The evolution of geadephagan chemical defense: A phylogenetic - - PowerPoint PPT Presentation
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
Chemical defense
Figure from Les Insectes, Hachette, Paris, 1883
Formica sp., Mt. Diablo, CA
Chemically defended Promecognathus laevissimus drives away Formica sp. ants, Mt. Diablo, CA
Thomas Eisner
Pherosophus image courtesy of the Institute of Physics
The bombardier beetle. Dramatic carabid beetle chemical defense made famous by Tom Eisner.
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
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
- 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
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
Carabidae
Worldwide 37,000 described species. All are thought to be chemically defended
Random images from Google search
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.
The system of cells and ducts in P. brunneomarginatus
Close up of secretory cells from the formic acid producing P. brunneomarginatus
- 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
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
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
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
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.
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
(Hydrocarbons) (Formic Acid) (Hydrocarbons Formic Acid)
The 2-phases are not miscible, but must be sprayed together
Anterior Dorsal Image of Platynus angustatus courtesy Igor Sokolov, LSU
(Formic Acid) (Hydrocarbons)
If the lipid phase floated to the top a climbing beetle would be disadvantaged Anterior Dorsal
(Formic Acid) (Hydrocarbons)
Tests suggest that Carabids spray a mix regardless
- f body orientation.
If the lipid phase floated to the top a climbing beetle would be disadvantaged Anterior Dorsal
(Formic Acid) (Hydrocarbons)
This does not happen!
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.
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.