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Octocoral Communities of the Deep Gulf of Mexico

Andrea M. Quattrini1, Cheryl Doughty1, Peter Etnoyer2, & Erik E. Cordes1 1Temple University & 2NOAA, Charleston SC : :

Clavularia rudis Anthomastus sp. Paragorgia sp.

Paracalyptrophora carinata ?biscaya Iridogorgia magnispiralis

2 Cnidaria: Anthozoa: Octocorallia

3 Overarching Goal To Determine Boundaries in the Deep GOM : from population differentiation to phylogenetic community structure

Determining pathways of genetic exchange will help effectively conserve deep coral habitat in the Gulf of Mexico

4 GOM Sites: Octocoral Collections 2008 – 2010

5 GOM Sites: Octocoral Collections 2008 – 2010

24 sites (250–2500 m): 380 specimens 6 mt msh: 44 haplotypes Bayesian Inference Node Support > 95% unless indicated Midpoint Rooting

Holaxonia- Alcyoniina

Calcaxonia

Scleraxonia-Alcyoniina

7

mt msh: 44 haplotypes Bayesian Inference Node Support > 95% unless indicated Midpoint Rooting

Holaxonia

8 P. citrina 77 Pterogorgia anceps Pinnigorgia flava M. bayeriM. flavida ? sp. P. multispina Paramuricea A P. placomus P. grandis P. multispina Paramuricea E Paramuricea E Paramuricea D P. ?biscaya Paramuricea sp. Paramuricea ?biscaya Paramuricea B Paramuricea C Paramuricea C Acanthogorgia sp. Echinomuricea sp. Acanthogorgia sp. 92 Acanthogorgia ?aspera Menella sp. Echinomuricea sp. Paraplexaura sp. Placagorgia sp. sp. ?Muriceides sp. 55 Bebryce sp. Lytreia sp. sp. Muriceides sp. Thrombophyton coronatum Muriceides sp. ?Muriceides sp.1 ?Muriceides sp. 2 ?Muriceides sp. 3 ?Muriceides sp. 4 ?Muriceides sp.5 Placogorgia tenuis Acanthogorgia ?armata Paracis sp. ?pallida Acanthogorgia sp. Acanthogorgia ?angustiflora N. ATL Villogorgia sp. 72 ventalina Pseudopterigia bipinnata Deep GOM Swiftia sp. virgulata Deep >50 m 54 Swiftia ?pacifica Shallow < 50 m ?Swiftia sp. 2 Plexauridae PACIFIC Swiftia ?pallida Deep >50m9 Thesea sp. Scleracis guadalupensis Shallow < 50m Widespread occurrence of Paramuricea in the N Atl, and bathymetric separation of sister spp in the GOM

P. multispina USNM P. placomus Paramuricea A Paramuricea A P. multispina Paramuricea E Paramuricea E Paramuricea D Paramuricea sp. Paramuricea ?biscaya B Paramuricea B Paramuricea C Paramuricea C 250–300 m Echinomuricea sp. 350–450 m 800–2400 m

P. ?biscaya Paramuricea Type C P. ?biscaya Paramuricea Type E 10 mt msh: 44 haplotypes Bayesian Inference Node Support > 95% unless indicated

Cheliodonisis aurantiaca mexicana

Calcaxonia

11 Cheliodonisis aurantiaca mexicana Narella cristata Narella abyssalis Parastenella sp. Primnoa wingi Candidella imbricata 78 Narella arbuscula 83 Narella paucilfora Arthogorgia sp. Arthogorgia sp. Calyptrophiora japonica Paracalyptrophora sp. Paracalyptrophora carinata C. a. mexicana Plumarella sp. Plumarella pellicuda Thourella grasshoffi Fanellia sp. Callogorgia americana Callogorgia sp. 1 Callogorgia gracilis Isidella sp. 72 Isidella sp. Isidella sp. Isidella sp. 1 Lepidisis caryophilla Isidella sp. 2 Acanella eburnea P. carinata Isididae sp. 2 Isididae Keratoisis sp. Keratoisis sp. Lepidisis sp. Isididae sp.1 Lepidisis sp. Keratoisis sp. Isididae nov. sp.2 Isididae genus nov. 1 (France, pers. comm) Isididae genus nov. sp.3 sp. Chrysogorgia sp. A Chrysogorgia sp. B Isididae Chrysogorgia sp. C Chrysogorgia sp. C Chrysogorgia sp. F Chrysogorgia sp. F N. ATL Chrysogorgia sp. D Chrysogorgia sp. E Deep GOM Chrysogorgia chryseisI. magnispiralis Deep >50 m Iridogorgia magnispiralis I.fontinalis Shallow < 50 m I. splendens 82 I. splendens PACIFIC Metallogorgia melanochris Deep >50 m Heliopora sp. 0.2 Shallow < 50 m Diversification of Callogorgia spp. in the GOM occurs along bathymetric and longitudinal gradients Fanellia sp.

