A functional classification for analysis of the structure of dung beetle assemblages

Abstract. 1. Regional scarabaeid dung beetle assemblages in southern Africa may contain over 100 species, ranging in live weight from 10 mg to 10 g. These show a wide variety of dung-use and reproductive strategies.
2. To facilitate analysis of these diverse assemblages, a system of classification analogous to guilds is proposed. Scarabaeid dung beetle species are allocated to one of seven functional groups (FGs) according to the way they use and disrupt dung. Each group therefore contains a set of species which are functional analogues of each other. This classification provides a conceptual framework within which to analyse the structure of dung beetle assemblages and the interactions between dung beetles and other dung-breeding species such as coprophagous flies.
3. There is a clear hierarchy of functional groups in their ability to compete for dung. Competitively dominant groups such as the large ball rollers (FG I) and fast-burying tunnellers (FG III) are mostly large, aggressive beetles which rapidly remove dung from the pad. The smaller ball rollers (FG II) are also effective competitors for dung. Subordinate groups are those which bury dung slowly over many days (FG IV and V) and those which breed inside the pad (FG VII, endocoprids). Kleptocoprids (FG VI) breed in dung buried by other beetles and so are not part of the hierarchy.
4. The use of this classification is illustrated by reference to three contrasting assemblages of dung beetles in a summer rainfall region of southern Africa. The potential of these beetles for biological control of dung-breeding flies is discussed.     

Duration and benefits of biparental brood care in the dung beetle Onthophagus vacca (Coleoptera: Scarabaeidae)

Abstract.

• 1 Single males, single females or pairs of dung beetles, Onthophagus vacca, were released on artificial small (100 g) or large (1000 g) dung pats in the laboratory. Emigrating beetles were trapped at 12 h intervals, and the number and size of the brood chambers were recorded after each replicate.
• 2 Emigration of males was delayed if females were present in the same dung pats, whereas emigration times of females were independent of the presence or absence of males.
• 3 A residency of 60 h proved to be a threshold value. Females emigrating before this time did not breed, whereas those emigrating later had built at least two brood chambers.
• 4 Females paired with males built more brood chambers than single females.
• 5 The reproductive success of pairs was not influenced by the size of the dung pats.

     

Dung burial strategies in some South African coprine and onitine dung beetles (Scarabaeidae: Scarabaeinae)

ABSTRACT.

• 1 The dung colonization and dung burial behaviour of twelve crepuscular/nocturnal tunnelling (paracoprid) species of beetles were examined in order to identify mechanisms which might facilitate resource (dung) partitioning. The species were selected from a diverse assemblage of dung beetles, the members of which coexist in the sandy-soil regions of Natal, South Africa.
• 2 The pattern of dung colonization in relation to dung age was examined in the field using baited pitfall traps. Some species, e.g. Onitis deceptor Peringuey, Catharsius tricornutus De Geer and Copris elphenor Klug, showed a marked preference for fresh dung (1–2 days old) whereas other species, e.g. O. viridulus Boheman and Copris fallaciosus Gillet, preferred older dung (3–7 days old).
• 3 Two distinct patterns of dung burial were recognized. In the Coprini, dung burial was complete within 24–48h of pad colonization, and the level of dung burial was similar in the laboratory and in the field. In the Onitini, dung burial occurred progressively over a 12-day period, although the timing of initiation of dung burial varied between species: in O. deceptor nearly all individuals had begun burial within 2 days of pad colonization, whereas only 20% of O. viridulus had commenced dung burial by that time. However, nearly all O. viridulus had buried substantial quantities of dung by day 12.
• 4 The mass of dung buried per pair by the larger coprine beetles (100–300 g) and onitine beetles (400–1000 g) suggests that there is potential for inter- and intraspecific competition, even in pads colonized by relatively few beetles. The colonization and use of dung of different ages are discussed as means of resource partitioning in relation to the relative abilities of species to compete for dung.

