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opencc-zeroDec 2019View details →

Wolbachia infection in wild mosquitoes (Diptera: Culicidae): Implications for transmission modes and host-endosymbiont associations in Singapore

Background: Wolbachia are intracellular bacterial endosymbionts found in most insect lineages. In mosquitoes, the influence of these endosymbionts on host reproduction and arboviral transmission has spurred numerous studies aimed at using Wolbachia infection as a vector control technique. However, there are several knowledge gaps in the literature and little is known about natural Wolbachia infection across species, their transmission modes, or associations between various Wolbachia lineages and their hosts. This study aims to address these gaps by exploring mosquito-Wolbachia associations and their evolutionary implications. Methods: We conducted tissue-specific polymerase chain reaction screening for Wolbachia infection in the leg, gut and reproductive tissues of wild mosquitoes from Singapore using the Wolbachia surface protein gene (wsp) molecular marker. Mosquito-Wolbachia associations were explored using three methods— tanglegram, distance-based, and event-based methods—and by inferred instances of vertical transmission and host shifts. Results: Adult mosquitoes (271 specimens) representing 14 genera and 40 species were screened for Wolbachia. Overall, 21 species (51.2%) were found positive for Wolbachia, including five in the genus Aedes and five in the genus Culex. To our knowledge, Wolbachia infections have not been previously reported in seven of these 21 species: Aedes nr. fumidus, Aedes annandalei, Uranotaenia obscura, Uranotaenia trilineata, Verrallina butleri, Verrallina sp. and Zeugnomyia gracilis. Wolbachia were predominantly detected in the reproductive tissues, which is an indication of vertical transmission. However, Wolbachia infection rates varied widely within a mosquito host species. There was no clear signal of cophylogeny between the mosquito hosts and the 12 putative Wolbachia strains observed in this study. Host shift events were also observed. Conclusions: Our results suggest that the mosquito-Wolbachia relationship is complex and that combinations of transmission modes and multiple evolutionary events likely explain the observed distribution of Wolbachia diversity across mosquito hosts. These findings have implications for a better understanding of the diversity and ecology of Wolbachia and for their utility as biocontrol agents.

opencc-zeroDec 2014View details →

Data from: Macroevolutionary persistence of heritable endosymbionts: acquisition, retention, and expression of adaptive phenotypes in Spiroplasma

The phylogenetic incongruence between insects and their facultative maternally transmitted endosymbionts indicates that these infections are generally short-lived evolutionarily. Therefore, long-term persistence of many endosymbionts must depend on their ability to colonize and spread within new host species. At least 17 species of Drosophila are infected with endosymbiotic Spiroplasma that have various phenotypic effects. We transinfected five strains of Spiroplasma from three divergent clades into Drosophila neotestacea to test their capacity to spread in a novel host. A strain that causes male killing in Drosophila melanogaster (its native host) also does so in D. neotestacea, even though these host species diverged 40–60 mya. A strain native to D. neotestacea (designated sNeo) and the two other strains of the poulsonii clade of Spiroplasma confer resistance to wasp parasitism, suggesting that this trait may be ancestral within this clade of Spiroplasma. Conversely, no strain other than sNeo conferred resistance to the sterilizing effects of nematode parasitism, suggesting that nematode resistance is a recently derived condition. The apparent addition of nematode resistance to a Spiroplasma lineage that already confers resistance to wasp parasitism suggests endosymbionts can increase the repertoire of traits conducive to their spread. The capacity of an endosymbiont to undergo maternal transmission and express adaptive phenotypes in novel hosts, without requiring a period of host–symbiont co-evolution, enables the spread of such symbionts immediately after the colonization of a new host. This could be critical for the macroevolutionary persistence of facultative endosymbionts whose sojourn times within individual host species are relatively brief.

opencc-zeroJun 2021View details →

Data from: Horizontal Transmission of the Heritable Protective Endosymbiont Hamiltonella defensa Depends on Titre and Haplotype

