Crustacea: Decapoda: Caridea: Palaemonidae)
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ZV-336 003-434 | fransen 04-01-2007 07:44 Pagina 3 Taxonomy, phylogeny, historical biogeography, and historical ecology of the genus Pontonia (Crustacea: Decapoda: Caridea: Palaemonidae) C.H.J.M. Fransen Fransen, C.H.J.M. Taxonomy, phylogeny, historical biogeography, and historical ecology of the genus Pontonia (Crustacea: Decapoda: Caridea: Palaemonidae). Zool. Verh. Leiden 336, 31.iii.2002: 1-433 figs. 1-257, plates 1-19.— ISSN 0024-1652, ISBN 90-73239-80-X. C.H.J.M. Fransen, Nationaal Natuurhistorisch Museum, P.O. Box 9517, 2300 RA Leiden, The Nether- lands (e-mail: [email protected]). Key words: Crustacea; Decapoda; Caridea; Pontoniinae; Pontonia; taxonomy; phylogeny; historical bio- geography; historical ecology. Species of the genus Pontonia Latreille, 1829, are distributed in tropical and subtropical waters around the world, living in association with either molluscan or ascidian hosts. In the present taxonomic revision, Pontonia sensu lato is divided into six genera: Pontonia sensu stricto; Ascidonia gen. nov., Rostronia gen. nov., Dactylonia gen. nov., Odontonia gen. nov., and Bruceonia gen. nov. A total of 29 species is described and figured, four of which are new to science: Pontonia pilosa spec. nov., Dactylonia holthuisi spec. nov., Odontonia rufopunctata spec. nov., and O. seychellensis spec. nov. The division in six genera is based on a stepwise phylogenetic analysis using morphological charac- ters, performed using PAUP and McClade software. The hypothesized phylogeny is used to analyse the historical biogeography of the shrimps. A deduc- tive method using Brooks Parsimony Analysis (BPA) is performed with regards to the Atlantic and East Pacific areas. The distribution patterns in the Indo-West Pacific are tested against the three sce- nario’s explaining the ‘East Indies Triangle of maximum diversity’. The historical ecology of the shrimps is analysed using their hypothesized phylogeny. In analogy with the historical biogeographical analyais, a deductive BPA method was applied, measuring the fit of the shrimp phylogeny on the hypothesized host phylogeny at different taxonomical levels of the hosts. Contents 1. Introduction ............................................................................................................................................ 5 2. Material and methods ........................................................................................................................ 5 3. Historical review .................................................................................................................................. 6 4. Morphology and terminology ....................................................................................................... 9 5. Phylogenetic analysis ......................................................................................................................... 9 5.1. Taxa .................................................................................................................................................. 9 5.2. Character state definition .................................................................................................... 10 5.3. Finding monophyletic groups .......................................................................................... 17 5.4. Intergeneric phylogenetic relations ................................................................................ 26 5.5. Intrageneric phylogenetic relations ................................................................................ 36 5.6. A complete cladogram .......................................................................................................... 41 5.7. Character evolution in retrospect .................................................................................... 42 6. Historical biogeography ................................................................................................................. 44 6.1. Biological characteristics ...................................................................................................... 44 6.2. Processes ...................................................................................................................................... 44 6.2.1. Vicariance ....................................................................................................................... 45 ZV-336 003-434 | fransen 04-01-2007 07:44 Pagina 4 4 Fransen. Revision of the genus Pontonia. Zool. Verh. Leiden 336 (2002) 6.2.2. Coalescence .................................................................................................................... 46 6.2.3. Dispersal .......................................................................................................................... 46 6.2.4. Non-allopatric speciation ........................................................................................ 46 6.2.5. Extinction ........................................................................................................................ 47 6.3. Distributional patterns ......................................................................................................... 47 6.3.1. The Atlantic and East Pacific area ....................................................................... 47 6.3.2. The Indo-West Pacific area ..................................................................................... 53 7. Historical ecology .............................................................................................................................. 56 7.1. Methodology ............................................................................................................................. 56 7.2. The hosts ..................................................................................................................................... 58 7.3. Results .......................................................................................................................................... 62 7.4. Conclusions and discussion ............................................................................................... 67 8. Systematic account ............................................................................................................................ 68 8.1. Key to the genera ..................................................................................................................... 68 8.2. Pontonia Latreille, 1829 ......................................................................................................... 69 8.2.1. Key to the species of Pontonia Latreille ............................................................. 70 8.2.2. Pontonia chimaera Holthuis, 1951 .......................................................................... 71 8.2.3. Pontonia domestica Gibbes, 1850 ............................................................................ 80 8.2.4. Pontonia longispina Holthuis, 1951 ....................................................................... 93 8.2.5. Pontonia manningi Fransen, 2000 ........................................................................ 102 8.2.6. Pontonia margarita Smith, 1869 ............................................................................ 119 8.2.7. Pontonia mexicana Guérin, 1856 .......................................................................... 131 8.2.8. Pontonia pilosa spec. nov. ....................................................................................... 147 8.2.9. Pontonia pinnae Lockington, 1878 ...................................................................... 158 8.2.10. Pontonia pinnophylax (Otto, 1821) .................................................................... 172 8.2.11. Pontonia simplex Holthuis, 1951 ....................................................................... 190 8.3. Ascidonia gen. nov. ............................................................................................................... 201 8.3.1. Key to the species of Ascidonia gen. nov. ....................................................... 202 8.3.2. Ascidonia californiensis (Rathbun, 1902) ........................................................... 203 8.3.3. Ascidonia flavomaculata (Heller, 1864) .............................................................. 215 8.3.4. Ascidonia miserabilis (Holthuis, 1951) ............................................................... 227 8.3.5. Ascidonia pusilla (Holthuis, 1951) ....................................................................... 236 8.3.6. Ascidonia quasipusilla (Chace, 1972) .................................................................. 248 8.4. Rostronia gen. nov. ................................................................................................................ 258 8.4.1. Rostronia stylirostris (Holthuis, 1951) ............................................................... 260 8.5. Dactylonia gen. nov. ............................................................................................................. 270 8.5.1. Key to the species of Dactylonia gen. nov. ..................................................... 271 8.5.2. Dactylonia anachoreta (Kemp, 1922) ................................................................... 271 8.5.3. Dactylonia ascidicola (Borradaile, 1898) ............................................................ 282 8.5.4. Dactylonia holthuisi spec. nov. ............................................................................