Strict Cospeciation Research Articles

Let Emerging Plant Diseases Be Predictable

A prevalent concept for colonization and evolution among plant pathogens and their hosts stems from a post-Darwinian paradigm rooted in the formalized assumption of “specialized parasitism.” Seminal studies on rust fungi of socioeconomic importance integrated such an evolutionary perspective driven by the assumption of strict coevolution among pathogens and their plant hosts. Following this fundamentally unfalsifiable assumption, theories regarding host-switching for parasites were dismissed. If colonization occurred, this process would depend upon the origin of specific and novel mutations that allow infections of previously unexploited hosts or host groups, the acquisition of a broader host range. After a specific mutation arose, parasites and hosts would be locked into an eventual evolutionary dead end (e.g., codified under Dietel’s Law). Accordingly, if the parasites are highly specialized (one parasite, one plant), then new associations are rare or otherwise unpredictable. Similar schools of thought became dominant for animal pathogens and were established during the same period (i.e., Müller’s rule, Fuhrmann’s rule, and Fahrenholz’s rule). Other studies that focused on plant pathogens took the one host–one parasite idea for granted and only tentatively included evolutionary insights in subsequent development of plant pathogen scientific frameworks. Later, emerging from neo-Darwinian views, the paradigm of strict cospeciation was conflated with the gene-for-gene rule postulated in 1956 and which has persisted among phytopathologists even to the present day. In a parallel history, conceptual development among plant pathologists and parasitologists has assumed that colonization is rare and cannot be predicted, given the dependence on the origin of the elusive special mutation. In contrast, current impacts and increasing frequency of emerging pathogens and epidemics across the globe, which influence health and food security, suggest that this historical approach fails in describing a complex biosphere in dynamic change. The Stockholm paradigm (SP) provides a powerful alternative to what may be regarded as the standard model of coevolutionary diversification. The SP creates a theoretical workbench from which emergence of new associations can be evaluated and predicted. The SP provides a new perspective in exploring the dynamics among the phytoplasmas, an emergent group of plant pathogens with substantial risk for food security. New insights are examined, pushing for resolution of the internal conflicts generated by assumptions of the standard coevolutionary model, which has dominated the scientific reasoning for more than a century of plant pathology research.

Molecular phylogeny of Asian Ardisia (Myrsinoideae, Primulaceae) and their leaf-nodulated endosymbionts, Burkholderia s.l. (Burkholderiaceae).

The genus Ardisia (Myrsinoideae, Primulaceae) has 16 subgenera and over 700 accepted names, mainly distributed in tropical Asia and America. The circumscription of Ardisia is not well-defined and sometimes confounded with the separation of some small genera. A taxonomic revision focusing on Ardisia and allies is necessary. In the Ardisia subgenus Crispardisia, symbiotic association with leaf-nodule bacteria is a unique character within the genus. The endosymbionts are vertically transmitted, highly specific and highly dependent on the hosts, suggesting strict cospeciation may have occurred in the evolutionary history. In the present study, we aimed to establish a phylogenetic framework for further taxonomic revision. We also aimed to test the cospeciation hypothesis of the leaf-nodulate Ardisia and their endosymbiotic bacteria. Nuclear ITS and two chloroplast intergenic spaces were used to reconstruct the phylogeny of Asian Ardisia and relatives in Myrsinoideae, Primulaceae. The 16S-23S rRNA were used to reconstruct the bacterial symbionts’ phylogeny. To understand the evolutionary association of the Ardisia and symbionts, topology tests and cophylogenetic analyses were conducted. The molecular phylogeny suggested Ardisia is not monophyletic, unless Sardiria, Hymenandra, Badula and Oncostemum are included. The results suggest the generic limit within Myrsinoideae (Primulaceae) needs to be further revised. The subgenera Crispardisia, Pimelandra, and Stylardisia were supported as monophyly, while the subgenus Bladhia was separated into two distant clades. We proposed to divide the subgenus Bladhia into subgenus Bladhia s.str. and subgenus Odontophylla. Both of the cophylogenetic analyses and topology tests rejected strict cospeciation hypothesis between Ardisia hosts and symbiotic Burkholderia. Cophylogenetic analyses showed general phylogenetic concordance of Ardisia and Burkholderia, and cospeciation events, host-switching events and loss events were all inferred.

