Corresponding author: Oliver M. Selz (
Academic editor: M. E. Bichuette
The alpha taxonomy of the endemic whitefish of lakes Brienz and Thun, Switzerland, is revised. We evaluate the status of seven known species:
Selz OM, Dönz CJ, Vonlanthen P, Seehausen O (2020) A taxonomic revision of the whitefish of lakes Brienz and Thun, Switzerland, with descriptions of four new species (Teleostei, Coregonidae). ZooKeys 989: 79–162.
The European whitefish (
Here, we revise the whitefish species of lakes Brienz and Thun, Switzerland. Whitefish can be found in the large pre-alpine lakes of France, Germany, Austria, and Switzerland, which historically harboured approximately 50 different species native to approximately 30 lakes in three major river drainages: the Rhine, the Danube, and the Rhone (
In this paper we compile and review morphological, genetic and ecological data for seven species of whitefish from the connected lakes Thun and Brienz, three of which are found in both lakes. Three of the species were previously described as
One of the previously described species,
One of the newly described species,
We studied the type material designated by
Type material of all currently valid species (based on
The different whitefish species in this study derive from different lakes, namely Lake Thun (
Contemporary samples of whole specimens from lakes Thun and Brienz were collected in the course of many projects of the Seehausen research group (Eawag and the University of Bern). Contemporary material (whole specimens and tissue samples) used here was collected in the years 2005, 2011, 2013, 2014 and
Sampling details for the fish collected in the years 2005, 2011, 2013, and 2014 can be found in the corresponding publications (
In the field the fish were identified to species level as good as possible. Sex, fresh mass (to the nearest 0.1g), ripeness (4 = not ripe; 5 = partially ripe, i.e., slow flow of egg sand sperm when stripped; 6 = ripe, i.e. eggs and sperm flow easily when stripped) and the presence of tubercles on the scales (modified from
The age of the specimens that were used in this study was determined in the lab by counting the annual growth rings of four scales under a confocal microscope following
Morphological measurements and counts on the old type material (N = 31) and on contemporary specimens (N = 340) were taken of 25 body, 19 head, and 4 gill characters with adigital calliper to the nearest 0.1 mm. Twelve meristic characters were counted. The measurements and counts were taken on the left body side of the fish, unless a specific character was missing or deformed, in which case that character was measured or counted on the right side of the fish. The mean of two measurements were taken for each character, whereby the difference between two measurements had to be less than 5%. If agreement was less good, the distance was measured again two times. The average inaccuracy between two measurements taken over all morphological characters was 1.4%. Not all measurements could be taken for several specimens since characters where damaged or absent, and we thus sometimes report incomplete character lists for certain specimens. This results in varying sample sizes for each character. All characters for which we had missing values were not retained in the multivariate ratio analyses (see below). The number of characters used for each analysis is explicitly mentioned in the results section. Most of the morphological and meristic characters follow
Morphological characters, their acronyms and a brief description of each character.
Morphological characters | Acronym | Description |
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Pelvic fin base | PelvFB | Length between insertions of fin |
Pelvic fin "spine" length | PelvFS | Length from upper insertion point of fin to tip of spine; the spine is actually an elongated scale structure |
Pelvic fin length | PelvF | Length from upper insertion point of fin to tip of longest branched ray |
Pectoral fin base | PecFB | Length between insertions of fin |
Pectoral fin 1 length | PecF1 | Length from upper insertion point of fin to tip of unbranched ray |
Pectoral fin 2 length | PecF2 | Length from upper insertion point of fin to tip of longest branched ray |
Dorsal fin base | DFB | Length between insertions of fin |
Length of anterior part of dorsal fin erected | DFAe | Length from anterior insertion point of fin to tip of longest unbranched ray, when fin is fully erected |
Length of anterior part of dorsal fin depressed | DFAd | Length from anterior insertion point of fin to tip of longest unbranched ray, when fin is depressed |
Length of posterior part of dorsal fin erected | DFPe | Length from posterior insertion point of fin to tip of most posterior branched ray, when fin is erected |
Anal fin base | AFB | Length between insertions of fin |
Length of anterior part of the anal fin | AFAe | Length from anterior insertion point of fin to tip of longest branched ray, when fin is fully erected |
Adipose fin base | AdFB | Length between insertions of fin |
Caudal fin length | CF | Length from the middle of hypural plate of the caudal fin (internally this is the expanded bones at the end of the backbone that support the caudal fin, externally where the lateral line scales end) to the tip of the longest unbranched ray either being on the dorsal or ventral part of the caudal fin |
Caudal peduncle depth | CD | Vertical distance between dorsal and ventral margins of the caudal peduncle at its narrowest part |
Caudal peduncle length | CL | Length from posterior insertion point of anal fin to the middle of the hypural plate of the caudal fin |
Length from anterior part of adipose fin to caudal fin base | PAdC | Length from anterior insertion point of adipose fin to the middle of the hypural plate of the caudal fin |
Dorsal head length | DHL | Length from tip of snout to most posterior part of the frontal head bone |
Prepelvic length | PreP | Length from tip of snout to anterior insertion point of pelvic fin |
Preanal length | PreA | Length from tip of snout to anterior insertion point of anal fin |
Standard length | SL | Length from tip of snout to the middle of the hypural plate of the caudal fin |
Total length | TL | Length from tip of snout to the tip of longest unbranched ray either being on the dorsal or ventral part of the caudal fin |
Predorsal length | PreD | Length from tip of snout to anterior insertion point of dorsal fin |
Body depth | BD | Vertical distance between dorsal and ventral margins of body from anterior insertion point of dorsal fin to anterior insertion of pelvic fin: not necessarily the greatest body depth |
Postdorsal length | PostD | Length from posterior insertion point of dorsal fin to middle of hypural plate of the caudal fin |
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Eye diameter | ED | Horizontal distance across the midline of the eye from the anterior to the posterior margin of the soft eye tissue |
Eye cavity | EC | Horizontal distance across the midline of the eye from the anterior margin of the eye socket to the posterior margin of the eye cavity |
Eye height | EH | Vertical distance across the midline of the eye from the dorsal margin of the eye cavity to the ventral margin of the eye cavity |
Eye socket | ES | Horizontal distance from the anterior margin of the eye socket to the most anterior point of the the posterior margin of the eye socket |
Postorbital length | PostO | Length from posterior margin of the eye to the most posterior point of the operculum |
Head length | HL | Length from the tip of snout to most posterior point of the operculum margin |
Head depth | HD | The transverse distance between margins at the widest point of the head. |
Head width | HW | Distance between the posterior margins of the left and right operculum |
Mouth width | MW | The transverse distance between margins of the upper and lower jaw |
Upper jaw length | UJ | Length from the tip of the snout to most posterior point of the upper jaw |
Lower jaw length | LJ | Length from the most anterior point of the lower jaw to the lower jaw insertion |
Lower jaw width | LJW | Length between the anterior left and right side of the lower jaw |
Uperr jaw width | UJW | Length between the posterior left and right point of the upper jaw |
Length of maxilla | M | Length from the most anterior point of the maxilla to the most posterior point of the maxilla |
Snout length | SN | Length from tip of snout to anterior margin of the eye |
Snouth depth | SD | Vertical distance from the upper to the lower margin of the rostral plate |
Snouth width | SW | Horizontal distance from the left to the right margin of the rostral plate |
Interorbital width | IOW | Distance between the anterior margin of the left and right eye cavity |
Internarial width | INW | Distance between the right and left nostrils |
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Upper arch length | UA | Length of the first hypobranchial (upper arch) from the most anterior point to the joint of the hypo- and ceratobranchial where the middle raker emerges |
Lower arch length | LA | Length of the first ceratobranchial (lower arch) from the most anterior point to the joint of the hypo- and ceratobranchial where the middle raker emerges |
Middle gill raker length | MGR | Length of the gill raker directly at the joint of the the upper and lower first arch, from the insertion of the gill raker to the tip of the gill raker |
Longest gill raker length | LGR | Length of the longest gill raker either on the upper and lower first arch, from the insertion of the gill raker to the tip of the gill raker |
Meristic characters, their acronyms and a brief description of each character.
Mersitic characters | Acronym | Description |
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Pelvic fin rays | PelvFR | Number of unbranched and branched rays |
Pectoral fin rays | PecFR | Number of unbranched and branched rays |
Dorsal fin rays | DFR | Number of unbranched and branched rays; the posteriormost dorsal rays are often borne from a single pterygiophore (the bones on which the rays articulate), in such a case the two rays are acounted as 2 rays, rudimentary unbranched rays in front of the fin are counted |
Anal fin rays | AFR | Number of unbranched and branched rays; the posteriormost anal rays are often borne from a single pterygiophore (the bones on which the rays articulate), in such a case the two rays are acounted as 2 rays, rudimentary unbranched rays in front of the fin are counted |
Lateral line scales | LS | Scales bearing the lateral-line column canal from the head to the end of the hybpural plate of the caudal peduncle |
Predorsal scales | PDS | Dorsal scales starting from the posterior end of the head to the anterior insertion of the dorsal fin |
Transverse dorsal scales | TDS | "Number of scale rows between anterior insertion of the dorsal fin and the lateral line, not accounting for the lateral line scale and the scale on the dorsal midline (in front of the dorsal fin) " |
Transverse anal scales | TAS | "Number of scale rows between anterior insertion of the anal fin and the lateral line, not accounting for the lateral line scale and the scale on the ventral midline (in front of the anal fin) " |
Transverse pelvic scales | TPS | "Number of scale rows between anterior insertion of the pelvic fin and the lateral line, not accounting for the lateral line scale and the scale on the ventral midline (in front of the pelvic fin)" |
Upper arch gill raker number | UGR | Number of gill rakers on first upper arch; all rakers including rudimentary developed rakers |
Lower arch gill raker number | LGR | Number of gill rakers on first lower arch; all rakers including rudimentary developed rakers and the middle raker |
Total gill raker number | total GR | Gill raker number of upper and lower arch combined |
The average sizes of fish from each species differ between lakes enough that for certain species such as for
Multivariate ratio analysis is a method that performs principal component analysis (PCA) and linear discriminant analysis (LDA) on morphological ratios (
For the development of a species identification key we used LDA analysis on all characters together and on subsets of only head or only body characters for all contemporary specimens from lakes Thun and Brienz separately to calculate the first two ratios of characters that best separate each of the species in each lake. This method also allows to estimate the extent of shape change with size (i.e., the contribution of allometry to these ratios) which is given as δ and describes how good shape discriminates in comparison to size (see
Genetic analysis of ten microsatellite loci were used for the Bayesian clustering algorithm program STRUCTURE (
From the targeted spawning fisheries (each at one spawning site in lakes Thun and Brienz along a depth gradient) a total of 663 individuals from Lake Thun had complete genotypes, ten individuals had one missing locus, and four individuals had two missing loci. A total of 284 individuals from Lake Brienz had complete genotypes, eighteen individuals had one missing locus, and four individuals had two missing loci. These individuals were assigned to the different species using the program STRUCTURE with reference populations of each species deriving from the study by
We used the reference panel from
Newer genomic findings by De-Kayne et al. (unpublished) suggest that whitefish from Lake Brienz, that have previously been assigned based on genetic analysis (see above;
The principal component analyses (shapePCA) on the morphological characters show that the type specimens of all previously described species
Principal Component Analysis showing that the types of the previously described species
Principal Component Analysis showing the morphospace of the contemporary whitefish species
Principal Component Analysis showing the morphospace of the contemporary whitefish species
No single character was sufficient to distinguish
The specimens from lakes Thun and Brienz of
In Lake Brienz
In Lake Thun
In Lake Brienz
In Lake Thun
General appearance is shown in Figure 4. Morphological and meristic characters of both sexes can be found in Table
Brienzlig, Brienzling; often the time of the year the fish is caught on the spawning grounds is added to the name and shows that this species has a very wide spawning season: Sommer-Brienzlig (for summer) or Winter-Brienzlig (for winter). This species was historically known by local fishermen and fisherwomen as white whitefish (German: "Weissfelchen", but also Albele and Albuli). The common name for this species today is Brienzling which has an ending that is known as a diminutive suffix.
Differential diagnosis against
For specimens in Lake Brienz smaller than 163.5 mm SL
The specimens from lakes Thun and Brienz of
In Lake Brienz
In Lake Thun
General appearance is shown in Figure 5. Morphological and meristic characters of both sexes can be found in Table
Balchen.
Differential diagnoses against
In Brienz
The specimens of
General appearance is shown in Figure 6. Morphological and meristic characters of both sexes can be found in Table
The name given to this species by
Tiefenalbock in Lake Thun and Felchen in Lake Brienz.
General appearance is shown in Figure 7. Morphological and meristic characters of both sexes can be found in Table
The specific epithet
None; this species was not recognized by local fishermen or fisherwomen as distinct from
General appearance is shown in Figure 8. Morphological and meristic characters of both sexes can be found in Table
The specific epithet
None. We suggest the German name «Brienzer Kleinbalchen»
General appearance is shown in Figure 9. Morphological and meristic characters of both sexes can be found in Table
It is important to note that native whitefish species of Lake Biel were only known to spawn in the winter months (
The adjective
Kropfer.
The wider underjaw of
General appearance is shown in Figure 10. Morphological and meristic characters of both sexes can be found in Tables
The name
Albock
Kilch
Sandfelchen.
Gangfisch.
Blaufelchen.
