Photo by Frank Magallanes (top) S. rhombeus (bottom) P. nattereriSubfamily Serrasalminae or Family Serrasalmidae?

By Frank Magallanes, OPEFE

 

Photo (left) demonstrate different body forms of the pirambeba S. rhombeus (top) P. nattereri (bottom). 

 

 

 

INTRODUCTION

Photo by Frank MagallanesThe systemic of the Characiformes has long been (and continues to be) a difficult task to undertake. They were recognized as a homogeneous group in 1844 by German ichthyologist's Müller and Trochel. The Viennese Kner and his student Steindachner (between 1858-1915) followed by describing accurately a number of species without paying much attention to the classification. It was not until much later that Carl H. Eigenmann (considered to be the Father of Characoidologist) established the natural classifications of characins from South America. His principal manuscripts were posted between 1910 and 1927. He was then followed by one of his students Dr. George S. Myers. Within recent modern times a series of anatomical studies was completed by S. Weitzman, T. Roberts, and others helped establish the critical position of several groups, which led to the recognition of many families within the suborder (a recognition that would not have been accepted during Eigenmann's time). The research revealed the importance, as well as the complexity, of the Characiformes, and is presently considered as a key-group among teleosts. Included in this huge grouping to be discussed specifically at this website are the Serrasalmin which includes the pacus, silver dollars, pirambebas and of course, the true piranhas. The first authoritative division of the Characoids into several families (16) was done by S. Weitzman, in Greenwood et al. (1966). The French ichthyologist J. Géry (1972) would modify this order in a later manuscript.

Piranha and Pirambeba have distinctive body differences which become apparent as maturity sets in. These differences are the key reason why South American natives call non-true piranhas by other names. It is only outside South America (or S. A. non-fishing city dweller's) that the name "piranha" is loosely applied to all species within the subfamily Serrasalminae.

HISTORICAL IMAGES COURTESY OF ADRIEN LEROY

 

 

GENERAL EXPLANATION

The name Serrasalminae means saw-salmon-family the saw or serration pertaining to the scutes (or serrated keel) found on the belly of these fishes. Both carnivorous (single row teeth per jaw) and the vegetarian (double row teeth per jaw) practice mimicry. Perhaps this has much to do with the ecological home they inhabit and survival.

The epithet "piranha" is perhaps the most over used common name on fishes that could not even be scientifically called piranhas. It has been used for Serrasalminae vegetarian fishes and other related forms. In order to properly understand what a piranha is, one must do some research into common names and how they are applied. The Piranha Book, edited by Dr. G. S. Myers, (pg 26, TFH Publications Inc.,1971) covers what the usage of common names should be defined as. I recommend the research student use that reference to understand common name usage.

I frequently use the loose term piranha when I am discussing the "carnivorous" group as a whole, since this is the name more closely associated with the fish. For specifics, I prefer the Brazilian pirambeba for the species not genus Pygocentrus. For the true piranhas placed in genus Pygocentrus I use the epithet Caribe or Piranha. Caribe pertains to the Spanish Venezuelan piranhas but is a much more loose application since many of the piranha-like forms are also called that. But if you get a native fishermen and try to pin him down he will simply distinguish the more innocuous species with another name or the true caribe as caribe. The same holds true for epithet piranha. The native fishermen will give another name for the piranha-like and use the name piranha for the most dangerous ones in genus Pygocentrus. One last thing, native fishermen do not use scientific names in describing their fishes, we do! So we must be careful when attaching a common name to a scientifically described fish.

It has been common practice for biologists, news media and laymen to describe vegetarian fishes (pacus and tambaqui) in the genera Colossoma or Piaractus as belonging to the "piranha family" they are not! Pacus, and piranhas are all members of the Characidae family that hosts well-over 2,000 species of fish. The Characidae family (loosely called tetras) are delineated into groups or subfamilies.