Two\u2019s company, three\u2019s a crowd: co-occurring pollinators and parasite species in Breynia oblongifolia (Phyllanthaceae)

BackgroundObligate pollination mutualisms (OPMs) are specialized interactions in which female pollinators transport pollen between the male and female flowers of a single plant species and then lay eggs into those same flowers. The pollinator offspring hatch and feed upon some or all of the developing ovules pollinated by their mothers. Strong trait matching between plants and their pollinators in OPMs is expected to result in reciprocal partner specificity i.e., a single pollinator species using a single plant species and vice versa, and strict co-speciation. These issues have been studied extensively in figs and fig wasps, but little in the more recently discovered co-diversification of Epicephala moths and their Phyllanthaceae hosts. OPMs involving Epicephala moths are believed occur in approximately 500 species of Phyllanthaceae, making it the second largest OPM group after the Ficus radiation (> 750 species). In this study, we used a mixture of DNA barcoding, genital morphology and behavioral observations to determine the number of Epicephala moth species inhabiting the fruits of Breynia oblongifolia, their geographic distribution, pollinating behavior and phylogenetic relationships.ResultsWe found that B. oblongifolia hosts two species of pollinator that co-occurred at all study sites, violating the assumption of reciprocal specificity. Male and female genital morphologies both differed considerably between the two moth species. In particular, females differed in the shape of their ovipositors, eggs and oviposition sites. Phylogenetic analyses indicated that the two Epicephala spp. on B. oblongifolia likely co-exist due to a host switch. In addition, we discovered that Breynia fruits are also often inhabited by a third moth, an undescribed species of Herpystis, which is a non-pollinating seed parasite.ConclusionsOur study reveals new complexity in interactions between Phyllantheae and Epicephala pollinators and highlights that host switching, co-speciation and non-pollinating seed parasites can shape species interactions in OPMs. Our finding that co-occurring Epicephala species have contrasting oviposition modes parallels other studies and suggests that such traits are important in Epicephala species coexistence.

Cophylogenetic assessment of New World warblers (Parulidae) and their symbiotic feather mites (Proctophyllodidae)

Host–symbiont relationships are ubiquitous in nature, yet evolutionary and ecological processes that shape these intricate associations are often poorly understood. All orders of birds engage in symbioses with feather mites, which are ectosymbiotic arthropods that spend their entire life on hosts. Due to their permanent obligatory association with hosts, limited dispersal and primarily vertical transmission, we hypothesized that the cospeciation between feather mites and hosts within one avian family (Parulidae) would be perfect (strict cospeciation). We assessed cophylogenetic patterns and tested for congruence between species in two confamiliar feather mite genera (Proctophyllodidae: Proctophyllodes, Amerodectes) found on 13 species of migratory warblers (and one other closely related migratory species) in the eastern United States. Based on COI sequence data, we found three Proctophyllodes lineages and six Amerodectes lineages. Distance‐ and event‐based cophylogenetic analyses suggested different cophylogenetic trajectories of the two mite genera, and although some associations were significant, there was little overall evidence supporting strict cospeciation. Host switching is likely responsible for incongruent phylogenies. In one case, we documented prairie warblers Setophaga discolor harboring two mite species of the same genus. Most interestingly, we found strong evidence that host ecology may influence the likelihood of host switching occurring. For example, we documented relatively distantly related ground‐nesting hosts (ovenbird Seiurus aurocapilla and Kentucky warbler Geothlypis formosa) sharing a single mite species, while other birds are shrub/canopy or cavity nesters. Overall, our results suggest that cospeciation is not the case for feather mites and parulid hosts at this fine phylogenetic scale, and raise the question if cospeciation applies for other symbiotic systems involving hosts that have complex life histories. We also provide preliminary evidence that incorporating host ecological traits into cophylogenetic analyses may be useful for understanding how symbiotic systems have evolved.

The diversity and evolution of nematodes (Pharyngodonidae) infecting New Zealand lizards.