1 | Caudal peduncle depth / upper jaw length ratio is 1.36–1.65 and caudal peduncle depth / maxilla length ratio is 1.77–2.24 |
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– | Caudal peduncle depth / upper jaw length ratio is 0.96–1.43 |
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2 | Total number of gill rakers 25–30 |
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– | Total number of gill rakers 31–35 |
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3 | Total number of gill rakers 15–27 |
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– | Total number of gill rakers 30–44 |
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4 | Colouration above the lateral line on the dorsum from a pale rose colouration to a pale brown colouration; no or few small pigmented dots on the edge of the scales or on the boundary of two scales on the flank; no pigmented dots on the dorsum |
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– | Colouration above the lateral line on the dorsum from a light to dark green and rarely a light olive; moderate to many dots on the edge of the scales or on the boundary of two scales on the flank and/or the dorsum |
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5 | Angle to body axis of the erected dorsal fin approx. 60–80° |
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– | Angle to body axis of the erected dorsal fin approx. 40–60° |
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1 | Total number of gill rakers 26–30 and erected dorsal fin length / upper jaw length ratio is 3.25–4.1 |
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– | Total number of gill rakers 32–42 and erected dorsal fin length / upper jaw length ratio is 2.14–3.19 |
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2 | Predorsal length / eye height ratio is 6.1–7.58 |
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– | Predorsal length / eye height ratio is 8.12–10.5 |
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3 | Body depth 19.6–25.1% SL, eye depth 23.1–28.3 % HL |
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– | Body depth 22.1–26.2% SL, eye depth 21.2–27.6% HL |
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Morphological and meristic data of
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165.0 | 173.2±51.9 | (101–271) | 221.4±16.4 | (177–271) | 224.4±14.9 | (205–271) | 216.6±18.1 | (177–254) | 122.1±11.0 | (101–164) | 122.9±13.5 | (101–164) | 120.9±6.4 | (108–129) |
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3.6 | 3.9±0.4 | (3.2–5.6) | 4.0±0.3 | (3.4–4.7) | 4.0±0.3 | (3.4–4.7) | 4.1±0.2 | (3.7–4.5) | 3.8±0.5 | (3.2–5.6) | 4.0±0.5 | (3.2–5.6) | 3.6±0.2 | (3.3–4.0) |
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na | 6.3±0.7 | (5.1–8.6) | 6.2±0.7 | (5.1–7.3) | 6.1±0.7 | (5.1–7.2) | 6.5±0.7 | (5.1–7.3) | 6.4±0.7 | (5.5–8.6) | 6.5±0.8 | (5.5–8.6) | 6.3±0.6 | (5.5–7.8) |
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14.1 | 17±1.1 | (14.7–20.9) | 16.7±0.9 | (14.8–18.7) | 16.5±0.6 | (15.2–18) | 17±1.1 | (14.8–18.7) | 17.3±1.1 | (14.7–20.9) | 17.5±1.4 | (14.7–20.9) | 17.1±0.7 | (15.3–18.1) |
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3.1 | 3.3±0.3 | (2.7–4.8) | 3.4±0.3 | (2.7–3.9) | 3.3±0.3 | (2.7–3.8) | 3.5±0.2 | (2.9–3.9) | 3.3±0.4 | (2.9–4.8) | 3.5±0.5 | (3.0–4.8) | 3.2±0.2 | (2.9–3.6) |
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na | 17.6±1.2 | (14.7–22.6) | 17.3±1.0 | (14.7–19.1) | 17.2±0.9 | (15.4–19.1) | 17.4±1.1 | (14.7–18.8) | 18.0±1.4 | (15.7–22.6) | 18.2±1.5 | (15.7–22.6) | 17.6±1.3 | (15.9–19.8) |
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na | 18.9±1.3 | (15.7–23.8) | 18.4±1.1 | (15.7–20.7) | 18.2±1 | (16.4–20.1) | 18.8±1.3 | (15.7–20.7) | 19.4±1.3 | (16.9–23.8) | 19.5±1.5 | (16.9–23.8) | 19.3±1 | (18.1–21.2) |
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11.7 | 12.1±0.9 | (10.7–16.2) | 11.9±0.6 | (10.8–13.6) | 11.8±0.6 | (10.8–13.3) | 12.0±0.7 | (11.0–13.6) | 12.2±1.1 | (10.7–16.2) | 12.6±1.2 | (10.8–16.2) | 11.8±0.9 | (10.7–13.2) |
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na | 19.1±1.4 | (16.1–24.7) | 18.5±1.2 | (16.1–21.2) | 18.3±0.9 | (16.1–19.6) | 18.9±1.5 | (16.7–21.2) | 19.7±1.5 | (17.3–24.7) | 20.2±1.6 | (17.8–24.7) | 19.1±0.9 | (17.3–20.8) |
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na | 20.1±1.4 | (17.1–26.6) | 19.7±1.1 | (17.1–21.9) | 19.5±0.9 | (17.3–21.4) | 20.0±1.3 | (17.1–21.9) | 20.6±1.5 | (18.3–26.6) | 21.0±1.7 | (18.3–26.6) | 20.0±0.9 | (18.9–21.7) |
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7 | 5.4±0.7 | (3.7–7.2) | 5.1±0.6 | (3.7–6.1) | 5.0±0.5 | (3.7–5.9) | 5.2±0.7 | (3.8–6.1) | 5.7±0.7 | (4.4–7.2) | 5.7±0.7 | (4.4–7.2) | 5.7±0.6 | (4.4–7.2) |
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13.1 | 12.9±1 | (10.5–17) | 12.7±0.8 | (11.1–14.9) | 12.4±0.6 | (11.1–13.3) | 13.1±0.8 | (12.0–14.9) | 13.0±1.2 | (10.5–17.0) | 13.3±1.3 | (11.5–17.0) | 12.7±1.0 | (10.5–14.9) |
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na | 12±0.9 | (9.7–14.4) | 12.3±0.8 | (10.9–14.0) | 12.0±0.7 | (10.9–13.4) | 12.7±0.9 | (11.3–14.0) | 11.7±1.0 | (9.7–14.4) | 11.8±1.0 | (9.7–14.4) | 11.6±1.0 | (9.7–13.8) |
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6.8 | 6.5±1.1 | (4.5–9.2) | 5.8±0.7 | (4.5–7.7) | 5.8±0.7 | (4.7–6.8) | 5.9±0.8 | (4.5–7.7) | 7.2±1.1 | (4.5–9.2) | 7.2±1.1 | (4.9–9.2) | 7.1±1.1 | (4.5–8.6) |
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na | 25±1.8 | (21.7–32.9) | 24±1.3 | (21.7–26.5) | 23.8±1.1 | (21.7–26.4) | 24.2±1.5 | (21.8–26.5) | 26±1.8 | (24–32.9) | 26.5±1.9 | (24.4–32.9) | 25.3±1.4 | (24–28.6) |
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6.9 | 7.2±0.5 | (6.3–10) | 7.1±0.3 | (6.4–7.9) | 7.1±0.4 | (6.4–7.9) | 7.2±0.3 | (6.8–7.7) | 7.3±0.6 | (6.3–10) | 7.5±0.8 | (6.6–10) | 7±0.3 | (6.3–7.4) |
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14.7 | 13.3±0.8 | (11.7–16) | 13.2±0.7 | (11.7–14.6) | 13.2±0.7 | (11.9–14.5) | 13.1±0.9 | (11.7–14.6) | 13.5±0.9 | (12.1–16.0) | 13.4±0.9 | (12.2–16.0) | 13.6±0.9 | (12.1–15.5) |
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18.9 | 19.8±1.5 | (16.9–24.2) | 18.9±1 | (16.9–20.9) | 18.8±0.9 | (16.9–20.9) | 19.3±0.9 | (17.3–20.4) | 20.7±1.4 | (17.7–24.2) | 20.5±1.8 | (17.7–24.2) | 20.9±0.6 | (19.6–21.8) |
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14.7 | 16.4±1.8 | (14.1–23.1) | 15.0±0.7 | (14.1–16.5) | 14.9±0.7 | (14.1–16.2) | 15.2±0.7 | (14.1–16.5) | 17.9±1.3 | (16.1–23.1) | 18.3±1.4 | (16.8–23.1) | 17.3±0.9 | (16.1–19.7) |
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49.2 | 51.9±2.4 | (48.4–67.7) | 52.0±1.6 | (49.5–56.0) | 52.1±1.7 | (49.5–56.0) | 51.9±1.4 | (49.5–54.3) | 51.8±3.2 | (48.4–67.7) | 52.2±3.9 | (49.1–67.7) | 51.4±1.7 | (48.4–54.0) |
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76.2 | 77.1±3.1 | (74–100) | 77.1±1.3 | (74.8–80.0) | 77.2±1.2 | (75.2–79.7) | 76.9±1.6 | (74.8–80.0) | 77.2±4.3 | (74.0–100.0) | 77.7±5.5 | (74.8–100.0) | 76.4±1.4 | (74.0–78.7) |
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45.5 | 47.2±2.2 | (43.1–61.2) | 46.5±1 | (43.1–48.4) | 46.6±0.9 | (44.9–48.4) | 46.2±1.1 | (43.1–47.7) | 48±2.7 | (45.8–61.2) | 48.5±3.3 | (45.8–61.2) | 47.3±1.3 | (46–51.3) |
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25.2 | 23.4±2.8 | (16.9–30.2) | 25.3±1.4 | (23.0–29.6) | 25.5±1.5 | (23.5–29.6) | 24.9±1.2 | (23.0–26.8) | 21.4±2.5 | (16.9–30.2) | 22.2±2.8 | (19.4–30.2) | 20.2±1.4 | (16.9–22.6) |
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43.8 | 44.3±2 | (41.2–56.2) | 44.5±1.5 | (41.8–48.3) | 44.6±1.4 | (41.8–46.6) | 44.3±1.7 | (43–48.3) | 44.1±2.5 | (41.2–56.2) | 44.1±3.1 | (41.5–56.2) | 44±1.3 | (41.2–45.8) |
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na | 121.3±2.1 | (116.6–130) | 120.7±1.8 | (116.6–123.7) | 120.5±1.8 | (116.6–123.7) | 121.1±1.8 | (118.4–123.7) | 122±2.2 | (118.2–130) | 122.1±2.3 | (119.6–130) | 121.7±2.2 | (118.2–125.5) |
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34.4 | 38.4±8.8 | (26–51.9) | 46.4±3.2 | (36.1–51.9) | 46.6±2.8 | (41.5–51.9) | 46.1±3.8 | (36.1–51.6) | 29.9±2.5 | (26–39.2) | 30.6±3 | (26–39.2) | 28.8±1 | (27.7–31.1) |
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25.4 | 22.8±2 | (18.7–27.2) | 24.2±1.3 | (20.9–27.2) | 24.1±1.3 | (20.9–26.3) | 24.4±1.3 | (22.5–27.2) | 21.2±1.4 | (18.7–25.2) | 21.3±1.2 | (19.7–23) | 21.1±1.6 | (18.7–25.2) |
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23.5 | 26.2±3.3 | (21.4–32) | 23.4±1.3 | (21.4–25.9) | 23.7±1.2 | (21.7–25.9) | 22.9±1.3 | (21.4–25.3) | 29.3±1.5 | (26.1–32) | 29.4±1.7 | (26.1–32) | 29.2±1.2 | (27.4–31.2) |
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27 | 29.9±3.7 | (23.1–36.8) | 26.6±1.4 | (23.1–29.2) | 26.9±1.4 | (24–29.2) | 26.2±1.4 | (23.1–28.1) | 33.3±1.6 | (30.4–36.8) | 33.2±1.9 | (30.4–36.8) | 33.5±1.2 | (31.5–36.2) |
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na | 26.1±2.9 | (20.8–30.6) | 23.5±1.3 | (20.8–26.2) | 23.7±1.4 | (20.8–26.2) | 23.4±1 | (21.4–25.1) | 28.7±1.1 | (26.5–30.6) | 28.7±1.2 | (26.5–30.6) | 28.9±0.9 | (27.1–30.4) |
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na | 3.4±0.7 | (2–4.9) | 3.2±0.8 | (2.0–4.9) | 3.1±0.8 | (2.0–4.5) | 3.3±1.0 | (2.1–4.9) | 3.5±0.5 | (2.5–4.8) | 3.5±0.5 | (2.9–4.8) | 3.5±0.5 | (2.5–4.3) |
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49.8 | 50.2±2.5 | (45.5–55.6) | 52±1.5 | (48.5–55.6) | 51.8±1.6 | (48.5–55.6) | 52.3±1.4 | (50.1–55.1) | 48.2±1.8 | (45.5–53.1) | 48.3±2 | (45.5–53.1) | 48.2±1.4 | (45.7–50.5) |
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75.5 | 68.3±4.5 | (59.2–80.3) | 71.8±2.9 | (66.9–80.3) | 71.5±3.2 | (66.9–80.3) | 72.2±2.4 | (67.1–75.8) | 64.6±2.3 | (59.2–69) | 64.4±2.4 | (59.2–68.8) | 64.8±2.3 | (61.3–69) |
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na | 10.4±0.8 | (8.8–12.2) | 10.3±0.7 | (8.8–12) | 10±0.6 | (8.8–11.2) | 10.8±0.7 | (9.4–12) | 10.5±0.8 | (8.8–12.2) | 10.5±0.8 | (8.9–12.2) | 10.5±0.8 | (8.8–11.7) |
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na | 31.7±1.5 | (28.6–34.9) | 31.2±1.6 | (28.8–34.7) | 30.9±1.5 | (28.8–33.9) | 31.8±1.7 | (29–34.7) | 32.1±1.3 | (28.6–34.9) | 31.8±1.1 | (28.6–33.4) | 32.6±1.3 | (30.3–34.9) |
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45.5 | 43.6±2.8 | (38.4–49.2) | 41.4±1.6 | (38.4–44.6) | 41.2±1.7 | (38.6–44.4) | 41.6±1.6 | (38.4–44.6) | 45.9±1.6 | (41.9–49.2) | 45.6±1.7 | (41.9–48.5) | 46.4±1.5 | (43.9–49.2) |
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na | 23.5±1.7 | (20.1–26.9) | 22.4±1.4 | (20.1–26.8) | 22.1±1.4 | (20.1–26.8) | 22.8±1.3 | (21.5–26.4) | 24.7±1.1 | (22.6–26.9) | 24.4±1.1 | (22.6–26.5) | 25.1±1.2 | (23.4–26.9) |
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na | 8.8±1.6 | (5.6–13.1) | 9.6±1.6 | (6–13.1) | 9.2±1.4 | (6–12.7) | 10.3±1.7 | (7.4–13.1) | 8±1 | (5.6–10.4) | 8.2±1.1 | (5.6–10.4) | 7.7±0.6 | (6.4–8.6) |
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na | 17.9±1.5 | (13.5–21.1) | 17.4±1.7 | (13.5–20) | 16.9±1.5 | (13.5–19.1) | 18.1±1.6 | (14.3–20) | 18.4±1.2 | (15.4–21.1) | 18.4±1.3 | (15.4–20.6) | 18.4±1.2 | (16.3–21.1) |
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48.1 | 48.5±4.3 | (37.5–56.1) | 50.7±3.3 | (43.5–56.1) | 50.6±3.2 | (45.1–56.1) | 50.7±3.6 | (43.5–55.1) | 46.2±4.1 | (37.5–54.7) | 45.7±3.7 | (37.5–51.7) | 46.9±4.5 | (41.3–54.7) |
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25.8 | 26.9±2.2 | (22.7–33.6) | 28.4±2 | (24.4–33.6) | 28.4±2 | (24.8–33.6) | 28.4±2.1 | (24.4–31.9) | 25.4±1.2 | (22.7–27.5) | 25.4±1.3 | (23.1–27.5) | 25.5±1.2 | (22.7–27.4) |
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na | 11.5±1.3 | (7.8–15.4) | 11.9±1.3 | (9.9–15.4) | 11.8±1.4 | (9.9–15.4) | 12.1±1.3 | (9.9–15) | 11±1.2 | (7.8–13.7) | 11.1±1.1 | (9–13.7) | 11±1.3 | (7.8–13) |
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12.2 | 11.7±2.4 | (7.5–17.5) | 13.3±2.2 | (8.6–17.5) | 13.1±2 | (9–16.5) | 13.5±2.4 | (8.6–17.5) | 10±1.3 | (7.5–12.3) | 9.9±1.2 | (8.4–12.3) | 10±1.5 | (7.5–12.2) |
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na | 24.5±2.2 | (19.1–28.9) | 25.4±1.9 | (20.9–28.7) | 24.9±1.9 | (20.9–28.7) | 26.2±1.8 | (22.7–28.4) | 23.5±2 | (19.1–28.9) | 23.4±2.1 | (19.1–28.9) | 23.6±1.9 | (19.8–27.4) |
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16.1 | 16±1.5 | (11.7–19.4) | 15.6±1.7 | (11.7–18.3) | 15.7±1.6 | (11.7–18.3) | 15.5±1.8 | (12–18.2) | 16.5±1.2 | (13.7–19.4) | 16.4±1.3 | (13.7–19.3) | 16.6±1.2 | (15.1–19.4) |
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17 | 17.7±1.6 | (14.1–21.8) | 17.2±1.5 | (14.1–20.3) | 17.3±1.6 | (14.1–20.3) | 17±1.4 | (14.7–18.9) | 18.2±1.5 | (14.9–21.8) | 18±1.6 | (14.9–21.8) | 18.6±1.2 | (16.8–21.5) |
