Morphological and biometric comparison of the scales of the barbels (Barbus Cuvier) of Spain

Morphological and Biometric Comparison of the Scalesof the Barbels (Barbus Cuvier) of SpainRafael Miranda* and MaCarmen Escala

Department of Zoology and Ecology, University of Navarra, Pamplona, Spain

ABSTRACT The scales of 386 specimens of the eightspecies of the genus Barbus present in Spain (B. bocagei,B. comiza, B. graellsii, B. guiraonis, B. haasi, B. meridi-onalis, B. microcephalus, and B. sclateri) were studied. Sixscales were extracted from each specimen, measurementstaken, and the number of radii noted. Indices were ob-tained from these data. Statistical analysis allowed twogroups to be established: 1) B. meridionalis and B. haasi,living in the upper stretches of rivers and distributed in

the north of Spain; and 2) the remaining species. Withinboth groups a latitudinal gradation in the elongation ofthe scales and in the proportion of radii of the lateral fieldswas observed. J. Morphol. 245:196–205, 2000.© 2000 Wiley-Liss, Inc.

KEY WORDS: Barbus; scales; morphology; lepidometricsmeasurements; radii; Spain

Scales have been used for morphological studies(Cockerel, 1914; Rousset, 1983) but few authorshave used them for taxonomic purposes (Maitland,1972; Morales-Nin and Fauquet, 1983; Elvira, 1988)or as a structural means of identifying the remainsof icthiophagous animals (Suter, 1989; Veldkamp,1995).

Kobayashi (1952) compared the scales of freshwa-ter fish from Japan and claimed that scales were thebest means of studying the affinity between fish, anopinion shared by Bertin (1958) and Rubın (1981),who considered scales as key components in thecurriculum vitae of fish. However, difficulties in theuse of scales for taxonomic and phylogenetic studieshave been observed. This is due to their contact withthe external medium, which can lead to multiplemodifications and to variation in shape, size, andstructures according to the area of the body and ageof the animal (Lippitsch, 1990; Pou and Gallego,1990).

The objective of the present study is to undertakea comparative biometric and morphological study ofthe scales of the different species of barbels presentin Spain with the aim of being able to differentiateand analyze the variations between species.

MATERIALS AND METHODS

In this study 386 specimens of the eight species ofbarbs present in Spain from 13 different localities(Table 1, Fig. 1) were analyzed. Six scales weretaken from each specimen: two from the anteriordorsal area near the beginning of the dorsal fin,another two from the posterior dorsal area, and twofrom the middle area of the lateral line (Fig. 2). Thescales were collected solely from the left side. They

were washed in water, cleaned with a stiff-bristledbrush, and mounted on slides without any type offixation for viewing by binocular magnifying glasses(Elvira, 1988). The scales thus mounted were la-beled to include data on the species, specimen num-ber, sample, and position of the scales on the body.Slides were subsequently stored in alphabetical or-der of the species and by numerical order of thespecimens. Throughout the study we followed, withslight modifications, the method proposed by Pouand Gallego (1990).

Initially, a morphological description of the scalesfrom each species was made. Subsequently, the fol-lowing data were noted (Fig. 3): 1) the number ofradii in the posterior field, anterior field, and in thedorsal and ventral lateral fields; 2) the number ofregenerating scales (for morphological and biometriccomparison these scales were excluded); 3) the num-ber of lobes in the anterior field; and 4) lepidometricmeasurements. These included: ML, maximumlength, the distance between the upper end of theposterior edge and the lower end of the anteriorfield; FD, focal distance, the length between theupper end of the posterior edge and the focus; MW,maximum width, the maximum distance betweenthe end of the upper and lower lateral edge.

The measurements were taken using a GENIUSdigitalizing tablet (HISKETC 1212 model; programIDRISI: Vector Digitalizing Module v. 3.03) and witha binocular magnifying glass with a ZEISS clearcamera to take the points to obtain measurements of

*Correspondence to: R. Miranda, Department of Zoology and Ecol-ogy, University of Navarra, E-31080 Pamplona, Spain.E-mail: [email protected]

JOURNAL OF MORPHOLOGY 245:196–205 (2000)

© 2000 WILEY-LISS, INC.

the scales. The precision obtained was 0.02 mm di-vided among the magnification factor of the lens (forexample, if the scale image projected was 20 timeslarger than the bone, the precision of the measure-ment would be 0.001 mm).

