Dealing with allometry in linear and geometric morphometrics: a taxonomic case study in the Leporinus cylindriformis group (Characiformes: Anostomidae) with description of a new species from Suriname

To achieve maximum efficacy, taxonomic studies that seek to distinguish amongst species must first account for allometric shape variation within species. Two recently developed software packages (SMATR and MorphoJ) offer regression-based allometric approaches that are notable for their statistical power and ease of use and that may prove highly useful to taxonomists working with linear or geometric morphometric data. We investigate species delimitation of the slender-bodied fishes in the Leporinus cylindriformis group using these programs and demonstrate the utility of the allometric corrections that they provide. Without allometric correction, many pairs of species are difficult to distinguish on the basis of morphometrics, but once regressions are used to account for marked allometric variation within species, most of the recognized species in this group can be readily distinguished with linear or geometric morphometrics, particularly using variation in the depth of the body. Both approaches returned congruent patterns of separation amongst putative species, but the geometric approach in MorphoJ distinguished amongst four more pairs of species than did the linear approach in SMATR and appears to provide slightly more statistical power. Based on distinctive morphometrics, meristics, and coloration, a highly elongate species of Leporinus from the Suriname, Corantijn, and Coppename rivers of Suriname is described herein as a new species, Leporinus apollo sp. nov. The unique L. cylindriformis holotype from Porto de Moz, Brazil differs in morphology, meristics, and pigmentation from specimens commonly referred to that species from the main basin of the Amazon; the latter specimens may represent an additional undescribed species. The L. cylindriformis holotype itself may represent a rare species or a specimen collected at the edge of its native range. Measurements of the holotype and paratype of Leporinus niceforoi, which were collected in the Amazonian slope of Colombia, differ substantially from similarly pigmented and putatively conspecific specimens from Amazonian portions of Ecuador and Peru. Recently collected specimens from Colombia are needed to determine whether the observed morphometric variation encompassed by the current concept of L. niceforoi indicates a morphocline within a single species, suggests the presence of multiple cryptic species, or results from shrinkage of the types. In all these cases, linear or geometric morphometric data can reliably differentiate amongst species, but only after one accounts for allometric shape variation. The new SMATR and MorphoJ software packages both offer easy and effective approaches to such allometrically informed taxonomy, and may prove useful to any systematist working on taxa that change shape as they grow. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162, 103–130.

[1]  W. Taylor,et al.  Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study , 1985 .

[2]  J. Stevens Applied Multivariate Statistics for the Social Sciences , 1986 .

[3]  F. Rohlf,et al.  Extensions of the Procrustes Method for the Optimal Superimposition of Landmarks , 1990 .

[4]  Rudolph J. Miller Comparative Morphology of Three Cyprinid Fishes: Notropis cornutus, Notropis rubellus, and the Hybrid, Notropis cornutus X Notropis rubellus , 1963 .

[5]  R. Winterbottom Systematics, osteology and phylogenetic relationships of fishes of the Ostariophysan subfamily Anostominae (Characoidei, Anostomidae) , 1980 .

[6]  W. Fink,et al.  The so-called cheirodontin fishes of Central America with descriptions of two new species (Pisces: Characidae) , 1974 .

[7]  A. Finstad,et al.  To what extent does ethanol and freezing preservation cause shrinkage of juvenile Atlantic salmon and European minnow , 2007 .

[8]  S. Gould Geometric Similarity in Allometric Growth: A Contribution to the Problem of Scaling in the Evolution of Size , 1971, The American Naturalist.

[9]  Miriam Leah Zelditch,et al.  Shape Analysis and Taxonomic Status of Pygocentrus Piranhas (Ostariophysi, Characiformes) from the Paraguay and Paraná River Basins of South America@@@Shape Analysis and Taxonomic Status of Pygocentrus Piranhas (Ostariophysi, Characiformes) from the Paraguay and Parana River Basins of South America , 1997 .

[10]  K. Mardia,et al.  Statistical Shape Analysis , 1998 .

[11]  T. P. Burnaby Growth-Invariant Discriminant Functions and Generalized Distances , 1966 .

[12]  Tomas Hrbek,et al.  Getting into Shape: An Empirical Comparison of Traditional Truss-Based Morphometric Methods with a Newer Geometric Method Applied to New World Cichlids , 2003, Environmental Biology of Fishes.

[13]  F. Rohlf,et al.  Morphometric Spaces, Shape Components and the Effects of Linear Transformations , 1996 .

[14]  S. Weinberg,et al.  Canonical Analysis when Number of Variables is Large Relative to Sample Size , 1976 .

[15]  C. Kimmel,et al.  Linked morphological changes during palate evolution in early tetrapods , 2009, Journal of anatomy.

[16]  Fishes of the Great Lakes Region. , 1959 .

[17]  G. Turner,et al.  Identification of the Lake Malawi Oreochromis (Nyasalapia) spp. using multivariate morphometric techniques , 1989 .

[18]  J. Huxley Problems of relative growth , 1932 .

[19]  R. Vari,et al.  Phylogenetic relationships within the South American fish family Anostomidae (Teleostei, Ostariophysi, Characiformes) , 2008 .

[20]  Matthew T. Wilson,et al.  Shrinkage correction and length conversion equations for Theragra chalcogramma, Mallotus villosus and Thaleichthys pacificus , 2005 .