Secondary endosymbionts of aphids have an important ecological and evolutionary impact on their host, as they provide resistance to natural enemies but also reduce the host's lifespan and reproduction. While secondary symbionts of aphids are faithfully transmitted from mother to offspring, they also have some capacity to be transmitted horizontally between aphids. Here we explore whether 11 isolates from 3 haplotypes of the secondary endosymbiont Hamiltonella defensa differ in their capacity for horizontal transmission. These isolates vary in the protection they provide against parasitoid wasps as well as the costs they inflict on their host, Aphis fabae. We simulated natural horizontal transmission through parasitoid wasps by stabbing aphids with a thin needle and assessed horizontal transmission success of the isolates from one shared donor clone into three different recipient clones. Specifically, we asked whether potentially costly isolates reaching high cell densities in aphid hosts are more readily transmitted through this route. This hypothesis was only partially supported. While transmissibility increased with titre for isolates from two haplotypes, isolates of the H. defensa haplotype 1 were transmitted with greater frequency than isolates of other haplotypes with comparable titres. Thus, it is not sufficient to be merely frequent—endosymbionts might have to evolve specific adaptations to transmit effectively between hosts.

opencc-zeroDec 2016View details →

Data from: Establishment and maintenance of aphid endosymbionts after horizontal transfer is dependent on host genotype

Animal-associated microbial communities have important effects on host phenotypes. Individuals within and among species differ in the strains and species of microbes that they harbour, but how natural selection shapes the distribution and abundance of symbionts in natural populations is not well understood. Symbionts can be beneficial in certain environments but also impose costs on their hosts. Consequently, individuals that can or cannot associate with symbionts will be favoured under different ecological circumstances. As a result, we predict that individuals within a species vary in terms of how well they accept and maintain symbionts. In pea aphids, the frequency of endosymbionts varies among host-plant-associated populations ('biotypes'). We show that aphid genotypes from different biotypes vary in how well they accept and maintain symbionts after horizontal transfer. We find that aphids from biotypes that frequently harbour symbionts are better able to associate with novel symbionts than those from biotypes that less frequently harbour symbionts. Intraspecific variation in the ability of hosts to interact with symbionts is an understudied factor explaining patterns of host–symbiont association.

opencc-zeroDec 2014View details →

Data from: Facultative endosymbionts mediate dietary breadth in a polyphagous herbivore

1. Intraspecific variation in dietary breadth can influence important ecological and evolutionary processes, yet the mechanisms generating this variation are usually unknown. Maternally-transmitted bacterial symbionts frequently infect insect herbivores, and many have been shown to mediate key ecological interactions. For polyphagous herbivores, infection with particular symbionts is often strongly correlated with feeding on particular plant species, suggesting that facultative symbionts might directly determine herbivore food plant specificity. However, previous tests of this hypothesis have returned inconsistent results, providing little empirical support for a causal relationship between facultative symbiont infection and dietary breadth. 2. Here, we investigate whether heritable facultative symbionts mediate dietary breadth in the polyphagous aphid, Aphis craccivora. We first determined that asexual clones of the aphid differ dramatically in performance across two leguminous food plants, locust and alfalfa, and could be considered biotypes with distinct ecological characteristics. The heritable symbiont Arsenophonus is strongly associated with locust-origin aphids. 3. We created experimental lines that share aphid genotypes but differed with respect to Arsenophonus infection status, and compared performance across three food plant species. Naturally Arsenophonus-infected clones performed 2-4× better on locust and up to 75% worse on two alternate plant species than uninfected controls, clearly demonstrating that Arsenophonus promotes specialization on locust. In both laboratory and field experiments, uninfected locust- and alfalfa-origin clones exhibited similar and modest performance on locust, indicating that the "locust-associated biotype" wouldn't exist without Arsenophonus. 4. We also hypothesized that moving Arsenophonus, via transinfection, to an alfalfa-origin lineage would improve performance on locust and serve to expand dietary breadth. Indeed, transinfection doubled aphid performance on locust, and halved aphid performance on alfalfa. However, because this aphid lineage naturally performs better on alfalfa, the transinfected symbiont functionally equalized aphid performance between locust and alfalfa, making the alfalfa biotype more generalized. Thus, the same symbiont can either reduce or expand dietary breadth, depending on host genotype. 5. Our results unequivocally demonstrate that symbiont gain or loss can instantaneously and substantially change the topology of food plant use in a polyphagous insect, modifying diet in ways that potentially influence the insect's ecological niche, evolutionary trajectory and pest status.

opencc-zeroDec 2013View details →

Data from: Ecologically differentiated, stress tolerant endosymbionts in the dinoflagellate genus Symbiodinium (Dinophyceae) Clade D are different species.

