UNDER CONSTRUCTION - SUBJECT TO CHANGE
Serrasalmidae
Serrulatus Type Group/Humeralis Group
1/1/2012
By Frank Magallanes
DISCUSSION
Since the creation of the genera Serrasalmus and Pristobrycon there have been numerous historical taxonomic problems with the species placed within it. Unlike Pygocentrus and Pygopristis where the number of species appears to be fixed and set as of 2011.
Presently there are 28 described species of Serrasalmus, many of which are morphologically similar. Historical authorities originally placed the piranhas as a group in genus Pygocentrus. Then Norman 1929, lumped them all (except Pygopristis denticulata) in genus Serrasalmus. Jacques Géry, the French Ichthyologist further delineated the piranhas into sub-generic ranking. In so doing, Géry recognized only 12 species of piranha in the entire continent of South America (Machado-Allison et al., 1996).
Some species collected from regions are assigned a scientific name without closer examination of the type. This can further present problems on what is actually being collected and catalogued. A few examples are shown below.
Pristobrycon aureus
Historical authorities placed Serrasalmo aureus as a synonym to S. eigenmanni, but today Pristobrycon aureus is considered a valid species under genus Pristobrycon. Pristobrycon aureus (Spix & Agassiz 1829), though not a member of the humeralis group, also fits closely in appearance to this group. However, there are no types known of this species, only a plate image (see below). It was originally proposed as a Serrasalmo and found in Lower portions of tributaries of lower Amazon River and Guyana rivers: Brazil and Guyana. Today Serrasalmo aureus is valid as Pristobrycon aureus (Spix & Agassiz 1829) (Jégu in Reis et al. 2003).
Pristobrycon calmoni
A few of the species placed within Serrasalmus and Pristobrycon are questionable because no types exist. In the case of Pristobrycon, this genus was erected by Carl Eigenmann 1915 and he used Pristobrycon calmoni as its type. The problem centers on the placement of P. striolatus which does not have ectopterygoid (palatine teeth) or a preanal spine in this genus. It remains unknown why Eigenmann placed this species in the genus. P. careospinus and P. maculipinnis also present problems because like striolatus, these two also lack these key features. The other members of genus Pristobrycon have the key features required of the genus. Freeman and colleagues (2007) have suggested creating a new genus with P. striolatus as the type. So far, this has not gone further than a suggestion. DNA is still in its infancy when it comes to piranhas and it might be years before more is known and decided by future authorities should a new genus be erected as suggested. The rest of the problem seems to be lack of better identifications and specimens.
Pristobrycon striolatus
Serrasalmus scapularis (Günther, A.. 1864) was described from the Essequibo River, Guyana. Géry, 1972 placed it as a synonym of Serrasalmus serrulatus (Valenciennes 1849), but a valid subspecies. This error in study created further taxonomic problems. S. scapularis is presently a synonym to Pristobrycon striolatus. Pristobrycon striolatus is found Amazon and Orinoco River basins and northern and eastern Guiana Shield rivers: Brazil, French Guiana, Guyana, Suriname and Venezuela.
Serrasalmus eigenmanni
Described below are several species reviewed along with their distribution. Most overlap their respective ranges. All these species present morphological similarities:
Serrasalmus eigenmanni (Norman, J. R.,1929), no types exist and is considered unique. These species has been bounced back and forth between Pristobrycon (Jégu & Santos 2001) and a Serrasalmus (Géry 1972, Machado-Allison 2002, Jégu in Reis et al. 2003, and Machado-Allison et al. 2009). Today this species is valid as Serrasalmus eigenmanni Norman 1929. Distributed over the Amazon River basin and northern and eastern Guiana Shield rivers: Brazil, French Guiana, Guyana, Suriname and Venezuela.
The species is difficult to identify in the field. Samples of S. eigenmanni resembles both S. striolatus and S. hollandi in preservative. However, neither species is found in Peru to my knowledge. Jegu, 1991 suggested that S. eigenmanni could qualify as a twin and synonym to S. hollandi ( 5 & 6, below).