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20 | 19.5±1.4 | (14.8–22.6) | 19.2±1.4 | (14.8–22.6) | 19.1±1.6 | (14.8–22.6) | 19.5±1.2 | (17.5–21.3) | 19.7±1.4 | (17.2–22.1) | 19.5±1.5 | (17.2–22.1) | 20±1.1 | (17.8–21.3) |
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37.8 | 36.4±2.7 | (29.6–42.2) | 35.4±1.7 | (32.3–39.6) | 35.4±1.9 | (32.3–39.6) | 35.4±1.4 | (32.8–37.5) | 37.5±3.1 | (29.6–42.2) | 36.8±3.6 | (29.6–42.2) | 38.7±1.9 | (35.3–42) |
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1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
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11 | 10 | (9–12) | 10 | (9–11) | 10 | (9–11) | 10 | (9–11) | 10 | (10–12) | 10 | (10–11) | 10 | (10–12) |
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1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
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16 | 16 | (14–17) | 16 | (14–17) | 16 | (14–17) | 16 | (15–17) | 16 | (15–17) | 16 | (15–17) | 16 | (15–17) |
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3 | 4 | (3–4) | 4 | (3–4) | 4 | (3–4) | 4 | (3–4) | 4 | (3–4) | 4 | (3–4) | 4 | (3–4) |
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11 | 10 | (9–12) | 10 | (9–12) | 10 | (9–12) | 10 | (9–12) | 10 | (9–12) | 10 | (9–12) | 10 | (9–11) |
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3 | 3 | (2–4) | 3 | (2–4) | 3 | (2–4) | 3 | (2–4) | 4 | (3–4) | 4 | (3–4) | 4 | (3–4) |
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14 | 12 | (10–14) | 12 | (10–13) | 12 | (10–13) | 12 | (11–13) | 12 | (10–14) | 12 | (11–14) | 12 | (10–14) |
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75 | 79 | (70–86) | 79 | (73–86) | 79 | (73–83) | 77 | (76–86) | 76 | (70–84) | 77 | (70–80) | 78 | (70–84) |
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na | 31 | (26–37) | 34 | (29–37) | 34 | (29–37) | 34 | (30–37) | 31 | (26–35) | 32 | (30–35) | 31 | (26–34) |
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9 | 9 | (7–10) | 9 | (8–10) | 9 | (8–10) | 9 | (9–10) | 8 | (7–9) | 8 | (7–9) | 8 | (7–9) |
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7 | 8 | (6–9) | 8 | (7–9) | 8 | (7–9) | 8 | (8–9) | 7 | (6–8) | 7 | (6–8) | 8 | (6–8) |
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8 | 9 | (7–9) | 9 | (8–9) | 9 | (8–9) | 9 | (8–9) | 8 | (7–9) | 8 | (7–9) | 8 | (7–8) |
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11 | 13 | (9–17) | 13 | (9–17) | 13 | (9–15) | 14 | (11–17) | 13 | (11–16) | 13 | (11–16) | 13 | (12–14) |
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23 | 25 | (20–29) | 24 | (20–28) | 24 | (20–28) | 26 | (22–27) | 25 | (22–29) | 25 | (22–29) | 27 | (24–28) |
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34 | 38 | (32–44) | 38 | (32–44) | 37 | (32–42) | 38 | (33–44) | 40 | (35–42) | 38 | (35–42) | 40 | (37–42) |
Morphological and meristic data of
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283.0 | 266.1±56.4 | (147-364) | 288.3±45.1 | (210-364) | 299.6±37.3 | (267-364) | 273.3±52.2 | (210-352) | 214.3±46.0 | (147-290) | 240.1±27.9 | (213-290) | 162.7±23.3 | (147-190) |
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4.5 | 4.2±0.4 | (3.1-5.0) | 4.4±0.3 | (3.8-5) | 4.3±0.4 | (3.8-5.0) | 4.4±0.3 | (4.0-4.8) | 3.7±0.3 | (3.1-4.0) | 3.9±0.1 | (3.7-4.0) | 3.5±0.5 | (3.1-4.0) |
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4.7 | 6.1±0.8 | (4.7-7.6) | 5.9±0.7 | (4.7-7.2) | 5.9±0.7 | (4.8-6.6) | 5.9±0.8 | (4.7-7.2) | 6.7±0.5 | (5.9-7.6) | 6.8±0.5 | (6.0-7.6) | 6.5±0.6 | (5.9-7.1) |
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18.1 | 16.7±1.1 | (14-18.2) | 16.6±0.9 | (14.7-18.2) | 16.2±0.9 | (14.7-17.3) | 17.2±0.6 | (16.3-18.2) | 16.9±1.5 | (14-18.1) | 16.9±1.6 | (14-18.1) | 16.8±1.4 | (15.2-18) |
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3.7 | 3.3±0.3 | (2.5-3.7) | 3.4±0.3 | (2.8-3.7) | 3.4±0.3 | (2.8-3.7) | 3.3±0.2 | (3.0-3.5) | 3.0±0.3 | (2.5-3.5) | 3.0±0.3 | (2.5-3.5) | 3.1±0.3 | (2.8-3.3) |
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18.7 | 16.2±1.2 | (13.6-18.7) | 16.2±1.3 | (13.6-18.7) | 16.0±1.4 | (13.6-18.6) | 16.5±1.1 | (15.1-18.7) | 16.3±1.1 | (13.9-17.1) | 16.4±1.3 | (13.9-17.1) | 16.1±0.9 | (15.1-16.8) |
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19.7 | 17±1.1 | (14.4-19.7) | 17±1.1 | (15.3-19.7) | 16.8±1.1 | (15.3-19.1) | 17.3±1.1 | (15.8-19.7) | 16.9±1.2 | (14.4-17.7) | 17±1.3 | (14.4-17.7) | 16.7±1.1 | (15.5-17.6) |
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12.9 | 12.8±0.9 | (10.7-14.7) | 12.9±0.9 | (11.5-14.7) | 12.9±0.9 | (11.7-14.4) | 12.9±0.9 | (11.5-14.7) | 12.6±1.1 | (10.7-14.3) | 12.6±1.1 | (10.7-14.0) | 12.7±1.4 | (11.7-14.3) |
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na | 19.7±1.2 | (16.4-23.0) | 19.5±0.8 | (17.6-20.6) | 19.3±0.8 | (17.6-20.5) | 19.7±0.8 | (18.6-20.6) | 20.3±1.8 | (16.4-23.0) | 20.1±1.9 | (16.4-21.5) | 20.9±1.9 | (19.5-23.0) |
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22.6 | 20.8±1.4 | (17.0-24.0) | 20.6±0.9 | (18.6-22.5) | 20.4±0.8 | (18.6-21.3) | 20.9±1.1 | (19.6-22.5) | 21.4±2.0 | (17.0-24.0) | 21.2±2.2 | (17.0-22.7) | 21.7±2.0 | (20.0-24.0) |
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5.6 | 5.0±0.6 | (4.0-6.5) | 4.8±0.5 | (4-5.8) | 4.8±0.6 | (4.0-5.8) | 4.9±0.4 | (4.2-5.5) | 5.3±0.7 | (4.3-6.5) | 5.2±0.6 | (4.3-5.9) | 5.5±1.1 | (4.4-6.5) |
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12.5 | 12.4±0.9 | (9.8-14.2) | 12.5±0.9 | (10.3-14.2) | 12.3±0.9 | (10.3-14.0) | 12.8±0.8 | (11.1-14.2) | 12.1±1.0 | (9.8-13.3) | 12.1±1.1 | (9.8-12.9) | 12.2±1.0 | (11.4-13.3) |
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14 | 12.3±0.9 | (9.8-13.8) | 12.2±0.7 | (10.5-13.3) | 11.9±0.7 | (10.5-12.7) | 12.5±0.7 | (10.9-13.3) | 12.5±1.3 | (9.8-13.8) | 12.6±1.4 | (9.8-13.8) | 12.4±1.3 | (11.2-13.7) |
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4.8 | 4.4±0.6 | (3.4-5.5) | 4.3±0.6 | (3.4-5.5) | 4.3±0.6 | (3.5-5.5) | 4.2±0.5 | (3.4-4.7) | 4.7±0.5 | (3.8-5.4) | 4.6±0.5 | (3.8-5.1) | 4.9±0.5 | (4.5-5.4) |
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na | 24±1.4 | (19.1-26.3) | 24±1 | (21.9-26) | 23.8±0.7 | (22.7-24.8) | 24.3±1.2 | (21.9-26) | 24.1±2.2 | (19.1-26.3) | 24±2.6 | (19.1-26.3) | 24.3±1.5 | (22.9-25.9) |
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8.8 | 8.1±0.5 | (6.7-8.9) | 8.2±0.4 | (7.6-8.9) | 8.1±0.3 | (7.6-8.6) | 8.3±0.4 | (7.8-8.9) | 7.8±0.6 | (6.7-8.6) | 7.8±0.6 | (6.7-8.6) | 7.8±0.8 | (7-8.4) |
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12.6 | 12.6±0.8 | (11.2-14.4) | 12.6±0.9 | (11.2-14.4) | 12.5±0.8 | (11.2-13.6) | 12.9±0.9 | (11.8-14.4) | 12.5±0.9 | (11.3-13.9) | 12.5±0.8 | (11.3-13.6) | 12.5±1.2 | (11.5-13.9) |
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18.4 | 17.7±1.1 | (15.3-19.5) | 17.8±1 | (15.7-19.4) | 17.8±1.1 | (15.7-19.4) | 17.8±1 | (16.2-19.4) | 17.6±1.3 | (15.3-19.5) | 17.3±1.4 | (15.3-19.5) | 18.3±1.1 | (17.2-19.3) |
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15.3 | 14.6±0.9 | (12.6-16.3) | 14.2±0.6 | (12.6-15.6) | 14.1±0.8 | (12.6-15.6) | 14.3±0.5 | (13.7-15.0) | 15.4±0.8 | (14.0-16.3) | 15.2±0.9 | (14.0-16.3) | 15.8±0.2 | (15.6-16.0) |
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53.6 | 51.5±2.3 | (42.7-55.2) | 51.9±1.8 | (48-55.2) | 51.8±1.8 | (49.3-55.2) | 52.1±1.9 | (48.0-54.5) | 50.5±3.1 | (42.7-53.0) | 50.1±3.8 | (42.7-53.0) | 51.4±0.7 | (50.6-51.9) |
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77.2 | 78.3±1.5 | (75.2-81.8) | 78.5±1.4 | (75.2-81.1) | 78.8±1.3 | (76.5-81.1) | 78.1±1.5 | (75.2-80.3) | 77.8±1.8 | (76.1-81.8) | 78.2±2.0 | (76.1-81.8) | 76.8±0.3 | (76.5-77.0) |
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45 | 47.8±2 | (39.9-50.3) | 48±1.5 | (45.5-50.3) | 48.3±1.5 | (45.5-50.3) | 47.6±1.7 | (45.6-50) | 47.3±3 | (39.9-50.2) | 47.1±3.6 | (39.9-50.2) | 47.8±1.6 | (46.3-49.5) |
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27.6 | 26.5±2.7 | (19.9-31.8) | 27.5±1.9 | (24.5-31.8) | 28.2±2.0 | (25.2-31.8) | 26.5±1.4 | (24.5-28.8) | 24.2±2.9 | (19.9-27.0) | 24.8±2.8 | (19.9-27.0) | 23.1±3.3 | (20.5-26.8) |
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43.7 | 42.6±2 | (34.7-46) | 42.7±1.3 | (38.3-43.9) | 42.3±1.5 | (38.3-43.9) | 43.2±0.5 | (42.1-43.8) | 42.3±3.3 | (34.7-46) | 42.3±4.1 | (34.7-46) | 42.5±0.9 | (41.4-43) |
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118 | 120.5±4.2 | (100-124.2) | 121±1.3 | (118.7-123.4) | 120.6±1.5 | (118.7-123.4) | 121.5±0.7 | (120.8-122.8) | 119.2±7.5 | (100-124.2) | 118.6±9.3 | (100-124.2) | 120.5±1.7 | (118.9-122.4) |
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58.6 | 54.2±10.6 | (31.1-73.1) | 58.3±8.6 | (41.9-73.1) | 60.2±6.8 | (53-73.1) | 55.7±10.4 | (41.9-72.2) | 44.6±8.7 | (31.1-58.3) | 49.5±4.7 | (45.7-58.3) | 34.9±6 | (31.1-41.8) |
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22.2 | 23.1 ± 1.6 | (20.1-26.4) | 23.5 ± 1.5 | (20.9-26.4) | 23.7 ± 1.5 | (21.4-26.4) | 23.2 ± 1.5 | (20.9-25.3) | 22.4 ± 1.6 | (20.1-25) | 22.8 ± 1.7 | (20.3-25) | 21.7 ± 1.4 | (20.1-22.7) |
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22.1 | 22.5 ± 2 | (19.5-27.2) | 21.7 ± 1.5 | (19.5-26.1) | 21.3 ± 1.1 | (19.5-22.9) | 22.3 ± 2 | (19.8-26.1) | 24.3 ± 1.7 | (21.8-27.2) | 23.3 ± 1.1 | (21.8-24.9) | 26.1 ± 1 | (25.4-27.2) |
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28.9 | 27.4 ± 1.7 | (24.5-31.5) | 26.7 ± 1.2 | (24.5-29) | 26.6 ± 1 | (24.9-28.3) | 26.9 ± 1.4 | (24.5-29) | 29 ± 1.9 | (26.4-31.5) | 28.1 ± 1.7 | (26.4-30.6) | 30.6 ± 0.8 | (29.9-31.5) |
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23.7 | 22.2 ± 1.8 | (19.1-27.1) | 21.4 ± 1.4 | (19.1-23.6) | 21.1 ± 1.2 | (19.1-23) | 21.9 ± 1.6 | (19.6-23.6) | 23.9 ± 1.5 | (22.4-27.1) | 23.3 ± 0.9 | (22.4-24.6) | 25.1 ± 2 | (23.2-27.1) |
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5.7 | 5.0 ± 0.8 | (3.3-6.3) | 5.1 ± 0.8 | (3.4-6.3) | 5.0 ± 0.7 | (3.7-6.0) | 5.2 ± 0.9 | (3.4-6.3) | 4.7 ± 0.8 | (3.3-5.8) | 4.6 ± 0.9 | (3.3-5.5) | 4.9 ± 0.8 | (4.3-5.8) |
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51.2 | 52.4 ± 1.5 | (48.9-55.4) | 52.6 ± 1.2 | (50.2-55.4) | 52.5 ± 1.3 | (50.8-55.4) | 52.8 ± 1.2 | (50.2-54.4) | 51.7 ± 1.9 | (48.9-54.8) | 52.2 ± 1.5 | (50.7-54.8) | 50.8 ± 2.6 | (48.9-53.8) |
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71.7 | 71 ± 3.7 | (65.5-79.6) | 71.6 ± 4 | (65.6-79.6) | 71.5 ± 4.6 | (65.6-79.6) | 71.8 ± 3.4 | (67.9-76.7) | 69.4 ± 2.5 | (65.5-73.2) | 69.7 ± 2.7 | (65.5-73.2) | 68.7 ± 2.3 | (66.3-70.8) |
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8.9 | 9.4 ± 0.5 | (8.4-10.4) | 9.4 ± 0.5 | (8.4-10.4) | 9.3 ± 0.6 | (8.4-10.4) | 9.4 ± 0.6 | (8.8-10.2) | 9.5 ± 0.5 | (8.7-10.2) | 9.6 ± 0.5 | (9.2-10.2) | 9.3 ± 0.5 | (8.7-9.7) |
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27.8 | 27.4 ± 1.5 | (24.3-30.1) | 27.7 ± 1.5 | (24.3-30.1) | 27.2 ± 1.5 | (24.3-29.4) | 28.4 ± 1.3 | (26.5-30.1) | 26.8 ± 1.3 | (25.4-29.1) | 26.5 ± 1 | (25.6-28.2) | 27.3 ± 1.8 | (25.4-29.1) |
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38.2 | 38.4 ± 1.7 | (33.8-41.4) | 38.6 ± 1.7 | (36.6-41.4) | 38.5 ± 1.8 | (36.6-41.4) | 38.7 ± 1.6 | (37-41.2) | 38.2 ± 1.8 | (33.8-39.4) | 38 ± 2.1 | (33.8-39.2) | 38.5 ± 1.3 | (36.9-39.4) |
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22.5 | 20 ± 1.2 | (16.6-22.5) | 20 ± 1.1 | (17.7-22.1) | 19.7 ± 0.9 | (18.4-21) | 20.3 ± 1.3 | (17.7-22.1) | 20 ± 1.6 | (16.6-22.5) | 19.6 ± 1.6 | (16.6-21.1) | 20.8 ± 1.5 | (19.5-22.5) |
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9.9 | 10.1 ± 1.4 | (7.2-12.9) | 10.5 ± 1.3 | (8.5-12.9) | 10.5 ± 1.4 | (8.5-12.9) | 10.6 ± 1.2 | (8.7-12.2) | 9.2 ± 1.1 | (7.2-10.5) | 9.2 ± 0.8 | (7.9-10) | 9.3 ± 1.9 | (7.2-10.5) |
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13.8 | 15.6 ± 1.1 | (13.7-17.6) | 15.6 ± 1.2 | (13.7-17.6) | 15.7 ± 1.1 | (14.1-17.6) | 15.5 ± 1.4 | (13.7-17.2) | 15.7 ± 0.9 | (14.6-17.6) | 15.6 ± 0.8 | (14.6-16.7) | 16 ± 1.3 | (15.3-17.6) |
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47.1 | 50 ± 4.5 | (39.2-59.5) | 51.3 ± 4.1 | (44.2-59.5) | 51.5 ± 5 | (44.2-59.5) | 51 ± 2.8 | (46.3-55.9) | 46.9 ± 4 | (39.2-52.3) | 47.4 ± 5 | (39.2-52.3) | 45.8 ± 0.4 | (45.4-46.1) |
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24.5 | 27.7 ± 2.2 | (22.4-32.5) | 28.3 ± 2.3 | (22.4-32.5) | 28 ± 2.4 | (22.4-32.4) | 28.7 ± 2.3 | (24.9-32.5) | 26.5 ± 1.2 | (24.9-28) | 26.5 ± 1.4 | (24.9-28) | 26.6 ± 1 | (25.5-27.4) |
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10.7 | 11.7 ± 1 | (9.5-14.1) | 12.1 ± 1 | (10.5-14.1) | 12 ± 1.1 | (10.5-14.1) | 12.1 ± 0.9 | (10.7-14) | 11 ± 0.7 | (9.5-11.9) | 10.9 ± 0.9 | (9.5-11.9) | 11.2 ± 0.4 | (10.9-11.6) |
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10.3 | 11.1 ± 2.2 | (7.3-15.7) | 12.1 ± 1.7 | (10.1-15.7) | 11.9 ± 1.8 | (10.1-15.4) | 12.4 ± 1.6 | (10.5-15.7) | 8.8 ± 1.2 | (7.3-10.6) | 9 ± 1.2 | (7.5-10.6) | 8.5 ± 1.4 | (7.3-10.1) |