For the description and measurements the scaleswere observed with the posterior field uppermostand the anteriolateral angles symmetrical to thelongitudinal axis of the scale.

On the basis of the preceding data the followingindices were obtained: MW/ML, the relationshipbetween the width and length, the degree of elon-gation of the scale; FD/ML, the position of thefocus relative to the length of the scale; TR/ML,the number of radii of the scale divided by thedistance ML; LF/PF, the number of radii in thelateral fields divided by the number of radii in theposterior field; and AF/PF, the number of radii in

TABLE 1. Species and number of specimens (N° ind.) captured in each sampling point (L.s.p.)

Species N°ind. River Basin Locality Province Date L.s.p.

Barbus bocagei 17 Arlanzon Duero Burgos Burgos 15/11/93 13 Arlanzon Duero Burgos Burgos 20/6/94 18 Arlanza Duero Mst. S. Pedro de Arlanza Burgos 29/9/95 2

B. comiza 11 Guadiana Merida (Emb. de Montijo) Badajoz 16/11/94 320 Guadiana Sanlucar de Guadiana Huelva 19/7/94 4

B. graellsii 45 Arga Ebro Falces Navarra 24/2/92 54 Esca Ebro Burgui Navarra 3/3/92 6

18 Elorz Ebro Otano Navarra 21/10/93 78 Aragon Ebro Caparroso Navarra 2/5/94 85 Araquil Ebro Ibero Navarra 24/2/92 9

B. guiraonis 21 Turia Domeno Valencia 22/6/94 1015 Tuejar Turia Tuejar Valencia 22/6/94 11

B. haasi 11 Esca Ebro Burgui Navarra 3/3/92 640 Esca Ebro Burgui Navarra 21/7/94 6

B. meridionalis 53 Anyet Muga Sant Climent Sescebes Gerona 25/3/94 12B. microcephalus 32 Guadiana Sanlucar de Guadiana Huelva 19/7/94 4B. sclateri 75 Bembezar Guadalquivir Hornachuelos Cordoba 22/7/94 13

Fig. 1. Map of distribution of the barbels in Spain (shaded areas) and sampling points (blackcircles). The numbers of the sampling points correspond to those listed under L.s.p. in Table 1.

197SCALES OF BARBELS OF SPAIN

the anterior field divided by the number of radii inthe posterior field.

Scales from young specimens were discarded, asthese specific individual differences were not ex-pressed. Consequently, the conclusions of this studyare applicable only to adult specimens.

In the comparison of the indices between species,ANOVA (F, parametric) and Kruskall-Wallis (H;nonparametric) and post-hoc multiple range Tukeytest were used. In the case of nonparametric sam-ples, two-to-two comparisons were made using theMann-Whitney test (z score) (Sokal and Rolf, 1969).

RESULTSMorphological Description

Pentagonal- and hexagonal-shaped scales. The fo-cus in these scales was situated below the center ofthe scale. Radii were present in all fields.

The delimitation between the lateral and anteriorfields was clear, in that there was a point of inflec-tion in the curvature of the circuli, while in theseparation of the lateral and posterior fields therewas no change in the curvature. Here the fieldscould be clearly differentiated either by the clear

Fig. 2. Areas where scales were extracted from left side of fish.

Fig. 3. Surface view of the scale areas with diagram of principal fields and lepidometric measurements.

198 R. MIRANDA AND M.C. ESCALA

separation of the radii in the lateral fields fromthose in the posterior field or by the change in thecurvature of the radii.

The circuli were perfectly defined in the anteriorand lateral fields, whereas in the posterior field theyappeared to be further apart and in the adult spec-imens these circuli were deformed and were appar-ent as small, long protuberances. The circuli werearranged parallel to the edges of all the fields. Theannuli appeared clearly marked in the anterior andlateral fields by a greater space between each (alight area) in the summer season and a smallerspace (a dark area) in the winter season (Morales-Nin, 1987) and by the different ordering of the cir-culi from the winter season onwards (Bagliniere andLe Louarn, 1987). The regenerating scales exhibiteda slightly granulated surface scored by numerousdeformed radii forming a network from which theradii extended toward the other fields.

