Altitudinal effects on habitat selection of dung beetles (Scarabaeoidea: Aphodiidae) in the northern Iberian peninsula

The effects or vegetation cover on the abundance and species richness of dung beetles (Aphodiidae) were examined in three altitudinal zones: lowland (520–600 m), mid-altitude (950–963 m) and highland (1230–1275 m). Pitfall traps baited with fresh cattle dung were placed in three vegetation types at each altitudinal zone: forest, heathland and pasture. Species richness was lower in forests than in the other vegetation types. The total abundance of beetles also varied with vegetation type, but the effect was dependent on altitude. Number of individuals was higher in pasture at highlands and in heathland and forest at mid-altitude and lowlands. An altitudinal dependence of distribution of individuals between vegetation types was also found for the more abundant species in the study area. These results could be explained if the local distribution of individuals between vegetation types was determined by microcli-matic factors, mainly temperature. Differences in altitude between sites are associated with temperature changes, determining the habitat selection of species at each altitude.     

Determinant variables of Iberian Peninsula Aphodiinae diversity (Coleoptera, Scarabaeoidea, Aphodiidae)

Aim The aims of this paper are to examine diversity–variability patterns for species of Aphodiinae (Coleoptera, Scarabaeoidea, Aphodiidae) on the Iberian Peninsula, and to determine the factors that influence their geographic distribution. Location Iberian Peninsula (Spain and Portugal). Methods Data from 30 studies and their bibliographies on species of Iberian Peninsula Aphodiinae were compiled. The reliability of the inventories was evaluated using parametric species richness estimators. In addition, a further 11 variables related to rarity, geographic distribution, or phylogenetic diversity were considered. Diversity variables were analysed using principal components analysis to reduce the number of dependent variables. Subsequently, the effect of differences in locality size among the 30 studies was eliminated by calculating and retaining the residuals of the curvilinear relationship of each diversity variable with the area. Generalized linear models were used to examine the relationships between diversity and 17 environmental variables. The diversity variables and their residuals were also subject to trend surface analysis in order to identify the relevance of spatially structured variables that had not been considered. The contribution of explanatory variables was determined through hierarchical variance analysis. Results Principal components analysis of biodiversity variables revealed that most of the variability could be explained using three biodiversity indexes: BI1, correlated positively with species richness, widely distributed species, frequent species, abundant species, species occurring in North Africa, Europe and the Iberian Peninsula, and phylogenetic diversity; BI2, correlated positively with numbers of infrequent and African–Iberian species; and, BI3, correlated positively with numbers of endemic, non‐abundant, European, and Iberian‐restricted species. A latitudinal disjunction emerged in BI1, with maximum scores at the north‐western and southern corners, while maximum BI2 scores were found throughout the south, and maximum BI3 scores in the north‐west. For BI1, it was climate that had the greatest influence, followed by lithology, and livestock presence. Geographic variables were the most significant for BI2, followed by climate and livestock presence. Finally, for BI3, climate variables were the most important, while geography, lithology and livestock presence had some relevance. Main conclusions The relevance of geographic variables indicates that other unaccounted‐for factors that are spatially structured could possibly explain additional variation in Aphodiinae diversity. These factors may be historic in nature, relating to the species groups, namely the Ibero‐European and the Mediterranean or Afro‐Iberian. The northern pattern could reflect the fact that the Iberian Peninsula acted as a colonization route and as a refuge during the glacial/interglacial cycles, while the southern pattern could be a consequence of the connection between the Iberian Peninsula and North Africa during the Messinian crisis, and/or a historic relationship in common, related to human activity.     

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.

     

Species turnover of monkey beetles (Scarabaeidae: Hopliini) along environmental and disturbance gradients in the Namaqualand region of the succulent Karoo, South Africa