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19.8 | 23.2 ± 2.2 | (18.4-27.2) | 23.5 ± 2.2 | (19.9-27.2) | 23 ± 2.5 | (19.9-27.2) | 24.1 ± 1.6 | (21.9-26.5) | 22.8 ± 2.2 | (18.4-25.6) | 23.7 ± 1.6 | (21.4-25.6) | 20.9 ± 2.2 | (18.4-22.8) |
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10.6 | 10.9 ± 1.4 | (8.3-15.2) | 11.3 ± 1.4 | (9.3-15.2) | 11.3 ± 1.4 | (9.3-15.2) | 11.3 ± 1.4 | (9.5-13.2) | 9.8 ± 1 | (8.3-11.2) | 9.9 ± 1.1 | (8.3-11.2) | 9.7 ± 0.8 | (8.7-10.2) |
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11.5 | 11.9 ± 1.2 | (10-15.2) | 12.3 ± 1.1 | (10.6-15.2) | 12.2 ± 1.2 | (10.6-15.2) | 12.5 ± 1 | (11.3-14.1) | 10.8 ± 0.7 | (10-12.3) | 10.9 ± 0.8 | (10-12.3) | 10.6 ± 0.6 | (10-11.2) |
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na | 18.5 ± 1.4 | (15.6-21.5) | 18.4 ± 1.3 | (15.6-20.7) | 18.6 ± 1.5 | (15.6-20.7) | 18.2 ± 1.1 | (16.1-19.7) | 18.7 ± 1.7 | (16.4-21.5) | 18.2 ± 1.4 | (16.4-20) | 19.8 ± 1.7 | (18-21.5) |
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35 | 33.9 ± 2.1 | (28.6-38.8) | 33.5 ± 1.8 | (28.6-36.3) | 34.1 ± 1.5 | (30.9-36.3) | 32.7 ± 1.9 | (28.6-35) | 34.7 ± 2.7 | (30.4-38.8) | 34.5 ± 2.7 | (30.4-38.2) | 35.1 ± 3.2 | (32.8-38.8) |
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1 | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) |
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11 | 11 | (10-11) | 11 | (10-11) | 11 | (10-11) | 11 | (10-11) | 10 | (10-11) | 10 | (10-11) | 11 | (10-11) |
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1 | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) |
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16 | 15 | (14-17) | 15 | (14-17) | 15 | (14-17) | 16 | (14-16) | 15 | (15-17) | 15 | (15-16) | na | (15-17) |
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3 | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (4-4) |
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11 | 11 | (10-13) | 11 | (10-13) | 11 | (10-13) | 11 | (10-12) | 11 | (10-11) | 11 | (10-11) | 11 | (11-11) |
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3 | 3 | (3-4) | 3 | (3-4) | 3 | (3-3) | 3 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) |
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12 | 12 | (10-14) | 12 | (10-14) | 12 | (10-14) | 12 | (10-13) | 11 | (11-13) | 12 | (11-13) | 11 | (11-11) |
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82 | 84 | (77-93) | 81 | (77-93) | 84 | (77-93) | 80 | (78-84) | 84 | (80-88) | 86 | (80-88) | na | (81-84) |
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39 | 36 | (32-42) | 36 | (32-42) | 36 | (32-42) | 33 | (33-38) | 33 | (32-42) | na | (33-42) | na | (32-37) |
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10 | 10 | (8-11) | 10 | (9-11) | 10 | (9-11) | 10 | (9-11) | 10 | (8-10) | 10 | (8-10) | 9 | (9-10) |
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8 | 8 | (7-9) | 8 | (8-9) | 8 | (8-9) | 8 | (8-9) | 8 | (7-8) | 8 | (7-8) | 8 | (7-8) |
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8 | 8 | (7-9) | 9 | (8-9) | 9 | (8-9) | 8 | (8-9) | 8 | (7-8) | 8 | (8-8) | 8 | (7-8) |
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10 | 10 | (8-11) | 10 | (8-11) | 10 | (9-11) | 11 | (8-11) | 9 | (9-11) | 9 | (9-11) | na | (9-11) |
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18 | 19 | (15-23) | 19 | (16-23) | 19 | (16-21) | 20 | (17-23) | 18 | (15-21) | 17 | (17-21) | na | (15-19) |
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28 | 29 | (25-34) | 30 | (25-34) | 28 | (26-32) | 30 | (25-34) | 28 | (26-30) | 29 | (27-30) | 28 | (26-28) |
Morphological and meristic data of
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154.5 | 207.5±35.2 | (132-288) | 230.2±21.2 | (191-288) | 226.6±14.3 | (191-245) | 234.9±27.8 | (202-288) | 184.8±31.7 | (132-244) | 195.9±29.7 | (141-244) | 177.3±31.6 | (132-225) |
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3.6 | 3.8±0.4 | (3.1-4.8) | 4.0±0.4 | (3.2-4.8) | 3.9±0.3 | (3.3-4.4) | 4.0±0.4 | (3.2-4.8) | 3.7±0.4 | (3.1-4.6) | 3.7±0.3 | (3.1-4.1) | 3.7±0.4 | (3.2-4.6) |
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5.4 | 6.2±0.8 | (3.9-8.0) | 6.2±0.9 | (3.9-8.0) | 6.3±0.8 | (4.6-7.4) | 6.1±1.0 | (3.9-8.0) | 6.2±0.8 | (3.9-7.4) | 6.2±0.8 | (3.9-7.0) | 6.2±0.9 | (3.9-7.4) |
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17.4 | 16.1±1 | (13.7-19.3) | 16.4±1.2 | (13.7-19.3) | 16.6±1 | (15.1-19.3) | 16.1±1.4 | (13.7-18.6) | 15.8±0.8 | (14.6-17.4) | 15.8±0.7 | (14.7-17.1) | 15.9±0.8 | (14.6-17.4) |
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3.1 | 3.2±0.3 | (2.7-3.8) | 3.3±0.3 | (2.8-3.8) | 3.2±0.3 | (2.8-3.7) | 3.4±0.3 | (2.9-3.8) | 3.1±0.2 | (2.7-3.4) | 3.0±0.2 | (2.7-3.4) | 3.1±0.2 | (2.7-3.4) |
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17.8 | 16.2±1.3 | (13.3-18.9) | 16.5±1.4 | (13.3-18.9) | 16.8±1.2 | (14.7-18.9) | 16.1±1.5 | (13.3-18.4) | 16.0±1.1 | (14.1-18.7) | 15.8±1.0 | (14.2-17.3) | 16.0±1.2 | (14.1-18.7) |
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18.4 | 17.2±1.4 | (13.8-20.6) | 17.7±1.5 | (13.8-20.6) | 18±1.4 | (15.5-20.6) | 17.2±1.7 | (13.8-19.8) | 16.8±1.1 | (14.9-19.7) | 16.5±0.9 | (14.9-17.9) | 17±1.2 | (15.3-19.7) |
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11.4 | 11.9±0.7 | (10.3-13.3) | 11.8±0.7 | (10.3-13.1) | 11.6±0.7 | (10.3-12.9) | 12.0±0.6 | (11.2-13.1) | 12.0±0.7 | (10.3-13.3) | 12.0±0.4 | (11.3-12.6) | 12.0±0.9 | (10.3-13.3) |
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na | 17.9±1.3 | (15.4-21.9) | 18.0±1.4 | (15.4-21.9) | 18.4±1.4 | (15.4-21.9) | 17.4±1.2 | (15.8-19.8) | 17.8±1.1 | (15.7-20.0) | 17.6±1.0 | (15.7-19.4) | 18.0±1.2 | (16.5-20.0) |
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19.1 | 19.1±1.3 | (16.7-23.5) | 19.3±1.4 | (16.7-23.5) | 19.7±1.4 | (16.7-23.5) | 18.8±1.3 | (17.0-21.3) | 18.8±1.1 | (17.1-21.1) | 18.8±0.9 | (17.1-20.3) | 18.8±1.3 | (17.1-21.1) |
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6.3 | 5.1±0.6 | (3.9-7.0) | 5.0±0.7 | (3.9-6.9) | 5.1±0.7 | (3.9-6.9) | 4.8±0.7 | (4.1-6.3) | 5.3±0.5 | (4.3-7.0) | 5.3±0.5 | (4.7-6.3) | 5.3±0.6 | (4.3-7.0) |
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12.1 | 12.3±0.9 | (10.6-15.1) | 12.6±1.0 | (10.7-15.1) | 12.5±1.0 | (10.7-14.0) | 12.8±1.1 | (11.7-15.1) | 11.9±0.8 | (10.6-13.3) | 12.0±0.6 | (11.4-13.0) | 11.9±0.8 | (10.6-13.3) |
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na | 11.5±0.9 | (9.8-13.9) | 11.9±0.9 | (10.2-13.9) | 12.1±0.9 | (10.5-13.9) | 11.7±1.0 | (10.2-13.3) | 11.1±0.6 | (9.8-12.7) | 11.0±0.5 | (9.8-11.9) | 11.1±0.7 | (10.2-12.7) |
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5.7 | 5.5±0.8 | (4.0-8.1) | 5.6±0.7 | (4.6-8.1) | 5.7±0.7 | (4.8-8.1) | 5.4±0.6 | (4.6-6.9) | 5.5±0.8 | (4.0-7.7) | 5.5±0.7 | (4.3-6.7) | 5.5±0.9 | (4.0-7.7) |
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na | 23.8±1.3 | (19.6-27.2) | 23.6±1.3 | (19.6-26.6) | 23.9±1.1 | (22.1-26.4) | 23.3±1.6 | (19.6-26.6) | 24±1.2 | (22.4-27.2) | 23.8±1.3 | (22.4-27.2) | 24.1±1.1 | (22.7-26.4) |
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7.7 | 7.2±0.3 | (6.7-8.5) | 7.1±0.3 | (6.7-8.5) | 7.1±0.4 | (6.7-8.5) | 7.2±0.2 | (6.9-7.7) | 7.3±0.3 | (6.8-8) | 7.3±0.3 | (6.8-7.8) | 7.4±0.3 | (6.9-8) |
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13.9 | 13.7±0.9 | (11.5-16.1) | 13.3±0.8 | (11.5-14.7) | 13.1±0.8 | (11.5-14.7) | 13.5±0.6 | (12.6-14.6) | 14.2±0.8 | (13.1-16.1) | 13.9±0.7 | (13.1-15.5) | 14.3±0.8 | (13.2-16.1) |
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19.1 | 18.9±1 | (16.8-22.2) | 18.8±1 | (16.8-22.2) | 18.6±1.1 | (16.8-22.2) | 19±0.8 | (17.8-20) | 19±1.1 | (16.9-20.8) | 18.7±1.1 | (16.9-20.4) | 19.1±1.1 | (17.2-20.8) |
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15.9 | 15.2±0.8 | (13.6-16.8) | 14.8±0.7 | (13.6-16.2) | 14.9±0.6 | (13.9-16.1) | 14.6±0.9 | (13.6-16.2) | 15.7±0.7 | (14.5-16.8) | 15.4±0.8 | (14.5-16.8) | 15.8±0.6 | (14.7-16.7) |
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52.1 | 52.0±1.5 | (47.5-55.5) | 51.8±1.7 | (47.5-55.1) | 52.2±1.7 | (47.5-55.1) | 51.3±1.6 | (48.5-53.8) | 52.1±1.3 | (48.8-55.5) | 52.7±1.4 | (51.3-55.5) | 51.8±1.1 | (48.8-53.9) |
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78.8 | 77.0±1.3 | (74.4-80.2) | 76.9±1.5 | (74.4-80.2) | 77.4±1.6 | (74.7-80.2) | 76.3±1.1 | (74.4-78.1) | 77.1±1.0 | (75.2-79.2) | 77.4±0.9 | (76.4-79.2) | 76.8±1.0 | (75.2-78.6) |
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48.4 | 46.8±1.3 | (41.5-49.1) | 46.8±1.5 | (41.5-49.1) | 46.5±1.7 | (41.5-48.8) | 47.2±1.1 | (44.8-49.1) | 46.8±1 | (43.4-49) | 46.7±1.4 | (43.4-49) | 46.9±0.7 | (45.9-48) |
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24.9 | 24.4±1.4 | (22.1-28.1) | 24.9±1.4 | (22.7-28.1) | 25.3±1.3 | (23.2-28.1) | 24.4±1.4 | (22.7-28.1) | 23.9±1.2 | (22.1-26.2) | 24.7±1.1 | (22.4-26.2) | 23.4±0.9 | (22.1-24.9) |
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45.4 | 44.5±1.4 | (41.6-50.7) | 44.9±1.7 | (41.6-50.7) | 44.5±2 | (41.6-50.7) | 45.3±1.2 | (43.4-47.1) | 44.2±0.9 | (42.5-45.8) | 44±0.8 | (42.5-45.6) | 44.3±1 | (42.5-45.8) |
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na | 121.1±1.8 | (117.3-126) | 120.7±1.9 | (117.3-124.2) | 121.3±1.7 | (117.3-124) | 120±1.9 | (117.4-124.2) | 121.5±1.8 | (118.8-126) | 121.3±2.1 | (119-126) | 121.6±1.5 | (118.8-124.2) |
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35.1 | 43.5±6.5 | (27.9-55.9) | 47.5±3.4 | (42-55.9) | 47.4±2.8 | (42.6-50.9) | 47.7±4.2 | (42-55.9) | 39.5±6.3 | (27.9-48.8) | 41.3±5.1 | (31.4-48.2) | 38.3±6.9 | (27.9-48.8) |
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19.9 | 23.6±1.9 | (18.2-27) | 24.3±1.6 | (18.2-27) | 24.2±2 | (18.2-27) | 24.5±1.1 | (23-26.5) | 22.8±1.9 | (18.5-26.4) | 23.2±1.4 | (21.4-25.5) | 22.5±2.2 | (18.5-26.4) |
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25.8 | 23.6±2 | (19.9-27.6) | 22.4±1.4 | (19.9-25.9) | 22.8±1.3 | (20.9-25.9) | 22±1.5 | (19.9-24.7) | 24.8±1.7 | (21.2-27.6) | 24.9±1 | (22.9-26.6) | 24.8±2.1 | (21.2-27.6) |
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31.8 | 27.6±2.2 | (23.2-33) | 26.3±1.6 | (23.2-29) | 26.6±1.5 | (23.6-29) | 25.9±1.7 | (23.2-28.6) | 29±1.9 | (25.3-33) | 29.1±0.9 | (27.5-30.6) | 28.9±2.4 | (25.3-33) |
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25.8 | 23.4±1.6 | (19.7-26.3) | 22.4±1.3 | (19.7-25.3) | 22.6±1.2 | (20.8-25.3) | 22.1±1.4 | (19.7-25) | 24.4±1.3 | (22.1-26.3) | 24.4±1.2 | (22.8-26.2) | 24.4±1.4 | (22.1-26.3) |
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3.6 | 4.3±1.2 | (1.7-6.8) | 3.6±1.1 | (1.7-5.9) | 3.3±0.9 | (1.7-5.2) | 4.1±1.1 | (2.1-5.9) | 4.9±0.9 | (3.4-6.8) | 4.9±0.8 | (4.0-6.8) | 4.9±1.1 | (3.4-6.5) |
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49.8 | 51.3±1.8 | (46.8-54.8) | 52.2±1.4 | (48.7-54.8) | 52±1.6 | (48.7-54.1) | 52.5±1.1 | (51.4-54.8) | 50.5±1.7 | (46.8-54) | 50.7±1 | (49.8-53) | 50.3±2.1 | (46.8-54) |
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68.3 | 69.6±3.1 | (63.6-78.6) | 70.7±3.3 | (65.5-78.6) | 70.6±3.7 | (65.5-78.6) | 70.8±2.7 | (66.3-74.8) | 68.5±2.6 | (63.6-73.2) | 68.8±2.8 | (65.7-73.2) | 68.3±2.5 | (63.6-72.2) |
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11.5 | 10.1±0.8 | (8.2-12.1) | 10±0.9 | (8.2-12.1) | 10.1±0.9 | (8.5-12.1) | 9.8±0.9 | (8.2-11) | 10.1±0.8 | (8.5-11.4) | 10±0.9 | (8.5-11.4) | 10.2±0.8 | (8.6-11.4) |
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30.5 | 30±1.4 | (27.6-34.1) | 30.5±1.5 | (28-34.1) | 30.8±1.6 | (28-34.1) | 30.2±1.3 | (28.2-33.1) | 29.5±1.3 | (27.6-32) | 29.4±1.2 | (28.1-31.2) | 29.6±1.3 | (27.6-32) |
|
38.4 | 41.6±2.6 | (36.9-48.4) | 40.7±2.1 | (36.9-46.1) | 41.3±2.1 | (37.4-46.1) | 39.8±1.8 | (36.9-42.3) | 42.6±2.6 | (37.6-48.4) | 43.1±2.3 | (40.6-47.8) | 42.2±2.9 | (37.6-48.4) |
|
26.6 | 21.8±1.3 | (18.5-25.6) | 21.8±1.6 | (18.5-25.6) | 22.1±1.4 | (19.8-25.6) | 21.5±1.7 | (18.5-25.1) | 21.7±1 | (18.7-24.2) | 22±0.8 | (21-23.8) | 21.5±1.1 | (18.7-24.2) |
|
8.1 | 9.3±1.2 | (6.7-12.4) | 9.5±1.2 | (6.7-12.4) | 9±1 | (6.7-10.5) | 10.1±1.1 | (8.7-12.4) | 9±1.1 | (6.7-10.9) | 9.4±1.2 | (6.9-10.9) | 8.8±1 | (6.7-10.4) |
|
15 | 17.7±1.3 | (14.7-20.4) | 17.5±1.3 | (14.7-20.4) | 17.6±1.5 | (14.7-20.4) | 17.5±1.1 | (16-19.6) | 17.8±1.3 | (14.7-19.7) | 17.4±1.6 | (14.7-19.7) | 18±1.1 | (15.8-19.7) |
|
45.2 | 49.8±3.1 | (42.3-57.2) | 51.1±3 | (45.8-56.6) | 51.5±3.4 | (45.8-56.6) | 50.6±2.3 | (47.8-54.1) | 48.5±2.7 | (42.3-57.2) | 48.8±3.5 | (42.3-57.2) | 48.3±2 | (44.8-52.8) |
|
24.5 | 27±1.5 | (22.8-31.5) | 27.7±1.6 | (23.6-31.5) | 27.2±1.4 | (23.6-29.7) | 28.3±1.6 | (25.4-31.5) | 26.4±1.1 | (22.8-28.8) | 26.6±0.9 | (25.4-28.8) | 26.2±1.3 | (22.8-28.5) |