We used an integrative genetics approach using sequences of (1) nuclear ribosomal rDNA (internal transcribed spacers and partial large subunit rDNA), (2) single-copy microsatellite nuclear DNA, (3) chloroplast-encoded 23S rDNA, (4) mitochondrial cytochrome b, and (5) repeat variation at eight microsatellite markers, to test the hypothesis that the stress-tolerant, 'morphologically cryptic' Clade D Symbiodinium (Dinophyceae) was composed of more than one species. Concordant phylogenetic and population genetic evidence clearly differentiate separately evolving, reproductively isolated lineages. We describe Symbiodinium boreum sp. nov. and S. eurythalpos sp. nov., two symbionts known to occur in colonies of the zebra coral, Oulastrea crispata (Scleractinia), which lives in turbid, marginal habitats extending from equatorial Southeast Asia to the main islands of Japan in the temperate northwest Pacific Ocean. Symbiodinium boreum was associated with O. crispata in temperate latitudes and S. eurythalpos was common to colonies in the tropics. The geographical ranges of both symbiont species overlapped in the subtropics where they sometimes co-occurred in the same host colony. Symbiodinium trenchii sp. nov. is also described. As a host-generalist symbiont, it often occurs in symbiosis with various species of Scleractinia possessing open (horizontal) modes of symbiont acquisition and is common to reef coral communities thriving in warm turbid reef habitats in the western Pacific Ocean, Indian Ocean, Arabian/Persian Gulf, Red Sea and western Atlantic (Caribbean). As is typical for dinoflagellates, S. boreum and S. eurythalpos were haploid, but microsatellite loci from field-collected and cultured S. trenchii often possessed two alleles, implying that a genome-wide duplication occurred during the evolution of this species. The recognition that Clade D Symbiodinium contains species exhibiting marked differences in host specificity and geographical distribution will yield greater scientific clarity about how stress-tolerant symbionts function in the ecological response of coral–dinoflagellate symbioses to global climate change.

opencc-zeroDec 2013View details →

Data from: Origin, acquisition and diversification of heritable bacterial endosymbionts in louse flies and bat flies

The γ-proteobacterium Arsenophonus and its close relatives (Arsenophonus and like organisms, ALOs) are emerging as a novel clade of endosymbionts, which are exceptionally widespread in insects. The biology of ALOs is, however, in most cases entirely unknown, and it is unclear how these endosymbionts spread across insect populations. Here, we investigate this aspect through the examination of the presence, the diversity and the evolutionary history of ALOs in 25 related species of blood-feeding flies: tsetse flies (Glossinidae), louse flies (Hippoboscidae) and bat flies (Nycteribiidae and Streblidae). While these endosymbionts were not found in tsetse flies, we identify louse flies and bat flies as harbouring the highest diversity of ALO strains reported to date, including a novel ALO clade, as well as Arsenophonus and the recently described Candidatus Aschnera chinzeii clade. We further show that the origin of ALO endosymbiosies extends deep into the evolutionary past of louse flies and bat flies, and that it likely played a major role in the ecological specialization of their hosts. The evolutionary history of ALOs is notably complex and was shaped by both vertical transmission and horizontal transfers with frequent host turnover and apparent symbiont replacement in host lineages. In particular, ALOs have evolved repeatedly and independently close relationships with diverse groups of louse flies and bat flies, as well as phylogenetically more distant insect families, suggesting that ALO endosymbioses are exceptionally dynamic systems.

opencc-zeroDec 2018View details →

Data from: Estimating costs of aphid resistance to parasitoids conferred by a protective strain of the bacterial endosymbiont Regiella insecticola

