Is the Family Characidae Monophyletic?

The first attempt at a cladistic analysis to include most lineages currently grouped in the "family" Characidae was undertaken by Lucena (1993). We reproduce his results below.

A phylogenetic analysis of the family Characidae, based in part on a classification proposed by Greenwood et al. (1966), is presented. The main findings are expressed as a strict consensus cladogram defined by 122 morphologial characters and 69 terminal taxa. The study suggested a monophyletic status for the family, but that remarkable modifications in characid nomenclature should take place in order to express the results obtained.

Molecular evidence (see Ortí and Meyer, 1997) does not lend support to these results.

              ==================================== Ctenolucius 
           ===|
        ===|  ==================================== Boulengerella 
        |  |
        |  ======================================= Hepsetus 
        |
        |     ==================================== Hoplias 
        |     |
     ===|  ===|  ================================= Erythrinus 
     |  |  |  ===|
     |  |  |     ================================= Hoplerythrinus 
     |  ===|
     |     |     ================================= Pyrrhulina 
     |     |  ===|
     |     ===|  ================================= Nannostomus 
     |        |
     |        ==================================== Lebiasina 
     |
     |        ==================================== Cynodon 
     |     ===|
     |  ===|  ==================================== Rhaphiodon 
     |  |  |
     |  |  ======================================= Acestrorhynchus 
     |  |
     |  |  ======================================= Agoniates 
     |  |  |
     |  |  |              ======================== Gnathocharax 
     |  |  |           ===|
     |  |  |        ===|  ======================== Hoplocharax 
     |  |  |     ===|  |
     |  |  |  ===|  |  =========================== Heterocharax 
     |  |  |  |  |  |
     |  |  |  |  |  ============================== Lonchogenys 
     |  |  |  |  |
     |  |  |  |  ================================= Roestes 
     |  |  |  |
     |  |  |  |                          ========= Roeboides p 
     |  |  |  |                       ===|
     |  |  |  |                    ===|  ========= Roeboides sp 
     |  |  |  |                 ===|  |
     |  |  |  |              ===|  |  ============ Eucynopotamus 
     |  |  |  |        ======|  |  |
     |  |  |  |        |     |  |  =============== R.myersi 
     |  |  |  |     ===|     |  |
     |  |  |  |  ===|  |     |  ================== Charax 
     |  |  |  |  |  |  |     |
     |  |  |  |  |  |  |     ===================== Cynopotamus 
     |  |  |  |  |  |  |
     |  |  |  |  |  |  |========================== Acestrocephalus 
     |  |  |  |  |  |  |
     |  |  |  |  |  |  =========================== Galeocharax 
     |  |  |  |  |  |
     |  |  |  |  |  ============================== Acanthocharax 
     |  |  |  |  |
     |  |  |  |  |  ============================== Roeboexodon 
     |  |  |  |  |  |
     |  |  |  |  |  |  =========================== Hemibrycon 
     |  |  |  |  |  |  |
     |  |  |  |  |  |  |              ============ Tetragonopterus 
     |  |  |  |  |  |  |        ======|
     |  |  |  |  |  |  |        |     ============ Markiana 
     |  |  |  |  |  |  |  ======|
     |  |  |  |  |  |  |  |     |================= Poptella 
     |  |  |  |  |  |  |  |     |
     |  |  |  |  |  |  |  |     ================== Moenkhausia gr 
     |  |  |  |  |  |  |  |
  ===|  |  |  |  |  |  |  |        =============== Piabina 
     |  |  |  |  |  |  |  |     ===|
     |  |  |  |  |  |  |  |     |  =============== Creagrutus 
     |  |  |  |  |  |  |  |=====|
     |  |  |  |  |  |  |  |     |  =============== Ctenobrycon 
     |  |  |  |  |  |  |  |     ===|
     |  |  |  |  |  |  |  |        =============== Iguanodectes 
     |  |  |  |  |  |  |  |
     |  |  |  |  |  |  |  |     ================== Phenacogaster 
     ===|  |  |  |  |  |  |=====|
        |  |  |  |  |  |  |     ================== Aphyocharax 
        |  |  |  |  |  |  |
        ===|  |  |  |  |  |     ================== Oligosarcus 
           ===|  |  |  |  |=====|
              |  |  |  |  |     ================== Exodon 
              |  |  |  |  |
              |  |  |  |  |     ================== Aphyodite 
              |  |  |  |  |     |
              ===|  |  |  |     |================= Astyanax 
                 |  |  |  |     |
                 |  |  |  |     |================= Bryconamericus 
                 |  |  |  |     |
                 |  |  |  |     |================= Cheirodon 
                 |  |  |  |     |
                 |  |  |  |     |================= Deuterodon 
                 |  |  |  |     |
                 |  |  |  |     |================= Diapoma 
                 ===|  |  |     |
                    ===|  |     |================= Hemigrammus 
                       ===|     |
                          |     |================= Holoshesthes 
                          |     |
                          |     |================= Hyphessobrycon 
                          |=====|
                          |     |================= Moenkhausia le 
                          |     |
                          |     |================= Odontostilbe 
                          |     |
                          |     |================= Othonocheirodus 
                          |     |
                          |     |================= Paragoniates 
                          |     |
                          |     |================= Parecbasis 
                          |     |
                          |     |================= Prionobrama 
                          |     |
                          |     |================= Probolodus 
                          |     |
                          |     |================= Prodontocharax 
                          |     |
                          |     ================== Pseudocorynopoma 
                          |
                          |     ================== Bryconops 
                          |     |
                          |     |================= Triportheus 
                          |     |
                          |     |     ============ Serrasalmus 
                          |     |     |
                          |     |     |     ====== Brycon 
                          ======|     |  ===|
                                |     |  |  ====== Chalceus 
                                ======|  |
                                      ===|  ====== Alestes 
                                         |  |
                                         ===|  === Hemigrammopetersius 
                                            ===|
                                               === Hydrocynus 

References

Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966. Phyletic studies of teleostean fishes, with a provisional classification of living forms. Bulletin of the American Museum of Natural History 131:339-354.

Lucena, C. A. S. d. 1993. Estudo filogenetico da familia Characidae com uma discussio dos grupos naturais propostos (Teleostei, Ostariophysi, Characiformes). Unpublished Ph.D. thesis, Universidade de Sao Paulo, Brasil.

Ortí, G.and A. Meyer. 1997. The radiation of characiform fishes and the limits of resolution of mitochondrial ribosomal DNA sequences. Syst. Biol. 46:75-100.