|
10.9 | 11.5±1.1 | (9.2-13.5) | 11.7±1 | (9.8-13.5) | 11.6±1.1 | (9.8-13.3) | 11.8±0.9 | (10.6-13.5) | 11.4±1.2 | (9.2-13.5) | 11.2±1 | (10.1-13.3) | 11.5±1.3 | (9.2-13.5) |
|
14.7 | 12±1.9 | (7.9-16) | 12.4±2.3 | (7.9-16) | 12.5±2.2 | (7.9-15.8) | 12.4±2.5 | (7.9-16) | 11.6±1.2 | (8.6-13.3) | 11.4±1.3 | (8.7-13.3) | 11.8±1.2 | (8.6-13.2) |
|
23.7 | 24±1.8 | (20.3-30.3) | 24.7±1.8 | (21-30.3) | 25.2±2 | (21-30.3) | 24.1±1.3 | (22.1-26.8) | 23.4±1.5 | (20.3-26.5) | 23.1±1.7 | (20.3-26.5) | 23.5±1.5 | (21-26) |
|
14.3 | 14.5±2 | (10.5-21.3) | 15.8±1.9 | (12.5-21.3) | 16.3±1.8 | (13.9-21.3) | 15±1.8 | (12.5-19.6) | 13.2±1.2 | (10.5-15) | 13±1 | (11.5-14.4) | 13.3±1.4 | (10.5-15) |
|
14.9 | 15.6±2.1 | (12.3-22.6) | 16.9±2 | (12.8-22.6) | 17.5±1.9 | (15.4-22.6) | 16.1±1.9 | (12.8-19.6) | 14.3±1.1 | (12.3-16.4) | 14.3±0.9 | (12.7-15.5) | 14.3±1.2 | (12.3-16.4) |
|
19 | 18.6±1.5 | (15.7-22.6) | 19.1±1.6 | (16.1-22.6) | 18.6±1.4 | (16.1-21.2) | 19.7±1.7 | (17.3-22.6) | 18.2±1.1 | (15.7-20.1) | 18±0.8 | (16.7-19.4) | 18.4±1.3 | (15.7-20.1) |
|
35.3 | 35.8±1.8 | (32.5-41.3) | 35.9±1.7 | (32.5-41.3) | 36.1±1.4 | (34-38.6) | 35.7±2.1 | (32.5-41.3) | 35.7±1.9 | (32.9-39.8) | 35.7±1.2 | (33.4-37.3) | 35.8±2.3 | (32.9-39.8) |
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1 | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) |
|
11 | 10 | (9-11) | 10 | (9-11) | 10 | (9-11) | 10 | (9-11) | 10 | (9-11) | 10 | (10-11) | 10 | (9-11) |
|
1 | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) |
|
16 | 16 | (14-17) | 16 | (14-17) | 16 | (14-17) | 16 | (15-17) | 16 | (14-17) | 16 | (14-17) | 16 | (14-17) |
|
4 | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) |
|
10 | 10 | (10-13) | 10 | (10-11) | 10 | (10-11) | 10 | (10-11) | 11 | (10-13) | 11 | (10-12) | 11 | (10-13) |
|
3 | 3 | (2-5) | 3 | (2-4) | 3 | (2-4) | 2 | (2-4) | 4 | (3-5) | 3 | (3-5) | 4 | (3-4) |
|
12 | 12 | (10-14) | 12 | (10-14) | 12 | (11-14) | 12 | (10-14) | 12 | (11-13) | 12 | (11-13) | 12 | (11-13) |
|
82 | 86 | (78-93) | 86 | (78-93) | 85 | (78-86) | 86 | (78-93) | 86 | (79-92) | 86 | (81-92) | 86 | (79-91) |
|
32 | 36 | (30-44) | 32 | (30-44) | 36 | (30-40) | 38 | (31-44) | 34 | (30-40) | 34 | (30-38) | 34 | (33-40) |
|
10 | 9 | (8-11) | 9 | (8-11) | 9 | (9-10) | 10 | (8-11) | 10 | (9-10) | 10 | (9-10) | 10 | (9-10) |
|
7 | 8 | (7-10) | 8 | (7-9) | 8 | (7-9) | 8 | (7-9) | 8 | (7-9) | 8 | (8-9) | 8 | (7-9) |
|
8 | 9 | (7-10) | 9 | (7-9) | 9 | (8-9) | 9 | (7-9) | 9 | (8-9) | 9 | (8-9) | 9 | (8-9) |
|
11 | 12 | (10-16) | 14 | (10-16) | 14 | (12-16) | 14 | (10-15) | 12 | (11-15) | 12 | (11-14) | 12 | (11-15) |
|
22 | 24 | (19-27) | 24 | (22-27) | 24 | (22-27) | 24 | (22-26) | 22 | (19-27) | 22 | (22-26) | 24 | (19-27) |
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33 | 38 | (32-43) | 38 | (32-43) | 38 | (34-43) | 38 | (32-40) | 35 | (32-40) | 37 | (33-38) | 39 | (32-40) |
Morphological and meristic data of
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301 | 275.3±29.4 | (211-323) | 276.5±36.9 | (234-301) | 274.9±29.2 | (211-323) |
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4.0 | 4.4±0.3 | (4.0-4.8) | 4.1±0.2 | (4.0-4.3) | 4.5±0.3 | (4.1-4.8) |
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5.7 | 6.2±0.5 | (5.3-6.9) | 6.2±0.4 | (5.7-6.5) | 6.2±0.6 | (5.3-6.9) |
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15.3 | 16.5±1.1 | (14.6-18.3) | 16.1±1.1 | (15.3-17.4) | 16.6±1.1 | (14.6-18.3) |
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3.2 | 3.4±0.3 | (3.1-3.8) | 3.2±0.2 | (3.1-3.4) | 3.4±0.3 | (3.1-3.8) |
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14.7 | 16.2±1.3 | (13.9-18.2) | 15.8±1.1 | (14.7-16.9) | 16.4±1.4 | (13.9-18.2) |
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15.2 | 17±1.3 | (15.2-19.1) | 16.2±1 | (15.2-17.2) | 17.3±1.3 | (15.5-19.1) |
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11.4 | 12.6±0.8 | (11.4-13.8) | 12.4±1.2 | (11.4-13.7) | 12.6±0.7 | (11.7-13.8) |
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16.2 | 18.8±1.7 | (16.2-21.2) | 18.4±2.3 | (16.2-20.9) | 19.0±1.6 | (16.2-21.2) |
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17.8 | 20.1±1.6 | (17.5-22.4) | 20.0±2.3 | (17.8-22.4) | 20.2±1.4 | (17.5-22.1) |
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4.8 | 4.8±0.7 | (3.9-6.3) | 4.6±0.2 | (4.4-4.8) | 4.9±0.7 | (3.9-6.3) |
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12.1 | 12.6±0.8 | (11.5-14.2) | 12.6±0.5 | (12.1-13.0) | 12.6±0.9 | (11.5-14.2) |
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11.2 | 12.4±1.0 | (10.8-13.7) | 12.3±1.3 | (11.2-13.7) | 12.4±0.9 | (10.8-13.5) |
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5 | 4.5±0.6 | (3.7-5.4) | 4.4±0.5 | (4.0-5.0) | 4.5±0.6 | (3.7-5.4) |
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23.6 | 23.4±1.2 | (22.2-25.9) | 23.7±1.2 | (22.4-24.9) | 23.3±1.3 | (22.2-25.9) |
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7.8 | 8±0.4 | (7.5-8.6) | 7.9±0.2 | (7.7-8.1) | 8±0.4 | (7.5-8.6) |
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13 | 13.0±0.7 | (11.4-14.0) | 13.5±0.5 | (13.0-14.0) | 12.9±0.7 | (11.4-13.9) |
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18.2 | 18±1 | (16.4-19.6) | 17.8±0.8 | (16.9-18.4) | 18±1.1 | (16.4-19.6) |
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13.6 | 14.0±0.7 | (13.2-15.1) | 14.3±0.7 | (13.6-14.9) | 13.9±0.7 | (13.2-15.1) |
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53.4 | 51.7±1.9 | (48.6-54.3) | 52.8±1.2 | (51.4-53.6) | 51.3±2.0 | (48.6-54.3) |
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78.1 | 77.5±0.9 | (75.0-78.4) | 78.0±0.6 | (77.4-78.4) | 77.3±0.9 | (75.0-77.9) |
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50 | 47.2±1.5 | (44.5-50) | 48.2±1.7 | (46.7-50) | 46.9±1.4 | (44.5-49.7) |
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30 | 27.0±1.5 | (24.6-30.0) | 28.0±1.8 | (26.5-30.0) | 26.7±1.3 | (24.6-28.7) |
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43.3 | 43.3±1.2 | (41.9-45.6) | 42.5±0.8 | (41.9-43.3) | 43.6±1.2 | (42-45.6) |
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120.1 | 119.6±2.3 | (115.3-122.5) | 119.5±0.6 | (118.8-120.1) | 119.6±2.7 | (115.3-122.5) |
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58.7 | 55.3±4.9 | (44.8-63.3) | 55.6±5.3 | (49.4-58.7) | 55.2±5.1 | (44.8-63.3) |
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22.2 | 23.2±1.7 | (20.5-26.3) | 23.5±1.2 | (22.2-24.6) | 23.1±1.9 | (20.5-26.3) |
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22.2 | 22±1.1 | (20.5-24.5) | 22.6±1.7 | (21.1-24.5) | 21.8±0.8 | (20.5-23) |
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25.5 | 26.2±1.2 | (24.2-27.8) | 26.3±1.3 | (25.5-27.8) | 26.2±1.2 | (24.2-27.4) |
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22.5 | 21.6±1.1 | (19.6-24.1) | 22.5±1.5 | (21-24.1) | 21.3±0.8 | (19.6-22) |
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5.1 | 4.8±0.6 | (3.9-5.6) | 4.9±0.5 | (4.3-5.2) | 4.8±0.7 | (3.9-5.6) |
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54.4 | 52.4±1.4 | (50.3-54.4) | 53±1.4 | (51.6-54.4) | 52.2±1.5 | (50.3-54.3) |
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72.1 | 72.1±2.1 | (68.9-76.3) | 72.8±0.9 | (72.1-73.8) | 71.8±2.4 | (68.9-76.3) |
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10.7 | 9.3±0.7 | (8.3-10.7) | 9.7±0.8 | (9.1-10.7) | 9.2±0.7 | (8.3-10.6) |
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27 | 27.3±1.4 | (25.2-30) | 27.3±0.7 | (26.9-28.1) | 27.3±1.6 | (25.2-30) |
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39.4 | 39±1.2 | (36.6-40.4) | 39.7±0.3 | (39.4-40) | 38.7±1.3 | (36.6-40.4) |
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19.7 | 19.7±1.2 | (18.1-21.8) | 19.4±0.7 | (18.6-19.9) | 19.8±1.3 | (18.1-21.8) |
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10.4 | 10±1.7 | (6.5-13.2) | 10.1±0.4 | (9.7-10.4) | 10±2 | (6.5-13.2) |
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15.8 | 16.7±1.1 | (15.3-18.9) | 16±0.8 | (15.3-17) | 16.9±1.1 | (15.7-18.9) |
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53.1 | 51.6±3.1 | (44.5-56.9) | 49±4.3 | (44.5-53.1) | 52.4±2.2 | (49.5-56.9) |
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29.6 | 27.6±2.3 | (23.8-31.2) | 27.9±2.2 | (25.4-29.6) | 27.5±2.4 | (23.8-31.2) |
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11.6 | 12.1±0.7 | (11-13.2) | 11.7±0.1 | (11.6-11.8) | 12.3±0.7 | (11-13.2) |
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14.3 | 11.9±1.4 | (9.7-14.3) | 12±2.3 | (9.7-14.3) | 11.9±1.1 | (10.1-13.6) |
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24.1 | 23±1.6 | (19.3-25) | 21.6±2.4 | (19.3-24.1) | 23.4±1.1 | (21.2-25) |
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11.3 | 11.5±1.7 | (9.1-14.3) | 11.3±1.1 | (10.2-12.4) | 11.5±1.9 | (9.1-14.3) |
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11.7 | 12.1±1.5 | (10-14.4) | 11.6±1.2 | (10.4-12.9) | 12.3±1.6 | (10-14.4) |
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19.6 | 18.6±0.6 | (17.8-19.8) | 18.9±0.6 | (18.4-19.6) | 18.6±0.6 | (17.8-19.8) |
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34.7 | 34.3±1.2 | (31.6-36.5) | 33.9±0.8 | (33-34.7) | 34.4±1.3 | (31.6-36.5) |
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1 | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) |
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10 | 10 | (10-12) | na | (10-12) | 10 | (10-12) |
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1 | 1 | (1-1) | 1 | (1-1) | 1 | (1-1) |
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15 | 15 | (14-16) | na | (14-16) | 15 | (15-16) |
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4 | 4 | (3-4) | 4 | (3-4) | 4 | (3-4) |
|
10 | 10 | (10-12) | 10 | (10-11) | 10 | (10-12) |
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3 | 3 | (3-3) | 3 | (3-3) | 3 | (3-3) |
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11 | 12 | (11-13) | 12 | (11-12) | 12 | (11-13) |
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78 | 78 | (78-87) | 78 | (78-80) | 85 | (78-87) |
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40 | 36 | (32-40) | na | (32-40) | 35 | (33-40) |
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9 | 10 | (8-10) | 9 | (9-10) | 10 | (8-10) |
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8 | 8 | (8-9) | 8 | (8-8) | 8 | (8-9) |
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8 | 9 | (8-9) | 8 | (8-8) | 9 | (8-9) |
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10 | 11 | (10-12) | 11 | (10-11) | 12 | (10-12) |
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20 | 20 | (19-23) | 20 | (20-21) | 21 | (19-23) |
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30 | 31 | (30-35) | na | (30-32) | 31 | (30-35) |
Morphological and meristic data of
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223.0 | 181.5±37.0 | (118–226) | 187.8±47.4 | (118–223) | 178.7±34.3 | (129–226) |
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4.1 | 3.7±0.6 | (2.8–4.8) | 3.6±0.6 | (2.8–4.2) | 3.8±0.6 | (2.9–4.8) |
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6.1 | 6.1±0.8 | (4.6–7.4) | 5.8±0.4 | (5.1–6.1) | 6.3±0.9 | (4.6–7.4) |
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15.2 | 15.6±1.1 | (14–17.5) | 15±0.5 | (14.6–15.7) | 15.9±1.2 | (14–17.5) |
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3.1 | 3.1±0.2 | (2.6–3.4) | 3.0±0.3 | (2.6–3.2) | 3.1±0.2 | (2.8–3.4) |
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16.0 | 15.9±1.6 | (13.9–20.1) | 15.4±0.7 | (14.5–16.0) | 16.2±1.8 | (13.9–20.1) |
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17.0 | 16.8±1.6 | (14–20.7) | 16.4±0.7 | (15.5–17) | 17±1.9 | (14–20.7) |
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12.3 | 11.8±0.8 | (10.4–12.9) | 11.7±0.4 | (11.3–12.3) | 11.9±1.0 | (10.4–12.9) |
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17.6 | 17.9±1.2 | (15.5–19.8) | 17.7±0.7 | (16.9–18.6) | 18.0±1.4 | (15.5–19.8) |
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18.7 | 18.6±1.5 | (15.3–20.8) | 18.3±0.4 | (17.8–18.7) | 18.8±1.8 | (15.3–20.8) |
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5.1 | 5.2±0.6 | (4.2–6.5) | 5.0±0.2 | (4.7–5.2) | 5.3±0.7 | (4.2–6.5) |
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13.7 | 12.4±0.9 | (11.1–13.7) | 12.9±1.1 | (11.4–13.7) | 12.2±0.7 | (11.1–13.6) |
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11.1 | 11.2±1.0 | (9.4–12.6) | 11.2±0.5 | (10.5–11.6) | 11.2±1.2 | (9.4–12.6) |
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5.1 | 5.5±0.8 | (4.0–7.1) | 5.2±0.3 | (5.0–5.7) | 5.6±0.9 | (4.0–7.1) |