Heritable bacterial endosymbionts are common in aphids (Hemiptera: Aphididae), and they can influence ecologically important traits of their hosts. It is generally assumed that their persistence in a population is dependent on a balance between the costs and benefits they confer. A good example is Hamiltonella defensa Moran et al., a facultative symbiont that provides a benefit by strongly increasing aphid resistance to parasitoid wasps, but becomes costly to the host in the absence of parasitoids. Regiella insecticola Moran et al. is another common symbiont of aphids and generally does not influence resistance to parasitoids. In the green peach aphid, Myzus persicae (Sulzer), however, one strain (R5.15) was discovered that behaves like H. defensa in that it provides strong protection against parasitoid wasps. Here we compare R5.15-infected and uninfected lines of three M. persicae clones to test whether this protective symbiont is costly as well, i.e., whether it has any negative effects on aphid life-history traits. Furthermore, we transferred R5.15 to two other aphid species, the pea aphid, Acyrthosiphon pisum (Harris), and the black bean aphid, Aphis fabae Scopoli, where this strain is also protective against parasitoids and where we could compare its effects with those of additional, non-protective strains of R. insecticola. Negative effects of R5.15 on host survival and lifetime reproduction were limited and frequently non-significant, and these effects were comparable or in one case weaker than those of R. insecticola strains that are not protective against parasitoid wasps. Unless the benefit of protection is counteracted by detrimental effects on traits that were not considered in this study, R. insecticola strain R5.15 should have a high potential to spread in aphid populations.

opencc-zeroDec 2017View details →

Data from: Camponotus floridanus ants incur a trade-off between phenotypic development and pathogen susceptibility from their mutualistic Endosymbiont Blochmannia

Various insects engage in microbial mutualisms in which the reciprocal benefits exceed the costs. Ants of the genus Camponotus benefit from nutrient supplementation by their mutualistic endosymbiotic bacteria, Blochmannia, but suffer a cost in tolerating and regulating the symbiont. This cost suggests that the ants face secondary consequences such as susceptibility to pathogenic infection and transmission. In order to elucidate the symbiont's effects on development and disease defence, Blochmannia floridanus was reduced in colonies of Camponotus floridanus using antibiotics. Colonies with reduced symbiont levels exhibited workers of smaller body size, smaller colony size, and a lower major-to-minor worker caste ratio, indicating the symbiont's crucial role in development. Moreover, these ants had decreased cuticular melanisation, yet higher resistance to the entomopathogen Metarhizium brunneum, suggesting that the symbiont reduces the ants' ability to fight infection, despite the availability of melanin to aid in mounting an immune response. While the benefits of improved growth and development likely drive the mutualism, the symbiont imposes a critical trade-off. The ants' increased susceptibility to infection exacerbates the danger of pathogen transmission, a significant risk given ants' social lifestyle. Thus, the results warrant research into potential adaptations of the ants and pathogens that remedy and exploit the described disease vulnerability.

opencc-zeroDec 2013View details →

Data from: New insights into the dynamics between reef corals and their associated dinoflagellate endosymbionts from population genetic studies.

The mutualistic symbioses between reef-building corals and micro-algae form the basis of coral reef ecosystems, yet recent environmental changes threaten their survival. Diversity in host-symbiont pairings on the sub-species level could be an unrecognized source of functional variation in response to stress. The Caribbean elkhorn coral, Acropora palmata, associates predominantly with one symbiont species (Symbiodinium 'fitti'), facilitating investigations of individual-level (genotype) interactions. Individual genotypes of both host and symbiont were resolved across the entire range of the species. Most colonies of a particular animal genotype were dominated by one symbiont genotype (or strain) that may persist in the host for decades or more. While Symbiodinium are primarily clonal, the occurrence of recombinant genotypes indicates sexual recombination is the source of this genetic variation, and some evidence suggests this happens within the host. When these data are examined at spatial scales spanning the entire distribution of A. palmata, gene flow among animal populations was an order of magnitude greater than among populations of the symbiont. This suggests that independent micro-evolutionary processes created dissimilar population genetic structures between host and symbiont. The lower effective dispersal exhibited by the dinoflagellate raises questions regarding the extent to which populations of host and symbiont can co-evolve during times of rapid and substantial climate change. However, these findings also support a growing body of evidence suggesting that genotype by genotype interactions may provide significant physiological variation; influencing the adaptive potential of symbiotic reef corals to severe selection.

