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Assassin bugs (Reduvioidea) are one of the most diverse (>7,000 spp.) lineages of predatory animals and have evolved an astounding diversity of raptorial leg modifications for handling prey. The evolution of these modifications is not well understood due to the lack of a robust phylogeny, especially at deeper nodes. We here utilize refined data from transcriptomes (370 loci) to stabilize the backbone phylogeny of Reduvioidea, revealing the position of major clades (e.g., the Chagas disease vectors Triatominae). Analyses combining transcriptomic and Sanger-sequencing datasets result in the first well-resolved phylogeny of Reduvioidea. Despite amounts of missing data, the transcriptomic loci resolve deeper nodes while the targeted ribosomal genes anchor taxa at shallower nodes, both with high support. This phylogeny reveals patterns of raptorial leg evolution across major leg types. Hairy attachment structures (fossula spongiosa), present in the ancestor of Reduvioidea, were lost multiple times within the clade. In contrast to prior hypotheses, this loss is not directly correlated with the evolution of alternative raptorial leg types. Our results suggest that prey type, predatory behavior, salivary toxicity, and morphological adaptations pose intricate and interrelated factors influencing the evolution of this diverse group of predators.
In The Arms Of An Assassin Download
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Next-generation sequencing technology has produced a wealth of genomic data for phylogenomic analyses, which now shed light on parts of the Tree of Life that were previously intractable based on morphology or molecular analyses using few loci30,31,32,33. Integrating genomic and Sanger-sequencing data has great potential to capitalize on existing datasets with broad taxon sampling and to increase branch support, but also poses challenges34. For instance, many phylogenomic studies are restricted to small numbers of taxa31,33,35, and integration with broad, but shallow datasets will then introduce extensive levels of missing data34. Integration of genomic and Sanger-sequencing data matrices in phylogenetic reconstruction has rarely been explored34.
In total, 52 terminals are included, 48 of which are Reduvioidea (47 Reduviidae; 1 Pachynomidae) and 4 outgroups (see Table S1). Among them, transcriptomic data were obtained from 4 outgroups and 19 reduvioids sampled across major lineages of Reduvioidea. Taxon sampling covers all major raptorial leg types within the Reduvioidea, allowing for examination of the evolution of raptorial legs.
Alignments of the amino acid (AA) sequences of the orthologous genes were conducted using MAFFT v7.20541 with the G-INS-i algorithm. Alignment masking was performed on each orthologous gene using ALISCORE v2.042 and ALICUT v2.343 with default settings. The alignments of orthologous genes were manually checked and the genes with apparent paralogs were removed from subsequent analyses. After alignment masking, the orthologous genes with
In addition, the 370-gene AA dataset was analyzed under a coalescent model to accommodate potential gene tree heterogeneity and discordance while also testing the robustness of our dataset under different analytical methods. The best-fitting model of protein evolution for each of the 370 AA genes was optimized using ProtTest v3.250. ML analyses were performed on each AA gene in RAxML using the AA substitution matrix selected by ProtTest and a GAMMA model of rate heterogeneity. A hundred bootstrap replicates were completed for each of the 370 genes, which were imported into STAR51 to estimate the species tree.
Other morphological and behavioral factors have almost certainly influenced the evolution of raptorial legs, ranging from prey specializations to the toxicity of the paralytic saliva. Certain assassin bugs lack raptorial legs altogether. For example, some Harpactorinae, Holoptilinae, and Tribelocephalinae possess slender legs without armature, fossula spongiosa, or sticky substances. However, some of these taxa show behaviors relaxing the constraint of tightly holding prey before the saliva immobilizes the prey item. The most prominent example of this behavior is seen in certain Holoptilinae, which have been reported to sedate ants with fluids released from an abdominal trichome10,15. Forefemoral armature observed in some vertebrate-blood feeding Triatominae (Fig. 3) may be retained plesiomorphic features reflecting the evolution of these bugs from typical predators. Interestingly, numerous species of Triatominae still possess a fossula spongiosa (Fig. 3). Lent and Wygodzinsky3 reported that the fossula is sexually dimorphic in Triatominae (i.e., larger in males than females), indicating that the function of this structure may have shifted from a role in prey capture to one associated with mating.