Host-parasite co-evolutionary studies can shed light on diversity and the processes that shape it. Molecular methods have proven to be an indispensable tool in this task, often uncovering unseen diversity. This study used two nuclear markers (18S rRNA and 28S rRNA) and one mitochondrial (cytochrome oxidase subunit I) marker to investigate the diversity of nematodes of the family Pharyngodonidae parasitizing New Zealand (NZ) lizards (lygosomine skinks and diplodactylid geckos) and to explore their co-evolutionary history. A Bayesian approach was used to infer phylogenetic relationships of the parasitic nematodes. Analyses revealed that nematodes parasitizing skinks, currently classified as Skrjabinodon, are more closely related to Spauligodon than to Skrjabinodon infecting NZ geckos. Genetic analyses also uncovered previously undetected diversity within NZ gecko nematodes and provided evidence for several provisionally cryptic species. We also examined the level of host-parasite phylogenetic congruence using a global-fit approach. Significant congruence was detected between gecko-Skrjabinodon phylogenies, but our results indicated that strict co-speciation is not the main co-evolutionary process shaping the associations between NZ skinks and geckos and their parasitic nematodes. However, further sampling is required to fully resolve co-phylogenetic patterns of diversification in this host-parasite system.

Phylogeographic analyses of bacterial endosymbionts in fig homotomids (Hemiptera: Psylloidea) reveal codiversification of both primary and secondary endosymbionts.

While obligate primary (P-) endosymbionts usually cospeciate with their insect hosts, less is known about codiversification of secondary (S-) endosymbionts that are generally considered facultative. Typically, insects of the superfamily Psylloidea harbour one P- (Carsonella) and at least one S-endosymbiont, thought to compensate for Carsonella genome reduction. Most codiversification studies have used phylogenies of psyllids and their endosymbionts across and within host families or genera, but few have explored patterns within species. We focussed on P- and S-endosymbionts of three Mycopsylla (Homotomidae) species to explore whether they have congruent phylogenies and within-species geographic structures. The P-endosymbiont Carsonella, a S-endosymbiont and Wolbachia all had 100% prevalence, while Arsenophonus was only found in one species at low prevalence. Congruent phylogenies of Mycopsylla and P-endosymbionts across populations and species support strict cospeciation.S-endosymbiont phylogenies were also congruent across host species but low genetic variation in the S-endosymbiont was not correlated with host phylogeography, possibly due to a shorter evolutionary association. Between species, Wolbachia and Mycopsylla phylogenies were incongruent, probably due to horizontal transmission events. Our study is the first to explore endosymbionts of Mycopsylla and further supports the codivergence of Psylloidea hosts and P-endosymbionts, with obligate host interactions for both P- and S-endosymbionts.

Codivergence of the primary bacterial endosymbiont of psyllids versus host switches and replacement of their secondary bacterial endosymbionts.

Coevolution between insects and bacterial endosymbionts contributes to the success of many insect lineages. For the first time, we tested for phylogenetic codivergence across multiple taxonomic scales, from within genera to superfamily between 36 psyllid species of seven recognised families (Hemiptera: Psylloidea), their exclusive primary endosymbiont Carsonella and more diverse secondary endosymbionts (S-endosymbionts). Within Aphalaridae, we found that Carsonella and S-endosymbionts were fixed in one Glycaspis and 12 Cardiaspina populations. The dominant S-endosymbiont was Arsenophonus, while Sodalis was detected in one Cardiaspina species. We demonstrated vertical transmission for Carsonella and Arsenophonus in three Cardiaspina species. We found strong support for strict cospeciation and validated the informative content of Carsonella as extended host genome for inference of psyllid relationships. However, S-endosymbiont and host phylogenies were incongruent, and displayed signs of host switching and endosymbiont replacement. The high incidence of Arsenophonus in psyllids and other plant sap-feeding Hemiptera may be due to repeated host switching within this group. In two psyllid lineages, Arsenophonus and Sodalis genes exhibited accelerated evolutionary rates and AT-biases characteristic of long-term host associations. Together with strict vertical transmission and 100% prevalence within host populations, our results suggest an obligate, and not facultative, symbiosis between psyllids and some S-endosymbionts.

