Our Seahorse Research
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We work on fish evolution in the seahorse family.
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Our Seahorse Research
- 1. A Multigene Phylogeny of Syngnathid Fishes Healy Hamilton California Academy of Sciences Norah Saarman University of California, David Harasti Santa Cruz Beth Moore, Graham Short, & W. Brian Simison California Academy of Sciences Martin Gomon Museum Victoria
- 2. Acknowledgments Dr. Jeff Leis and Mark McGrouther Australian Museum Dianne Bray, Museum Victoria Ralph Foster, South Australian Museum David Harasti, Dept. of Primary Industries, NSW SEAHORSE Clive Roberts, Carl Struthers SLEUTH Museum of New Zealand, Te Papa, Tongarewa Dr. David Neely, Dave Catania, Jens Vindum, Jim Evans California Academy of Sciences Dr. E. O. Wiley and Andy Bentley National History Museum, The University of Kansas Dr. Mark Erdmann, Conservation International Dr. Gerry Allen, Western Australian Museum Institute for Research & Development, New Caledonia Pierre LaBoute, Noumea, New Caledonia Dr. Robert Browne, Dr. David Muirhead, Kevin Smith Inshore Fish Group Dr. Theodore W. Pietsch and Katherine Pearson University of Washington
- 3. Outline: A Multigene Phylogeny of Syngnathid Fishes 1. Introduction: why syngnathids? 2. Issues in current syngnathid taxonomy 3. Molecular approach to address syngnathid systematics 4. Results: molecular phylogenetic Photos: hypotheses, mapping of non-molecular David Harasti characters 5. Conclusions and next steps
- 4. Introduction: Why we study syngnathid fishes • Syngnathid fishes: a diverse, unique and IUCN Syngnathid listings charismatic group DD VU EN CR Seahors es 23 9 1 • over 50 genera and at Pipefishes 5 1 least 278 valid species of Seadragons 2 Pipehorses 5 pipefishes, seahorses, DD = Data Deficient sea dragons, and VU = Vulnerable EN = Endangered pipehorses CR = Critically Endangered Syngnathidae are characterized by: unique gills absence of pelvic fins dermal plates in rings unusual reproductive strategies, including sex role reversal and male brooding, with complex and varied brood-pouch morphology
- 5. Hippocampus whitei Graham Short
- 6. Hippocampus abdominalis Graham Short
- 9. Weedy Sea Dragon Phyllopteryx taeniolatus Graham Short
- 10. Leafy Sea Dragon Phycodurus eques Graham Short
- 11. Pygmy pipehorse Idiotropiscis lumnitzeri Graham Short
- 12. Spotted pipefish Stigmatopora argus Graham Short
- 13. Briggs Crested pipefish Histiogamphelus briggsi Graham Short
- 14. Bent stick pipefish Trachyrhampus bioarctatus David Harasti
- 15. The Problem: Systematic relationships unresolved • Unresolved systematic relationships at multiple levels • Generic and specific boundaries are often uncertain • evolutionary relationships at the species, genus and subfamily level are largely unresolved • order Gasterosteiformes / Syngnathiformes debated • many new species remain to be described • Many genera are monotypic, yet no study has measured their degree of evolutionary uniqueness Conservation and management of syngnathids is undermined by their challenging alpha taxonomy and poorly resolved systematics
