Larvae of Ceratocanthidae and Hybosoridae (Coleoptera: Scarabaeoidea): study of morphology, phylogenetic analysis and evidence of paraphyly of Hybosoridae

Abstract. Larvae of the scarabaeoid genera Germarostes Paulian, Cyphopisthes Gestro, Paulianostes Ballerio, Ceratocanthus White, Pterorthochaetes Gestro, Madrasostes Paulian, Astaenomoechus Martínez & Pereira (Ceratocanthidae) and Hybosorus Macleay, Phaeochrous Castelnau, and Anaides Westwood (Hybosoridae) are described, keyed and illustrated with fifty‐seven drawings. A phylogenetic analysis of these two families based on larval morphology is presented. Fifty‐four larval morphological and three biological characters from twenty‐seven taxa revealed nineteen equally parsimonious cladograms. The monophyly of (Ceratocanthidae + Hybosoridae) is supported by four unambiguous unique synapomorphies: dorsal medial endocarina on cranium extended anteriorly into frontal sclerite; presence of large membranous spot on apical antennomere; labium dorsally with four pores in centre (secondarily reduced to two pores in some groups); and presence of stridulatory organ on fore‐ and middle legs (secondarily reduced in some groups). Our analysis suggests that the family Hybosoridae is paraphyletic with respect to Ceratocanthidae. The clade comprising the hybosorid genera Hybosorus and Phaeochrous is the sister group of the remaining Hybosoridae plus Ceratocanthidae. It is supported by two unambiguous synapomorphies: two apical antennomeres completely joined and the stridulatory organ represented by seven to nine large teeth anteriorly on the middle leg. The hybosorid genus Anaides is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus and Phaeochrous) and the ceratocanthid genus Germarostes is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus, Phaeochrous and Anaides). The ceratocanthid genera Cyphopisthes, Astaenomoechus, Paulianostes, Pterorthochaetes, and Madrasostes constitute a sister group to the hybosorid genus Cryptogenius and are supported by the presence of two reversions: two dorsal pores on labium and completely reduced stridulatory organs on fore‐ and middle legs.     

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.     

Germ band differentiation in the stomatopod Gonodactylaceus falcatus and the origin of the stereotyped cell division pattern in Malacostraca (Crustacea)

We analysed aspects of the embryonic development of the stomatopod crustacean Gonodactylaceus falcatus focusing on the cell division in the ectoderm of the germ band. As in many other malacostracan crustaceans, the growth zone in the caudal papilla is formed by 19 ectoteloblasts and 8 mesoteloblasts arranged in rings. These teloblasts give rise to the cellular material of the largest part of the post-naupliar germ band in a stereotyped cell division pattern. The regularly arranged cells of the genealogical units produced by the ectoteloblast divide twice in longitudinal direction. The intersegmental furrows form within the descendants of one genealogical unit in the ectoderm. Hence, embryos of G. falcatus share some features of the stereotyped cell division pattern with that in other malacostracan crustaceans, which is unique among arthropods. In contrast to the other malacostracan taxa studied so far, stomatopods show slightly oblique spindle direction and a tilted position of the cells within the genealogical units. The inclusion of data on Leptostraca suggests that aspects of stereotyped cell divisions in the germ band must be assumed for the ground pattern of Malacostraca. Moreover, Stomatopoda and Leptostraca share the lateral displacement of cells during the mediolateral divisions of the ectodermal genealogical units in the post-naupliar germ band. The Caridoida within the Eumalacostraca apomorphically evolved the strict longitudinal orientation of spindle axes and cell positions, reaching the highest degree of regularity in the Peracarida. The phylogenetic analysis of the distribution of developmental characters is the prerequisite for the analysis of the evolution of developmental patterns and mechanisms.     

Roles of environmental variables and land usage as drivers of dung beetle assemblage structure in mopane woodland

Colophospermum mopane woodland covers large areas of dry lowland savanna in southeastern Africa. Dominant land usage is conservation (45%) with the remainder mostly modified by farming. Dung beetle responses to environment (dung type, habitat, weather) and land usage (conservation, farming, mining) were examined at Phalaborwa (23.9431°S 31.1411°E) in the Phalaborwa‐Timbavati Mopaneveld, South Africa. Partitioning of gamma species richness and diversity showed lower alpha values in mine areas than in farm and conserved areas. However, between‐land usage differences in species richness, alpha diversity, abundance and biomass, showed lower significance than those between dung type and different weather. At two sampling scales, three multivariate techniques variously separated assemblages according to land usage, dung type and weather. Analysis of 21 mean samples separated clusters according to dung type (Canonical Correspondence Analysis, CCA) or mine assemblages, conserved plus farm assemblages on pig plus elephant, or cattle dung (NMDS, Factor Analysis) with shared variance of >80% and unique variance of 16–18% per cluster. In analysis of 188 samples (CCA), each overlapping dung type cluster was offset in ordinal space with congruent patterns of separation according to land usage and weather (drier days distant from moister days; conserved plus farm areas distant from early succession mine areas, which were distant from disturbed and later succession mine areas). Mining, dung types, and moist conditions were the strongest contributors to between‐assemblage differences. Compared with conserved areas, dung beetle diversity is appreciably altered by mining but only slightly altered by intensive game farming or livestock ranching with subsistence agriculture.