The evolution of geadephagan chemical defense: A phylogenetic - - PowerPoint PPT Presentation

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The evolution of geadephagan chemical defense: A phylogenetic - - PowerPoint PPT Presentation

The evolution of geadephagan chemical defense: A phylogenetic understanding of the diversity and stasis of compounds and gland structures in carabid beetles Kipling Will, Aman Gill - University of California, Berkeley Athula Attygalle - Stevens


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The evolution of geadephagan chemical defense:

A phylogenetic understanding of the diversity and stasis of compounds and gland structures in carabid beetles

Kipling Will, Aman Gill - University of California, Berkeley Athula Attygalle - Stevens Institute of Technology, Hoboken Wendy Moore - University of Arizona, Tucson Tanya Renner - San Diego State University, San Diego

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Chemical defense

Figure from Les Insectes, Hachette, Paris, 1883

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Formica sp., Mt. Diablo, CA

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Chemically defended Promecognathus laevissimus drives away Formica sp. ants, Mt. Diablo, CA

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Thomas Eisner

Pherosophus image courtesy of the Institute of Physics

The bombardier beetle. Dramatic carabid beetle chemical defense made famous by Tom Eisner.

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Images courtesy of (1) Thomas Eisner; (2,4) Alex Wild, & (3) Carabidae.Org

Diversity of quinone producers

  • 1. Pherosophus sp
  • 2. Metrius contractus
  • 3. Clivina fossor
  • 4. Chlaenius tricolor
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The NSF funded Carabid Q project team

Aman Gill UC Berkeley Wendy Moore

  • U. of AZ

Tanya Renner San Diego State

Athula Attygalle Stevens Inst. of Technol. Hoboken

The evolution of Geadephagan chemical defense: Molecular evolution and functional validation of genes essential to quinone production in two bombardier beetle species using RNAi. Dr. Aman Gill The Paussus L. (Carabidae: Paussinae) of Madagascar: An explosive adaptive radiation of myrmecophilous beetles. Dr. James Robertson and

  • Dr. Wendy Moore

Molecular phylogeny of the flanged bombardier beetles (Carabidae: Paussinae). Dr. Wendy Moore

Related talks at ICE

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  • 1. Genetic basis of chemical production
  • 2. Biosynthetic pathways leading to final

products

  • 3. Evolutionary transitions of compounds,

pathways, and genes.

  • 4. Environmental and selective correlates

Project objectives: Telling the story of chemical defense of carabid beetles

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Talk Outline

  • Brief background carabid beetle chemical

defense

  • Structure of the gland system
  • Summary of knowledge and sampling for

defensive chemicals in carabids

  • Phylogenetic distribution of selected chemical

classes

  • Implications of a 2-phase system
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Carabidae

Worldwide 37,000 described species. All are thought to be chemically defended

Random images from Google search

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Replete defensive gland reservoirs

Open abdomen, dorsal view of Platynus brunneomarginatus

All Adephagan beetles have a pair of pygidial glands. Gland reservoirs may

  • ccupy 20-30% of the

space in the abdomen.

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The system of cells and ducts in P. brunneomarginatus

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Close up of secretory cells from the formic acid producing P. brunneomarginatus

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  • 545 species from across the family
  • covering 59 of the ~75 tribes
  • Approximately 250+ compounds
  • 18 classes of compounds

Current understanding of carabid defensive gland products

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Carabidae Chemical Classes [species/tribes]

I: Hydrocarbons [105/33] II: Ketones and non-Aromatic Aldehydes [66/14] III: Aliphatic Alcohols [10/9] IV: Esters [34/18] V: Formic Acid [161/27] VI: Higher Saturated Carboxylic Acids [183/34] VII: Unsaturated Carboxylic Acids [168/26] VIII: Phenols [33/3] IX: Aromatic Aldehydes [102/6] X: Aromatic Alcohols [2/2] XI: Aromatic Esters [13/12] XII: Aromatic Acids [2/2] XIII: Quinones [37/7] XIV: Sulfur Compounds [2/1] XV: Terpenes [6/4] XVI: Aromatic Ethers [1/1] XVII: Nitrogen Heterocyclic [2/2] XVIII: Cyanides [1/1]

Classes in black from Moore (1979), in red newly recognized Nomius pygmaeus Psydrus piceus

Image, Joyce Gross

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Unsaturated Carboxylic Acids Hydrocarbons Higher Saturated Carboxylic Acids

Complementary distribution of Hydrocarbons and Carboxylic Acids

present absent Unk/Amb

Adephaga ML tree from 6 gene analysis, D.R. Maddison, et al. unpubl. BToL data

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Formic Acid Quinone

Correlated distribution

  • f Hydrocarbons with

Formic Acid and Quinone

present absent Unk/Amb

Hydrocarbons Adephaga ML tree from 6 gene analysis, D.R. Maddison, et al. unpubl. BToL data

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Agra sp., Carabidae Maquipucuna reserve, Pichincha, Ecuador. Image by Alex Wild

  • A huge diversity of “ground

beetles” are arboreal.

  • Arboreal habitat exposes

them to predation.

  • Most arboreal species

produce formic acid & hydrocarbons.

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One gland reservoir per gland the produce the same compounds Two phases occur together (aqueous/hydrophobic) Formic acid/Hydrocarbons Beetles live in a three dimensional habitat with lots of potential predators

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(Hydrocarbons) (Formic Acid) (Hydrocarbons Formic Acid)

The 2-phases are not miscible, but must be sprayed together

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Anterior Dorsal Image of Platynus angustatus courtesy Igor Sokolov, LSU

(Formic Acid) (Hydrocarbons)

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If the lipid phase floated to the top a climbing beetle would be disadvantaged Anterior Dorsal

(Formic Acid) (Hydrocarbons)

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Tests suggest that Carabids spray a mix regardless

  • f body orientation.
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If the lipid phase floated to the top a climbing beetle would be disadvantaged Anterior Dorsal

(Formic Acid) (Hydrocarbons)

This does not happen!

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Formic Acid Hydrocarbons A dorsal lobe on the reservoir in some taxa and at least one

  • ther mechanism in
  • ther taxa holds the

lipid phase near the efferent duct. We don’t know how the second case works, yet.

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Conclusions

  • 1. Using formic acid with hydrocarbons as a defensive

chemical mix was probably key for carabids to move into the canopy habitat, a factor driving the Harpalinae radiation.

  • 2. In addition to these chemicals, there needs to be an

effective storage and delivery system for a 2-phase

  • admix. Carabids have evolved the dorsal lobe and at

least one other undetermined mechanism to do this.

  • 3. Our telling of the detailed story of chemical defense
  • f carabid beetles is just beginning.
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This project is supported by funding from the National Science Foundation, DEB1556957 DEB1556813, DEB1556931, DEB1556898