Species turnover of monkey beetle (Scarabaeidae: Hopliini) assemblages along disturbance and environmental gradients was examined at three sites within the arid, winter rainfall Namaqualand region of the succulent Karoo, South Africa. At each site two study plots with comparable vegetation and soils but contrasting management (grazing) histories were chosen, the disturbed sites having fewer perennial shrubs and generally more annuals and bare ground. Beetles collected using coloured pan-traps showed a consistently higher abundance in disturbed sites. Lepithrix, Denticnema and Heterochelus had higher numbers in disturbed plots, while Peritrichia numbers were lower in disturbed areas. Measures of species richness and diversity were consistently higher in the undisturbed sites. Distinctive assemblages of monkey beetles and plants occurred at each site. A high compositional turnover ( diversity) was recorded for both monkey beetles and plants along a rainfall gradient; between-site diversity values ranged from 0.7 to 0.8 (out of a maximum of 1.0). Species turnover of beetles was higher between the disturbed sites along the environmental gradient than the corresponding undisturbed sites. The high monkey beetle species turnover is probably linked to the high plant species turnover, a distinctive feature of succulent Karoo landscapes. Monkey beetles are useful indicators of overgrazing disturbance in Namaqualand, as their pollinator guilds are apparently disrupted by overgrazing. A shift away from perennial and bulb pollinator guilds towards those favouring weedy annuals was observed in disturbed areas. The consequences to ecosystem processes due to the effects of disturbance on monkey beetle communities and the role of monkey beetles as indicators of disturbance is discussed, as well as the implications of disturbance on monkey beetle pollination guilds.     

Structure and function of the mandibles of adult dung beetles (Coleoptera : Scarabaeidae)

The mouthparts of adult dung beetles (Coleoptera : Scarabaeidae) are adapted for manipulation of soft, pasty food—usually the excrement of vertebrates. Nutrients are derived primarily from micro-organisms contained in the food and these must be broken up before ingestion. The mandibles, particularly the molar lobes, are designed to finely grind these particulates; the molae function as a mortar-pestle system, which mills the organic “grist” contained in the food by a combination of squeezing and grinding actions. The ability of the molae to finely grind food particles resides primarily in the structure of the molar surface, which consists of a series of ridges bearing rows of submicroscopic scrapers (“tritors”). The incisor lobes of the mandibles scrape food from the surfaces of the galeae and lacinae, which bring food into the preoral cavity. The structure of the mouthparts of Canthon pilularius (L.), which is described in detail, is basically the same as that of 10 other species examined. Those of Aphodius and Geotrupes are similar in some respects to those of scarabaeines but are obviously less well specialized for microphagy.     

Seasonal distribution and prevalence of dung beetles (Coleoptera: Scarabaeidae) in Kurukshetra,Northern India

A total of 8306 individuals as 29 species from 3 subfamilies (Hybosorinae, Aphodiinae, Scarabaeinae) were trapped by pitfall bait trapping. The maximum number of species of Scarabaeinae (68.97%) and number of individuals of Aphodiinae (56.84%) take part in the formation of a community. The number of species, number of individuals, diversity and dominance changes throughout the season. Seasonal prevalence of 82.76% of species and 50.81% of individuals was found in June. Aphodius campestris was found to be more attracted to the pitfall bait traps. The beetle community found in May shows the maximum Shannon's Diversity Index (2.46) but Margalef's Diversity Index was highest in August (3.06). Environmental factors play an important role in the occurrence and abundance of dung beetles. In the present study non-significant but positive correlations between temperature and species richness, abundance and diversity were found. Percent relative humidity showed a positive correlation with abundance only, and negative correlations with species richness and diversity.     

Phylogeography of the Namib Desert dung beetles Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scarabaeidae)

Aim Namib biogeography in many instances remains reliant on advanced and detailed systematic studies. This study attempts to combine molecular phylogenetic data, geology and palaeo‐climatic data to (i) resolve the relationships of the 13 morphological species of Scarabaeus (Pachysoma) and (ii) relate their evolution to past climatic and geological events. Location South Africa and Namibia. Methods Sequencing of a 1197 bp segment of the mitochondrial cytochrome oxidase I (COI) gene of the 13 species within Scarabaeus (Pachysoma) was undertaken. Analyses performed included Parsimony and Maximum Likelihood as well as imposing a molecular clock. Results The molecular phylogeny showed strong support for 11 of the 13 morphological species. The remaining two species, S. (P.) glentoni and S. (P.) hippocrates, formed a complex and could not be assigned specific status on the basis of the COI gene phylogeny. Strong support for the three species formerly classified within the genus Neopachysoma was consistently obtained. The subgenus appears to have arisen c. 2.9 Ma. Species within the subgenus arose at different times, with the common ancestor to Neopachysoma and the hippocrates complex having evolved 2.65 and 2.4 Ma, respectively. Scarabaeus (P.) denticollis, S. (P.) rotundigenus, S. (P.) rodriguesi and S. (P.) schinzi are some of the youngest species, having diverged between 2 million and 600,000 years ago. Main conclusions Scarabaeus (Pachysoma) is a derived monophyletic clade within the Scarabaeini. The subgenus appears to be young in comparison with the age of the Namib Desert, which dates back to the Miocene (c. 15 Ma). The psammophilous taxa are shown to disperse with their substratum and habitat, barchan dunes. Clear south/north evolutionary gradients can be seen within the species of this subgenus, which are consistent with the unidirectional wind regime. Species with a suite of mostly plesiomorphic characters have a southerly distribution while their derived psammophilous relatives have central to northern Namib distributions. Major rivers such as the Orange, Buffels and Holgat appear to be gene barriers to certain species as well as areas of origin of speciation events.     