The lateral fields were extensive and triangular.The radii in these fields crossed the surface diago-nally from the anterior field, curving toward thefocus in their origin. The anterior field was rhomboi-dal or triangular in shape with numerous radii. Thenumber of lobes varied from six to two, with three orfour being the most frequent number. The posteriorfield was bell-shaped and was the widest part of thescale. The edge was slightly undulated and rounded.The interior part of the posterior field exhibited aseries of bony structures in a network. These werenot well developed, the result of the disorganizationof the circuli during growth. These bony structuresoccupied a more or less triangular area (Rubın,1981). On the surface of the outer part of the fieldmany dark pigmentations were present, some ofwhich were round and dark, while others were star-shaped and lighter in color.

Species DescriptionsBarbus bocagei Steindachner, 1865 (Fig. 5A).

Scales were hexagonally shaped, a little longer thanwide. The focus was situated in the center of the

scale or a little below. The anterior field was exten-sive and exhibited numerous well-developed radii,many of which were primary. The lobules present inthis field were clearly marked and were smooth inoutline.

Barbus comiza Steindachner, 1865 (Fig. 5B).Scales were hexagonally shaped, generally of equalwidth and length. The focus was situated in thecenter of the scale or a little below. The surface ofthe scale was scored by numerous radii, many ofwhich were ramified, secondary, and/or deformed.Of the scales studied, 31.91% were regenerating.The harsh conditions of the area inhabited by thespecimens studied and the sole presence of adults inthe sample may explain this high percentage(Bereiter-Hahn and Zylberberg, 1993).

The lateral fields were extensive and the edgeswere parallel and slightly convex. The radii in thesefields, as is the case in the anterior field, were poorlydeveloped, generally following a twisting path, andwere very short. The anterior field was extensiveand scored by numerous radii. The lobules wereclearly marked, especially in the lateral zone. Theposterior field was highly pigmented.

Barbus graellsii Steindachner, 1866 (Fig. 5C). Thescales were slightly elongated, although some scalesthat were wider than they were long could also ap-pear. The lateral field exhibited numerous radii onthe surface. The anterior field was scored by manyradii that were ramified, secondary, or twisting. Thecirculi of the posterior field had been deformedthroughout growth, in such a way that in adultspecimens they were faint. The lobules of the edgewere clearly marked, especially in the lateral zone.

Barbus guiraonis Steindachner, 1866 (Fig. 5D).Scales very similar to those of B. graellsii. However,they exhibited a larger number of radii and thecirculi in the adult specimens appeared clearlymarked in the posterior field and less deformed thanthose in the scales of B. graellsii.

Barbus haasi Mertens, 1924 (Fig. 5E). Scales werefairly long hexagonally or pentagonally shaped. Theanterior fields exhibited numerous radii, most ofwhich were primary and straight. The lateral fieldswere elongated, with many radii on the surface. Thelateral lobules of the anterior field were well devel-oped, whereas the central lobules were littlemarked, giving a rectangular aspect to the anteriorarea of the scale.

The circuli of the posterior field had not disap-peared or been deformed during growth and werecontinuous with the circuli of the lateral fields. Somepoorly developed bony structures could be seen inthe inner area of the field.

Barbus meridionalis Risso, 1810 (Fig. 5F). Scaleswere lengthened, hexagonal in shape, and with thefocus situated clearly below the center of the scale.The surface of the scale appeared to be scored bynumerous radii in all its fields. These were uni-formly distributed, a characteristic that Bagliniere

TABLE 2. Multiple range test (Tukey method) for MW/MLindex by species*

Species N

Homogeneous groups(alpha 5 0.05)

1 2 3

B. meridionalis 192 0.667B. haasi 256 0.674B. guiraonis 187 0.756B. graellsii 354 0.769B. sclateri 344 0.824B. microcephalus 191 0.837B. bocagei 140 0.839B. comiza 184 0.844

Significance level 0.995 0.885 0.441

*Indices for homogeneous groups are averages. N, number ofdata.


Fig. 4. Notched box and whisker plots of indices obtained. BB, Barbus bocagei; BC, B. comiza;BG, B. graellsii; BH, B. haasi; BI, B. guiraonis; BM, B. meridionalis; BR, B. microcephalus; BS, B.sclateri.


Figure 5. (Continued)


Fig. 5. Scales of each corporal zone of different species of genus Barbus. A: B. bocagei. B: B.comiza. C: B. graellsii. D: B. guiraonis. E: B. haasi. F: B. meridionalis. G: B. microcephalus. H: B.sclateri.


and Le Louarn (1987) used to distinguish thesescales from those of B. barbus.

