Robert M. Plowes

Specificity Between Lactobacilli And Hymenopteran Hosts Is The Exception Rather Than The Rule

ABSTRACT Lactobacilli (Lactobacillales: Lactobacillaceae) are well known for their roles in food fermentation, as probiotics, and in human health, but they can also be dominant members of the microbiota of some species of Hymenoptera (ants, bees, and wasps). Honey bees and bumble bees associate with host-specific lactobacilli, and some evidence suggests that these lactobacilli are important for bee health. Social transmission helps maintain associations between these bees and their respective microbiota. To determine whether lactobacilli associated with social hymenopteran hosts are generally host specific, we gathered publicly available Lactobacillus 16S rRNA gene sequences, along with Lactobacillus sequences from 454 pyrosequencing surveys of six other hymenopteran species (three sweat bees and three ants). We determined the comparative secondary structural models of 16S rRNA, which allowed us to accurately align the entire 16S rRNA gene, including fast-evolving regions. BLAST searches and maximum-likelihood phylogenetic reconstructions confirmed that honey and bumble bees have host-specific Lactobacillus associates. Regardless of colony size or within-colony oral sharing of food (trophallaxis), sweat bees and ants associate with lactobacilli that are closely related to those found in vertebrate hosts or in diverse environments. Why honey and bumble bees associate with host-specific lactobacilli while other social Hymenoptera do not remains an open question. Lactobacilli are known to inhibit the growth of other microbes and can be beneficial whether they are coevolved with their host or are recruited by the host from environmental sources through mechanisms of partner choice.

Introducing Phorid Fly Parasitoids of Red Imported Fire Ant Workers from South America to Texas: Outcomes Vary by Region and by Pseudacteon Species Released

Abstract Since the first report on introductions to Texas of Pseudacteon decapitating flies, a variety of participants have released flies in a range of sites. The expansions of Pseudacteon populations have been systematically and widely monitored. Before 2002, the widely released initial species P. tricuspis Borgmeier did not become established. Severe drought in 1996–2001 and host-size-dependent sex ratio were proposed constraints in establishing this species. In recent years, however, these limitations have been lifted in some areas by favorable weather, irrigation of release sites, and/or by use of a smaller Pseudacteon species, P. curvatus Borgmeier, not reliant on larger fire ant workers to produce females. Beginning in 2002, the USDA-APHIS collaboration with USDA-ARS and Texas Cooperative Extension programs began to supplement release sites in Texas beyond those initiated by the University of Texas, Austin phorid fly project. In 2005, private citizens began to participate in the spread of Pseudacteon to new sites. By fall 2006, P. tricuspis, expanding from releases between 1999 and 2001, was found on more than 3 million hectares of Central and Coastal Texas, while P. curvatus, with its later start, is only now beginning to expand at some sites. Pseudacteon that established more easily in mesic and moderate climates has difficulty surviving unfavorable weather in South Texas. However, two sites where flies “failed” to become established were revealed to be false negatives after the record rains of summer 2007. Starting in late 2006, the first releases of P. obtusus Borgmeier in North America established, and three to five additional species are being released.

Indirect competition facilitates widespread displacement of one naturalized parasitoid of imported fire ants by another.

Species abundances in natural systems are usually close to some equilibrium, making mechanisms that maintain or prevent species coexistence difficult to discern. Biological control projects provide an opportunity to observe systems transition between equilibriums as a result of the influence of the newly introduced species. In the southeastern United States and Texas, species of phorid fly parasitoids are being sequentially introduced as control agents for imported fire ants. The first two species introduced, Pseudacteon tricuspis and P. curvatus, partition the host niche based upon body size and co-exist broadly in their native range in Argentina, indicating they would form a co-existing and complementary suite of parasitoids in North America. This study examines the interaction between these parasitoids at multiple temporal and spatial scales. Surprisingly, data at all scales reveal that as P. curvatus establishes at a site it competitively displaces P. tricuspis. However, the speed of this reduction appears to differ between ecoregions, suggesting that the rate of displacement depends on environment. At the site where P. curvatus has been established the longest, this population interaction approaches complete displacement. Tests of potential mechanisms causing this displacement reveal that direct competition for host workers alters the operational sex ratio of the P. tricuspis population, but the strength of this effect is insufficient to explain the displacement. Experiments reveal the operation of a strong, indirect effect whereby locally common species preempt reproductive opportunities from rarer species by inducing host behavioral defenses. Finally, a re-examination of published data from their native range reveals that a previously overlooked negative relationship between the densities of these two species also exists there, suggesting that the same processes as those reported here also operate in South America.

Dynamic expansion in recently introduced populations of fire ant parasitoids (Diptera: Phoridae).

Combating invasive species requires a detailed, mechanistic understanding of the manner and speed with which organisms expand their ranges. Biological control efforts provide an opportunity to study the process of species invasions and range expansions under known initial conditions. This study examines the rate, pattern and mechanisms of spread for two populations of the biological control agent Pseudacteon tricuspis, phorid-fly parasitoids of imported fire ants. We employ a trap-based survey method that detects phorid flies in low-density populations, and provides data on abundance. This technique allows us to differentiate between continuous population spread and effective long-distance dispersal and to examine density gradients of phorid flies across the expanding population front. We find that occupied sites in front of the leading edge of continuous populations were common; forming small populations we refer to as satellite populations. Satellite populations are tens of kilometers from the nearest possible source. Wind governs the dynamics of spread in these two central Texas populations. Population edges expanding with the wind exhibited a higher frequency of effective long-distance dispersal than did populations expanding into the wind. This enhanced effective long-distance dispersal rate translated into a five times faster rate of spread for population edges traveling with the wind. This planned invasion shares many characteristics in common with unplanned species invasions including: protracted establishment phase during which densities were below detection thresholds, and slow initial spread immediately after establishment followed by rapid, accelerating spread rates as population sizes grew.

