Synopsis of Antitrogus Burmeister (Coleoptera: Scarabaeidae: Melolonthini)

Abstract The genus Antitrogus Burmeister is widely distributed in eastern Australia and, because some species are significant pests of sugarcane, pastures and pineapples, it has attracted considerable research interest. This paper reviews the systematics, biology, ecology and management of the genus. The morphology of the 22 known species is described and a further species, A. gubbi sp. n. from south-eastern Queensland which was previously confused with A. robertsi Britton, is described. These species clearly divide into two species-groups: (i) those from south-eastern Queensland, eastern New South Wales and south-eastern Australia, which sometimes have closely related species pairs; and (ii) a more diverse group of species from central and northern Queensland. The latter is linked to A. villosus Allsopp from western Victoria, and this may indicate that further species await discovery in western New South Wales. Known life cycles are of 1 or 2 years. Adults fly for short periods following rain in spring or summer and do not feed. Adult females tend to disperse poorly, and this, coupled with ephemeral and localised rainfall, has probably contributed to speciation. Larvae feed on the roots of grasses and other similar plants. Insecticides are widely used to control them in sugarcane, but are too expensive for use on pastures. Integrating new controls with an emphasis on cultural controls is seen as the best way of managing these pest species.     

Effect of Soil Moisture on Oviposition by Childers Canegrub, Antitrogus parvulus Britton (Coleoptera: Scarabaeidae)

Ovipositing Childers beetles, Antitrogus parvulus Britton, exhibit preference for soils of certain moisture levels. In a choice trial, A. parvulus laid most eggs in soil at approximately—74 kPa (near field capacity), no or very few eggs in soil near wilting point or drier, and few eggs in soil above field capacity (approximately -5.7 kPa). Under no-choice conditions most females retained their eggs if soil was near wilting point, oviposited immediately in soil near field capacity, and tended to extend egg-laying in soil above field capacity. Females in very wet soil laid many eggs when soil was changed to near field capacity within 3 d. After 3 d there was very little egglaying response to a change to soil near field capacity.     

Insect parasites of scarabs from sugarcane fields in southern Queensland (Coleoptera: Scarabaeidae)

Data on parasitism of scarabs were collected by sampling sugarcane in the Bundaberg region of southern Queensland during 1993–1997. The scoliid Campsomeris tasmaniensis (Saussure) parasitised third instars of the melolonthines Antitrogus parvulus Britton, Lepidiota crinita Brenske, Lepidiota negatoria Blackburn and Lepidiota noxia Britton, and of the ruteline Anoplognathus porosus (Dalman). Levels of parasitism were low (< 2%). Third instars of A. porosus were also parasitised by the dexine tachinid Rutilia sp. Puparia of Prosena sp. (Tachinidae: Dexiinae) were found in soil with larvae of L. noxia . Larvae of Prosena sp. are parasites of scarab larvae elsewhere, and are probably parasites of scarabs in southern Queensland. Adults of A. parvulus , L. crinita and L. noxia were parasitised by the tachinine tachinid Palpostoma sp. at levels of 2–18.4%.     

Overwintering,soil distribution and phenology of Childers canegrub,Antitrogus parvulus (Coleoptera: Scarabaeidae) in Queensland sugarcane

In this study, the question of whether Childers canegrub, Antitrogus parvulus (Britton) overwinters in the subsoil was addressed. Irrigated fields of sugarcane were sampled during a 2-year period near Bundaberg in southern Queensland. Antitrogus parvulus overwintered as second and third instars at each of three sites. During autumn and winter third instars of different allochronic (separated in age by 12 months) populations occurred together and could not be readily separated. Field-collected third instars were reared on ryegrass and separated into two age groups based on the date of pupation. Third instars in the first year of their life cycle (young third instars) remained at shallow depth (100-200 mm) and did not overwinter in the subsoil as once thought. Minimum temperatures during winter were 13-16 degrees C and did not prevent young third instars from feeding and gaining weight. Third instars in their second and final year moved downwards from late summer and pupated in the subsoil at 293-425 mm in spring. General phenology was as previously reported with first instar larvae occurring from January until April, second instars from January until November and third instar larvae throughout the year. Prepupae and pupae were found between October and December and adults occurred in soil during November and January. Batches of eggs occurred at a mean depth of 350 mm. First and second instars occurred predominantly at relatively shallow (100-200 mm) depths in the soil profile. All stages tended to be most common under rows of sugarcane rather than in the interrow.     