The edges of the lateral fields were straight in thescales of the middle area of the lateral line and inthe posterior dorsal area and slightly convex in thescales of the anterior dorsal area. The posterior fieldwas well developed and was clearly larger than therest of the fields.

As was the case with the scales of Barbus haasi,the bony structures of the posterior field were poorlydeveloped. The circuli in this field were not de-formed or faint and were continuous with the circuliof the lateral fields.

Barbus microcephalus Almaca, 1967 (Fig. 5G).Scales were hexagonal, of equal width and length.The focus was situated in or slightly below the cen-ter of the scale. Of the scales studied, 33.3% wereregenerating scales (see B. comiza).

Both the anterior and lateral fields exhibited asmall number of radii, most of which were deformedor short and did not reach to the edge of the scale. Inthe anterior field this edge was slightly undulating.

As in the scales of Barbus comiza, those of B.microcephalus exhibited numerous deformationswith necrosis on the surface of the scale and theposterior field was highly pigmented.

Barbus sclateri Gunther, 1868 (Fig. 5H). Scaleswere elongated, hexagonal in shape, and with thefocus situated below the center of the scale. Fewradii were present on the surface, especially in thelateral fields. The radii in these fields were generallyshort and poorly developed.

Comparative Biometric Study

Study of the MW/ML index (Table 2, Fig. 4) al-lowed three groups to be established (F7,1840 5120,17, P , 0.001): 1) meridionalis group (Barbusmeridionalis and B. haasi) with clearly elongatedscales; 2) B. graellsii and B. guiraonis with lesselongated scales; and 3) the remaining species (B.

bocagei, B. sclateri, B. comiza and B. microcephalus)with scales of equal width and length.

It is worth noting that the elongated scales corre-spond to the two species of mountain barbs thatinhabit stretches of rapid water, while the widerscales correspond to species such as Barbus comizaor B. microcephalus that usually inhabit slowly mov-ing water or reservoirs.

The FD/ML index (Table 3, Fig. 4) clearly distin-guished (F7,1610 5 111,96, P , 0.001) the scales ofBarbus meridionalis, in which the focus was foundto be well below the center of the scale, as a result ofthe well-developed posterior field. In the remainingspecies, two groups could be established: 1) scaleswith the focus in the center (B. microcephalus, B.bocagei, B. comiza, and B. sclateri); 2) scales withthe focus below the center (B. graellsii, B. guiraonis,and B. haasi).

The comparative analysis of the TR/ML index (Ta-ble 4, Fig. 4) separated three groups of greater tolower number of radii (H 5 431.60, P , 0.001): 1)Barbus meridionalis and B. comiza with scales withnumerous radii on the surface (B. comiza with nu-merous radii in the anterior and posterior fields,which were generally short, poorly marked, and didnot reach to the edge and B. meridionalis, withnumerous radii in all fields); 2) B. bocagei, B. haasi,and B. guiraonis; and 3) B. graellsii, B. microceph-alus, and B. sclateri.

The LF/PF index (Table 5, Fig. 4) separated (H 5713.27, P , 0.001) the scales of Barbus haasi (thelateral fields scored by numerous straight radii)from the others, which may, in turn, be separatedinto two groups: B. graellsii and B. meridionalis andthe remaining species. Here the scales exhibit veryfew radii in the lateral fields. Radii were generallypoorly developed.

The scales of Barbus bocagei studied exhibited adeveloped anterior field scored by numerous radiiand the AF/PF index (Table 6, Fig. 4) differentiated(H 5 458.37, P , 0.001) these scales from those ofthe remaining species where, although separationswere also apparent, no clear tendency could be ob-served.