Salivary toxicity may be another factor influencing the evolution of raptorial leg features. Saliva with higher toxicity may paralyze and kill prey faster and could remove constraints on raptorial legs. Studies have shown that assassin bugs without fossula spongiosa immobilize their prey faster than those with fossula spongiosa12, supporting this hypothesis. We conclude that understanding morphological adaptations to prey capture in assassin bugs will benefit from additional research. Studies should focus on carefully teasing apart the complex relationships between prey specialization, predatory behavior, and function of salivary compounds.
Accession numbers: rDNA sequences from the transcriptomic data have been deposited in GenBank with the accession numbers KT231814-KT231896. The matrices, corresponding partition files and gene trees are available from the Dryad Digital Repository:
J.Z., E.R.L.G., M.F., W.H., C.W. and R.M. designed the research. J.Z., E.R.L.G., M.F. and W.H. conducted extractions. J.Z., E.R.L.G., M.F. and C.W. analyzed the data. K.W. and K.J. performed sequencing. W.H., D.R.S. and C.W. contributed specimens. K.J., R.M. and C.W. contributed funds. All authors contributed in writing of the manuscript.
It is a female, and she is a contract assassin who belongs to an old secret clan that only teaches the deadly arts to its few members. She is their most adorable clan member and can bypass any suspicion using the power of her cuteness. But make no mistake, she is also their most skilled assassin.
For this creature, I imagined a being from another dimension, call it a demon if you will, had entered a human. It finds its own shape within the human body, but the more humans it consumes, the larger and more twisted it grows.
I am always aware of the physical workings of my creatures, as I want them to exist in a real world, however alien or horrific they may look. This means, constantly questioning the design in terms of practical appearance.
Why did the creature develop like that? What kind of environment does it live in and what role does this creature play? Predator or prey? For this particular beast I imagined that an alien tribe learned that if they were the first thing the beast saw when it hatched, it would develop a lifelong loyalty towards that creature. So each beast rider had to catch their own egg first and raise the beast to be able to ride and hunt with it. This is a combination of traits we find in real animals.
Now is probably the most exciting time for someone like me, whose profession it is to design for fantasy and science fiction. We are watching film-quality TV shows like Game of Thrones and complex games have never been as high in quality and as complex in narrative. Easy grounds to find inspiration.
Assassination is a term thought to be derived from "Hashish", a drug similar to marijuana, said to have been used by Hasan-Dan-Sabah to induce motivation in his followers, who were assigned to carry out political and other murders, usually at the cost of their lives.
It is here used to describe the planned killing of a person who is not under the legal jurisdiction of the killer, who is not physically in the hands of the killer, who has been selected by a resistance organization for death, and who has been sele cted by a resistance organization for death, and whose death provides positive advantages to that organization.
Assassination is an extreme measure not normally used in clandestine operations. It should be assumed that it will never be ordered or authorized by any U.S. Headquarters, though the latter may in rare instances agree to its execution by membe rs of an associated foreign service. This reticence is partly due to the necessity for committing communications to paper. No assassination instructions should ever be written or recorded. Consequently, the decision to employ this technique must nearly always be reached in the field, at the area where the act will take place. Decision and instructions should be confined to an absolute minimum of persons. Ideally, only one person will be involved. No report may be made, but usually the act will be pr operly covered by normal news services, whose output is available to all concerned.
Murder is not morally justifiable. Self-defense may be argued if the victim has knowledge which may destroy the resistance organization if divulged. Assassination of persons responsible for atrocities or reprisals may be regarded as just puni shment. Killing a political leader whose burgeoning career is a clear and present danger to the cause of freedom may be held necessary.
The techniques employed will vary according to whether the subject is unaware of his danger, aware but unguarded, or guarded. They will also be affected by whether or not the assassin is to be killed with the subject hereafter, assassinations in which the subject is unaware will be termed "simple"; those where the subject is aware but unguarded will be termed "chase"; those where the victim is guarded will be termed "guarded." 152ee80cbc
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