This document discusses bioluminescent symbioses between marine organisms and bacteria. It summarizes evidence from multiple studies that refutes the assumption that hosts and bioluminescent bacterial symbionts have coevolved. Specifically, it finds a lack of genetic adaptations for the symbiosis, independent development of light organs in hosts, and an ability for horizontal transfer of symbionts between host individuals and species. This suggests the symbiotic relationships do not involve tight co-speciation of hosts and bacteria.

1. 1
symbiosis phylogenetics
bioluminescence

2. Bioluminescent symbioses:
analysis of the host-symbiont
specificity and codivergence using a
phylogenetic approach
Patricia S. San Jose
The Marine Science Institute
University of the Philippines
2

3. reciprocal exploitations:
costs and benefits
3Herre, EA, Knowlton, N, Mueller, UG, and Rehner, SA. 1999. “The evolution of mutualisms: exploring the paths between conflict
and cooperation”. Trends in Ecology & Evolution 14: 49-53.
Evolution of mutualisms: trends, models

4. alignment of interest
4Herre, EA, Knowlton, N, Mueller, UG, and Rehner, SA. 1999. “The evolution of mutualisms: exploring the paths between conflict
and cooperation”. Trends in Ecology & Evolution 14: 49-53.
Evolution of mutualisms: trends, models

5. There is no general theory of mutualism…;
underlying problems revolve around explicitly
defining vague terms, such as “alignment of
interest”, and employing biologically realistic
currencies (i.e., costs and benefits) at
biologically relevant scales of organizations.
5Herre, EA, Knowlton, N, Mueller, UG, and Rehner, SA. 1999. “The evolution of mutualisms: exploring the paths between conflict
and cooperation”. Trends in Ecology & Evolution 14: 49-53.

6. conflict, cooperation, constraint
6Herre, EA, Knowlton, N, Mueller, UG, and Rehner, SA. 1999. “The evolution of mutualisms: exploring the paths between conflict
and cooperation”. Trends in Ecology & Evolution 14: 49-53.
Evolution of mutualisms: elements where factors
maintaining stability of mutualisms operate on

7. 7
transmission
genotypic
uniformity
spatial
structure
restricted
options
Herre, EA, Knowlton, N, Mueller, UG, and Rehner, SA. 1999. “The evolution of mutualisms: exploring the paths between conflict
and cooperation”. Trends in Ecology & Evolution 14: 49-53.
Evolution of mutualisms: factors that maintain the stability
of mutualisms across evolutionary timescales

8. coevolution: reciprocal evolutionary
change
8
Thompson J. 1989. “Concepts of Coevolution”. Trends in Ecology & Evolution 4: 179-183.
The phylogenetic approach towards understanding
the stability of mutualisms: codivergence

9. cophylogenetic analyses: patterns
9Charleston MA and SL Perkins. 2006. “Traversing the tangle: Algorithms and applications for cophylogenetic studies”. Journal of
Biomedical Informatics 39: 62-71.
The phylogenetic approach towards understanding
the stability of mutualisms: cophylogeny

10. bioluminescence in the marine
environment:
the rule rather than the exception
10Widder, EA. 2010. “Bioluminescence in the Ocean: Origins of Biological, Chemical, and Ecological Diversity”. Science DOI:
10.1126/science.1174269.
The nature of bioluminescent symbioses

11. 11
light
production
anatomical
adaptations
specificity
lack of genetic
adaptations
horizontal
transmission
Dunlap PV , Ast JC, Kimura S, Fukui A, Yoshino, T, and Endo, H. 2007. “Phylogenetic analysis of host-symbiont specificity and
codivergence in bioluminescent symbioses”. Cladistics 23: 507-532.
The nature of bioluminescent symbioses:
the conflict

12. 12Dunlap PV , Ast JC, Kimura S, Fukui A, Yoshino, T, and Endo, H. 2007. “Phylogenetic analysis of host-symbiont specificity and
codivergence in bioluminescent symbioses”. Cladistics 23: 507-532.

13. 13
lack of
metabolic
contribution
Claes MF and PV Dunlap. 2000. “Aposymbiotic Culture of the Sepiolid Squid Euprymna scolopes: Role of the Symbiotic Bacterium
Vibrio fischeri in Host Animal Growth, Development, and Light Organ Morphogenesis”. Journal of Experimental Zoology 286: 280-
296
The nature of bioluminescent symbioses:
non-molecular data

14. 14
independent
development
of light organ
Claes MF and PV Dunlap. 2000. “Aposymbiotic Culture of the Sepiolid Squid Euprymna scolopes: Role of the Symbiotic Bacterium
Vibrio fischeri in Host Animal Growth, Development, and Light Organ Morphogenesis”. Journal of Experimental Zoology 286: 280-
296
Thacker CE and DM Roje. 2009. “Phylogeny of cardinalfishes (Teleostei: Gobiiformes: Apogonidae) and the evolution of visceral
bioluminescence”. Molecular Phylogenetics and Evolution 52: 735-745
The nature of bioluminescent symbioses:
non-molecular data

15. 15
lack of ability to
discriminate
among symbionts
Kaeding AJ, Ast JC, Pearce MM, Urbanezyk, H, Kimura S, Endo H, Nakamura M, and Dunlap PV. 2007. “Phylogenetic Diversity and
Cosymbiosis in the Bioluminescent Symbioses of ‘Photobacterium mandapamensis’”. Applied and Environmental Microbiology 73:
3173-3182
The nature of bioluminescent symbioses:
non-molecular data

16. 16
horizontal
transfer
Wadda M, Azuma N, Mizuno N, Kurokura H. 1999. “Transfer of symbiotic luminous bacteria from parental Leiognathus nuchalis to
their offspring”. Marine Biology 135: 683-687
Wollenberg MS and EG Ruby. 2012. “Phylogeny and fitness of Vibrio fischeri from the light organs of Euprymna scolopes in two
Oahu, Hawaii populations”. The ISME (International Society for Microbial Ecology) Journal 6: 352-362
Dunlap PV, Davis KM, Tomiyama S, Fujino M, Fukui A. 2008. “Developmental and Microbiological Analysis of the Inception of
Bioluminescent Symbiosis in the Marine Fish Nuchequula nuchalis (Perciformes: Leognathidae)”. Applied and Environmental
Microbiology 74: 7471-7481
Dunlap PV, Kojima Y, Nakamura S, Nakamura M. 2009. “Inception of formation and early morphogenesis of the bacterial light
organ of the sea urchin cardinalfish, Siphamia versicolor”. Marine Biology 156: 2011-2020
The nature of bioluminescent symbioses:
non-molecular data

17. assumption of coevolution: refuted
17

18. 18