- 16. The problem Only two recognized sub-families, the Hippocampinae and the Syngnathinae (Paxton 1975) (Whitley, 1952) A. Hippocampinae: seahorses B. Syngnathinae: all others; is paraphyletic and includes 54 includes only one genus with genera and 233 valid species 38 - 50 species (debated) • Seadragons (Solegnathinae • Tail-brooding Pipefishes (original Syngnathinae) • Abdominal-brooding Pipefishes (Doryrhamphinae and Nerophinae) • Pipehorses (originally included in Hippocampinae)
- 17. Our Approach: Methods Morphological characters have proven problematic Molecular studies to date (Wilson et al., 2001 & 2003) suffer from limited taxon and molecular character sampling. WE HAVE ADDED: 20 new genera • Total of 59 taxa (37 genera) 4 new genetic markers, including 2 nuclear fragments • 6 markers total, ~4500 aligned bps • Mitochondrial; CO1, ND1, 12s, 16s • Nuclear; RP1 intron of S7, Myh 6
- 18. Our Approach: Increased Taxon Sampling Previous Generic Assignment Species Wilson (2003) This Study Previous Generic Assignment Species Wilson (2003) This Study Acentronura 5 1 Kimblaeus 1 Idiotropiscis 1 1 Leptoichthys 1 1 Anarchopterus 2 Leptonotus 3 Apterygocampus 1 Lissocampus 5 Bhanotia 3 Maroubra 2 1 Bryx 5 Micrognathus 7 1 Bulbonaricus 3 Microphis 21 2 Campichthys 4 Minyichthys 4 Choeroichthys 6 2 Mitotichthys 4 1 Corythoichthys 10 3 2 Nannocampus 4 Cosmocampus 14 1 Nerophis 3 1 1 Doryichthys 5 1 Notiocampus 1 Doryrhamphus 14 1 3 Penetopteryx 2 Dunkerocampus 1 1 1 Phoxocampus 3 Enneacampus 2 Phycodurus 1 1 Entelurus 1 1 1 Phyllopteryx 1 1 1 Festucalex 6 1 Pseudophallus 3 1 Filicampus 1 1 Pugnaso 1 1 1 Halicampus 12 1 Siokunichthys 5 1 Haliichthys 1 1 Solegnathus 5 1 2 Heraldia 1 Stigmatopora 3 4 3 Hippichthys 5 1 Stipecampus 1 1 Hippocampus 45 9 9 Syngnathoides 1 1 Histiogamphelus 2 2 Syngnathus 32 14 3 Hypselognathus 2 1 1 Trachyrhamphus 3 1 Ichthyocampus 2 Urocampus 2 1 1 Kaupus 1 1 1 Vanacampus 4 2 4 Total 54 genera (16 monotypic) 278 species 16 genera 37 genera
- 19. Results: Bayesian consensus tree Aulostomus chinensis 1 Aulostomus maculatus Fistularia petimba Solenostomus cyanopterus Choeroichthys brachysoma 1 Choeroichthys sculptus 1 Doryichthys boaja Outgroups 1 Microphis doecata 1 1 Microphis ocellatus Doryrhamphus dactyliophorus 1 1 0.99 Dunckerocampus chapmani 1 Doryrhamphus excises 1 0.51 Doryrhamphus jansii Maroubra perserrata Leptoichthys fistularis 1 1 Entelerus aequoreus Abdominal-brooding pipefish Nerophis spp 1 Corythoichthys amplexus Corythoichthys intestinalis 1 1 Stigmatopora argus 1 0.99 Stigmatopora narinosa Stigmatopora nigra 1 Phycodurus eques Phyllopteryx taeniolatus 0.79 1 Solegnathus hardwicki 1 1 Solegnathus spinosissimus Syngnathoides biaculeatus Stipecampus cristatus Seadragons Cosmocampus elucens Micrognathus andersonii 1 1 1 Syngnathus louisianae 1 1 Syngnathus fuscus 1 Syngnathus exilis 0.85 Pseudophallus brasiliensis Hippicthys penicillus 1 0.71 Siokunichthys nigrolineatus Urocampus carinirostris 1 Festucalex cinctus Histiogamphelus briggsi 0.72 0.65 1 Histiogamphelus cristatus Hypselognathus rostratus Tailbrooding pipefish 1 1 1 Mitotichthys semistriatus Kaupus costatus 0.66 Pugnaso curtirostris 1 0.91 1 Vanacampus vercoi Vanacampus margaritifer 1 Vanacampus phillipi 1 Vanacampus poecilolaemus 1 Halicampus sp 0.52 Hippocampus abdominalis 0.95 Hippocampus trimaculatus Hippocampus whitei Seahorses 1 Hippocampus fuscus 1 1 Hippocampus ingens 1 Hippocampus zosterae Hippocampus bargibanti 1 Hippocampus denise 1 1 Hippocampus pontohi Filicampus tigiris 1 1 Trachyrhamphus bioarctatus Haliichthys taeniophorus 0.65 1 Acentronura tentaculata Pipehorses Idiotropiscis lumnitzeri