Temporal coexistence of dung-dweller and soil-digger dung beetles (Coleoptera, Scarabaeoidea) in contrasting Mediterranean habitats

The western part of the Mediterranean basin is a transitional biogeographical region for the distribution of the representatives of the main guilds of dung beetles; towards the south, Aphodiinae (dung-dwellers) become scarce, whereas northwards Scarabaeinae (soil-diggers) progressively disappear. The number of species in local dung beetle assemblages is enhanced by this double faunistic contribution. Annual dung beetle assemblages were sampled in two sub-Mediterranean sites, which differed by 600 m in elevation, in order to determine the phenological dynamics related to the way of using dung (dung-dwellers/Aphodiinae vs. soil-diggers/Scarabaeinae and Geotrupinae). Aphodiids were active all year round, although they were affected by summer drought and, at high elevation, by the length of the cold season. This reduced activity was related to an impoverishment of Aphodiinae and to reduced temporal segregation between species. In contrast, soil-diggers were not active all year round and showed different species assemblages in the two sites. An extension of the activity period of these beetles was observed due to the occurrence of cold resistant species at high elevation. Our results suggested that the occurrence of soil-diggers seemingly did not affect the seasonality of dung-dwellers; their local abundance showed no negative correlation and, most importantly, phenological differences between dung-dwellers were always significantly higher than the seasonal differences between dwellers and diggers.     

Raptor conservation in the Northern Cape Province,South Africa

Anderson, M.D. 2000. Raptor conservation in the Northern Cape Province, South Africa. Ostrich 71 (1 & 2): 25–32.

The semi-arid Northern Cape is the largest and most sparsely populated province in South Africa. One of the main economic activities is sheep farming, and the properties are generally very large (>5 000 ha). A heterogeneous environment results in many habitats for a wide diversity of raptors. Fifty-one different species have been recorded, of which 37 are resident, 21 are common, and eight species are listed in the South African Red Data Book—Birds. As habitat degradation is limited, besides the areas immediately adjacent to the perennial rivers, raptors are still commonly encountered in many areas. Conflict with small-stock farming, however, results in some birds being inadvertently or directly persecuted each year. Two farmer questionnaire postal surveys (1976 and 1992) indicated a decrease in range and abundance of certain scavenging species, such as the Tawny Eagle. The more recent survey also indicated that most stock mortalities occurred in the mountainous areas, with the Black Eagle being the apparent culprit. In the Northern Cape there are both positive and negative anthropogenic effects on raptors. For example, 14 species have been recorded breeding on man-made structures, while 18 species have been documented to drown in farm reservoirs. Through the activities of the Raptor Conservation Group, the Northern Cape Nature Conservation Service and the Northern Cape Raptor Conservation Forum various threats are being addressed. Raptor conservation activities include: monitoring of raptor breeding populations, road censuses, farmer extension, establishment of raptor conservancies, rehabilitation of injured raptors, issuing of certificates to land-owners who have breeding raptors on their properties, presentation of the annual Gariep Raptor Conservationist Award, production and distribution of raptor pamphlets and booklets, modification of farm reservoirs to prevent raptor drownings, modification of powerlines to prevent electrocutions, and establishment of “vulture restaurants”. Through these efforts, the status of many raptor species is improving and species such as the Bateleur and Lappet-faced Vulture are becoming increasingly common.     

