Regular inbreeding by sib-mating is one of the most successful ecological strategies in the bark beetle family Scolytinae. Within this family, the many species (119) in Coccotrypes are found breeding in an exceptional variety of untraditional woody tissues different from bark and phloem. Species delineation by morphological criteria is extremely difficult, however, as in most other inbreeding groups of beetles, perhaps due to the unusual evolutionary dynamics characterizing sib-mating organisms. Hence, we here performed a phylogenetic analysis using molecular data in conjunction with morphological data to better understand morphological and ecological evolution in this sib-mating group. We used partial DNA sequences from the nuclear gene EF-alpha and the mitochondrial genes 12S and CO1 to elucidate patterns of morphological evolution, haplotype variation, and evolutionary pathways in resource use. Sequence variation was high among species and far above that expected at the species level (e.g., 19% for CO1 within Coccotrypes advena). The tendency for exhaustive sequence variation at deeper nodes resulted in ambiguous reconstructions of the deepest splits. However, all results suggested that species with the broadest diets were clustered in a single derived position-another piece of evidence against specialization as a derived evolutionary feature. (C) 2002 Elsevier Science (USA). All rights reserved.

The weevil subfamily Scolytinae includes at least seven groups of regularly sib-mating species with extremely female-biased offspring sex ratios. The enigmatic SE-Asian bark beetle genus Ozopemon (25 spp.) belong to the most diversified clade (>1400 spp.) of such ancient inbreeding lineages. While males of all sib-mating scolytines are flightless, and are usually dwarfed versions of their larger sisters, the existence of males in Ozopemon has been a controversial issue. Some strangely modified male beetles, with fully developed aedeagus, strongly flattened pronotum and bead, and 10-segmented larviform abdomen, were first described as males of O.brownei, but were later assigned to the Histeridae. With the new evidence provided here, based on DNA sequence data from mitochondrial and nuclear gene partitions, and examination of genitalic characters, we re-assign these males, as well as males for two more species, to Ozopemon. Neoteny evolved close to the origin of sib-mating and possibly haplodiploidy, but the transition to neoteny occurred separately from all other inbreeding dryocoetine and xyleborine beetles. The neotenic development of these males is the first known example in Coleoptera, and several remarkable morphological modifications demonstrate an ontogenetic transformation series from female to males of different species. We discuss possible scenarios for the evolution of neoteny, precocity and fighting characteristics in these male beetles, in the light of W. D. Hamilton's 'ideal biofacies' of extreme inbreeding. (C) 2002 The Linnean Society of London.

Beetles in the weevil subfamilies Scolytinae and Platypodinae are unusual in that they burrow as adults inside trees for feeding and oviposition. Some of these beetles are known as ambrosia beetles for their obligate mutualisms with asexual fungi-known as ambrosia fungi-that are derived from plant pathogens in the ascomycete group known as the ophiostomatoid fungi. Other beetles in these subfamilies are known as bark beetles and are associated with free-living, pathogenic ophiostomatoid fungi that facilitate beetle attack of phloem of trees with resin defenses. Using DNA sequences from six genes, including both copies of the nuclear gene encoding enolase, we performed a molecular phylogenetic study of bark and ambrosia beetles across these two subfamilies to establish the rate and direction of changes in life histories and their consequences for diversification. The ambrosia beetle habits have evolved repeatedly and are unreversed. The subfamily Platypodinae is derived from within the Scolytinae, near the tribe Scolytini. Comparison of the molecular branch lengths of ambrosia beetles and ambrosia fungi reveals a strong correlation, which a fungal molecular clock suggests spans 60 to 21 million years. Bark beetles have shifted from ancestral association with conifers to angiosperms and back again several times. Each shift to angiosperms is associated with elevated diversity, whereas the reverse shifts to conifers are associated with lowered diversity. The unusual habit of adult burrowing likely facilitated the diversification of these beetle-fungus associations, enabling them to use the biomass-rich resource that trees represent and set the stage for at least one origin of eusociality.

Haplodiploidy is a highly unusual genetic system that has arisen at least 17 times in animals of varying lifestyles, but most of these haplodiploid lineages remain relatively poorly known. In particular, the ecological and genetic circumstances under which haplodiploidy originates have been difficult to resolve. A recent molecular-phylogenetic study has resolved the phylogenetic position of the haplodiploid clade of scolytine beetles as the sister group of the genus Dryocoetes. Haplodiploid bark beetles are remarkable in that the entire clade of over 1300 species are apparently extreme (sib-mating) inbreeders, most of which cultivate fungi for food while some attack phloem, twigs or seeds. Here we present a much more detailed molecular-phylogenetic study of this clade. Using partial sequences of elongation factor 1-alpha and the mitochondrial small ribosomal subunit (12S), we reconstructed the phylogeny for 48 taxa within the haplodiploid clade, as well as two species of the diplodiploid sister genus Dryocoetes. Results indicate that the genus Ozopemon is the basal lineage of the haplodiploid clade. Since Ozopemon, Dryocoetes and other outgroups are phloem-feeding, this strongly suggest that haplodiploidy and inbreeding evolved in a phloem feeding ancestor. Following the divergence of Ozopemon there is a series of extremely short internodes near the base of the clade, suggesting a very rapid rate of diversification in early Miocene (based on fossil evidence and sequence divergence). Among the many substrates for breeding and food resources utilized within this species-rich clade, the cultivation of yeast-like ambrosia fungi in tunnels deep into the wood predominates (nearly 90% of the species). The number of transitions to feeding on such fungi was few, possibly only one, and is perhaps an irreversible transition. The habit of feeding on fungi cultured in xylem makes it possible for the beetles to use a great variety of plant taxa. This extreme resource generalism, in conjunction with the colonization advantage conferred by haplodiploidy and inbreeding, may have promoted the rapid diversification of this clade. (C) 2000 The Linnean Society or London.

The beetle family Scolytidae includes several groups having regular sib-mating and extremely female-biased sex ratios. Two such groups are known to include haplodiploid species: (i) the tribe Xyleborini and (ii) Coccotrypes and related genera within the tribe Dryocoetini. Relationships of these groups have been controversial. We analysed elongation factor 1-alpha (852 bp) and cytochrome oxidase 1 (1179 bp) sequences for 40 species. The most-parsimonious trees imply a single origin of haplodiploidy uniting Xyleborini (approximately 1200 species) and sib-mating Dryocoetini (approximately 160 species). The sister-group of the haplodiploid clade is the outcrossing genus Dryocoetes. The controversial genus Premnobius is outside the haplodiploid clade. Most haplodiploid scolytids exploit novel resources, ambrosia fungi or seeds, but a few have the ancestral habit of feeding on phloem. Thus, scolytids provide the dearest example of W. D. Hamilton's scenario for the evolution of haplodiploidy (life under bark leading to inbreeding and hence to female-biased sex ratios through haplodiploidy) and now constitute a unique opportunity to study diplo-diploid and haplodiploid sister-lineages in a shared ancestral habitat. There is some evidence of sex determination by maternally inherited endosymbiotic bacteria, which may explain the consistency with which female-biased sex ratios and close inbreeding have been maintained.