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Philip S Ward

Publications and source records attributed to Philip S Ward.

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Evaluating alternative hypotheses for the early evolution and diversification of ants.

Ants are the world's most diverse and ecologically dominant eusocial organisms. Resolving the phylogeny and timescale for major ant lineages is vital to understanding how they achieved this success. Morphological, molecular, and paleontological studies, however, have presented conflicting views on early ant evolution. To address these issues, we generated the largest ant molecular phylogenetic data set published to date, containing approximately 6 kb of DNA sequence from 162 species representing all 20 ant subfamilies and 10 aculeate outgroup families. When these data were analyzed with and without outgroups, which are all distantly related to ants and hence long-branched, we obtained conflicting ingroup topologies for some early ant lineages. This result casts strong doubt on the existence of a poneroid clade as currently defined. We compare alternate attachments of the outgroups to the ingroup tree by using likelihood tests, and find that several alternative rootings cannot be rejected by the data. These alternatives imply fundamentally different scenarios for the early evolution of ant morphology and behavior. Our data strongly support several notable relationships within the more derived formicoid ants, including placement of the enigmatic subfamily Aenictogitoninae as sister to Dorylus army ants. We use the molecular data to estimate divergence times, employing a strategy distinct from previous work by incorporating the extensive fossil record of other aculeate Hymenoptera as well as that of ants. Our age estimates for the most recent common ancestor of extant ants range from approximately 115 to 135 million years ago, indicating that a Jurassic origin is highly unlikely.

Animals↗

Ants.

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Animals↗

The role of opportunity in the unintentional introduction of nonnative ants.

A longstanding goal in the study of biological invasions is to predict why some species are successful invaders, whereas others are not. To understand this process, detailed information is required concerning the pool of species that have the opportunity to become established. Here we develop an extensive database of ant species unintentionally transported to the continental United States and use these data to test how opportunity and species-level ecological attributes affect the probability of establishment. This database includes an amount of information on failed introductions that may be unparalleled for any group of unintentionally introduced insects. We found a high diversity of species (232 species from 394 records), 12% of which have become established in the continental United States. The probability of establishment increased with the number of times a species was transported (propagule pressure) but was also influenced by nesting habit. Ground nesting species were more likely to become established compared with arboreal species. These results highlight the value of developing similar databases for additional groups of organisms transported by humans to obtain quantitative data on the first stages of the invasion process: opportunity and transport.

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Energy gradients and the geographic distribution of local ant diversity.

Geographical diversity gradients, even among local communities, can ultimately arise from geographical differences in speciation and extinction rates. We evaluated three models--energy-speciation, energy-abundance, and area--that predict how geographic trends in net diversification rates generate trends in diversity. We sampled 96 litter ant communities from four provinces: Australia, Madagascar, North America, and South America. The energy-speciation hypothesis best predicted ant species richness by accurately predicting the slope of the temperature diversity curve, and accounting for most of the variation in diversity. The communities showed a strong latitudinal gradient in species richness as well as inter-province differences in diversity. The former vanished in the temperature-diversity residuals, suggesting that the latitudinal gradient arises primarily from higher diversification rates in the tropics. However, inter-province differences in diversity persisted in those residuals--South American communities remained more diverse than those in North America and Australia even after the effects of temperature were removed.

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