[21]  F. Rohlf,et al.  A revolution morphometrics. , 1993, Trends in ecology & evolution.

[22]  H. Hotelling The most predictable criterion. , 1935 .

[23]  C. Sturmbauer,et al.  Assessment of traditional versus geometric morphometrics for discriminating populations of the Tropheus moorii species complex (Teleostei: Cichlidae), a Lake Tanganyika model for allopatric speciation , 2008 .

[24]  Pierre Jolicoeur,et al.  The multivariate generalization of the allometry equation , 1963 .

[25]  R. Gonzalez Applied Multivariate Statistics for the Social Sciences , 2003 .

[26]  Gerald R. Smith,et al.  Fishes of the Great Lakes Region, Revised Edition , 2004 .

[27]  D. Schluter,et al.  The Analysis of Biological Data , 2008 .

[28]  W. Rice ANALYZING TABLES OF STATISTICAL TESTS , 1989, Evolution; international journal of organic evolution.

[29]  C. C. Fernandes,et al.  Largest of All Electric-Fish Snouts: Hypermorphic Facial Growth in Male Apteronotus hasemani and the Identity of Apteronotus anas (Gymnotiformes: Apteronotidae) , 2002, Copeia.

[30]  Chuanmin Hu,et al.  The dispersal of the Amazon and Orinoco River water in the tropical Atlantic and Caribbean Sea: Obse , 2004 .

[31]  A. Günther Catalogue of the fishes in the British museum. Volume 3, volume 3 / by Dr Albert Günter , 2009 .

[32]  P. Bail,et al.  Atlas des poissons d'eau douce de Guyane , 1997 .

[33]  P. Rincón,et al.  Big fish, small fish: still the same species. Lack of morphometric evidence of the existence of two sturgeon species in the Guadalquivir river , 2000 .

[34]  A. Leviton,et al.  Standards in herpetology and ichthyology : Part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology , 1985 .

[35]  C. Nunn,et al.  Allometric Slopes and Independent Contrasts: A Comparative Test of Kleiber’s Law in Primate Ranging Patterns , 2000, The American Naturalist.

[36]  J. Mol,et al.  The fish fauna of Brokopondo Reservoir, Suriname, during 40 years of impoundment , 2007 .

[37]  F. Muller‐Karger,et al.  The dispersal of the Amazon's water , 1988, Nature.

[38]  Ø. Hammer,et al.  PAST: PALEONTOLOGICAL STATISTICAL SOFTWARE PACKAGE FOR EDUCATION AND DATA ANALYSIS , 2001 .

[39]  W. Fink Revision of the Piranha genus Pygocentrus (Teleostei, Characiformes) , 1993 .

[40]  F. Rohlf,et al.  Geometric morphometrics: Ten years of progress following the ‘revolution’ , 2004 .

[41]  G. L. Buffon,et al.  Histoire Naturelle des Poissons , 2009 .

[42]  Geographic and environmental variation in Bryconops sp. cf. melanurus (Ostariophysi: Characidae) from the Brazilian Pantanal , 2006, Ichthyological Research.

[43]  Bryconops magoi and Bryconops collettei (Characiformes: Characidae), two new freshwater fish species from Venezuela, with comments on B. caudomaculatus (Günther) , 2005 .

[44]  Fred L. Bookstein,et al.  Morphometric Tools for Landmark Data. , 1998 .

[45]  M. Westoby,et al.  Bivariate line‐fitting methods for allometry , 2006, Biological reviews of the Cambridge Philosophical Society.

[46]  Geraldo Mendes dos Santos Aspectos de sistemática e morfologia de Schizodon fasciatus Agassiz, 1829, Rhytiodus microlepis Kner, 1859 e Rhytiodus argenteofuscus Kner, 1829 (Osteichthyes, Characoidei, Anostomidae) do lago Janauacá - Amazonas () , 1980 .

[47]  G. Cuvier Les reptiles, les poissons, les mollusques et les annélides , 1969 .

[48]  Robert R. Miller,et al.  Systematic Position of the Lake Trout, Salvelinus namaycush , 1954 .

[49]  A. Gunther Catalogue of the physostomi : containing the families siluridæ, characinidæ, haplochitonidæ, sternoptychidæ, scopelidæ, stomiatidæ, in the collection of the British Museum , 1981 .

[50]  Fred L. Bookstein,et al.  MULTIVARIATE DISCRIMINATION BY SHAPE IN RELATION TO SIZE , 1981 .

[51]  D. Penny The comparative method in evolutionary biology , 1992 .

[52]  C. Hubbs,et al.  A revision of the black basses (Micropterus and Huro) with descriptions of four new forms , 1940 .

[53]  J. Marr The Use of Morphometric Data in Systematic, Racial and Relative Growth Studies in Fishes' , 1955 .

[54]  Scott Roberts,et al.  The fish , 1998, SIGGRAPH '98.

[55]  F. Bookstein,et al.  Morphometrics in Evolutionary Biology. , 1986 .

[56]  M. Zelditch,et al.  Phylogenetic Analysis of Ontogenetic Shape Transformations: A Reassessment of the Piranha Genus Pygocentrus (Teleostei) , 1995 .

[57]  J. Birch,et al.  Comparing Wing Shape of Bats: The Merits of Principal-Components Analysis and Relative-Warp Analysis , 1997 .