a latitudinal gradient: Implications for future impacts of climate change

Abstract The structure of free‐living arthropod communities on the foliage of Acacia falcata was assessed along an extensive latitudinal gradient in eastern Australia. We hypothesized that abundance and biomass of arthropods within feeding groups would increase from temperate latitudes towards the tropics. We also hypothesized that the ratio of carnivores to herbivores would be consistent along the latitudinal gradient. Three sites at each of four latitudes, spanning 9° and 1150 km (Batemans Bay, Sydney, Grafton, Gympie in Australia), were sampled every season for 2 years, using pyrethrum knockdown. Abundance and biomass (based on dry weight) of arthropods within eight feeding groups were measured. The relative size of the feeding groups, and the ratio of carnivores to herbivores were then compared among latitudes and seasons. We found no consistent north to south (tropical to temperate) change in feeding group structure in terms of abundance. A weak latitudinal trend was evident for predator biomass, consisting of a reduction from north to south, but no significant trends in biomass for other feeding groups were found. Relative abundance and relative biomass of both carnivores and herbivores, as well as the ratio of carnivores to herbivores were consistent among latitudes. Finally, we compared a subset of these data to arthropod communities found on congeneric host species at individual sites along the latitudinal gradient. Overall, 68% of comparisons showed no significant differences in abundance or biomass within different feeding groups between host plants and among latitudes. We conclude that arthropod communities show consistencies among latitudes and between congeneric host species, in terms of feeding group and trophic structure. These results have implications for predicting the impacts of future climate change on arthropod communities.     

Molecular evidence for the gnathobasic derivation of arthropod mandibles and for the appendicular origin of the labrum and other structures

 Mandibles are feeding appendages functioning as ”jaws” in the arthropod groups in which they occur. Which part of this appendage is involved in food manipulation (limb tip versus limb base), has been used to suggest phylogenetic relationships among some of the major taxa of arthropods (myriapods, crustaceans, and insects). As a way to independently verify the conclusions drawn from previous morphological analyses, we have studied the expression pattern of the gene Distal-less (Dll), which specifies the distal part of appendages. Our results show, in contrast to the traditional view, that both insect and crustacean adult mandibles are gnathobasic, handling food with the basal portion of the appendage. Furthermore, as is evident by the reduction in the number of Dll-expressing cells in the later developmental stages, adult diplopod jaws are also gnathobasic. Thus, jaws of all mandibulates (myriapods, crustaceans, and insects) seem to have a similar gnathobasic structure. We have also found that Dll is expressed in the labra of all arthropod taxa examined, suggesting that this structure is of appendicular derivation. Additionally, the spinnerets and book lungs of spiders, long considered on other grounds to be modified appendages, express Dll, confirming this interpretation. This study shows that, in addition to their use in phylogenetic and population genetic studies, molecular markers can be very useful for inferring the origins of a particular morphological feature. Received: 12 January 1998 / Accepted: 23 March 1998     

DNA metabarcoding quantifies the relative biomass of arthropod taxa in songbird diets: Validation with camera‐recorded diets

Ecological research is often hampered by the inability to quantify animal diets. Diet composition can be tracked through DNA metabarcoding of fecal samples, but whether (complex) diets can be quantitatively determined with metabarcoding is still debated and needs validation using free‐living animals. This study validates that DNA metabarcoding of feces can retrieve actual ingested taxa, and most importantly, that read numbers retrieved from sequencing can also be used to quantify the relative biomass of dietary taxa. Validation was done with the hole‐nesting insectivorous Pied Flycatcher whose diet was quantified using camera footage. Size‐adjusted counts of food items delivered to nestlings were used as a proxy for provided biomass of prey orders and families, and subsequently, nestling feces were assessed through DNA metabarcoding. To explore potential effects of digestion, gizzard and lower intestine samples of freshly collected birds were subjected to DNA metabarcoding. For metabarcoding with Cytochrome Oxidase subunit I (COI), we modified published invertebrate COI primers LCO1490 and HCO1777, which reduced host reads to 0.03%, and amplified Arachnida DNA without significant changing the recovery of other arthropod taxa. DNA metabarcoding retrieved all commonly camera‐recorded taxa. Overall, and in each replicate year (N = 3), the relative scaled biomass of prey taxa and COI read numbers correlated at R = .85 (95CI:0.68–0.94) at order level and at R = .75 (CI:0.67–0.82) at family level. Similarity in arthropod community composition between gizzard and intestines suggested limited digestive bias. This DNA metabarcoding validation demonstrates that quantitative analyses of arthropod diet is possible. We discuss the ecological applications for insectivorous birds.