Between-group differences in the Iberian dung beetle species-area relationship (Coleoptera: Scarabaeidae)

Between-group α- and β-diversity differences were derived from species-area relationships fitted to field data. The accuracy of spatial richness variation predictions based on area size was also checked. The log-log model (log S = c + z log A) was found to be the best-fit linear model, with slopes (z) ranging from 0.089 to 0.142. Between-group comparisons of z (slope) and q (intercept) parameters, using the S = q + cA^z curvilinear regression model, corroborated early results, indicating a lower β-diversity (slope) for Scarabaeinae than for Geotrupinae and Aphodiinae. The latter group, probably more sensitive to environmental heterogeneity, should contribute more to species richness in large areas. α-Diversity is greater for Aphodiinae, more relevant to local diversity (1 km²), than for Scarabaeinae and considerably greater for these two groups than for Geotrupinae. As earlier results show that the richness of a single dung pat is rather more a function of the Scarabaeinae species pool, richness on dung pat scales is probably due more to the between-dropping mobile Scarabaeinae, while Aphodiinae contribute mainly to local and regional pool richness. Nearly 88 % of the total richness variance is explained by area size. This percentage decreases to 37 % when the spatial structure of area size and species number are extracted. The corresponding figures for Scarabaeinae, Aphodiinae and Geotrupinae are 44, 22 and 31 %, respectively.     

Phylogeography and population genetics of the endangered Amazonian manatee, Trichechus inunguis Natterer, 1883 (Mammalia, Sirenia)

We used mitochondrial DNA control region sequences to examine phylogeography and population differentiation of the endangered Amazonian manatee Trichechus inunguis. We observe lack of molecular differentiation among localities and we find weak association between geographical and genetic distances. However, nested clade analysis supports restricted gene flow and/or dispersal with some long-distance dispersal. Although this species has a history of extensive hunting, genetic diversity and effective population sizes are relatively high when compared to the West Indian manatee Trichechus manatus. Patterns of mtDNA haplotype diversity in T. inunguis suggest a genetic disequilibrium most likely explained by demographic expansion resulting from secession of hunting and enforcement of conservation and protective measures. Phylogenetic analysis of T. manatus and T. inunguis haplotypes suggests that T. inunguis is nested within T. manatus, effectively making T. manatus a paraphyletic entity. Paraphyly of T. manatus and recent divergence times of T. inunguis and the three main T. manatus lineages suggest a possible need for a taxonomic re-evaluation of the western Atlantic Trichechus.     

Entomopathogenic nematodes as potential biological control agents of Popillia japonica (Coleoptera,Scarabaeidae) in Piedmont Region (Italy)

In 2014, the Japanese beetle Popillia japonica (Coleoptera: Scarabaeidae) was first spotted in northern Italy in the Nature Park of the Ticino Valley, its first detection in continental Europe. This polyphagous invasive species has the potential to cause serious losses to horticulture and agriculture. Particularly for its management in a Nature Park, environmentally friendly strategies are necessary. To develop baseline data for a biological control approach to the Italian outbreak of P. japonica, we conducted laboratory and field experiments testing several indigenous and commercial strains of the entomopathogenic nematode (EPN) species Heterorhabditis bacteriophora and Steinernema carpocapsae against P. japonica larvae. In the laboratory, strains of H. bacteriophora caused greater mortality (ranging from 57% to 100%) than those of S. carpocapsae (3% to 77%). In micro‐plot field tests carried out at three different times, the most virulent indigenous EPN strain, H. bacteriophora ItH‐LU1, showed again the best results ranging from 44% to 93% against young larvae. Finally, in a large‐plot field trial, the commercial H. bacteriophora product provided 46% larval mortality. This study shows that H. bacteriophora strains have good potential as biological control agents of larvae of the invasive P. japonica in northern Italy.     