     

Can taxonomic distinctness assess anthropogenic impacts in inland waters? A case study from a Mediterranean river basin

1. It is increasingly recognised that adequate measures of biodiversity should include information on the ‘relatedness’ of species within ecological assemblages, or the phylogenetic levels at which diversity is expressed. Taxonomic distinctness measures provide a series of indices to achieve this, which are independent of sample size. Taxonomic distinctness has been employed widely in marine systems, where it has been suggested that this index can provide a reliable measure of anthropogenic impact. 2. We tested the behaviour of three related taxonomic distinctiveness indices (Average Taxonomic Distinctness, Δ⁺; Variation in Taxonomic Distinctness, Λ⁺; and Total Taxonomic Distinctness, sΔ⁺) in relation to putative levels of anthropogenic impact in inland waters and their potential utility in environmental monitoring, using an extensive data set for aquatic beetles from the south‐east of the Iberian Peninsula. 3. Taxonomic distinctness measures were not able to identify human disturbance effects and there were no clear relationships between these new biodiversity measures and the disturbance level recorded at individual localities. Furthermore, the taxonomic distinctness measures used were apparently less sensitive to the effects of anthropogenic impact than other diversity metrics, such as species richness and rarity. 4. We conclude that taxonomic distinctness indices may not always perform as well as other metrics in the assessment of environmental quality. In addition, taxonomic distinctness measure should be interpreted with caution, as their performance and ability to detect anthropogenic disturbance may depend on the phylogenetic structure of sampled taxa within a region, and their evolutionary and ecological history.     

Area Networks for Conserving Iberian Insects: A Case Study of Dung Beetles (col., Scarabaeoidea)

Possibilities to create an efficient network of areas for the conservation of Iberian dung-beetle biodiversity were investigated. Data were taken from BANDASCA, a 15,740 record dung-beetle database with information for 101,996 specimens. The greatest species richness and phylogenetic diversity are found in the Iberian Central System cordilleras, while those of range-size rarity are in the southernmost corner of the Iberian Peninsula and in the coastal plains of the Guadalquivir basin. Additionally, broad regional score trends of these variables reach a maximum in a smaller northeastern, and a larger southeastern region (Baetic and sub-Baetic mountains). Major Central and Southern Spain dung-beetle hotspots contain many of the UTM grid cells which delimit the near-minimum-set of areas hosting maximum dung-beetle biodiversity scores. They are viewed as the nucleus of a potential network of reserves. A UTM grid cell (30TXK1) where Onthophagus albarracinus, an endemic and demographically rare species inhabits the Sierra of Albarracín and Montes Universales, was detected as an irreplaceable area. Because of recent land use changes and rapid human-induced habitat transformation threatening dung-beetle fauna, caution should be taken not to recommend an in memoriam network of areas no longer hosting a fauna extinct today. The abandoning of traditional herding management in Europe threatens the preservation of grasslands where dung-feeding beetles play a major ecological role in maintaining pasture quality and livestock health. A return to more traditional herding methods would require stable dung-beetle populations. Any insect conservation planning in Spain would benefit from more biogeographical information for other insect groups.     

Spatial Variation in Fish Assemblages across small Mediterranean Drainages: Effects of Habitat and Landscape Context

Much debate about assemblage organization in stream fish may stem from analysing the effects of both local and large-scale processes on assemblage attributes over whole geographic regions. This study addresses this issue, by examining the contribution of local habitat attributes and landscape context to fish assemblage variation across small Mediterranean drainages in southern Portugal. Fish abundance and species composition was estimated in 28 sites, across 10 drainages, in both a dry year (1999) and a wet year (2001), and related to two sets of variables reflecting habitat and landscape characteristics. Fish showed responses to both sets of variables with variance partitioning indicating that landscape context had important effects on species richness whereas habitat attributes were the primary determinants of local fish abundance. In general, high species richness was associated with larger drainage area and higher rainfall variability, whereas variation in species abundances mostly reflected the influence of width, depth, conductivity, current velocity, substrate size and emergent vegetation. The relative contributions of both landscape context and habitat attributes to species richness and abundance were generally lower in 1999 than in 2001, with much less diversified species–habitat relationships being found in the former dry year. These results point to the dynamic nature of assemblage organization, emphasizing the importance of innovative, multi-scale approaches in advancing our knowledge of fish assemblage structure in Mediterranean streams.     