- 20. Results Bayesian consensus tree Maximum parsimony strict consensus Aulostomus chinensis 1 Aulostomus maculatus Aulostomus chinensis Aulostomus maculatus Fistularia petimba Fistularia petimba Solenostomus cyanopterus Solenostomus cyanopterus Choeroichthys brachysoma 100 Choeroichthys brachysoma 1 Choeroichthys sculptus 67 Choeroichthys sculptus Doryichthys boaja 100 Doryrhamphus dactyliophorus 1 1 1 1 Microphis doecata Dunckerocampus chapmani Microphis ocellatus 100 Doryichthys boaja Doryrhamphus dactyliophorus 88 90 Microphis doecata 1 1 Dunckerocampus chapmani Microphis ocellatus 0.99 100 96 1 Doryrhamphus excises 96 Doryrhamphus excises 1 Doryrhamphus jansii 0.51 Doryrhamphus jansii Maroubra perserrata 95 Maroubra perserrata Leptoichthys fistularis 100 Entelerus aequoreus 1 Nerophis spp Entelerus aequoreus 1 Leptoichthys fistularis Nerophis spp Corythoichthys amplexus 100 1 Corythoichthys amplexus Corythoichthys intestinalis 98 Corythoichthys intestinalis 96 Stigmatopora argus 1 1 Stigmatopora argus 100 100 Stigmatopora narinosa 1 0.99 Stigmatopora narinosa Stigmatopora nigra Stigmatopora nigra 100 Phycodurus eques Phycodurus eques 100 Phyllopteryx taeniolatus 1 Phyllopteryx taeniolatus 100 80 Solegnathus hardwicki 0.79 Solegnathus spinosissimus 1 Solegnathus hardwicki 67 1 1 Solegnathus spinosissimus Syngnathoides biaculeatus Syngnathoides biaculeatus Stipecampus cristatus Stipecampus cristatus 100 Acentronura tentaculata Idiotropiscis lumnitzeri Cosmocampus elucens Filicampus tigiris 85 Micrognathus andersonii 100 Trachyrhamphus bioarctatus 1 1 1 Syngnathus louisianae Haliichthys taeniophorus 1 1 Syngnathus fuscus Cosmocampus elucens 1 Syngnathus exilis 100 Micrognathus andersonii 0.85 100 Pseudophallus brasiliensis Syngnathus louisianae 100 Hippicthys penicillus Syngnathus fuscus 1 99 0.71 Siokunichthys nigrolineatus Syngnathus exilis Urocampus carinirostris 98 Pseudophallus brasiliensis Festucalex cinctus 79 Hippicthys penicillus 1 Siokunichthys nigrolineatus 0.72 1 Histiogamphelus briggsi Histiogamphelus cristatus Festucalex cinctus 1 0.65 1 Hypselognathus rostratus L=20652 68 99 Histiogamphelus briggsi Histiogamphelus cristatus 1 Mitotichthys semistriatus 90 90 Kaupus costatus Kaupus costatus 0.66 Pugnaso curtirostris CI=0.228 100 Hypselognathus rostratus 1 Mitotichthys semistriatus 0.91 1 Vanacampus vercoi 98 Pugnaso curtirostris 1 Vanacampus margaritifer RI=0.499 Vanacampus vercoi Vanacampus phillipi Vanacampus margaritifer 1 100 Vanacampus poecilolaemus 100 Vanacampus phillipi 1 Halicampus sp Vanacampus poecilolaemus 0.52 Hippocampus abdominalis Halicampus sp Hippocampus trimaculatus Hippocampus abdominalis 0.95 Hippocampus whitei 100 Hippocampus fuscus 1 Hippocampus fuscus 100 Hippocampus ingens 1 1 Hippocampus ingens Hippocampus trimaculatus 1 Hippocampus zosterae Hippocampus whitei 100 Hippocampus bargibanti Hippocampus zosterae 1 Hippocampus bargibanti 1 Hippocampus denise 100 Hippocampus pontohi Hippocampus denise 1 Filicampus tigiris Hippocampus pontohi 1 Urocampus carinirostris 1 Trachyrhamphus bioarctatus 0.65 Haliichthys taeniophorus Acentronura tentaculata 1.1 1 Idiotropiscis lumnitzeri 0.2