Phylogeny and biogeography of the Japanese rhinoceros beetle,Trypoxylus dichotomus (Coleoptera: Scarabaeidae) based on SNP markers

The Japanese rhinoceros beetle Trypoxylus dichotomus is one of the largest beetle species in the world and is commonly used in traditional Chinese medicine. Ten subspecies of T. dichotomus and a related Trypoxylus species (T. kanamorii) have been described throughout Asia, but their taxonomic delimitations remain problematic. To clarify issues such as taxonomy, and the degree of genetic differentiation of Trypoxylus populations, we investigated the genetic structure, genetic variability, and phylogeography of 53 specimens of Trypoxylus species from 44 locations in five Asian countries (China, Japan, Korea, Thailand, and Myanmar). Using specific‐locus amplified fragment sequencing (SLAF‐seq) techniques, we developed 330,799 SLAFs over 114.16M reads, in turn yielding 46,939 high‐resolution single nucleotide polymorphisms (SNPs) for genotyping. Phylogenetic analysis of SNPs indicated the presence of three distinct genetic groups, suggesting that the various subspecies could be treated as three groups of populations. PCA and ADMIXTURE analysis also identified three genetic clusters (North, South, West), which corresponded to their locations, suggesting that geographic factors were important in maintaining within population homogeneity and between population divergence. Analyses of SNP data confirmed the monophyly of certain subspecies on islands, while other subspecies (e.g., T. d. septentrionalis) were found to be polyphyletic and nested in more than one lineage. AMOVA demonstrated high level of differentiation among populations/groups. Also, pairwise F_ST values revealed high differentiation, particularly between South and West, as well as between North and South. Despite the differentiation, measurable gene flow was inferred between genetic clusters but at varying rates and directions. Our study demonstrated that SLAF‐seq derived markers outperformed 16S and COII sequences and provided improved resolution of the genetic differentiation of rhinoceros beetle populations from a large part of the species’ range.     

Effect of agronomic factors on the spatial distribution of greyback canegrub, Dermolepida albohirtum (Waterhouse) (Coleoptera: Scarabaeidae), in sugarcane in the Burdekin River Irrigation Area

Abstract A series of studies were undertaken examining the factors that impact on the spatial distribution of greyback canegrub infestations in sugarcane in the Burdekin region of Queensland. Historic records of damage by greyback canegrub showed that sugarcane blocks planted or harvested earlier than surrounding blocks were more likely to be damaged than blocks planted or harvested later. The derived hypothesis that sugarcane height may be the primary determinant of where damage occurs was confirmed in field studies. The tallest sugarcane blocks at the time of oviposition consistently had the highest grub populations. There was no difference in the distribution of damage between the cultivars Q96 and Q117 and the age of these crops also had no impact. The finding that sugarcane height is the primary determinant of where damage occurs on Burdekin farms opens the possibility of using it as a tool to manipulate where canegrubs oviposit and the development of a range of associated cultural control strategies.     

Review of the subgenus Polyphylla (Granida) from continental Asia (Coleoptera, Scarabaeidae, Melolonthinae)

A review of Polyphylla Harris, 1841, species belonging to the subgenus Granida Motschulsky, 1861, from continental Asia is presented. One new species is described from Thailand: Polyphylla (Granida) simonisp. n.Polyphylla (Granida) nikodymi de Wailly, 1993, is recorded from Thailand for the first time. Polyphylla (Granida) minor Nomura, 1977, is recorded from Yunnan (China) for the first time. The previously unknown female of Polyphylla (Granida) phongsali Zídek, 2006, is described.     

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.     

Genetic variation and origin of red turpentine beetle (Dendroctonus valens LeConte) introduced to the People's Republic of China

Abstract 1 The red turpentine beetle, Dendroctonus valens LeConte, is a recent New World introduction to the People's Republic of China. An outbreak of these beetles has infested over 0.5 million hectares of pine forests. 2 Efforts are underway to suppress this outbreak using biological control measures. However, the wide distribution in the native range of D. valens suggests regional variation of the beetle's biology, predators, and parasitoids. Thus, knowledge of the origin of these beetles can help devise precise and effective control measures. 3 A portion of the mitochondrial cytochrome oxidase subunit I gene was sequenced for 218 individuals from 32 populations throughout the native range of D. valens and in China. 4 Haplotype diversity was high. A total of 131 haplotypes were found and Jukes–Cantor corrected nucleotide difference ranged from 0 to 16%. Haplotype diversity ranged from 0.53 to 0.98 and unique haplotypes were found in most populations. 5 Parsimony and statistical parsimony analyses of these haplotypes support the hypothesis that the introduction of D. valens to China was recent and originated from the Pacific North‐west of the U.S.A. 6 In addition, the high haplotype diversity also suggests a large or multiple introductions. However, based on the genetics of the beetle's reproductive behaviour, this diversity may also be explained by a limited number of individuals or introductions.     

