Geometric morphometric analysis of cyclical body shape changes in color
               pattern variants of <i>Cichla temensis</i> Humboldt, 1821 (Perciformes:
               Cichlidae) demonstrates reproductive energy allocation

Reiss, Paul; Grothues, Thomas M.

doi:10.1590/1982-0224-20140030

Paul Reiss

Graduate Program in Ecology and Evolution, 14 College Farm Road, Rutgers University, New Brunswick, New Jersey 08902, EUA. preiss@acuteangling.com (corresponding author)

Thomas M. Grothues

Rutgers University Marine Field Station, 800 c/o 132 Great Bay Blvd., Tuckerton, NJ 08087, EUA. grothues@marine.rutgers.edu

Figures (5)
Tables (2)

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Figure 1.
Configuration of landmarks (red dot within yellow circles) as applied to all color pattern variation (CPV) grades of Cichla temensis. (1) the anterior point of the skull; (2) the anterior edge of the base of the first dorsal spine; (3) the center of the base of the 10th dorsal spine; (4) the base of the first long ray of the second dorsal fin; (5) the base of the 9th ray of the second dorsal fin; (6) the posterior point of termination of the second dorsal fin; (7) the posterior base of the anal fin; (8) the base of the 7th anal fin ray; (9) the anterior insertion of the anal fin; (10) the cloaca; (11) the ventral termination of the scale row emanating from the insertion of the 10th dorsal spine; (12) the ventral termination of the scale row emanating from the anterior point of the insertion of the pelvic fin.

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Figure 2.
Thin-plate-spline deformation grids depicting (a) overall body shape differences between extremes of CPV grades (magnified 3x); (b) the increase in area at the bases of the second dorsal and anal fins between extremes of CPV grades (magnified 3x).

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Figure 3.
Biplot of the uniform components in each direction (UniX and UniY) of morphometrical differences in 80 specimens of Cichla temensis in 4 color variation patterns (CPV) as measured by 9 Thin Plate Spline (TPS) distortion variables (V1-V9). Colored numbers indicate the CPV grade of individuals. The total spread of scores among individuals of each CPV are indicated by an envelope (solid line polygon) calculated as the minimum convex hull for that group. Position in the plot relative to other individuals indicates the degree of similarity in morph. Vectors point in the direction of gradient change for that TPS variable and the magnitude indicates the strength of the gradient. Angles between vectors indicate the TPS interset correlations.

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Figure 4.
Height to length ratio of color pattern variants of Cichla temensis. The average height/SL ratio increased with CPV grade from a low of 0.2694 for CPV Grade 1 (paca), 0.2725 for CPV Grade 2, 0.2771 for CPV Grade 3 and 0.2822 for CPV Grade 4 (açu).

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Figure 5.
Relative mean GSI vs. relative mean HSI of color pattern variants of Cichla temensis. Points for GSI represent the mean value for each CPV grade as compared to the range encountered. Points for HSI represent the mean value for each CPV grade compared to the range encountered.

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Table 1.
Guide to color pattern variation (CPV) grades and local names of Cichla temensis.

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Table 2.
Relative Warp Analysis of Cichla temensis. Singular values (SV) and percent explained for each relative warp (principle axis). The first relative warp was significantly different from random (v2 1/4 7.6539, 2.1387; P < 0.05), indicating that much of the biologically significant individual body shape differences can be explained on this component.

CPV grade	Grade 1	Grade 2	Grade 3	Grade 4
Common name	paca	n/a (called paca)	n/a (called açu)	açu
Description	Cryptically patterned, dark grey/brown background, prominent series of light spots in four loosely delimited horizontal rows. Total absence to trace of gold/brass on ventral edge of operculum and torso. Dorsal fins dark with white spots, recapitulating body pattern. Ventral fins dark brown to dark red. Postorbital markings and vertical bars 1, 2 and 3 almost invisible against dark background.	Significant gold/brass color radiating from lower edges of operculum extends to torso, encompassing pelvic fins and ventral lateral region anterior to vertical bar 2. Dorsal and posterior torso dark grey/brown. Dorsal fins dark, ventral lower fins become red. Black vertical bars 1, 2 and 3 equal in prominence to background of white spots. Postorbital markings emerge from background.	Torso dominated by gold/brass color below lateral line. Dorsal fins remain dark while ventral fins become a lighter, brighter red. Black vertical bars become visibly dominant on torso. Light lateral spots remain visible, but contrast against torso diminished. Postorbital markings now clearly visible.	Brightly colored 3-barred variant. Torso bright yellow -gold color except for dorsal extremes. Dorsal fins and upper portion of caudal fin blue ranging to bright, electric blue. Ventral fins bright red to orange. Vertical black bars 1, 2 and 3 sharply demarcated, dominating pattern. Light spots mostly absent or trace. Postorbital markings sharp and distinct.

Relative warp (principle axis)	Singular value (SV)	% explained	Cumulative %
1	0.17896	20.59%	20.59%
2	0.17634	20.00%	40.59%
3	0.15128	14.72%	55.31%
4	0.12616	10.24%	65.54%

Geometric morphometric analysis of cyclical body shape changes in color pattern variants of Cichla temensis Humboldt, 1821 (Perciformes: Cichlidae) demonstrates reproductive energy allocation