TABLE 3. Multiple range test (Tukey method) for FD/ML indexby species*

Species N

Homogeneous groups (alpha 50.05)

1 2 3 4

B. microcephalus 150 0.615B. bocagei 124 0.616B. comiza 151 0.621B. sclateri 292 0.639B. graellsii 337 0.649 0.649B. guiraonis 158 0.653B. haasi 218 0.661B. meridionalis 188 0.715

Significance level 0.694 0.349 0.092 1.000


TABLE 4. Analysis of averages of TR/ML index by comparisonof two samples (Mann-Whitney test)*

Species N

Homogeneous groups (alpha 5 0.05)

1 2 3 4

B. sclateri 292 10.475B. microcephalus 150 10.549B. graellsii 337 10.592B. haasi 218 11.374B. guiraonis 158 11.436B. bocagei 124 11.590B. comiza 151 13.064B. meridionalis 188 13.820



DISCUSSION

The scales on different parts of the body werecompared separately (Miranda, 1997) and the in-traspecific variability is significant, according toother authors (Lippitsch, 1990; Pou and Gallego,1990). However, in this study the scales of the dif-ferent areas of the body were considered togetherwith the purpose of studying interspecific variabil-ity, independently of the corporal area.

The scales of the mountain barbs are clearlydifferent from those of other Spanish species.Within this group the scales of Barbus meridiona-lis are those that are best differentiated from theother species. This is consistent with the clearphylogenetic separation of these species that Al-maca (1985) includes in the cyclolepis group, andwhich were supposedly differentiated before theformation of the Pyrenees (Almaca, 1990; Doadrio,1990).

The scales of Barbus guiraonis present clear sim-ilarities with those of B. graellsii and, although cer-tain differences exist in the scales studied, thesewere not considered to be sufficiently consistent toallow differentiation of the scales of these two spe-cies.

The scales of Barbus bocagei, B. sclateri, B.comiza, and B. microcephalus present many similar-ities to each other. Within this group, the scalesstudied of B. bocagei can be clearly differentiateddue to the considerable development of the radii inall fields, while the differences between the remain-

ing three species are more questionable. This simi-larity is consistent with the hypothetical paralleldevelopment of the bocagei (B. bocagei and B. scla-teri) and the microcephalus (B. microcephalus andB. comiza) groups in the Iberian Peninsula (Ba-narescu, 1990).

The distribution of the means of the LF/PF index(Table 4) is noteworthy, as the higher values corre-spond to the species from the north of the Peninsulaand the lowest values to those from the south. Theradii are organs of flexibility and function as thearticulations of the scale, giving an idea of the de-gree of activity of the fish (Creaser, 1926; cited inJohal et al., 1984). Therefore, a greater number ofradii in the lateral fields corresponds to the moremobile species (Taylor, 1914), characteristic ofmountain streams (Barbus haasi and B. meridiona-lis) and a lower number corresponds to species suchas B. sclateri, B. microcephalus, and B. comiza thatlive in slowly moving waters or reservoirs (Doadrioet al., 1991).

This north–south variation can also be seen in theMW/ML and FD/ML index and the degree of defor-mation and pigmentation of the posterior field.Thus, the scales of the mountain barbs are the mostelongated, with the focus well below the center of thescale, and have less deformation of the posteriorfield while the scales of the species from the micro-cephalus group are the widest, with centric focus,and exhibit deformation and intense pigmentationof the posterior field.

TABLE 5. Analysis of averages of index LF/PF by comparison of two samples (Mann-Whitney test)*

Species N


1 2 3 4 5 6

B. sclateri 292 0.383B. microcephalus 150 0.404B. comiza 151 0.557 0.557B. bocagei 124 0.624 0.624B. guiraonis 158 0.637B. meridionalis 188 0.674B. graellsii 337 0.704B. haasi 218 1.037

*Indices for homogeneous groups are averages. N, number of data.

TABLE 6. Analysis of averages of index AF/PF by comparison of two samples by Mann-Whitney test*

Species N


1 2 3 4 5 6

B. microcephalus 150 0.597B. meridionalis 188 0.642B. comiza 151 0.745B. sclateri 292 0.822B. graellsii 337 0.849B. guiraonis 158 0.865B. haasi 124 0.892B. bocagei 218 1.046

*Indices for homogeneous groups are averages. N, number of data.


To summarize, the morphological and biometriccomparison of the scales of the barbels of Spain doesnot allow one to differentiate the species. However,scales of mountain barbels are shown to be differentfrom those of the other species, according to its phy-logenetic and taxonomic separation. On the otherhand, the analysis shows a north–south variation inthe scales of the barbels. This kind of analysis doesnot try to be a taxonomic or phylogenetic tool, but itmight be an interesting contribution to these fieldsand to others related to the habitat and behavior ofthese species.

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Morphological and biometric comparison of the scales of the barbels (Barbus Cuvier) of Spain

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