Host responses to cycles of climate change shape parasite diversity across North America's Intermountain West

Host-parasite cospeciation, in which parasite divergence occurs in response to host divergence, is commonly proposed as a driver of parasite diversification, yet few empirical examples of strict cospeciation exist. Host-parasite co-evolutionary histories commonly reflect complex mosaics of cospeciation, dispersal, lineage extinction and other phenomena. The episodic host-switching model of parasite diversification accounts for complexity by suggesting that diversification and faunal assembly is a consequence of fluctuation between environmental disruption and environmental stability. The phylogeographic predictions of the strict cospeciation and episodic host-switching models were tested using the North American pika/parasite assemblage, with a primary focus on the American pika, Ochotona princeps (Richardson, 1828), and a suite of its endoparasitic cestodes and nematodes. This approach integrating phylogeographic and demographic methods with inferences drawn from species distribution modelling revealed that the parasite community of pikas has been shaped by climate-driven range fluctuation of hosts and bouts of geographic and host colonization by parasites associated with transitions between glacial and interglacial phases.

Pattern of association between endemic Hawaiian fruit flies (Diptera, Tephritidae) and their symbiotic bacteria: Evidence of cospeciation events and proposal of “Candidatus Stammerula trupaneae”

Several insect lineages have evolved mutualistic association with symbiotic bacteria. This is the case of some species of mealybugs, whiteflies, weevils, tsetse flies, cockroaches, termites, carpenter ants, aphids and fruit flies. Some species of Tephritinae, the most specialized subfamily of fruit flies (Diptera: Tephritidae), harbour co-evolved vertically transmitted, bacterial symbionts in their midgut, known as “Candidatus Stammerula spp.”. The 25 described endemic species of Hawaiian tephritids, plus at least three undescribed species, are taxonomically distributed among three genera: the cosmopolitan genus Trupanea (21 described spp.), the endemic genus Phaeogramma (2 spp.) and the Nearctic genus Neotephritis (2 spp.). We examined the presence of symbiotic bacteria in the endemic tephritids of the Hawaiian Islands, which represent a spectacular example of adaptive radiation, and tested the concordant evolution between host and symbiont phylogenies.We detected through PCR assays the presence of specific symbiotic bacteria, designated as “Candidatus Stammerula trupaneae”, from 35 individuals of 15 species. The phylogeny of the insect host was reconstructed based on two regions of the mitochondrial DNA (16S rDNA and COI-tRNALeu-COII), while the bacterial 16S rRNA was used for the symbiont analysis. Host and symbiont phylogenies were then compared and evaluated for patterns of cophylogeny and strict cospeciation. Topological congruence between Hawaiian Tephritinae and their symbiotic bacteria phylogenies suggests a limited, but significant degree of host–symbiont cospeciation. We also explored the character reconstruction of three host traits, as island location, host lineage, and host tissue attacked, based on the symbiont phylogenies under the hypothesis of cospeciation.

Cophylogenetic interactions between marine viruses and eukaryotic picophytoplankton

BackgroundNumerous studies have investigated cospeciation (or cophylogeny) in various host-symbiont systems, and different patterns were inferred, from strict cospeciation where symbiont phylogeny mirrors host phylogeny, to complete absence of correspondence between trees. The degree of cospeciation is generally linked to the level of host specificity in the symbiont species and the opportunity they have to switch hosts. In this study, we investigated cophylogeny for the first time in a microalgae-virus association in the open sea, where symbionts are believed to be highly host-specific but have wide opportunities to switch hosts. We studied prasinovirus-Mamiellales associations using 51 different viral strains infecting 22 host strains, selected from the characterisation and experimental testing of the specificities of 313 virus strains on 26 host strains.ResultsAll virus strains were restricted to their host genus, and most were species-specific, but some of them were able to infect different host species within a genus. Phylogenetic trees were reconstructed for viruses and their hosts, and their congruence was assessed based on these trees and the specificity data using different cophylogenetic methods, a topology-based approach, Jane, and a global congruence method, ParaFit. We found significant congruence between virus and host trees, but with a putatively complex evolutionary history.ConclusionsMechanisms other than true cospeciation, such as host-switching, might explain a part of the data. It has been observed in a previous study on the same taxa that the genomic divergence between host pairs is larger than between their viruses. It implies that if cospeciation predominates in this algae-virus system, this would support the hypothesis that prasinoviruses evolve more slowly than their microalgal hosts, whereas host switching would imply that these viruses speciated more recently than the divergence of their host genera.