Supplementary material 12 from: Paez-Triana L, Herrera G, Vega L, Garcia-Corredor D, Pulido Medellín MO, Paniz-Mondolfi A, Muñoz M, Ramírez JD (2023) Metagenomic exploration of endosymbionts and pathogens in the tropical lineage of Rhipicephalus sanguineus sensu lato (s.l.) ticks in Colombia. Metabarcoding and Metagenomics 7: e109085. https://doi.org/10.3897/mbmg.7.109085

p-values obtained between departments and sexes within each department using the Wilcoxon test on the relative abundances of pathogens and endosymbionts

Supplementary material 3 from: Paez-Triana L, Herrera G, Vega L, Garcia-Corredor D, Pulido Medellín MO, Paniz-Mondolfi A, Muñoz M, Ramírez JD (2023) Metagenomic exploration of endosymbionts and pathogens in the tropical lineage of Rhipicephalus sanguineus sensu lato (s.l.) ticks in Colombia. Metabarcoding and Metagenomics 7: e109085. https://doi.org/10.3897/mbmg.7.109085

Significant differences in the relative abundances of assigned reads in Bacteria and Archaea between departments and sexes of the various genera found in the samples

Supplementary material 7 from: Paez-Triana L, Herrera G, Vega L, Garcia-Corredor D, Pulido Medellín MO, Paniz-Mondolfi A, Muñoz M, Ramírez JD (2023) Metagenomic exploration of endosymbionts and pathogens in the tropical lineage of Rhipicephalus sanguineus sensu lato (s.l.) ticks in Colombia. Metabarcoding and Metagenomics 7: e109085. https://doi.org/10.3897/mbmg.7.109085

Metadata and information regarding the genomes included in the analysis for endosymbiont and pathogen identification

Supplementary material 5 from: Paez-Triana L, Herrera G, Vega L, Garcia-Corredor D, Pulido Medellín MO, Paniz-Mondolfi A, Muñoz M, Ramírez JD (2023) Metagenomic exploration of endosymbionts and pathogens in the tropical lineage of Rhipicephalus sanguineus sensu lato (s.l.) ticks in Colombia. Metabarcoding and Metagenomics 7: e109085. https://doi.org/10.3897/mbmg.7.109085

Results of dereplication, as well as phylogenomics analysis and pangenome exploration of all samples (including longer MAGs)

Supplementary material 10 from: Paez-Triana L, Herrera G, Vega L, Garcia-Corredor D, Pulido Medellín MO, Paniz-Mondolfi A, Muñoz M, Ramírez JD (2023) Metagenomic exploration of endosymbionts and pathogens in the tropical lineage of Rhipicephalus sanguineus sensu lato (s.l.) ticks in Colombia. Metabarcoding and Metagenomics 7: e109085. https://doi.org/10.3897/mbmg.7.109085

p-values obtained between departments and sexes within each department using the Wilcoxon test on the relative abundances of each genus (Bacteria and Archaea)

opencc-zeroDec 2021View details →

Microbiome diversity and reproductive incompatibility induced by the prevalent endosymbiont Arsenophonus in two species of African cassava Bemisia tabaci whiteflies

This dataset contains data from two-part experiments described in the paper: "El Hamss, H., Ghosh, S., M. N., M., Delatte, H., & Colvin, J. (2021). Microbiome diversity and reproductive incompatibility induced by the prevalent endosymbiont Arsenophonus in two species of African cassava Bemisia tabaci whiteflies. Ecology and Evolution, 00, 1–10. https://doi.org/10.1002/ece3.840". The experiment investigates the effects of Arsenophonus on whitefly reproduction and microbiome diversity. In the first experiment ("crossing experiment"), the effect of Arsenophonus is studied on number of eggs, nymphs, males, and females on different whitefly species at intraspecies level (SSA1-SG3A+ with SSA1-SG3A-) and at interspecies level (SSA2A+ with SSA1-SG3A- and SSA1-SG3A+). Arsenophonus infection A+ means the presence of Arsenophonus in related whitefly colony, A- means the absence of Arsenophonus. In the second experiment ("microbiome diversity"), whitefly colonies of crossed parents and generated progenies were sequenced to check Arsenophonus threshold (number of reads) and the presence of other bacteria in those crossed whiteflies. The main results of this work are: (i) Arsenophonus did not induce reproductive incompatibility within SSA1-SG3 but reduced the number of eggs, nymphs and female ratios, (ii) complete RI was observed between SSA1-SG3 and SSA2 indicating the lack of gene flow between the two whitefly species, (iii) many new 'other bacteria' in SSA B. tabaci have been identified, whose role remains to be investigated.

opennotspecifiedNov 2017View details →

Fig. 2 in Absence of co-phylogeny indicates repeated diatom capture in dinophytes hosting a tertiary endosymbiont