A review of the scarab beetle tribe Aphodiini (Coleoptera,Scarabaeidae) of the fauna of Russia

An annotated list of 183 species of the dung-beetle tribe Aphodiini of the Russian fauna is given. The highest species diversity is characteristic of the southern mountainous regions of the country, the richest local faunas being known from the Lower Volga area. The greatest numbers of species are known from the steppe and broadleaved forest zones and from the nemoral types of altitudinal zonation of vegetation. The bulk of the fauna is formed by the species with wide ranges; 111 species occur in two or more landscape zones or altitudinal zonation types. Six groups of faunistically similar zoogeographic provinces are distinguished within the territory of Russia.     

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.

     

Conservation strategies for poorly surveyed taxa: a dung beetle (Coleoptera, Scarabaeidae) case study from southern Africa

Despite being the focus of an international research effort spanning decades, the spatial distribution of southern African scarab beetles remains poorly documented. As well as reinforcing the magnitude of the challenge facing biodiversity scientists, this raises real concerns about best practice conservation strategies in the absence of detailed distribution information. However, dung beetles appear to be well represented in established conservation areas. This apparent contradiction could be ascribed to anthropogenic transformation, successful conservation efforts, the presence of dung generalists and reserve-biased or mesic-biased dung beetle collection efforts. It is suggested that all of the above contribute to the observed pattern to varying degrees. The implications of selecting areas that are either rich in species, contain rare species or contain taxonomically distinct species from a group whose taxonomy is well known but for which inadequate distribution data exist are explored. Best practice, in the face of inadequate data, appears to revolve around a subtle interplay between advantages and disadvantages associated with data interpolation techniques, reserve selection algorithms that use criteria more robust than database rarity (such as taxonomic distinctiveness) and the long-term economic costs of proceeding with the data at hand versus investing in biological surveys.     

Phylogeography of the brown hare (Lepus europaeus) in Europe: a legacy of south‐eastern Mediterranean refugia?

Aim We analysed the population genetics of the brown hare (Lepus europaeus) in order to test the hypothesis that this species migrated into central Europe from a number of late glacial refugia, including some in Asia Minor. Location Thirty‐three localities in Greece, Bulgaria, Italy, Croatia, Serbia, Poland, Switzerland, Austria, France, Germany, the Netherlands, Spain, the United Kingdom, Turkey and Israel. Methods In total, 926 brown hares were analysed for mitochondrial DNA (mtDNA) variation by restriction fragment length polymorphism (RFLP) performed on polymerase chain reaction‐amplified products spanning cytochrome b (cyt b)/control region (CR), cytochrome oxidase I (COI) and 12S–16S rRNA. In addition, sequence analysis of the mtDNA CR‐I region was performed on 69 individuals, and the data were compared with 137 mtDNA CR‐I sequences retrieved from GenBank. Results The 112 haplotypes detected were partitioned into five phylogeographically well‐defined major haplogroups, namely the ‘south‐eastern European type haplogroup’ (SEEh), ‘Anatolian/Middle Eastern type haplogroup’ (AMh), ‘European type haplogroup, subgroup A’ (EUh‐A), ‘European type haplogroup, subgroup B’ (EUh‐B) and ‘Intermediate haplogroup’ (INTERh). Sequence data retrieved from GenBank were consistent with the haplogroups determined in this study. In Bulgaria and north‐eastern Greece numerous haplotypes of all five haplogroups were present, forming a large overlap zone. Main conclusions The mtDNA results allow us to infer post‐glacial colonization of large parts of Europe from a late glacial/early Holocene source population in the central or south‐central Balkans. The presence of Anatolian/Middle Eastern haplotypes in the large overlap zone in Bulgaria and north‐eastern Greece reveals gene flow from Anatolia to Europe across the late Pleistocene Bosporus land‐bridge. Although various restocking operations could be partly responsible for the presence of unexpected haplotypes in certain areas, we nevertheless trace a strong phylogeographic signal throughout all regions under study. Throughout Europe, mtDNA results indicate that brown hares are not separated into discernable phyletic groups.