Cophylogeny of Quill Mites from the GenusSyringophilopsis(Acari: Syringophilidae) and their North American Passerine Hosts

Species of Syringophilopsis quill mites are found in the flight feathers of passerine birds. A phylogeny of species from this genus infecting North American passerines was inferred from the mitochondrial cytochrome oxidase I gene and the nuclear 28S ribosomal RNA gene. Based on the large genetic distance among lineages, the genus appears to be composed of several cryptic species. A reconciliation analysis of these mites and their avian hosts indicates a limited, but significant, degree of cophylogeny. However, strict cospeciation is not found to be occurring in this system.

Données récentes sur les hantavirus et perspectives de recherche

Les virus du genre Hantavirus sont les seuls représentants de la famille des Bunyaviridae qui ne sont pas transmis par des vecteurs arthropodes mais par des petits mammifères. Les hantavirus transmis par les rongeurs (Ordre Rodentia) ont été les premiers découverts en raison de leur pouvoir pathogène chez l’homme. Les premières études phylogénétiques ont suggéré une coévolution entre chaque hantavirus et son espèce de rongeur réservoir. Toutefois, l’exploration d’autres réservoirs animaux a montré que des hantavirus circulent aussi chez les insectivores (Ordre Soricomorpha) et les chauvessouris (Ordre Chiroptera), sans qu’aucune transmission pathogène à l’homme n’ait pu être mise en évidence jusqu’à aujourd’hui. Des observations naturelles de cocirculation d’un même hantavirus au sein de plusieurs espèces de rongeurs sympatriques, ainsi que de nouvelles données phylogénétiques qui soulignent des changements d’hôte (host-switching) entre hantavirus très proches, remettent actuellement en cause la cospéciation stricte. De même, l’observation plus fine de cas cliniques suggère de modérer le dogme d’une maladie distincte chez l’homme, les hantavirus de l’Ancien Monde (Europe-Asie) provoquant une fièvre hémorragique à syndrome rénal (FHRS) et ceux du Nouveau Monde (Amériques) conduisant à une hantavirose à syndrome cardio-pulmonaire (HSCP). Ces points sont discutés car ils ouvrent d’importantes perspectives en matière de recherche transdisciplinaire, depuis l’immunologie comparée chez les mammifères jusqu’à la modélisation de la dynamique des réservoirs dans leur milieu naturel, en passant par la sociologie des populations à risque. Les données récentes sur la circulation et le pouvoir pathogène des hantavirus en Europe et dans le monde sont aussi présentées, ainsi que les nouveaux outils d’investigation sérologique et génétique permettant la découverte sans a priori de nouveaux virus «dormants» dans des réservoirs et l’évaluation de leur potentiel d’émergence chez l’homme.

Evolutionary history of nematodes associated with sweat bees

Organisms that live in close association with other organisms make up a large part of the world’s diversity. One driver of this diversity is the evolution of host-species specificity, which can occur via reproductive isolation following a host-switch or, given the correct circumstances, via cospeciation. In this study, we explored the diversity and evolutionary history of Acrostichus nematodes that are associated with halictid bees in North America. First, we conducted surveys of bees in Virginia, and found six halictid species that host Acrostichus. To test the hypothesis of cospeciation, we constructed phylogenetic hypotheses of Acrostichus based on three genes. We found Acrostichus puri and Acrostichus halicti to be species complexes comprising cryptic, host-specific species. Although several nodes in the host and symbiont phylogenies were congruent and tests for cospeciation were significant, the host’s biogeography, the apparent patchiness of the association across the host’s phylogeny, and the amount of evolution in the nematode sequence suggested a mixture of cospeciation, host switching, and extinction events instead of strict cospeciation. Cospeciation can explain the relationships between Ac. puri and its augochlorine hosts, but colonization of Halictus hosts is more likely than cospeciation. The nematodes are vertically transmitted, but sexual transmission is also likely. Both of these transmission modes may explain host-species specificity and congruent bee and nematode phylogenies. Additionally, all halictid hosts come from eusocial or socially polymorphic lineages, suggesting that sociality may be a factor in the suitability of hosts for Acrostichus.