Serrasalmus hollandi
Serrasalmus hollandi, Eigenmann, C. H.. 1915 was described from the Rio Guaporé at Maciél, Brazil. The Holotype is unique (FMNH 56978/ex CM5792). This species presents taxonomic problems on its identity. Jégu & Santos 2001 placed it valid as Serrasalmus hollandi Eigenmann 1915, Lasso et al., 2001, Jegu in Reis et al., 2003, Machado-Allison et al., 2009, Hubert & Renno 2010 concurred. The distribution of the species questionable in its range. Presently it is found in the Madeira River basin and ? Guyana rivers: Brazil and Guiana (?).
Holotype, Serrasalmus hollandi 6 |
S. hollandi? juvenile Rio San Martin, Bolivia 7 |
S. hollandi? adult collected Rio San Martin, Bolivia 8 |
Serrasalmus humeralis
S.
humeralis (Valenciennes, A., in Cuvier & Valenciennes 1850) is
problematic since its type is unique (a radiograph image). The image below for
example are specimens catalogued (CAS 71217) as Serrasalmus humeralis.
Previous historical authorities have placed different synonym's under this
species. Most of which either had a tail band or no humeral blemish. The only
known image of S. humeralis is a colored plate drawing by Castelnau 1850
(see 9 below). According to current research, this species is found; Amazonas,
Brazil (evidently in error for Araguay River, Tocantins basin). Gery, 1972
considered it valid as Serrasalmus humeralis. Ortega & Vari 1986,
Géry et al. 1991, Planquette et al. 1996, Jégu in Reis et al. 2003 all concur
it is valid as Serrasalmus humeralis Valenciennes 1850. Distribution
range is; Amazon River basin: Bolivia, Brazil and Peru. As a personal
note; I have only found images purported to be S. humeralis but
incredibly similar to Serrasalmus manueli (Fernández-Yépez, A., &
Ramírez, M. V.,1967). Included in my discovery are a couple photos identified
by Nicholas Hubert, IRD France, as S. humeralis VIEW. As can be seen, no humeral
blemish. More on this interesting species and its problems can be read here. In Zoological record, Volume 101, Issue
14, Part 3 - Issue 20, Part 3, page 83... The key to
subgenera, S. (
In different occasions, we were able to examine topotypes of Serrasalmus eigenmanni Norman (Guyana), the type of S. humeralis and exemplary topotypes of the Araguia, and many examplary Amazonia of the type, in particular basin of the Rio Madeira and Xingu. We were not able to find any differentiating character allowing separating eigenmanni from humeralis, defined by the type of the Araguaia (well figured by Castelnau, that does not show no marginal band to the tail one) and by the samples collected since in the Araguay (cf. Géry, 1979 pls. VII and VIII). The synonym of S. eigenmanni is therefore more probable. During a recent expedition (7.1989) in the Arataye, flow Approuague, and after the compilation of this work, MM. Boujard, Meunier and Easter discovered a sympatric form of S. humeralis which one could qualify as a twin one if it were not recognizable (in the field only) to its different flows. Examined in the laboratory, the three harvested specimens revealed themselves effectively very similar by the morphometric characters, but they seemed a little more lengthened and could be placed, by way of hypothesis, to S. hollandi. It is necessary to await a more abundant collection and specific knowledge of the live coloration thanks in part to. Also, this form is not fully understood in the account of the types of the Oyapock and Approuague.
N.B. - Serrasalmus rhombeus was not collected or in the Oyapock or in the Approuague: Its ecological niche maybe occupied by S. humeralis. Variance of the same order were noted in certain coastal rivers of Guyana (to see conclusion), and we inspire to go back over the description and the geographic distribution of these types in a future work (Jegu, 1991).
Castelnau Plate Image, Serrasalmus humeralis 9 |
Serrasalmus humeralis Holotype image MNHN A-9735 EDUCATIONAL USE ONLY |
Serrasalmus humeralis (CAS 71217) 10 |
Re-description of S. humeralis, Géry, 1963 11 |
Serrasalmus nalseni
Serrasalmus nalseni, Fernández-Yépez, A.,1969 is described from the Río Uracoa, 9°10'N, 62°27'W, Estado Monagas, Venezuela. The original Holotype (MACLPI 68697) is lost. However a neotype has been erected (MBUCV-V-35389). The species is valid and a new re-description was completed (Machado-Allison et al. 2009).
It is currently valid as Serrasalmus nalseni Fernández-Yépez 1969. Its distribution is Orinoco River basin, Venezuela.