Inferring species decline from collection records: roller dung beetles in Italy (Coleoptera, Scarabaeidae)

The decline of roller dung beetles (Coleoptera, Scarabaeidae) in Italy, at national and regional level, was described using a database of both literature and unpublished data, since the late of 19th century. The probability of finding roller species was assessed for each decade of the 20th century in order to detect changes in their distribution and status. An analysis of decline was performed using a complementary approach of different extinction indexes, according to IUCN criteria. During the 20th century, both the absolute number of records and the UTM cells where roller beetles were found increased clearly as a consequence of the escalation of entomological surveys. Nevertheless, comparing R species (rollers) with all the scarab dung beetles from the first to the last quarter of the 20th century, the relative frequency of roller records decreased by 31%, while the relative number of occupied UTM cells dropped by 23%. Six roller species showed a significant decline in Italy from the first to the second part of the 20th century, and nine in the last 30years. Other two species disappeared from the majority of the northern regions. Finally, the values of all the extinction indexes were strongly correlated and detected a high risk of extinction in Italy for six species. A gradual change in land use during the last three decades, consisting of a reduced extension of grasslands in favour of either intensive agriculture or reforestation after abandonment of livestock raising, was likely the main factor inducing decline and local extinction of roller species.     

Modelling the species richness distribution for French Aphodiidae (Coleoptera, Scarabaeoidea)

The species richness distribution of the French Aphodiidae was predicted using Generalized Linear Models to relate the number of species to spatial, topographic and climate variables. The entire French territory was studied, divided into 301 0.72×0.36 degree grid squares; the model was developed using 66 grid squares previously identified as well sampled. After eliminating nine outliers, the final model accounted for 74.8% of total deviance with a mean Jackknife predictive error of 10.5%. Three richest areas could be distinguished: the western head (Brittany), southwestern France, and, to a lesser extent, the northeastern region. Sampling effort should now be focused on the western head, where no square was correctly sampled, and on southwestern France, which was recognised as a diversity hotspot, both for Aphodiidae and for Scarabaeidae. The largest fraction of variability (37%) in the number of species was accounted for by the combined effect of the three groups of explanatory variables. After controlling for the effect of significant climate and topographic variables, spatial variables still explain 27% of variation in species richness, suggesting the existence of a spatial pattern in the distribution of species richness (greater diversity in western France) that can not be explained by the environmental variables considered here. We hypothesize that this longitudinal spatial pattern is due to the relevance of a western colonization pathway along the glacial-interglacial cycles, as well as by the barrier effect played by the Alps.     

Major species of the dung beetles (Superfamily: Scarabaeoidea) in three ecological systems in western Saudi Arabia

Members of the dung beetles superfamily Scarabaeiodea (families: Scarabaeidae and Aphodiidae) represent a major part of the ground‐dwelling fauna in a natural (fallow) and two agroecosystems in Western Saudi Arabia. The most abundant species include Pentodon algerinum Herbst and Aphodius assectora Balth. during the main activity period October‐May. Statistical analysis showed that there were no significant differences in the population density of both species between the three ecosystems under investigation. A pronounced peak of P. algerinum observed during November in the vegetable agroecosystem whereas a sharp peak observed during March in the natural (fallow) ecosystem. A. assectora was the dominant species with a sharp peak during February‐April in the vegetable agroecosystem which might be attributed to the abundance of decaying organic matter that resulted from crop harvesting. P. algerinum is active during early and late seasons October‐December and February‐April in the vegetable agroecosystem and natural (fallow) ecosystem respectively, whereas A. assectora is active during late season, February‐April in the vegetable agroecosystem. Low population numbers of both species were recorded in the datepalm agroecosystem which might be attributed to the predatory pressure exerted by residents of both Avian and vertebrate predators having continued presence and feeding on a high protein diet of immature and adult stages of both beetles.     

Floristic diversity in the Cape Flora of South Africa

Comprising a land area of ca 90000 km2, less than 4% of the total land area for the Southern African subcontinent, the Cape Floristic Province is one of the world's richest areas in terms of botanical diversity for its size. An estimated 8650 species of vascular plants occur in this area, about 65% of which are endemic. This is about 42% of the estimated total for all of southern Africa. The number of species packed into so small an area is remarkable for the temperate zone, and compares closely with species totals for areas of comparable size in the wet tropics. The Cape Region consists of a mosaic of sandstone and shale substrates that give rise to soils of quite different types, and in addition local areas of limestone add to the edaphic diversity. Climates across the region are extremely variable, and the predominant orographic rainfall pattern ranges from 2000mm locally to less than 100mm, often with extremely steep gradients, the result of a mountainous landscape. The edaphic diversity resulting from a mosaic of different soils is compounded by sharp local gradients in precipitation that creates an unusual number of local habitats. A feature of the some of the soils in the Region is low nutrient levels and many of the plants on such soils have low seed dispersal capabilities, a factor important in explaining the high levels of local endemism. Species richness in the Cape Region is hypothesized to have resulted from the presence of a complex mosaic of diverse habitats and steep ecological gradients against a background of relatively stable climate and geology after the mediterranean climate was established there sometime after the beginning of the Pliocene. A local or ecological mode of speciation may have been more important under these conditions than allopatric speciation.     