The urban fire ant paradox: native fire ants persist in an urban refuge while invasive fire ants dominate natural habitats

In contrast to the widespread extirpation of native fire ants (Solenopsis geminata) across southern US following the invasion by imported red fire ants (S. invicta), some residential areas of Austin form unexpected refuges for native fire ants. Ironically, these urban environments provide refuges for the native fire ants while adjacent natural habitats have been overrun by invasive fire ants. Resistance to invasive fire ants in these urban areas occurs mainly in older residential properties constructed prior to the S. invicta invasion, while more recent construction has allowed establishment by S. invicta. The invasive ability of S. invicta is often attributed to escape from parasitoids and efficient dispersal of polygyne multiple queen colonies. Here we also show the importance of landscape parameters in the invasion process, where low levels of disturbance and continuous plant cover in older residential areas form possible barriers to colonization. Dense leaf cover (high NDVI) was also found to be associated with native ant refuges. Long term residential land ownership may have resulted in lower recent disturbance levels and increased plant cover that support refuges of native fire ants.

A Review of Pseudacteon (Diptera: Phoridae) That Parasitize Ants of the Solenopsis geminata Complex (Hymenoptera: Formicidae)

ABSTRACT Some phorid flies of the genus Pseudacteon Coquillett parasitize workers of Solenopsis geminata (F.) complex fire ants. The group is found to comprise at least 21 species of flies, of which 11 are new to science and described here: P. amuletum, P. andinus, P. annulus, P. catarinae, P. deltoides, P. hippeus, P. kungae, P. laticarinatus, P. palomita, P. quinni, and P. robustus. An identification key to the females is included. This expansion of known Pseudacteon species parasitizing the S. geminata complex makes it an interesting system for comparison with Pseudacteon flies that parasitize fire ants of the S. saevissima (Smith) complex.

The introduction of the fire ant parasitoid Pseudacteon nocens in North America: challenges when establishing small populations

Several species of parasitoid phorid flies (Pseudacteon spp., Diptera: Phoridae) have been released into the United States as potential biological control agents for the red imported fire ant, Solenopsisinvicta Buren (Hymenoptera: Formicidae). Here we report the first successful introduction and spread of Pseudacteonnocens Borgmeier at a site in Texas, USA. Pseudacteon nocens is an important natural enemy since it is a widespread and often abundant parasitoid of S. invicta in Argentina, where it attacks larger fire ant workers eliciting a strong defensive response. Several years of effort to establish this species previously failed, and here we provide a model to better understand the likelihood of founding new populations when introducing sequential batches of flies in field or laboratory cultures. We also report on a novel method of establishing new populations of phorids in the field using pupae burial boxes to overcome constraints of releasing adult flies or infected worker ants.

Molecular diversity of the microsporidium Kneallhazia solenopsae reveals an expanded host range among fire ants in North America.

Kneallhazia solenopsae is a pathogenic microsporidium that infects the fire ants Solenopsis invicta and Solenopsis richteri in South America and the USA. In this study, we analyzed the prevalence and molecular diversity of K. solenopsae in fire ants from North and South America. We report the first empirical evidence of K. solenopsae infections in the tropical fire ant, Solenopsis geminata, and S. geminata×Solenopsis xyloni hybrids, revealing an expanded host range for this microsporidium. We also analyzed the molecular diversity at the 16S ribosomal RNA gene in K. solenopsae from the ant hosts S.invicta, S. richteri, S. geminata and S. geminata×S. xyloni hybrids from North America, Argentina and Brazil. We found 22 16S haplotypes. One of these haplotypes (WD_1) appears to be widely distributed, and is found in S. invicta from the USA and S. geminata from southern Mexico. Phylogenetic analyses of 16S sequences revealed that K. solenopsae haplotypes fall into one of two major clades that are differentiated by 2-3%. In some cases, multiple K. solenopsae haplotypes per colony were found, suggesting either an incomplete homogenization among gene copies within the 16S gene cluster or multiple K. solenopsae variants simultaneously infecting host colonies.

Myrmecomorba nylanderiae gen. et sp. nov., a microsporidian parasite of the tawny crazy ant Nylanderia fulva

A new microsporidian genus and species, Myrmecomorba nylanderiae, is described from North American populations of the tawny crazy ant, Nylanderia fulva. This new species was found to be heterosporous producing several types of binucleate spores in both larval and adult stages and an abortive octosporoblastic sporogony in adult ants. While microsporidia are widespread arthropod parasites, this description represents only the fifth species described from an ant host. Molecular analysis indicated that this new taxon is phylogenetically closely allied to the microsporidian family Caudosporidae, a group known to parasitize aquatic black fly larvae. We report the presence of 3 spore types (Type 1 DK, Type 2 DK, and octospores) with infections found in all stages of host development and reproductive castes. This report documents the first pathogen infecting N. fulva, an invasive ant of considerable economic and ecological consequence.

Pseudacteon notocaudatus and Pseudacteon obtusitus (Diptera: Phoridae), two new species of fire ant parasitoids from South America.

Ongoing studies in South America of phorid flies of the genus Pseudacteon Coquillett 1907 have revealed two further new species in this genus that are described here: P. obtusitus and P. notocaudatus. Both species are parasitoids of Solenopsis (F.) fire ants.