Serrasalmus serrulatus
S. serrulatus geographical range maybe synonymous with any one of the species discussed above or even a totally different species. In the present scientific literature S. serrulatus morphology is similar to it's congeners; S. eigenmanni, S. hollandi, S. humeralis and S. nalseni.
Serrasalmus serrulatus, has historically been difficult to delineate within populations and to positively identify it. The problem rests with the growth pattern which is between a Serrasalmus and a Pristobrycon. S. gouldingi also slightly appears to have the same issues of growth pattern. The species gouldingi is more Pristobrycon-like. And there are questions on what its relationship is to S. manueli.
Valenciennes (A.) in Cuvier & Valenciennes 1850 originally placed serrulatus as a Pygopristis. In Fowler (1911) Proceedings of the Academy of Natural Sciences of Philadelphia, Volume 63 (page 428-432) mentions problems distinguishing it from Serrasalmo coccogenis ( = Pristobrycon calmoni ). The authority thought it... may be identical, though the writer does not mention any dark blotches on the side of the body, and yet he does say there are some traces of the dark shoulder-spot. Géry 1972:221, Ortega & Vari 1986:9, Jégu in Reis et al. 2003:192, López et al. 2003:32, Menni 2004:78) revised and placed serrulatus in a different genus (Serrasalmus). Presently, Serrasalmus serrulatus is a valid species with its distribution questionable in some localities; Amazon River basin and ? Essequibo River basin: Amazon (?), Brazil, Guyana (?) and Peru; Argentina.
During rest or stressful periods (such as capture) Serrasalmus serrulatus takes on additional markings on the fin, namely the caudal fin which can show a faint caudal band on the edge. During its growth the shape of the body changes from Serrasalmus-like to Pristobrycon-like along with its body markings. These changes are what has confused authorities in the past in naming piranha fish from South American rivers. As one can ponder, all these species overlap each others range and appear similar.
Interestingly enough, the humeral blemish is not fixed on any of these species.
Note the changes in the body form and markings:
All 3 same fish showing growth. Collected in Rio Nanay, Peru
Photos by David M. Schleser, Nature's Images, Inc.
Caught - Serrasalmus shape. Approximately 2.50 SL. Note "V" of basal. 27 |
3 inches Body spotting pattern is modified from original 28 |
5 inches - Pristobrycon type body Note faint terminal band on caudal fin - stressed. 29 |
Serrasalmus species depicting growth changes and coloration. Aquarium specimen
Photos below by Steve Bennet, United Kingdom
Another Serrasalmus species, photos by Steve Bennet, United Kingdom
REFERENCES
Eigenmann, C. H. 1915 (Mar.) The Serrasalminae and Mylinae. Ann. Carnegie Mus. v. 9 (nos. 3-4): 226-272, Pls. 44-58.
Géry, J. 1972 (19 Dec.) Poissons characoďdes des Guyanes. I. Généralités. II. Famille des Serrasalmidae. Zool. Verh. (Leiden) No. 122: 1-250, Pls. 1-16.
Géry, J. 1976 (18 Mar.) Les genres de Serrasalmidae (Pisces, Characoidei). Bull. Zool. Mus. Univ. Amst. v. 5 (no. 6): 47-54.
Géry, J. 1977 Characoids of the world. T. F. H. Publications, Neptune City, New Jersey. Characoids World: 1-672.
MACHADO-ALLISON, Antonio and FINK, William., 1996 - Los Peces Caribes de Venezuela, Diagnosis, Claves, Aspectos Ecologicos Y Evolutivos, pg. 29-31.
FINK, William and MACHADO-ALLISON, Antonio., Three new species of piranhas from Brazil and Venezuela. Ichthyological Explorations of Freshwaters, 3(1):55-71. 1992.
Taphorn, D. C. 1992 The characiform fishes of the Apure River drainage, Venezuela. Biollania Edición Especial - No. 4. Monografias Cientificas del Museo de Ciencias Naturales, UNELLEZ -- Guanara, estado Portuguesa, Venezuela. 1-537.
Freeman, B., Nico, L. G., Ostentoski, M. Jelks, H. J. & Collins, T. M. 2007. Molecular systemics of Serrasalmidae: Deciphering the identities of piranha species and unraveling their evolutionary histories. Zootaxa 1484: 1 -38
California Academy of Sciences. On-line: http://research.calacademy.org