The effects of rainforest canopy loss on arboreal dung beetles in Borneo: implications for the measurement of biodiversity in derived tropical ecosystems

We examine the effect of selective timber extraction, and corresponding forest canopy loss, on arboreal dung beetles in the tropical rainforests of Sabah, Malaysian Borneo. Changes in vertical distribution of beetles are measured through differences in the abundance of beetles in ground-based pitfall traps in primary, logged and plantation forest. Previous research has demonstrated that arboreal dung beetles are not generally collected in pitfall traps in primary forest, but are present in large numbers above the ground in canopy vegetation: the presence of arboreal beetles in pitfalls in areas of reduced or modified canopy cover may therefore reflect a response to the absence or modification of their usual habitat, and the proliferation of these beetles nearer to the ground. In this paper, statistically significant differences are found in the abundance of beetles in ground pitfall traps from logged forest compared to primary forest. Results show that virtually no arboreal dung beetles are recorded in primary forest traps, with an increased abundance of arboreal dung beetles in traps from logged and plantation forest, with 1.72% of the total number of arboreal beetles recorded from primary forest, 22.32% from logged forest, and 75.96% from plantation forest. The presence of arboreal dung beetles in plantations demonstrates that arboreal dung beetles can survive outside their normal habitat, and we relate these observations to adaptations to upper rainforest canopy conditions, and proliferation of these microclimatic conditions in man-made habitats. Results are also discussed in terms of their relevance to the measurement of species richness and diversity in logged and other derived ecosystems, where mixing of the ground-based and arboreal faunas occurs.     

Factors influencing duiker dung decay in north‐east Gabon: are dung beetles hiding duikers?

We analysed seasonality of dung decay time and the influence of rainfall, leaf fall, beetle attack and moisture on dung decay. Our study was carried out at the Ipassa Reserve, north-east Gabon. We compared the seasonality of dung decay with the seasonality of dung beetle abundance and guild structure. Dung beetle activity was the main factor influencing dung decay in our study site. Decay time was the highest during the main dry season (3.4 days) and the lowest during the short rainy season (0.7 days). Dung decay time was closely related to dung beetle abundance, especially to the abundance of nocturnal beetles. We discuss the implications of such results for duiker survey methods based on dung pellet counts in areas where decay time is extremely short or unknown.     

Phylogenetic relationships of Iberian Aphodiini (Coleoptera: Scarabaeidae) based on morphological and molecular data

A phylogeny of Iberian Aphodiini dung beetles was reconstructed based on morphological and molecular data. The data set included a total of 84 variable characters from wing venation, mouthparts, genitalia, and external morphology, as well as mitochondrial partial cytochrome c oxidase I (COI), complete tRNA-Leu (UUR), and partial cytochrome c oxidase II (COII) gene nucleotide sequences (1210 positions). Phylogenetic trees based on molecular data were relatively more resolved than those based on morphological characters. The Bayesian analysis of combined molecular and morphological data provided resolution not achieved by each data set separately. Ammoecius and Aphodius are the first lineages that branch off from the tree, followed by Acrossus, Nimbus, and Heptaulacus. The remaining studied taxa are recovered in a more derived clade that lacks internal resolution. Reconstructed trees based on molecular data showed relatively short internal nodes that were weakly supported. Such pattern may reflect a rapid radiation at the origin of the tribe Aphodiini, but also saturation of mutational changes. Several tests were conducted to discern between both competing hypotheses, as well as to assess the effect of incomplete taxon sampling.     

Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.)

Climatic and biogeographical associations of southern African dung beetles (Scarabaeinae, Coprinae) were analysed from a collection amassed between 1971 and 1986. Endemism to Africa south of 15°S was much greater in southwesterly climates (winter rainfall, bimodal spring/autumn rainfall, arid late summer rainfall) than to the north‐east in the moist, mid‐summer rainfall region. Major biogeographical groups centred to the south‐west comprised predominantly southern African endemics, Western/Eastern Cape coast, Karoo, Karoo/Namib, northern Namibia and the south‐western Kalahari. Biogeographical groups centred on the south‐eastern highlands and the subtropical east coast (mid‐summer rainfall) also comprised predominantly southern African endemics. All other major groups centred to the north‐east in the mid‐summer rainfall region comprised predominantly species with widespread tropical biogeographical affiliations, pan‐southern Africa but centred in the east, pan‐mid‐summer rainfall region, eastern mid‐summer rainfall region, tropical east Zimbabwe/central Moçambique, subtropical/tropical game reserves (non‐ruminant dung specialists). There were cross‐climatic differences in taxonomic composition of the fauna. Within the winter rainfall region, percentage species composition of Scarabaeinae was greater whereas that of the coprine tribe, Onthophagini, was lower than within the other three climatic regions. Percentage species richness of most other tribes of Coprinae differed little between most climates but that of Scarabaeinae declined from west to east (Canthonini, Scarabaeini), east to west (Sisyphini) or to either side of the late summer rainfall region (Gymnopleurini).