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23.8 | 24.1±1.1 | (22.6–26.3) | 23.2±0.5 | (22.6–23.8) | 24.5±1.1 | (22.7–26.3) |
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7.3 | 7.3±0.3 | (6.7–7.7) | 7.1±0.4 | (6.7–7.5) | 7.4±0.2 | (7.1–7.7) |
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13.9 | 13.8±1.0 | (12.2–15.8) | 14.0±0.4 | (13.7–14.6) | 13.7±1.1 | (12.2–15.8) |
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18.8 | 19.1±0.7 | (17.9–20.7) | 19±0.4 | (18.6–19.4) | 19.1±0.9 | (17.9–20.7) |
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15.0 | 15.6±0.7 | (14.6–16.8) | 15.4±0.6 | (15.0–16.3) | 15.7±0.7 | (14.6–16.8) |
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48.6 | 51.1±1.7 | (47.8–54.0) | 50.9±1.8 | (48.6–52.8) | 51.2±1.8 | (47.8–54.0) |
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75.3 | 77.1±1.5 | (74.3–79.5) | 76.2±1.7 | (74.3–78.2) | 77.5±1.3 | (75.4–79.5) |
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46.2 | 47.5±1.7 | (43.9–49.4) | 47.2±1.1 | (46.2–48.2) | 47.6±2 | (43.9–49.4) |
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24.6 | 22.6±1.7 | (19.6–25.1) | 22.7±2.5 | (20.5–25.1) | 22.6±1.5 | (19.6–24.2) |
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45.9 | 44.1±1.1 | (42.4–45.9) | 44.6±1.3 | (43–45.9) | 43.9±1.1 | (42.4–45.5) |
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122.0 | 121.5±1.9 | (117.8–124.4) | 121.2±2.5 | (117.8–123.8) | 121.6±1.7 | (119.2–124.4) |
|
45.4 | 38.7±7.3 | (26.7–47.4) | 39.2±8.5 | (26.7–45.4) | 38.5±7.3 | (28.3–47.4) |
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25.6 | 23.3±1.8 | (20.5–26.3) | 23.6±2.1 | (21.1–25.6) | 23.2±1.7 | (20.5–26.3) |
|
24.4 | 25.3±1.6 | (23.1–28.3) | 25.2±1.6 | (24.2–27.6) | 25.3±1.7 | (23.1–28.3) |
|
27.8 | 29±2.3 | (25.6–32.9) | 28.8±3.1 | (25.6–32.9) | 29.1±2.1 | (26.5–32.7) |
|
22.0 | 24.4±1.4 | (22–27.2) | 23.9±1.7 | (22–26.2) | 24.7±1.3 | (23–27.2) |
|
3.5 | 4.7±1.2 | (3.3–7.2) | 4.8±1.4 | (3.5–6.5) | 4.7±1.2 | (3.3–7.2) |
|
50.9 | 50.7±1.1 | (48.2–52.3) | 49.8±1.5 | (48.2–51.1) | 51.1±0.7 | (50.3–52.3) |
|
75.2 | 68.5±3.3 | (64.4–75.2) | 69.8±4.4 | (65.2–75.2) | 67.9±2.8 | (64.4–73.1) |
|
9.7 | 9.9±0.9 | (8.5–10.9) | 9.4±0.8 | (8.5–10.3) | 10.1±0.8 | (8.6–10.9) |
|
30.2 | 29.5±1.6 | (27.1–32) | 29±1.8 | (27.1–30.8) | 29.6±1.6 | (27.3–32) |
|
42.9 | 42.2±1.5 | (40.5–45.7) | 43.2±1.7 | (42–45.7) | 41.8±1.2 | (40.5–43.7) |
|
23.4 | 21±2.4 | (15.4–24) | 21±3.9 | (15.4–24) | 21.1±1.7 | (18.3–23.8) |
|
7.0 | 8.8±1.4 | (6.4–11.6) | 7.4±0.9 | (6.4–8.6) | 9.4±1.2 | (8–11.6) |
|
18.0 | 17.8±1.2 | (15.7–20.2) | 17.6±0.6 | (16.7–18) | 17.8±1.4 | (15.7–20.2) |
|
52.1 | 48.1±3.1 | (44.1–52.4) | 48.5±4 | (44.1–52.1) | 47.9±2.9 | (44.1–52.4) |
|
28.4 | 26.2±1.9 | (22.8–30.7) | 26.3±1.7 | (25–28.4) | 26.1±2.1 | (22.8–30.7) |
|
9.7 | 11.1±0.8 | (9.7–12.6) | 10.8±0.7 | (9.7–11.2) | 11.3±0.9 | (10–12.6) |
|
14.1 | 11.5±1.2 | (10.1–14.1) | 11.9±2 | (10.1–14.1) | 11.3±0.6 | (10.5–12.4) |
|
25.9 | 23.4±1.6 | (20.2–26.1) | 23.3±2.3 | (20.2–25.9) | 23.5±1.3 | (21.4–26.1) |
|
13.5 | 13.5±1.3 | (10.9–15.1) | 13.7±1.6 | (11.6–15.1) | 13.3±1.3 | (10.9–14.9) |
|
13.9 | 14 .7±1.6 | (12.1–16.8) | 14.8±2.2 | (12.1–16.8) | 14.7±1.4 | (13–16.7) |
|
20.4 | 18.5±1.7 | (15.3–20.5) | 19.6±0.8 | (18.5–20.4) | 18±1.7 | (15.3–20.5) |
|
40.4 | 35.5±2 | (33–40.4) | 37.2±2.5 | (35–40.4) | 34.8±1.4 | (33–37.5) |
|
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|
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|
1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
|
10 | 10 | (9–11) | 10 | (9–10) | 10 | (10–11) |
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1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
|
15 | 15 | (15–17) | 15 | (15–17) | 15 | (15–17) |
|
4 | 4 | (3–4) | 4 | (4–4) | 4 | (3–4) |
|
12 | 11 | (10–13) | 11 | (10–12) | 10 | (10–13) |
|
4 | 4 | (3–4) | 4 | (4–4) | 4 | (3–4) |
|
13 | 12 | (11–13) | 13 | (11–13) | 12 | (12–12) |
|
89 | 86 | (80–91) | 89 | (80–91) | 86 | (80–88) |
|
36 | 35 | (32–40) | na | (34–37) | 32 | (32–40) |
|
9 | 9 | (7–10) | 9 | (7–9) | 9 | (8–10) |
|
8 | 8 | (7–8) | 8 | (7–8) | 8 | (7–8) |
|
8 | 8 | (8–9) | 8 | (8–8) | 8 | (8–9) |
|
14 | 14 | (11–14) | 13 | (13–14) | 12 | (11–14) |
|
25 | 24 | (20–25) | 24 | (24–25) | 23 | (20–25) |
|
39 | 37 | (32–39) | 37 | (37–39) | 32 | (32–38) |
Morphological and meristic data of
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194.0 | 223.3±26.7 | (188–316) | 248.7±42.2 | (188–316) | 216.3±16 | (188–241) |
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|
4.4 | 4.2±0.3 | (3.6–5.0) | 4.2±0.2 | (4–4.5) | 4.2±0.4 | (3.6–5) |
|
7.2 | 6.0±0.8 | (4.0–7.2) | 5.7±1 | (4–6.8) | 6.1±0.7 | (4.8–7.2) |
|
16.9 | 17.7±1.1 | (15.1–19.6) | 17.3±0.9 | (16.5–18.9) | 17.9±1.1 | (15.1–19.6) |
|
3.5 | 3.7±0.2 | (3.2–4.3) | 3.6±0.2 | (3.2–3.8) | 3.7±0.2 | (3.4–4.3) |
|
16.9 | 18.4±1.1 | (16.6–21.0) | 18.1±1.3 | (16.6–19.8) | 18.5±1 | (16.8–21) |
|
17.8 | 20.2±1.3 | (17.7–23.2) | 19.9±1.5 | (17.7–22.1) | 20.2±1.3 | (17.8–23.2) |
|
12.6 | 12.5±0.9 | (10.5–14.5) | 12.3±0.7 | (11.3–13.4) | 12.5±1 | (10.5–14.5) |
|
18.7 | 19.5±1.4 | (15.9–21.9) | 18.7±1.9 | (15.9–21.6) | 19.7±1.2 | (17–21.9) |
|
20.6 | 20.7±1.3 | (17.5–23.2) | 19.9±1.4 | (17.5–21.5) | 20.9±1.2 | (18.3–23.2) |
|
5.1 | 5.0±0.5 | (3.9–6.1) | 5.1±0.4 | (4.5–5.6) | 5±0.6 | (3.9–6.1) |
|
13.5 | 13.2±1.0 | (10.8–15.3) | 13.4±0.8 | (12.1–14.4) | 13.1±1.1 | (10.8–15.3) |
|
13.6 | 13.3±1.0 | (10.9–14.7) | 12.8±1 | (10.9–13.9) | 13.4±0.9 | (11.3–14.7) |
|
5.1 | 5.3±0.6 | (3.8–6.3) | 5.3±0.4 | (4.6–5.8) | 5.2±0.6 | (3.8–6.3) |
|
24.1 | 24.5±1.4 | (21.8–27.8) | 24.3±2 | (21.8–27.8) | 24.6±1.3 | (22.2–27.8) |
|
7.5 | 7.3±0.3 | (6.5–7.9) | 7.5±0.2 | (7.2–7.8) | 7.3±0.3 | (6.5–7.9) |
|
12.5 | 11.8±0.7 | (10.2–13.0) | 12±0.8 | (10.9–13) | 11.8±0.7 | (10.2–13) |
|
16.9 | 18.3±1.1 | (15.8–20.1) | 18.5±0.9 | (17.1–19.6) | 18.2±1.1 | (15.8–20.1) |
|
16.4 | 16.4±0.6 | (15.5–18.4) | 16.2±0.5 | (15.5–16.7) | 16.5±0.6 | (15.7–18.4) |
|
55.2 | 54.2±1.5 | (51.2–58.1) | 53.3±1.2 | (51.2–54.1) | 54.5±1.4 | (52.1–58.1) |
|
79.2 | 78.4±1.4 | (75.0–80.6) | 77.8±1.3 | (75.8–79.4) | 78.6±1.4 | (75–80.6) |
|
48.5 | 48.3±1.3 | (45.8–51.1) | 47.8±1.8 | (45.8–50) | 48.5±1.2 | (46.9–51.1) |
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24.4 | 24.2±1.4 | (22.1–27.6) | 25.4±1.3 | (24–27.6) | 23.9±1.2 | (22.1–26.6) |
|
40.6 | 42.5±1.5 | (38.9–44.5) | 43.2±1.4 | (41.3–44.5) | 42.3±1.5 | (38.9–44.4) |
|
122.2 | 121.3±1.7 | (117.3–125.6) | 120.5±1.1 | (118.9–121.8) | 121.5±1.8 | (117.3–125.6) |
|
41.2 | 48.9±5.5 | (39.8–66.2) | 54.1±8.7 | (39.8–66.2) | 47.4±3.2 | (41.2–53.7) |
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23.6 | 23.5±0.8 | (21.8–24.8) | 23.3±0.6 | (22.5–24) | 23.6±0.8 | (21.8–24.8) |
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23.3 | 23.8±1.4 | (21.3–26.2) | 23.7±1.5 | (21.9–25.7) | 23.8±1.4 | (21.3–26.2) |
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30.9 | 29.2±1.4 | (26.2–32.1) | 28.2±1.6 | (26.2–31.1) | 29.5±1.3 | (26.9–32.1) |
|
24.5 | 23.6±0.9 | (21.8–25.5) | 23.2±0.7 | (21.9–23.9) | 23.7±0.9 | (21.8–25.5) |
|
5.7 | 4.6±0.8 | (3.0–5.9) | 4.3±0.9 | (3.5–5.9) | 4.7±0.7 | (3–5.7) |
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51.1 | 50.9±1.4 | (48–54) | 52.2±1.8 | (49.2–54) | 50.6±1 | (48–52.1) |
|
78.3 | 71.8±2.8 | (65.9–78.3) | 73±2.1 | (69.4–75.7) | 71.5±2.9 | (65.9–78.3) |
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11.2 | 10±0.8 | (8.5–11.7) | 10±0.5 | (9.4–10.7) | 10±0.9 | (8.5–11.7) |
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29.1 | 28.7±1.2 | (26.4–30.6) | 28.1±1.3 | (26.4–30) | 28.9±1.1 | (26.8–30.6) |
|
41.4 | 39.9±1.7 | (37–43.6) | 39.1±1.4 | (37–40.9) | 40.1±1.8 | (37.2–43.6) |
|
24 | 20.7±1.2 | (17.3–24) | 20.1±1.4 | (17.3–21.2) | 20.8±1.1 | (18.7–24) |
|
10.1 | 10±0.8 | (8.1–11.3) | 9.7±0.6 | (8.8–10.7) | 10±0.8 | (8.1–11.3) |
|
17.6 | 15.8±1.3 | (12.5–17.8) | 15.3±1.6 | (13.7–17.3) | 16±1.1 | (12.5–17.8) |
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57.3 | 52.4±3.3 | (46.7–58.6) | 53.1±3.9 | (46.7–58.6) | 52.2±3.1 | (47.4–57.7) |
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28.7 | 28.1±1.2 | (26.1–30.3) | 28.9±1.4 | (26.5–30.3) | 27.9±1.1 | (26.1–29.5) |
|
11.1 | 11.1±1 | (8.2–13.3) | 11.7±1.1 | (10.3–13.3) | 10.9±1 | (8.2–12.5) |
|
9.3 | 11.7±2.2 | (7.8–16.2) | 12.7±0.7 | (11.4–13.6) | 11.5±2.4 | (7.8–16.2) |
|
28.9 | 26±1.7 | (22.7–29.2) | 25.2±1.5 | (22.7–27.4) | 26.2±1.7 | (22.8–29.2) |
|
10 | 9.2±1.1 | (7.6–11.7) | 9.4±1.2 | (7.6–10.9) | 9.2±1.1 | (8–11.7) |
|
10.7 | 10.1±1.2 | (7.8–12.4) | 10.5±1.6 | (7.8–12.4) | 9.9±1.1 | (8.1–12.3) |
|
19.6 | 18±1.8 | (15.5–21.8) | 18.7±2.4 | (15.5–21.8) | 17.8±1.6 | (15.5–21.2) |
|
35.8 | 34.3±1.8 | (30.3–37.7) | 35.1±2.1 | (32.9–37.7) | 34.1±1.7 | (30.3–36.6) |
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1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
|
10 | 10 | (9–11) | 10 | (10–11) | 10 | (9–11) |
|
1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
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16 | 16 | (13–17) | 16 | (16–16) | 16 | (13–17) |
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5 | 4 | (3–5) | 4 | (3–4) | 4 | (3–5) |
|
10 | 10 | (9–12) | 11 | (10–12) | 10 | (9–11) |
|
5 | 3 | (2–5) | 2 | (2–4) | 3 | (2–5) |
|
11 | 12 | (11–14) | 12 | (12–14) | 12 | (11–13) |
|
83 | 84 | (76–90) | 83 | (80–89) | 84 | (76–90) |
|
34 | 34 | (32–38) | 32 | (32–37) | 34 | (32–38) |
|
9 | 9 | (8–10) | 9 | (8–10) | 9 | (8–10) |
|
8 | 8 | (6–8) | 8 | (8–8) | 8 | (6–8) |
|
8 | 8 | (7–9) | 9 | (8–9) | 8 | (7–9) |
|
8 | 9 | (5–10) | 7 | (6–10) | 9 | (5–9) |
|
13 | 14 | (10–18) | 17 | (11–18) | 14 | (10–18) |
|
21 | 21 | (15–27) | na | (18–27) | 21 | (15–26) |
Morphological and meristic data of
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239.5 | 237.3±21.2 | (197–278) | 235.5±26.5 | (197–254) | 237.6±20.8 | (197–278) |
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|
5.1 | 4.1±0.4 | (3.5–5.1) | 3.7±0.4 | (3.5–4.3) | 4.1±0.4 | (3.5–5.1) |
|
7 | 6.2±0.7 | (4.6–7.5) | 6.2±0.3 | (5.7–6.4) | 6.2±0.7 | (4.6–7.5) |
|
17.4 | 16±0.9 | (14.3–17.5) | 15.6±1.2 | (14.6–17.2) | 16.1±0.8 | (14.3–17.5) |
|
3.6 | 3.4±0.2 | (3.1–4.0) | 3.2±0.1 | (3.1–3.4) | 3.5±0.2 | (3.1–4) |
|
17.4 | 15.9±1.1 | (13.8–18.2) | 15.6±1.8 | (14.1–18.1) | 16±1 | (13.8–18.2) |
|
18.6 | 16.9±1.3 | (15–19.7) | 16.5±2.2 | (15–19.7) | 17±1.1 | (15–19) |
|
14.8 | 12.4±0.8 | (11.2–14.8) | 12.3±0.8 | (11.5–13.4) | 12.4±0.9 | (11.2–14.8) |
|
20.9 | 18.1±1.2 | (15.7–20.9) | 17.8±1.5 | (15.8–19.1) | 18.1±1.2 | (15.7–20.9) |
|
21.7 | 19.3±1.1 | (17.0–21.7) | 19.1±1.1 | (18.0–20.3) | 19.3±1.2 | (17–21.7) |
|
5.5 | 5.0±0.5 | (4.0–6.1) | 4.9±0.5 | (4.2–5.3) | 5±0.5 | (4–6.1) |
|
13.6 | 12.6±0.6 | (11.3–13.6) | 12.6±0.6 | (11.9–13.3) | 12.6±0.6 | (11.3–13.6) |
|
13 | 11.6±0.8 | (9.2–13.0) | 11.4±0.5 | (11.0–12.2) | 11.7±0.9 | (9.2–13) |
|
4.5 | 4.7±0.7 | (3.7–6.2) | 4.8±0.6 | (4.2–5.6) | 4.7±0.7 | (3.7–6.2) |
|
24 | 23.3±0.9 | (21.5–25.1) | 23.2±0.2 | (23–23.4) | 23.4±1 | (21.5–25.1) |
|
7.5 | 7.6±0.4 | (7.1–8.3) | 7.8±0.3 | (7.6–8.2) | 7.6±0.4 | (7.1–8.3) |
|
11.7 | 12.8±0.6 | (11.7–14.2) | 12.4±0.4 | (11.9–12.8) | 12.9±0.6 | (11.7–14.2) |
|
15 | 18.1±1.2 | (15–20.1) | 17.6±1 | (16.2–18.4) | 18.2±1.2 | (15–20.1) |
|
14.9 | 15.2±0.6 | (13.8–16.1) | 14.9±0.9 | (14.0–15.9) | 15.2±0.5 | (13.8–16.1) |
|
50.3 | 52.6±1.6 | (49.1–56.8) | 51.9±0.5 | (51.4–52.4) | 52.7±1.8 | (49.1–56.8) |
|
78.5 | 77.7±1.2 | (75.3–80.3) | 77.1±0.5 | (76.5–77.6) | 77.8±1.3 | (75.3–80.3) |
|
45.4 | 47.5±1.4 | (45–50.7) | 47.5±1.1 | (46.3–48.6) | 47.5±1.4 | (45–50.7) |
|
25.6 | 24.7±1.6 | (20.7–28.1) | 26.1±1.6 | (24.4–28.1) | 24.4±1.5 | (20.7–26.7) |
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41.2 | 43±1.3 | (40.3–45.6) | 42.2±1.6 | (41–44.3) | 43.1±1.3 | (40.3–45.6) |
|
123.2 | 120.6±1.7 | (116–123.2) | 119±2.5 | (116–121.5) | 120.8±1.4 | (118.2–123.2) |
|
49 | 49.9±4 | (41.5–58.4) | 48.5±4.7 | (41.5–51.3) | 50.1±3.9 | (41.5–58.4) |
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23.4 | 23.9±1.4 | (20.5–27) | 22.6±1.8 | (20.5–24.6) | 24.1±1.3 | (21.6–27) |
|
23.2 | 23.7±0.8 | (21.6–25.5) | 23.8±0.8 | (22.6–24.4) | 23.7±0.9 | (21.6–25.5) |
|
27.4 | 27.7±1 | (26–29.6) | 28.6±1 | (27.2–29.6) | 27.6±0.9 | (26–28.8) |
|
22.8 | 22.9±0.9 | (21.7–24.8) | 23.6±1.1 | (22.2–24.8) | 22.8±0.8 | (21.7–24.5) |
|
4.9 | 4.7±0.8 | (3.2–6.4) | 5.6±0.9 | (4.8–6.4) | 4.5±0.6 | (3.2–6.1) |
|
51 | 50.9±1.5 | (48.5–54.1) | 52±1.8 | (49.8–54.1) | 50.7±1.4 | (48.5–53) |
|
69.8 | 69.1±2.4 | (65.1–74.9) | 69.8±2 | (67.8–72.5) | 68.9±2.5 | (65.1–74.9) |
|
9.7 | 9.8±0.7 | (8.1–11.4) | 9.8±0.6 | (8.8–10.3) | 9.8±0.8 | (8.1–11.4) |
|
28.8 | 29.4±1.2 | (26.7–30.9) | 30.1±0.8 | (29.3–30.8) | 29.2±1.3 | (26.7–30.9) |
|
40.5 | 40.9±1.7 | (38.6–47) | 40.5±1.1 | (39–41.5) | 41±1.8 | (38.6–47) |