Phylogenetic analyses reveal reliable morphological markers to classify mega‐diversity in Onthophagini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae)

Based largely on homoplastic characters of external morphology, the current systematics of the tribe Onthophagini and allied dung beetle lineages is unstable, contradictory, and thus inefficient. A number of recently proposed molecular phylogenies conflict strongly with each other and with formal classification, and none of them provides new tools for the improvement of dung beetle systematics. We explored the source of these inconsistencies by performing an independent, morphology‐based phylogenetic analysis of the “Serrophorus complex”, one of the most systematically confusing knots among the onthophagines, that involves 52 species from various genera of Onthophagini and allied tribes. The phylogenetic pattern revealed conflicts with existing classifications and with most of the earlier molecular phylogenies. However, it was largely congruent with the molecular phylogeny (Evolution 2005, 59 , 1060), using the largest gene sampling thus far. All current competing phylogenetic hypotheses were evaluated against each other, and the degree of their biogeographic plausibility was used as an additional evaluative criterion. Of the 91 morphological characters involved in our analyses, traits belonging to the endophallic sclerites of the aedeagus had a very strong phylogenetic signal. Terminology of these endophallic characters was established and their morphology was studied in detail, illustrated, and presented as a tool for further practical use. The enormous variety of shapes of the lamella copulatrix within the Onthophagini and allies present a methodological problem in character coding for phylogenetic analyses. Based on the performance of alternative coding approaches, it is argued that a seemingly less informative absence/presence coding scheme would be a better choice. The phylogenetic structure of the Serrophorus complex has been largely resolved, and some taxonomic changes improving its systematics are recommended.
© The Willi Hennig Society 2011.     

Reproductive properties of two sympatric dung beetles,Aphodius haroldianus andA. Elegans (Coleoptera: Scarabaeidae)

1. Reproductive properties of two sympatric dung beetles, Aphodius haroldianus and A. elegans, were studied at a pasture in the central Japan.
2. Overwintered adults of A. haroldianus came flying to dung pats from May to early August and bred from June to July. A. elegans overwintered as larvae, new adults came to dung pats from mid May to mid June. After aestivation, they reproduced from October to November.
3. A. haroldianus was a species with low fecundity and large food reserve for larvae. Female of this species had short duration of residence and laid fewer number of eggs in one dung pat. On the other hand, A. elegans was a species with high fecundity and small food reserve for larvae. Female of this species had long duration of residence and laid larger number of eggs in one dung pat.
4. The daily egg production in A. haroldianus reached a peak (0.40 eggs/female/day) at middle of oviposition period, but, that in A. elegans reached a peak (6.49 eggs/female/ day) at the beginning of oviposition period. The mortality of A. haroldianus female occurred after the daily egg production reached a peak and then rapidly increased, whereas that of A. elegans began soon after the beginning of oviposition and then gradually increased.

     

Discontinuous gas-exchange cycles in Scarabaeus dung beetles (Coleoptera: Scarabaeidae): mass-scaling and temperature dependence.

Although discontinuous gas exchange cycles (DGC) are known from many insects, the effects of body size and temperature on DGC have not been widely examined. Here, these effects are investigated in five Scarabaeus dung beetle species from mesic and xeric habitats. The investigation tests two hypotheses: that previous estimates of the scaling exponents for the DGC and its characteristics are more broadly applicable to insects, and that, in response to temperature, both DGC frequency and the quantity of CO2 emitted during the open (O) phase (O-phase emission volume) are modulated. Like previous workers, we find that V&d2;co2 scaled as mass0.968 and that O-phase emission volume scaled as mass0.833. However, temperature-associated increases in .Vco2 (Q10's of 2.19-2.65) were modulated mostly by increases in DGC frequency since O-phase volumes remained constant across temperature. Flutter (F)-phase and O-phase durations were closely coupled to DGC duration, although the relationship between closed (C)-phase duration and DGC duration was less pronounced. We show that ventilation phase coefficients, previously considered a measure of the proportional duration of each phase of the DGC, calculated from the slopes of these relationships are a measure of change in phase duration with change in DGC duration and not a measure of the way in which total DGC duration is apportioned among phases. We suggest that proportions be used to estimate the contribution of each of the phases to the total duration of the DGC.