Flyway homogenisation or differentiation? Insights from the phylogeny of the sandpiper (Charadriiformes: Scolopacidae: Calidrinae) wing louse genus Lunaceps (Phthiraptera: Ischnocera)

The wing louse genus Lunaceps, is the most speciose chewing louse (Phthiraptera) genus inhabiting sandpipers (Charadriiformes: Calidrinae) and is known from almost all sandpiper species. The hosts follow specific flyways from the Arctic breeding grounds to wintering locations in the southern hemisphere, and often form large mixed-species flocks during migration and wintering. We estimated a phylogeny of Lunaceps based on three mitochondrial loci, supporting monophyly of the genus but revealing extensive paraphyly at the species level. We also evaluated the relative importance of flyway differentiation (same host species having different lice along different flyways) and flyway homogenisation (different host species having the same lice along the same flyway). We found that while the lice of smaller sandpipers and stints show some evidence of flyway homogenisation, those of larger sandpipers do not. No investigated host species migrating along more than one flyway showed any evidence of flyway differentiation. The host–parasite associations within Lunaceps are in no case monophyletic, rejecting strict cospeciation.

Strict cospeciation of devescovinid flagellates and Bacteroidales ectosymbionts in the gut of dry‐wood termites (Kalotermitidae)

The surface of many termite gut flagellates is colonized with a dense layer of bacteria, yet little is known about the evolutionary relationships of such ectosymbionts and their hosts. Here we investigated the molecular phylogenies of devescovinid flagellates (Devescovina spp.) and their symbionts from a wide range of dry-wood termites (Kalotermitidae). From species-pure flagellate suspensions isolated with micropipettes, we obtained SSU rRNA gene sequences of symbionts and host. Phylogenetic analysis showed that the Devescovina spp. present in many species of Kalotermitidae form a monophyletic group, which includes also the unique devescovinid flagellate Caduceia versatilis. All members of this group were consistently associated with a distinct lineage of Bacteroidales, whose location on the cell surface was confirmed by fluorescence in situ hybridization. The well-supported congruence of the phylogenies of devescovinids and their ectosymbionts documents a strict cospeciation. In contrast, the endosymbionts of the same flagellates ('Endomicrobia') were clearly polyphyletic and must have been acquired independently by horizontal transfer from other flagellate lineages. Also the Bacteroidales ectosymbionts of Oxymonas flagellates present in several Kalotermitidae belonged to several distantly related lines of descent, underscoring the general perception that the evolutionary history of flagellate-bacteria symbioses in the termite gut is complex.

Speciation in fig wasps

1. There are over 700 species of fig trees in the tropics and several thousand species of fig wasps are associated with their syconia (inflorescences). These wasps comprise a monophyletic family of fig pollinators and several diverse lineages of non‐pollinating wasps. The pollinator larvae gall fig flowers, while larvae of non‐pollinating species either initiate their own galls or parasitise the galls of other wasps.2. A single fig species has one to four pollinator species and also hosts up to 30 non‐pollinating wasp species. Most wasps show a high degree of host‐plant specificity and are known from only a single fig species. However, in some cases wasps may be shared across closely related fig species.3. There is impressive morphological co‐evolution between figs and fig wasps and this, combined with a high degree of partner specificity, led to the expectation that figs and pollinators have cospeciated extensively. Comparison of deep phylogenies supports long‐term codivergence of figs and pollinators, but also suggests that some host shifts have occurred.4. Phylogenies of more closely related species do not match perfectly and may even be incongruent, suggesting significant roles for processes other than strict cospeciation. Combined with recent evidence on host specificity patterns, this suggests that pollinator wasps may often speciate by host shifts between closely related figs, or by duplication (the wasp speciates but the fig doesn't). The frequencies and biological details of these different modes of speciation invite further study.5. Far less is known about speciation in non‐pollinating fig wasps. Some lineages have probably co‐evolved with figs and pollinators for most of the evolutionary history of the symbiosis, while others appear to be more recent colonisers. Many species appear to be highly host‐plant specific, but those that lay eggs through the fig wall without entering the syconium (the majority of species) may be subject to fewer constraints on host shifting than pollinators. There is evidence for substantial host shifting in at least one genus, but also evidence for ecological speciation on the same host plant by niche shifts in other cases.6. Finally, recent work has begun to address the issue of ‘community phylogeny’ and provided evidence for long‐term co‐divergence of multiple pollinating and non‐pollinating wasp lineages with their host figs.