- 21. Results: Bayesian consensus tree Aulostomus chinensis Outgroups 1 Aulostomus maculatus Fistularia petimba Solenostomus cyanopterus Choeroichthys brachysoma 1 Choeroichthys sculptus Doryichthys boaja 1 1 1 Microphis doecata 1 Microphis ocellatus Doryrhamphus dactyliophorus 1 1 0.99 Dunckerocampus chapmani 1 Doryrhamphus excises 1 0.51 Doryrhamphus jansii Maroubra perserrata Leptoichthys fistularis 1 1 Entelerus aequoreus Nerophis spp 1 Corythoichthys amplexus Corythoichthys intestinalis 1 1 Stigmatopora argus 1 0.99 Stigmatopora narinosa Stigmatopora nigra Gastrophori; abdominal-brooding fish 1 Phycodurus eques Phyllopteryx taeniolatus 0.79 1 Solegnathus hardwicki 1 1 Solegnathus spinosissimus Syngnathoides biaculeatus Stipecampus cristatus Cosmocampus elucens Micrognathus andersonii 1 1 1 Syngnathus louisianae 1 1 Syngnathus fuscus Brood pouch 1 Syngnathus exilis 0.85 anus Pseudophallus brasiliensis Hippicthys penicillus 1 0.71 Siokunichthys nigrolineatus Urocampus carinirostris 1 Festucalex cinctus 0.72 1 Histiogamphelus briggsi Histiogamphelus cristatus 0.65 Hypselognathus rostratus 1 1 1 Mitotichthys semistriatus Kaupus costatus 0.66 Pugnaso curtirostris 1 0.91 1 1 Vanacampus vercoi Vanacampus margaritifer Urophori; tail-brooding fish Vanacampus phillipi 1 Vanacampus poecilolaemus 1 Halicampus sp 0.52 Hippocampus abdominalis Hippocampus trimaculatus 0.95 Hippocampus whitei 1 Hippocampus fuscus 1 1 Hippocampus ingens Brood pouch 1 Hippocampus zosterae Hippocampus bargibanti 1 Hippocampus denise anus 1 1 Hippocampus pontohi Filicampus tigiris 1 1 Trachyrhamphus bioarctatus 0.65 Haliichthys taeniophorus Acentronura tentaculata 1 Idiotropiscis lumnitzeri 0.2
- 22. Results: “Doryrhamphinae”: Flagtail pipefishes Mostly freshwater pipefishes Flagtail pipefishes DAVID HARASTI Dunkerocampus Doryrhamphus
- 23. Results “Solegnathinae”: Sea Dragon clade 100 Prehensile tails David Harasti Solegnathus Sygnathoides Haliichthys taeniophorus “Ribboned” sea dragon or pipefish
- 24. All photos Results: Who is sister to seahorses? David Harasti Prehensile tails Haliichthys Filicampus Trachyramphus
- 25. Conclusions • Our molecular phylogeny is in agreement with Wilson(2003). Much revisionary work is needed at multiple taxonomic levels. • The division of syngnathids into sub-families Syngnathinae and Hippocampinae is not supported; Syngnathinae is paraphyletic • Maximum Parsimony and Bayesian phylogenetic analyses reproduce similar sets of clades, but differ in resolution of their arrangement • Multiple origins of characters such as prehensile tails, freshwater habit, and fin presence/absence • The seahorse's sister group is still unresolved
- 26. Future research in syngnathid systematics • Further mapping of morphological characters to elucidate their origins and patterns in syngnathids • Broader intraspecific geographic sampling • Additional nuclear markers and increased axon sampling (21 genera to go!) These future steps will further advance a sufficiently robust phylogeny to support syngnathid taxonomy, systematics, biogeography, and conservation.