Fig. 2 Polyphyletic endosymbionts of Durinskia, Galeidinium and Kryptoperidinium with closest relatives among free-ranging diatoms. Maximum likelihood (ML) tree (−ln = 14,945) of all 77 Bacillariaceae operational taxonomic units available, derived from the comparison of concatenated rRNA and plastid sequences. Highlighted endosymbiont taxa were scattered along up to ten only distantly related lineages over the Bacillariaceae phylogeny. Note that accessions of neither Durinskia nor Kryptoperidinium constituted monophyletic groups. Within such lineages, endosymbionts were usually closer related to free-ranging diatoms than to other endosymbionts (though statistical support was not always high). Some endosymbionts were phylogenetically isolated from free-living diatoms exhibiting long branches (e.g. D. kwazulunatalensis N.Yamada, Sym & T.Horig., D. oculata and G. rugatum Tam. & T.Horig.). Branch lengths are drawn to scale, with the scale bar indicating the number of nt substitutions per site. The numbers on the branches are statistical support values (ML bootstrap values, values <50 are not shown, posterior probabilities below branches, values <90 are not shown; asterisks indicate maximal support)

opennotspecifiedNov 2017View details →

Fig. 1 in Absence of co-phylogeny indicates repeated diatom capture in dinophytes hosting a tertiary endosymbiont

Fig. 1 Polyphyletic endosymbionts of freshwater Unruhdinium with closest relatives among free-ranging diatoms. Maximum likelihood (ML) tree (−ln = 21,700) of all 37 Stephanodiscaceae operational taxonomic units available, derived from the comparison of concatenated rRNA and plastid sequences. Endosymbiont taxa are highlighted. Note that we did not observe any contradictory tree topologies between separated analyses of nuclear and plastid loci (not shown). Endosymbionts of Unruhdinium segregated into two distinct lineages. Four identical sequences of different U. penardii (Lemmerm.) Gottschling strains clustered together (94LBS, .98BPP) and built a well-supported (99LBS, 1.00BPP) sister group relationship with free-living Discostella nipponica (Skvortzov) A.Tuji & D.M.Williams (strain TNS:AL-5776). A second lineage (100LBS, 1.00BPP) did not include endosymbiont sequences of U. cf. kevei (G.X.Liu & Z.Y.Hu) Gottschling and U. jiulongense (H. Gu) Gottschling only, but also those of the free-living strains L435 and LO4-2. All sequences of this lineage were nearly identical. Branch lengths are drawn to scale, with the scale bar indicating the number of nt substitutions per site. The numbers on the branches are statistical support values (ML bootstrap values, values <50 are not shown, posterior probabilities below branches, values <90 are not shown; asterisks indicate maximal support)

opencc-zeroFeb 2020View details →

Data from: Dinoflagellates with relic endosymbiont nuclei as models for elucidating organellogenesis

Nucleomorphs are relic endosymbiont nuclei so far found only in two algal groups, cryptophytes and chlorarachniophytes, which have been studied to model the evolutionary process of integrating an endosymbiont alga into a host-governed plastid (organellogenesis). However, past studies suggested that DNA transfer from the endosymbiont to host nuclei had already ceased in both cryptophytes and chlorarachniophytes, implying that the organellogenesis at the genetic level has been completed in the two systems. Moreover, we have yet to pinpoint the closest free-living relative of the endosymbiotic alga engulfed by the ancestral chlorarachniophyte or cryptophyte, making it difficult to infer how organellogenesis altered the endosymbiont genome. To counter the above issues, we need novel nucleomorph-bearing algae, in which endosymbiont-to-host DNA transfer is on-going and for which endosymbiont/plastid origins can be inferred at a fine taxonomic scale. Here, we report two previously undescribed dinoflagellates, strains MGD and TGD, with green algal endosymbionts enclosing plastids as well as relic nuclei (nucleomorphs). We provide evidence for the presence of DNA in the two nucleomorphs and the transfer of endosymbiont genes to the host (dinoflagellate) genomes. Furthermore, DNA transfer between the host and endosymbiont nuclei was found to be in progress in both the MGD and TGD systems. Phylogenetic analyses successfully resolved the origins of the endosymbionts at the genus level. With the combined evidence, we conclude that the host-endosymbiont integration in MGD/TGD is less advanced than that in cryptophytes/chrorarachniophytes, and propose the two dinoflagellates as models for elucidating organellogenesis.