Callogorgia americana americana Callogorgia americana delta Callogorgia sp. 1 250-300 m Callogorgia gracilis 300-400 m 0.1 400-900 m

C. a. americana C. a. delta Callogorgia sp. 1 Callogorgia gracilis 13 Diversification of Callogorgia spp. in the GOM occurs along bathymetric and longitudinal gradients Fanellia sp.

Callogorgia americana americana Callogorgia americana delta Callogorgia sp. 1 250-300 m Callogorgia gracilis 300-400 m 0.1 400-900 m

14 Community Phylogenetics Community: “Co-occurring assemblages of trophically similar ” (Hubbell 2005)

. Framework that integrates evolutionary history and community ecology

.“The pattern of phylogenetic relatedness of species distributions within and among communities” (Cavender-Bares et al. 2009)

. Test community assembly hypotheses: historical, niche, vs. neutral processes

15 Phylogenetic Relatedness Differs from a Null Model? Clustering Random Overdispersion

16 Non-Random Phylogenetic Structure in Two Depth Zones

SES: Clustered Random Overdispersed

COI+igr+msh extended barcode (McFadden et al. 2010), Bayesian Inference, antipatharia outgroup 17 Phylogenetic Turnover Between Depth Zones

Similarity MC338 DC673 60% DC583 GC852 80% Depth (m) 300-400 m 400-550 m 800-900 m AT47 1,350-1,450 m MC751 2.150-2,450 m MC118 VK826

GC338 VK906 GB299

GB535 WFL3 WFL1 GC235 WFL2 VK862

COI+igr+msh extended barcode (McFadden et al. 2010), Bayesian Inference, antipatharia outgroup 18 Summary Depth-related patterns observed in the phylogenetic structure of octocorals

Clades restricted to depth zone, but also shallow & deep sister taxa • Evidence for deep to shallow water diversification?

Further bathymetric separation of sister species in the GOM • Callogorgia diversification down slope?

Phylogenetic community structure changes with depth in the GOM • Phylogenetic turnover ca. 900–1,300 m

Community assembly processes may differ between depth zones • Competitive interactions (Overdispersion) to environmental filtering (Clustering) Processes?

Isididae Paragorgia sp. Paramuricea ?biscaya Chrysogorgia sp. 19 Acknowledgments R. Falco, M. Rittinghouse, T. Shank, C. Fisher, J. Lunden, K. Ilves, C. Morrison, D. Ruiz, W. Cho, A. Demopoulos, L. Wickes, M. Nizinski, C. Ames, S. Ross, T. Casazza, J. McClain, R. Kulathinal, A. Freestone, J. Thoma, S. Herrera, J. Sanchez, S. Cairns, S. Morton and TDI-Brooks

Funding: BOEMRE, NOAA-OER,USGS, NSF RAPID, Temple University, & the Dr. Nancy Foster Scholarship Program

20 References

Cavender-Bares, J., K.H. Kozak, P.V. Fine, and S.W. Kembel. 2009. The merging of community ecology and phylogenetic biology. Ecology Letters 12:693–715. Hubbell, S.P. 2005. Neutral theory in community ecology and the hypothesis of functional equivalence. Functional Ecology 19:166–172. McFadden, C., J. Sanchez, S. France. 2010. Molecular phylogenetic insights into the evolution of Octocorallia: A review. Integr. Compar. Biol., doi:10.1093/icb/icq056.