There have been several unsuccessful attempts to split the sub-family into 2 groups; Mylinae (pacus and such) and the Serrasalminae (piranhas and associated forms) during the course of ichthyologic history. Norman (1929) lumped the both groups into one subfamily naming it Serrasalmoninae, but got the spelling wrong (should have been Serrasalminae.) His basis was certain characters found on both species. But the problem was much deeper than simple character assimilation. It was the advent of Phylogenetic and DNA evidence which now prevents such delineation. DnA research has also been used to separate the species with results proving confounding. Some species merged to the surprise of experts. A good case is the species Catoprion mento (the wimple piranha), this species is closest (sister) to Pygocentrus in genetic terms. Ichthyologists over the centuries have kept this particular species separate in its own ranking, but with genetics it puts it closer to the true piranhas. The wimple piranha is a good example of a species having the surname "piranha" while it is NOT a piranha by scientific definition. Most recently, genus Metynnis was discovered to be closely aligned with genus Pygocentrus using Phylogenetic and DNA sequencing.

Other species like P. denticulata , which is a piranha, have unique, specialized teeth which help it remove seed husks, much like a pacu's teeth.

TWO SCIENTIFIC OPPOSING VIEWS REGARDING CLASSIFICATION

JACQUES GÉRY

Family Serrasalmidae Classification: Dr. Jacques Géry

 

The French ichthyologist Jacques Géry (1972), modified the group into a new family and sub-family adding a sub-genera which further delineated the piranhas, pirambebas, and the vegetarian silver dollars and pacus;

  • Class: Osteichthyes

  • Superorder: Teleostei

  • Order: Cypriniformes

  • Suborder: Characoidei

  • Family: Serrasalmidae

  • Sub-family: Serrasalminae

  • genus Serrasalmus

  • sub-genus Pristobrycon

  • sub-genus Pygopristis

  • sub-genus Pygocentrus

  • sub-genus Serrasalmus

  • sub-genus Taddyella

  • Sub-family: Catoprioninae

  • genus Catoprion

  • Sub-family: Mylinae

  • genus Colossoma

  • genus Mylossoma

  • genus Myleus

  • sub-genus Myloplus

  • sub-genus Paramyloplus

  • sub-genus Prosomyleus

  • sub-genus Myleus

  • genus Uiaritichthys

  • genus Metynnis

  • genus Acnodon

  • genus Mylesinus

Géry split the group based on several factor's listed below

 

  1. Teeth variable, usually in more than one row on upper jaw (exceptions occur in Serrasalmidae).

  2. Anal fin moderate or long, with at least 3 unbranched rays and 10 branched ones (with a few exceptions in certain regressed species).

  3. Scales usually cycloid (with some exceptions) with circuli of caudal (apical) zone parallel or even divergent with axis of body (except Serrasalmidae).

  4. Maxilla reduced, not toothed; dorsal fin long, with at least 16 rays; usually a predorsal spine and a series of ventral spines (serrae) (body very compressed, usually disciform; teeth variable according to diet; scales small, the circuli concentric).

 

ANTONIO MACHADO-ALLISON & WILLIAM L. FINK

SCIENTIFIC CLASSIFICATION PIRANHAS - use this link.

 

  • Kingdom: Animalia

  • Phylum: Chordata

  • Subphylum: Vertebrata

  • Superclass: Osteichthyes

  • Class: Actinopterygii

  • Subclass: Neopterygii

  • Infraclass: Teleostei

  • Superorder: Ostariophysi

  • Order: Characiformes

  • Family: Characidae

  • Subfamily: Serrasalminae

 

Subfamily Serrasalminae Classification - Dr. William L. Fink & Dr. Antonio Machado-Allison

 

1. The most recent evaluation of the number of species, valid names, alimentary habits and their Phylogenetic relationship with other groups was investigated by Fink and Machado-Allison (Fink, 1978; Machado-Allison, 1982a, 1985).
Their present findings do not substantiate the further splitting as revised by Géry (1972). The reasoning behind this is because Phylogeny is the now the standard norm. The basic issue is that many of the older classifications, accepted non-monophyletic groups as valid to be named.