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21.1 | 21.8±1 | (19.4–23.8) | 21.9±0.9 | (21.3–23.2) | 21.8±1 | (19.4–23.8) |
|
9.8 | 8.6±1.3 | (6–11.3) | 9±1.1 | (7.9–10.5) | 8.6±1.3 | (6–11.3) |
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17.1 | 16±1.5 | (13.1–18.1) | 16.2±1.3 | (14.7–17.6) | 15.9±1.5 | (13.1–18.1) |
|
53.8 | 49.6±3.2 | (43.9–56.2) | 51.1±2.4 | (48.2–53.8) | 49.4±3.3 | (43.9–56.2) |
|
25.8 | 27±2.1 | (21.3–31.5) | 27.1±1.4 | (25.1–28.1) | 27±2.3 | (21.3–31.5) |
|
10.8 | 11.7±1 | (9.5–13.4) | 11.7±1.2 | (10.5–13.3) | 11.7±1 | (9.5–13.4) |
|
12.4 | 12.2±1.2 | (9.2–14.3) | 12.5±1 | (11–13.4) | 12.1±1.2 | (9.2–14.3) |
|
24 | 22.8±2.1 | (18.2–27.5) | 23.7±2.7 | (20.4–26.3) | 22.6±2 | (18.2–27.5) |
|
13.1 | 13.4±1.6 | (9.1–16.6) | 14.4±1.9 | (11.9–16.6) | 13.2±1.6 | (9.1–15.1) |
|
15 | 14.5±1.4 | (11.4–16.9) | 15.7±1.2 | (14.4–16.9) | 14.3±1.3 | (11.4–16.3) |
|
19.7 | 18.1±1.6 | (13.5–20.3) | 18.5±1.6 | (16.2–19.8) | 18.1±1.6 | (13.5–20.3) |
|
36.7 | 34.9±1.7 | (32.3–38.9) | 35.9±2.1 | (34–38.9) | 34.8±1.7 | (32.3–38.4) |
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1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
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11 | 10 | (9–12) | 10 | (10–11) | 11 | (9–12) |
|
1 | 1 | (1–1) | 1 | (1–1) | 1 | (1–1) |
|
15 | 15 | (13–16) | 16 | (15–16) | 15 | (13–16) |
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3 | 4 | (3–4) | 3 | (3–4) | 4 | (3–4) |
|
12 | 10 | (9–12) | 11 | (10–12) | 10 | (9–12) |
|
3 | 3 | (2–4) | 3 | (3–3) | 3 | (2–4) |
|
13 | 11 | (11–13) | 12 | (11–13) | 11 | (11–13) |
|
80 | 84 | (79–88) | 85 | (84–85) | 80 | (79–88) |
|
34 | 34 | (33–42) | 34 | (34–41) | 35 | (33–42) |
|
10 | 10 | (9–10) | 10 | (9–10) | 10 | (9–10) |
|
8 | 8 | (8–9) | 8 | (8–8) | 8 | (8–9) |
|
9 | 8 | (8–9) | 9 | (8–9) | 8 | (8–9) |
|
13 | 13 | (10–15) | na | (10–15) | 13 | (10–14) |
|
20 | 24 | (20–26) | 24 | (21–24) | 24 | (20–26) |
|
33 | 36 | (30–40) | 35 | (34–36) | 36 | (30–40) |
Types of the Lake Constance species, Switzerland
The first- and second-best ratios retrieved from the LDA ratio extractor of either head or body characters (see Table
Characters | Species comparison | Best ratios | Range species 1 | Range species 2 | Standard distance | δ (Shape vs. size) |
---|---|---|---|---|---|---|
head + body | 1: CD/UJ * | 0.96–1.29 | 1.36–1.65 | 18.09 | 0.1 | |
2: AdFB/ES | 5.54–13.54 | 2.87–5.94 | 17.49 | 0.1 | ||
body | 1: CD/DHL * | 0.44–0.54 | 0.54–0.62 | 5.98 | 0.26 | |
2: DFB/AdFB | 1.6–2.66 | 2.31–4.02 | 4.86 | 0.31 | ||
head | 1: UJ/ES | 6.48–16.01 | 4.52–7.93 | 6 | 0.22 | |
2: HL/UJ | 2.88–3.47 | 3.33–4.11 | 5.26 | 0.25 | ||
head + body | 1: TL/EH | 21.93–27.57 | 21.91–29.46 | 3.07 | 0.08 | |
2: PelvF/PecF1 | 0.87–1.03 | 0.87–1.07 | 2.89 | 0.08 | ||
body | 1: PecF1/TL | 0.12–0.16 | 0.11–0.15 | 1.4 | 0.19 | |
2: DFAe/DFAd | 0.88–1 | 0.9–0.98 | 1.18 | 0.21 | ||
head | 1: EH/HL | 0.21–0.26 | 0.2–0.25 | 1.63 | 0.08 | |
2: LJW/ES | 2.14–7.79 | 1.59–8.18 | 1.35 | 0.1 | ||
head + body | 1: CD/UJ * | 0.96–1.29 | 1.36–1.55 | 13.8 | 0.12 | |
2: AdFB/ES | 5.54–13.54 | 3.31–6.31 | 13 | 0.12 | ||
head | 1: HL/UJ | 2.88–3.47 | 3.33–3.97 | 5.3 | 0.21 | |
2: LJ/ES | 8.25–20.33 | 6.65–12.45 | 4.59 | 0.23 | ||
head + body | 1: CL/EC * | 1.97–2.87 | 1.56–2.09 | 13.19 | 0.03 | |
2: DHL/M | 2.77–3.53 | 3.21–3.79 | 12.79 | 0.03 | ||
body | 1: CL/DHL | 0.75–1.04 | 0.61–0.82 | 4.43 | 0.06 | |
2: CD/BD | 0.26–0.31 | 0.28–0.34 | 3.38 | 0.07 | ||
head | 1: EC/UJ | 0.74–0.95 | 0.87–1.05 | 5.02 | 0.1 | |
2: SW/ES | 3.22–9.67 | 2.31–5.26 | 4.05 | 0.12 | ||
head + body | 1: AdFB/ES | 5.54–13.54 | 3.31–6.5 | 9.13 | 0.06 | |
2: CD/UJW | 1.14–1.79 | 1.4–2 | 8.69 | 0.07 | ||
body | 1: PecF1/CD | 2.13–2.76 | 1.8–2.39 | 4.5 | 0.11 | |
2: DFB/AdFB | 1.6–2.66 | 1.98–3.45 | 3.88 | 0.13 | ||
head | 1: UJW/ES | 5.27–13.65 | 3.22–7.96 | 4.19 | 0.14 | |
2: ED/UJ | 0.66–0.84 | 0.74–0.9 | 3.51 | 0.16 | ||
|
|
|
|
|
|
|
head + body | 1: CD/PostD * | 0.17–0.21 | 0.14–0.17 | 22.73 | 0.07 | |
2: DFAe/UJ | 3.14–3.93 | 2.43–3.41 | 22.33 | 0.07 | ||
body | 1: CD/PostD * | 0.17–0.21 | 0.14–0.17 | 8.98 | 0.17 | |
2: DFAe/DHL | 1.26–1.55 | 1.02–1.36 | 7.9 | 0.19 | ||
head | 1: HD/UJ | 2.34–2.9 | 2.13–2.57 | 3.86 | 0.3 | |
2: MW/ES | 1.47–3 | 1.82–6.16 | 3.15 | 0.34 | ||
head + body | ED/EC | 0.74–0.9 | 0.74–0.9 | 8.07 | 0.05 | |
CD/CL | 0.6–0.75 | 0.54–0.7 | 8.02 | 0.05 | ||
body | 1: DFAe/AFAe | 1.5–1.83 | 1.43–1.62 | 5.7 | 0.06 | |
2: PelvFS/DFAe | 0.24–0.36 | 0.29–0.37 | 5.58 | 0.07 | ||
head | 1: EC/SW | 1.47–2.13 | 1.43–1.7 | 2.45 | 0.16 | |
2: ED/EC | 0.74--0.9 | 0.79–0.9 | 2.2 | 0.18 | ||
head + body | 1: CD/DHL * | 0.54–0.62 | 0.4–0.49 | 19.86 | 0.07 | |
2: PecF2/CF | 0.63–0.82 | 0.74–0.90 | 19.01 | 0.07 | ||
body | 1: CD/DHL * | 0.54–0.62 | 0.4–0.49 | 9.31 | 0.15 | |
2: PecF2/CF | 0.63–0.82 | 0.74–0.90 | 7.32 | 0.19 | ||
head | 1: EH/PostD | 0.09–0.11 | 0.11–0.15 | 4.32 | 0.21 | |
2: SD/UJW | 0.35–0.51 | 0.30–0.44 | 3.93 | 0.23 | ||
head + body | 1: CD/LJ | 0.95–1.11 | 0.79–1 | 65.21 | 0.02 | |
2: CF/M * | 5.55–6.55 | 4.4–5.57 | 65.13 | 0.02 | ||
body | 1: CD/DHL | 0.54–0.62 | 0.46–0.58 | 4.69 | 0.25 | |
2: DFAe/DFPe | 3.39–4.72 | 2.84–4.54 | 3.91 | 0.29 | ||
head | 1: PostO/M | 2.4–3 | 2.17–2.56 | 4.26 | 0.21 | |
2: HD/MW | 6.57–8.7 | 6.02–8.87 | 3.65 | 0.24 | ||
|
|
|
|
|
|
|
head + body | 1: CD/UJ * | 1.02–1.34 | 1.36–1.55 | 33.96 | 0.04 | |
2: PelvF/PAdC | 0.73–1 | 0.84–0.1 | 33.71 | 0.04 | ||
body | 1: CD/PostD * | 0.14–0.17 | 0.17–0.20 | 6.34 | 0.22 | |
2: DHL/BD | 0.5–0.7 | 0.45–0.58 | 5.37 | 0.25 | ||
head | 1: HD/UJ | 2.13–2.57 | 2.42–2.83 | 4.41 | 0.23 | |
2: HW/LJW | 3.17–6.12 | 3.72–5.1 | 3.25 | 0.29 | ||
head + body | 1: CL/EC | 1.84–2.98 | 1.56–2.09 | 10.03 | 0.02 | |
2: DHL/UJ | 2.11–2.70 | 2.32–2.92 | 9.54 | 0.02 | ||
body | 1: CL/DHL | 0.76–1.04 | 0.61–0.82 | 4.44 | <0.01 | |
2: DFPe/CD | 0.56–0.87 | 0.56.0.82 | 3.2 | <0.01 | ||
head | 1: EC/SW | 1.32–1.73 | 1.63–2.38 | 5.05 | 0.08 | |
2: UJ/UJW | 1.04–1.50 | 1–1.29 | 4.28 | 0.09 | ||
head + body | 1: CD/PostD | 0.14–0.17 | 0.16–0.2 | 8.3 | 0.05 | |
2: ED/SW | 1.08–1.5 | 1.3–1.79 | 8 | 0.05 | ||
body | 1: CD/PostD | 0.14–0.17 | 0.16–0.2 | 3.66 | 0.07 | |
2: AFAe/DHL | 0.69–0.9 | 0.61–0.93 | 2.93 | 0.09 | ||
head | 1: ED/SW | 1.08–1.5 | 1.3–1.79 | 3.05 | 0.15 | |
2: MW/ES | 1.82–6.16 | 1.4–3.02 | 2.45 | 0.18 | ||
|
|
|
|
|
|
|
head + body | 1: CD/DHL * | 0.53–0.63 | 0.4–0.49 | 23.9 | 0.05 | |
2: CL/IOW | 2.05–2.69 | 1.69–2.28 | 23.3 | 0.05 | ||
body | 1: CD/DHL * | 0.53–0.63 | 0.4–0.49 | 9.13 | 0.14 | |
2: PecF2/DFAe | 0.76–0.96 | 0.82–1.21 | 7.44 | 0.17 | ||
head | 1: SW/UJW | 0.65–0.80 | 0.54–0.69 | 5.9 | 0.12 | |
2: EH/PostO | 0.36–0.47 | 0.41–0.52 | 5.37 | 0.13 | ||
head + body | 1: CD/M * | 1.86–2.24 | 1.4–1.9 | 160.64 | <0.01 | |
2: PostD/LJ | 4.96–5.9 | 4.65–5.43 | 160.6 | <0.01 | ||
body | 1: CD/DHL | 0.53–0.63 | 0.46–0.58 | 4.46 | 0.23 | |
2: PelvF/DHL | 1.08–1.26 | 0.95–1.16 | 3.83 | 0.26 | ||
head | 1: ED/HD | 0.29–0.33 | 0.31–0.37 | 4.54 | 0.13 | |
2: HL/M | 4.6–5.53 | 4.21–5.17 | 3.41 | 0.17 | ||
|
|
|
|
|
|
|
head + body | 1: PecF2/CD | 2.37–3.16 | 1.91–2.59 | 13.46 | 0.01 | |
2: LJ/UJW | 1.34–1.86 | 1.54–2.27 | 13.12 | 0.01 | ||
body | 1: PecF2/CD | 2.37–3.16 | 1.91–2.59 | 4.58 | 0.05 | |
2: DHL/TL | 0.13–0.15 | 0.12–0.13 | 3.48 | 0.06 | ||
head | 1: M/UJW | 0.69–0.94 | 0.8–1.22 | 4.45 | 0.02 | |
2: EC/LJ | 0.66–0.81 | 0.58–0.72 | 3.88 | 0.02 | ||
|
|
|
|
|
|
|
head + body | 1: CD/UJ * | 1.36–1.65 | 0.96–1.43 | 5.34 | 0.24 |
(a) PelvFS, PecF1, DFAd, DFAe, DFPe, TL, SL, EH, SD, SW, INW, IOW (b) PelvFS, PecF1, DFAd, TL (c) PelvFS, PecF1, DFAd, TL,EH (d) PelvFS, PecF1, DFAd, TL, EH, ES, EC
The first- and second-best ratios retrieved from the LDA ratio extractor of either head or body characters (see Table
Characters | Species comparison | Size range | Best ratios | Range species 1 | Range species 2 | Standard distance | δ (Shape vs. size) |
---|---|---|---|---|---|---|---|
head + body |
|
<163.5mm | 1: PreA/LJ * | 6.33-7.44 | 9.24-9.97 | 27.13 | 0.04 |
2: AFAe/M | 1.65-2.25 | 2.58-2.63 | 25.94 | 0.04 | |||
body |
|
<163.5mm | 1: PecF2/DFAd * | 0.81-1.06 | 0.78-0.8 | 9.97 | 0.14 |
2: DHL/PreD | 0.34-0.42 | 0.32-0.34 | 9.4 | 0.15 | |||
head |
|
<163.5mm | 1: HD/LJ | 1.30-1.55 | 1.77-1.92 | 15.43 | 0.02 |
2: IOW/UJW | 0.89-1.30 | 1.20-1.26 | 14.14 | 0.02 | |||
head + body |
|
<163.5mm | 1: PecF2/PreA * | 0.22-0.28 | 0.2-0.22 | 5.78 | 0.16 |
2: DHL/PreP | 0.31-0.38 | 0.30-0.32 | 4.49 | 0.2 | |||
body |
|
<163.5mm | 1: PecF2/PreA * | 0.22-0.28 | 0.2-0.22 | 6.76 | 0.17 |
2: DHL/TL | 0.13-0.18 | 0.13-0.14 | 5.7 | 0.19 | |||
head |
|
<163.5mm | 1: UJ/ES * | 6.81-12.42 | 4.51-6.15 | 8.63 | 0.12 |
2: EH/HL * | 0.27-0.31 | 0.23-0.27 | 7.3 | 0.14 | |||
head + body |
|
<163.5mm | 1: PreD/LJ * | 3.99-4.68 | 5.05-5.57 | 47.9 | 0.01 |
2: M/ES * | 5.35-9.76 | 3.31-4.37 | 47.63 | 0.01 | |||
body |
|
<163.5mm | 1: PecF2/PreD * | 0.36-0.45 | 0.29-0.32 | 15.95 | 0.06 |
2: DHL/TL | 0.13-0.18 | 0.13-0.14 | 9.91 | 0.05 | |||
head |
|
<163.5mm | 1: LJ/ES * | 9.62-17.28 | 6.01-6.49 | 12.51 | 0.05 |
2: HL/UJ | 2.87-3.5 | 3.19-3.6 | 8.87 | 0.04 | |||
|
|
|
|
|
|
|
|
head + body |
|
>163.5mm | 1: AFAe/UJ * | 1.96-2.5 | 1.66-1.96 | 26.08 | 0.04 |
2: CL/PreA | 0.14-0.18 | 0.17-0.21 | 26.46 | 0.04 | |||
body |
|
>163.5mm | 1: AFae/TL | 0.1-0.11 | 0.09-0.1 | 13.41 | 0.11 |
2: CL/PreA | 0.14-0.18 | 0.17-0.21 | 13.41 | 0.11 | |||
head |
|
>163.5mm | 1: HL/UJ * | 3.55-3.93 | 3.13-3.55 | 11.51 | 0.07 |
2: LJW/UJW | 0.33-0.44 | 0.38-0.55 | 11.02 | 0.07 | |||
head + body |
|
>163.5mm | 1: CD/SW * | 2.25-2.64 | 1.82-2.04 | 34.25 | 0.02 |
2: LJW/UJW * | 0.33-0.44 | 0.45-0.55 | 33.91 | 0.02 | |||
body |
|
>163.5mm | 1: DFAe/PAdC * | 1.11-1.32 | 0.96-1.16 | 18.53 | 0.07 |
2: CD/AFB | 0.61-0.68 | 0.52-0.62 | 18.31 | 0.07 | |||
head |
|
>163.5mm | 1: LJW/UJW * | 0.33-0.44 | 0.45-0.55 | 7.44 | 0.08 |
2: PostO/UJ | 1.8-2.12 | 1.57-1.86 | 6.78 | 0.08 | |||
|
|
|
|
|
|
|
|
head + body |
|
>163.5mm | 1: CL/PAdC | 0.71-0.86 | 0.66-0.76 | 7.46 | 0.08 |
2: BD/LJ | 2.44-3.05 | 2.31-2.82 | 7.36 | 0.08 | |||
body |
|
>163.5mm | 1: CL/PAdC | 0.71-0.86 | 0.66-0.76 | 6.04 | 0.1 |
2: CF/BD | 0.87-1.13 | 0.93-1.12 | 5.92 | 0.1 | |||
head |
|
>163.5mm | 1: ED/M | 1.03-1.19 | 1.04-1.57 | 3.58 | 0.18 |
2: HW/UJW | 1.89-2.23 | 1.88-2.33 | 3.38 | 0.19 | |||
|
|
|
|
|
|
|
|
head + body |
|
<163.5mm | 1: DFAd/LJ * | 2.57-2.58 | 1.6-2.1 | 23.47 | 0.03 |
2: AdFB/PAdC | 0.26-0.28 | 0.21-0.42 | 22.66 | 0.03 | |||
head + body |
|
<163.5mm | 1: PostD/EH * | 5.47-6.93 | 7.5-8.9 | 48.36 | 0.02 |
2: UJW/ES | 4.88-9.3 | 3.41-5.31 | 48.13 | 0.02 | |||
head + body |
|
>163.5mm | 1: DFAe/UJ * | 3.28-4.1 | 2.58-3.19 | 24.71 | 0.05 |
2: CD/SW * | 2.25-2.64 | 1.76-2.27 | 24.37 | 0.05 | |||
|
|
|
|
|
|
|
|
head + body |
|
100-290 | 1: PreD/LJ * | 3.99-4.68 | 5.6-6.81 | 22.86 | 0.13 |
2: DFAe/UJ * | 2.14-2.79 | 3.25-4.1 | 21.65 | 0.14 | |||
head |
|
100-290 | 1: HD/UJ * | 1.87-2.2 | 2.38-2.78 | 14.39 | 0.18 |
2: LJ/IOW * | 1.53-1.99 | 1.33-1.57 | 13.25 | 0.19 | |||
head + body |
|
100-290 | 1: PreP/EH * | 6.56-7.98 | 8.94-11.43 | 15.95 | 0.13 |
2: CL/UJ | 1.44-2.02 | 1.93-2.72 | 15.09 | 0.14 | |||
head + body |
|
100-290 | 1: PreD/EH * | 6.1-7.58 | 8.12-10.32 | 50.86 | 0.04 |
2: CL/LJ | 0.99-1.45 | 1.38-1.65 | 50.6 | 0.04 | |||
head |
|
100-290 | 1: EH/HL * | 0.27-0.31 | 0.22-0.27 | 9.33 | 0.18 |
2: LJ/ES | 9.62-17.28 | 6.08-12.43 | 8.57 | 0.22 | |||
|
|
|
|
|
|
|
|
head + body |
|
100-290 | 1: DFAd/LJW * | 9.84-14.82 | 6.05-8.91 | 20.72 | 0.03 |
2: DHL/LJ * | 1.84-2.22 | 1.63-1.82 | 20.47 | 0.02 | |||
body |
|
100-290 | 1: PecF2/DFAd * | 0.74-0.85 | 0.85-1.03 | 87.52 | <0.01 |
2: CD/PostD | 0.17-0.2 | 0.15-0.18 | 87.48 | <0.01 | |||
head |
|
100-290 | 1: HD/LJW * | 6.72-9.39 | 5.23-6.66 | 11.94 | 0.04 |
2: HL/LJ * | 2.54-2.96 | 2.19-2.47 | 11.61 | 0.04 | |||
head + body |
|
100-290 | 1: DFAe/UJ * | 3.25-4.1 | 2.45-3.17 | 18.98 | 0.03 |
2: PecF2/AFAe | 1.24-1.47 | 1.37-1.63 | 18.63 | 0.03 | |||
body |
|
100-290 | 1: PecF2/DFAe | 0.77-0.89 | 0.87-1.02 | 9.25 | 0.08 |
2: AFAe/PostD | 0.27-0.32 | 0.22-0.29 | 8.71 | 0.08 | |||
head |
|
100-290 | 1: LJW/UJW | 0.33-0.47 | 0.37-0.55 | 5.62 | 0.08 |
2: HL/UJ | 3.43-3.93 | 3.13-3.63 | 4.98 | 0.08 | |||
|
|
|
|
|
|
|
|
head + body |
|
100-290 | 1: AFB/BD | 0.45-0.67 | 0.44-0.58 | 28.19 | <0.01 |
2: PreD/M | 8.87-14.85 | 9.13-11.41 | 28.16 | <0.01 | |||
body |
|
100-290 | 1: AFB/BD | 0.45-0.67 | 0.44-0.58 | 2.76 | 0.05 |
2: PreP/PreA | 0.58-0.65 | 0.57-0.64 | 2.51 | 0.05 | |||
head |
|
100-290 | 1: ED/M | 1.04-1.57 | 1.03-1.28 | 1.72 | 0.1 |
2: SN/MW | 2.09-2.63 | 1.78-2.87 | 1.52 | 0.1 | |||
|
|
|
|
|
|
|
|
head + body |
|
100-290 | 1: PreD/EH * | 6.1-7.58 | 8.12-10.5 | 10.89 | 0.16 |