Evolutionary Relationships of “ Candidatus Endobugula” Bacterial Symbionts and Their Bugula Bryozoan Hosts

Ribosomal gene sequences were obtained from bryozoans in the genus Bugula and their bacterial symbionts; analyses of host and symbiont phylogenetic trees did not support a history of strict cospeciation. Symbiont-derived compounds known to defend host larvae from predation were only detected in two out of four symbiotic Bugula species.

Cospeciation between the nasal mite Ptilonyssus sairae (Acari: Rhinonyssidae) and its bird hosts

Abstract The parasitic nasal mite Ptilonyssus sairae Castro has been classified as one species, but we find that significant sequence divergence exists between host-specific mite populations. The low level of gene flow between conspecifics is probably a result of the parasites' low rate of dispersal among different bird host species. The internal transcribed spacer (ITS) of the nuclear ribosomal RNA gene from P. sairae populations was sequenced and used for phylogenetic inference and subsequently compared to their passerine bird host phylogeny. In addition to finding restricted gene flow between host-associated parasites, the reconciliation of host and parasite gene trees reveals statistically significant levels of strict cospeciation between bird hosts and host-specific mite populations. Our findings suggest that cryptic speciation may be common in this group of mites.

Deep mtDNA divergences indicate cryptic species in a fig-pollinating wasp

BackgroundFigs and fig-pollinating wasps are obligate mutualists that have coevolved for ca 90 million years. They have radiated together, but do not show strict cospeciation. In particular, it is now clear that many fig species host two wasp species, so there is more wasp speciation than fig speciation. However, little is known about how fig wasps speciate.ResultsWe studied variation in 71 fig-pollinating wasps from across the large geographic range of Ficus rubiginosa in Australia. All wasps sampled belong to one morphological species (Pleistodontes imperialis), but we found four deep mtDNA clades that differed from each other by 9–17% nucleotides. As these genetic distances exceed those normally found within species and overlap those (10–26%) found between morphologically distinct Pleistodontes species, they strongly suggest cryptic fig wasp species. mtDNA clade diversity declines from all four present in Northern Queensland to just one in Sydney, near the southern range limit. However, at most sites multiple clades coexist and can be found in the same tree or even the same fig fruit and there is no evidence for parallel sub-division of the host fig species. Both mtDNA data and sequences from two nuclear genes support the monophyly of the "P. imperialis complex" relative to other Pleistodontes species, suggesting that fig wasp divergence has occurred without any host plant shift. Wasps in clade 3 were infected by a single strain (W1) of Wolbachia bacteria, while those in other clades carried a double infection (W2+W3) of two other strains.ConclusionOur study indicates that cryptic fig-pollinating wasp species have developed on a single host plant species, without the involvement of host plant shifts, or parallel host plant divergence. Despite extensive evidence for coevolution between figs and fig wasps, wasp speciation may not always be linked strongly with fig speciation.

Early molecular investigations of lichen-forming symbionts: 1986-2001*.

From the mid-1980s the symbionts in lichen associations, heterotrophic fungi and photosynthetic algae or cyanobacteria, were the subject of increasing numbers of molecular investigations. Many of the studies examined the phylogenetic placement of the individual symbiotic partners with their free-living relatives, refining their nomenclature and classification. Resulting phylogenies permitted the mapping of transitions to and from lichenization and stimulated discussion of the relative ease of gaining and losing symbiotic lifestyles. Comparing symbiont phylogenies both rejected strict cospeciation and mirrored phylogenies and hinted at more complex forces of coevolution, including symbiont switching and selection. Studies at the species and population levels examined patterns of species delimitation and geographic dispersion and processes such as gene flow, self-fertilization, and founder effect. Significant genetic variation often was associated with mobile elements, group I and spliceosomal introns. This review examines the influence of molecular investigation on lichenology during this first 15 years.