2. There is evidence that some "pacus" are more closely related to piranhas than other "pacus" ie; that some pacus share a more recent common ancestor with piranhas than with the other pacus.

3. That makes the group "pacus" non-monophyletic (actually paraphyletic). In relation to today's modern Phylogenetic classifications, the older methods of determining classification (or ranking) is outdated.

4. The older classifications were not built with the Phylogenetic philosophy in mind. Right now the evidence on pacu/piranha relationships is equivocal and its doesn't seem to be useful to revise the previous classifications.

5. The ranks are entirely arbitrary--there is no scientific basis for any ranking procedure, so accepting a group as a subfamily or family is entirely a matter of taste.

 

CURRENT SERRASALMINAE SUBFAMILY GENERIC RANKING - 2007

Pygocentrus

Serrasalmus 

Pristobrycon

Pygopristis

Catoprion

Metynnis

Mylossoma

Piaractus

Colossoma

Myleus

Myloplus

Mylesinus

Acnodon

Utiaritichthys

Tometes

Ossubtus

 

 

 

 

 

ABSTRACT

Phylogeny of the Serrasalminae (Characiformes) based on mitochondrial DNA sequences

Previous work (Ortí et al. 1995) based on DNA sequences of mitochondrial (mt) rRNA genes showed three main groups within the subfamily Serrasalminae:

(1) a basal clade of herbivores (Colossoma, Mylossoma, Piaractus);

(2) the "Myleus" clade (Myleus, Mylesinus, Tometes);

(3) the "piranha" clade (Serrasalmus, Pygocentrus, Pygopristis, Pristobrycon, Catoprion, Metynnis). The genus Acnodon was placed as the sister taxon of clade (1+2). However, poor resolution within each clade was obtained due to low levels of variation among rRNA sequences.

D-LOOP PHYLOGENY

Complete sequences of the hypervariable mtDNA D-loop are now presented for a total of 40 taxa representing all genera in the subfamily to address intragroup relationships. Phylogenetic analyses of these sequences identify the same groupings as before and provide further evidence to support the following observations:

(a) the genera Serrasalmus and Pristobrycon are paraphyletic and form a group that also includes Pygocentrus;

(b) Catoprion, Pygopristis, and Pristobrycon striolatus form a well supported clade, sister to the group described above in 'a';

(c) distinction of subgenera within Myleus (i.e., Myleus, Prosomyleus, Myloplus) is not supported;

(d) Mylesinus and Myleus are paraphyletic, since Tometes sp. is the sister taxon of Mylesinus paraschomburgkii and Mylesinus paucisquamatus is most closely related to other species of Myleus.

Present taxonomic structure as accepted by the U.S. systematists et al. (note the different and much shorter structure from the Géry classification above):

The Tree of Life which is Phylogenetic based is controversial in many ways. It establishes a new order of animals that would not have been accepted by previous 19th and early 20th century authors.

The levels at the generic (binomen) are not subdivided into lower ranking than genus. That is the main difference with the Géry classification method which does not use Phylogenetic methods in placing his fishes and the fact he uses subgeneric ranking.

This cladogram places several vegetarian fishes into the "piranha" clade. Unthinkable during Eigenmann's time.

 

Contributors and Advisers

  1. Fink, William L.

  2. Orti, Guillermo

  3. Petry, Paulo

 

REFERENCES

  1. Nelson, Joseph S. 1994,  Fishes of the World, Third Edition xvii + 600, ISSN: 0-471-54713-1, John Wiley and Sons, New York, NY.

  2. Wayne C. Starnes, Characiformes - Research Curator of Fishes, North Carolina State Museum of Natural Sciences, Research Laboratory, 4301 Reedy Creek Rd., Raleigh, NC, 27607.  

 

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UPDATED: 06/14/2007