2: CL/UJ | 1.44-2.02 | 1.85-2.72 | 9.79 | 0.17 | |||
head + body |
|
100-290 | 1: DFAe/UJ * | 3.25-4.1 | 2.14-3.19 | 9.59 | 0.11 |
2: LJW/UJW | 0.33-0.47 | 0.34-0.55 | 8.98 | 0.12 |
(a) PelvS, PecF1, DFAd, DFAe, DFPe, TL, SL, EH, SD, SW, INW, IOW (b) PelvFS, PelvFB, PecFB, DFPe, TL, EH, ED, SD, IOW (c) PelvFB, PelvFS, PelvF, PecF1, DFB, DFAe, DFPe, AFB, AFAe, AdFB, CF, PAdC, PreP, PreA, SL, TL, BD, PostD, DHL, ED, EH, ES, PostO, HD, MW, SN, SD, SW, IOW, INW (d) PecFB, PelvFB, PelvF, PelvFS, PecF1, DFAe, DFPe, AFAe, AdFB, PreP, CF, TL, PostD (e) ED, EH, HD, SD, SW, INW (f) PelvFB, PelvFS, PelvF, PecFB, DFAe, DFAd, DFPe, CF, PreP, SL, TL, ED, EH, MW, SD, SW, IOW, INW, ES (g) PelvFS, PecF1, PecFB, DFAd, DFPe, SL, TL, ED, EH, INW, CF (h) PelvS, PecF1, DFAd, DFAe, DFPe, TL, SL, EH, SD, SW, INW, IOW (i) PelvFS (j) PelvFB, PelvFS, PelvF, PecF1,DFAe, DFPe, AFB, AFAe,CF, SL, TL, BD, ED, EH, ES, PostO, HD, MW, SN, SD, SW, IOW, INW (k) PelvFS, PecF1, CF, SL, TL (l) PelvS, TL, INW
Morphological and meristic data of
Morphological characters |
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Non-types (N=10) | Holotype | Paratypes (N=3) | Syntypes (N=7) | Non-type | |||||||
Mean±StDev | Range | Mean±StDev | Range | Mean±StDev | Range | ||||||
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10 | 220.4±36.8 | (169–292) | 296.0 | 3 | 301.3±12.5 | (289–314) | 7 | 213.9±12.4 | (193–235) | 301 |
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10 | 4.1±0.2 | (3.7–4.4) | 3.9 | 3 | 4.4±0.3 | (3.9–4.6) | 7 | 3.8±0.3 | (3.3–4.2) | 3.8 |
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10 | 6.1±0.4 | (5.3–6.7) | 5.4 | 3 | 5.7±0.5 | (5.2–6.1) | 7 | 5.7±0.7 | (4.8–6.9) | 6.5 |
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10 | 17.1±1.2 | (15.4–19.1) | 17.3 | 3 | 17.3±0.7 | (16.8–18.1) | 7 | 16.5±0.9 | (15.2–17.6) | 15.4 |
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10 | 3.4±0.3 | (2.9–3.9) | 3.3 | 3 | 3.4±0.2 | (3.2–3.5) | 7 | 3.2±0.4 | (2.8–3.9) | 3 |
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10 | 16.8±1.1 | (14.8–18.9) | 14.4 | 3 | 16.8±0.5 | (14.4–17.2) | 7 | 16.4±1.2 | (15.1–18.1) | 16 |
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10 | 18.2±1.5 | (16.8–20.3) | 15.7 | 3 | 17.4±0.8 | (15.7–18) | 7 | 17.1±1 | (15.6–18.4) | 17 |
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10 | 11.9±0.7 | (10.7–12.8) | 12.2 | 3 | 12.2±1.1 | (11.0–13.1) | 7 | 11.6±0.7 | (10.8–12.4) | 11.2 |
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10 | 19.3±1.3 | (17.6–21.6) | 18.9 | 3 | 19.2±1.2 | (18.0–20.3) | 7 | 18.2±1.2 | (16.6–19.6) | 16.6 |
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10 | 20.4±1.1 | (19.0–22.2) | 20.2 | 3 | 20.5±1.3 | (19.3–21.9) | 7 | 19.2±1.3 | (17.2–20.5) | 18.2 |
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10 | 5.5±0.7 | (4.8–7.0) | 5.2 | 3 | 5.5±0.2 | (5.2–5.7) | 7 | 5.2±0.6 | (4.4–5.9) | 4.6 |
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10 | 12.4±0.8 | (11.4–13.4) | 11.9 | 3 | 11.5±1.1 | (10.7–12.7) | 7 | 12.3±1.3 | (10.6–14.2) | 12.5 |
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10 | 12.3±1.0 | (10.7–13.9) | 13.2 | 3 | 13.3±0.5 | (12.9–13.8) | 5 | 12.1±1.3 | (10.8–13.9) | 11.1 |
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10 | 5.6±0.4 | (4.9–6.1) | 5.7 | 3 | 5.0±1.3 | (3.7–6.2) | 7 | 5.3±0.3 | (4.9–5.8) | 4 |
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9 | 23.2±1.9 | (20.8–25.6) | na | 2 | 24±0.1 | (24–24.1) | 3 | 22.6±1.2 | (21.8–24) | 23.8 |
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10 | 7.4±0.4 | (6.7–8.2) | 7.7 | 3 | 8.1±0.1 | (7.7–8.2) | 7 | 7.4±0.4 | (6.9–8) | 7.4 |
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10 | 12.9±0.8 | (11.5–13.9) | 14.4 | 3 | 12.9±0.8 | (12.0–14.4) | 7 | 13.8±1.3 | (12.4–16.5) | 13.2 |
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10 | 18.5±0.6 | (17.4–19.3) | 19.6 | 3 | 17.2±2.2 | (14.6–19.6) | 7 | 18.9±1.1 | (17.6–20.2) | 17.8 |
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10 | 16.8±0.8 | (15.4–18.1) | 15.1 | 3 | 15.1±0.2 | (14.8–15.3) | 7 | 15.7±0.8 | (14.4–16.5) | 14.5 |
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10 | 52.7±1.4 | (50.4–54.1) | 49.5 | 3 | 50.6±0.5 | (49.5–51.0) | 7 | 51.7±1.7 | (48.1–53.1) | 50.7 |
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10 | 77.9±1.4 | (76.0–80.4) | 75.0 | 3 | 79.2±0.9 | (75.0–80.3) | 7 | 76.8±0.9 | (75.7–78.3) | 77.4 |
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10 | 48.4±0.9 | (46.8–49.6) | 47.9 | 3 | 49.2±0.5 | (47.9–49.6) | 7 | 47±1 | (45.8–48.5) | 47.3 |
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10 | 25.9±1.9 | (22.9–29.6) | 24.4 | 3 | 26.2±0.8 | (24.4–27.1) | 7 | 23.5±1.9 | (21.0–26.9) | 23.5 |
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10 | 43±1.5 | (40.6–45.4) | 44.8 | 3 | 42.2±1.7 | (40.4–44.8) | 7 | 43±1.4 | (41.6–45.7) | 44 |
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9 | 120.2±3 | (115.1–124.3) | na | 2 | 121.5±2.9 | (119.4–123.5) | 3 | 119.2±0.7 | (118.9–120) | 120.6 |
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10 | 50.3±7.1 | (41.6–62.4) | 61.8 | 3 | 63.7±3.9 | (59.6–67.2) | 7 | 47.5±3.2 | (42.6–51.3) | 64.7 |
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10 | 22.4±0.7 | (21.1–23.1) | 23.4 | 3 | 23.4±1.6 | (21.6–24.6) | 7 | 21.7±2.7 | (18–25.6) | 24 |
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10 | 21.1±1.4 | (19.4–23) | 19.6 | 3 | 17.7±0.4 | (17.3–19.6) | 7 | 24.1±1.7 | (21.3–26.1) | 18.9 |
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10 | 26.9±1.2 | (25.4–29.3) | 25.7 | 3 | 25±0.8 | (24.1–25.7) | 7 | 28.9±2 | (25.4–30.8) | 23.9 |
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10 | 21.3±0.6 | (20.5–22.6) | 20.8 | 3 | 19.6±0.9 | (18.8–20.8) | 7 | 23.2±2.1 | (19.5–25.6) | 19 |
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10 | 4.8±0.7 | (3.5–5.6) | 4.9 | 3 | 5.2±0.5 | (4.6–5.5) | 7 | 3.9±0.8 | (2.7–4.6) | 5.1 |
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10 | 52.8±1 | (51.5–54.4) | 55.7 | 3 | 54±1 | (53–55.7) | 7 | 50.2±1.9 | (48.5–53.2) | 53.4 |
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10 | 74.2±3.2 | (69.9–80.6) | 68.2 | 3 | 72.6±3 | (68.2–75) | 7 | 68.6±4.8 | (61.6–76.3) | 67.6 |
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10 | 9.8±0.6 | (9–11.2) | 10.4 | 3 | 10.5±0.5 | (10–11) | 6 | 10.1±0.8 | (8.7–11.1) | 10.6 |
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10 | 26.8±1.2 | (24.6–29) | 27.2 | 3 | 29.3±1 | (27.2–30.1) | 7 | 30.3±2.3 | (26.7–33.8) | 28.8 |
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10 | 36.6±1.4 | (34.3–39.1) | 37.8 | 3 | 38.7±0.6 | (37.8–39.1) | 7 | 42.2±2 | (40–44.4) | 43.5 |
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10 | 18.9±1.3 | (17.3–21.7) | 21.1 | 3 | 19.7±0.9 | (18.6–21.1) | 5 | 23.1±1.9 | (20.1–24.7) | 22 |
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10 | 10.2±0.8 | (9.3–11.9) | 9.7 | 3 | 10.9±1.3 | (9.7–12.3) | 7 | 7.4±1.2 | (5.5–9.5) | 6.8 |
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10 | 15.1±1.6 | (12.3–17.6) | 14.9 | 3 | 17.8±0.7 | (14.9–18.5) | 7 | 15.6±1.2 | (14.1–17.4) | 15 |
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10 | 56.1±4.3 | (46.7–62.3) | 51.8 | 3 | 50.8±0.5 | (50.5–51.8) | 7 | 41.6±1.5 | (39.3–43.3) | 45.5 |
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10 | 28.4±1.7 | (26.2–31.6) | 29.6 | 3 | 29.7±1 | (28.8–30.8) | 7 | 26.1±1.6 | (23.8–28.9) | 24.2 |
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10 | 11.9±0.7 | (10.7–12.7) | 12.0 | 3 | 13.7±0.1 | (12–13.8) | 5 | 11.9±1.3 | (10.7–14.1) | 12.7 |
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10 | 7.7±1 | (6.8–9.9) | 7.8 | 3 | 8.1±0.3 | (7.8–8.5) | 5 | 7.8±1 | (6.4–8.8) | 8.1 |
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10 | 25.2±1.2 | (23.1–26.8) | 24.9 | 3 | 26.4±0.8 | (24.9–27.2) | 7 | 21.6±1.9 | (18.6–24.6) | 22.7 |
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9 | 6.9±1.3 | (4.1–8.7) | 9.9 | 2 | 10.2±0.6 | (9.8–10.6) | 4 | 12.5±1.4 | (11.6–14.7) | 10.8 |
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9 | 8.2±1.4 | (6.7–10.6) | 10.9 | 2 | 11.5±0.6 | (10.9–12) | 4 | 14.6±1.2 | (13.3–16.1) | 11.3 |
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na | na | na | na | na | na | na | na | na | na | na |
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na | na | na | na | na | na | na | na | na | na | na |
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10 | 1 | (1–1) | 1 | 3 | 1 | (1–1) | 7 | 1 | (1–1) | 1 |
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10 | 11 | (9–11) | 11 | 3 | 11 | (11–11) | 7 | 10 | (10–11) | 12 |
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10 | 1 | (1–1) | 1 | 3 | 1 | (1–1) | 7 | 1 | (1–1) | 1 |
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10 | 13 | (12–14) | 12 | 3 | 12 | (12–14) | 7 | 15 | (14–15) | 16 |
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10 | 4 | (3–4) | 4 | 3 | 4 | (4–4) | 7 | 4 | (4–4) | 4 |
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10 | 10 | (9–10) | 9 | 3 | 10 | (9–10) | 7 | 9 | (9–10) | 10 |
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10 | 3 | (3–4) | 3 | 3 | 3 | (3–3) | 7 | 3 | (3–4) | 4 |
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10 | 11 | (10–12) | 10 | 3 | 10 | (10–11) | 7 | 11 | (10–13) | 13 |
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10 | 78 | (76–82) | 82 | 3 | na | (82–90) | 7 | 80 | (73–80) | 84 |
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10 | 33 | (31–35) | 36 | 3 | na | (36–44) | 7 | 32 | (32–36) | 34 |
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10 | 10 | (9–10) | 10 | 3 | 10 | (10–11) | 7 | 9 | (9–10) | 10 |
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10 | 8 | (7–9) | 8 | 3 | 9 | (8–9) | 7 | 7 | (7–9) | 8 |
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10 | 8 | (8–9) | 8 | 3 | 9 | (8–9) | 7 | 7 | (7–9) | 8 |
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9 | 7 | (7–9) | 9 | 2 | na | (9–12) | 4 | 14 | (12–14) | 11 |
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9 | 10 | (9–12) | 13 | 2 | na | (13–19) | 4 | 24 | (22–24) | 23 |
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9 | 19 | (16–21) | 22 | 2 | na | (22–31) | 4 | 36 | (36–38) | 34 |
Illustrations of specimens of each species from Lake Thun. From top to bottom:
Continued.
Phylogeographic studies have shown that the pre-alpine whitefish are a monophyletic clade, most closely related to whitefish from northern Europe. The clade is of hybrid origin involving two glacial lineages that must have come into secondary contact several hundred thousand years after their separation. Independent events of intra-lacustrine speciation led to a series of adaptive radiations in each major lake system of the northern pre-Alps (
Based on genetic, morphological and ecological data at least two species from the Lake Thun-Brienz radiation, namely
Lakes Thun and Brienz in the Bernes Highlands today harbour the most speciose pre-alpine whitefish radiation. These lakes have also suffered the least anthropogenic pressures of all the large pre-alpine lakes in Switzerland. Species delineation and description in such rich radiations require an integrative approach to taxonomy, combining morphology with population genetics and ecology and extensive contemporary and historical specimen collections. Such work is also much needed for conservation-minded fisheries management because, as we have shown here and others before us (
Many thanks to the commercial fisherwomen and fishermen of lakes Thun, Brienz, and Biel and the cantonal fisheries authorities of Bern for their help, hospitality, and sharing of knowledge about the whitefish species of all three lakes. Especially the fisherwomen and fishermen Edith Klopfenstein, Kurt Klopfenstein, Elsbeth Abegglen, Beat Abegglen and Stefan Dasen and the cantonal fisheries warden of Bern, Benjamin Gugger, have been essential in many aspects and especially so in the collection of specimens of the rare species from all three lakes. We would also like to thank the people who helped in the field (Marta Reyes, Pascal Reichlin, Felix Vögtli, Jana Jucker, Andreas Taverna, Dr. Marcel Häsler, Jonas Walker, Dr. Philine Feulner, Dr. Miguel Leal, Dr. Jaime Mauricio-Anayas, Hélène Hefti, Dr. Florian Moser, Andrin Krähenbühl, David Frei, Dr. Kotaro Kagawa). We are grateful to Dr. Rudolf Müller in introducing OMS into the methods of how to read and age scales. We thank Dr. Lukas Rüber and the Natural History Museum in Bern for the adoption and curation of the Seehausen/Steinmann whitefish collection and the Natural History Museum in Geneva for granting us access to their collection. Many thanks to Verena Kälin (
Tables S1-S8, Figures S1-S11
measurements