Table Of ContentAMAZONIANA XVI (3/4): 539-563 Kiel, Dezember 2001
Temporal fluctuations in scale counts and body proportions of
Amazonian riparian lizards (Cnemidophorus, Sauria, Teiidae)
by
P.E. Vanzolini-
Prof. Dr. Paulo E. vanzolini, Museu de Zoologia, universidade de São paulo, Av
Nazaré, 481,04263-000 São Paulo/SP, Brazll.
(Accepted for publication: November, 2000).
Abstract
Abrupt but short-lived changes in number of ventral rows of scales and of dorsal granules, as well as in
relative length of tail, tibia and head, were found in widely separated riverine populations of Amazonian
Cnemidophorus, both parthenogenetic and bisexual. The explanation adopted is that offered by CARSON
(1985) for a similar case in Hawaiian Drosophila: presence of considerable intra-clonal variability and
sensitivity to stabilizing selection.
Keywords: Lizards, riverine, parthenogenetic, variability, selection, Teiidae, Sauria, Amazon,
Neotropics.
Resumo
Mudanças abruptas, mas de curta duração, foram encontradas no númsro de fileiras de escamas ventrais
e de grânulos dorsais, bem como no comprimento relativo da cauda, da tibia e da cabeça, em populações
ribeirinhas, bissexuais e partenogenéticas, de lagartos do gênero Cnemìdophorus na Amazonia brasileira.
A explicação adotada é aquela oferecida por CARSON (1985) para um caso similar de Drosophila no
Hawaii: presença de considerável variabilidade genética intra-clonal e sensibilidade a seleção estabiliza-
dora.
.Dedicated
to Prof. Dr. Harald Sioli on the occasion ofhis 90th anniversary
ISSN 0065-6755/200115391 @ MPI für Limnologie, AG Tropenökologie, Plön; INpA, Manaus
539
Introduction Characters
In the years 1965 Io 1977, this Museum operated the "Expedição Permanente da I examined four meristic characters in the analysis: (i) the number of transverse rows of ventral scales,
Amazônia" (EPA), which consisted of a barge for living and working, the "Garbe", counted between the interbrachial series and the anal flap; (ii) the total number offemoral pores; (iii) the
towed by a small river boat, the "Lindolpho R. Guimarães". Scientific crews rotated, number of infradigital lamellae on toe IV, of either side; (iv) the number of dorsal granules, counted
and the expedition would stop on the riverside and collect, including purchase of transversely at midbody. These are characters routinely used in teiid taxonomy, and known to be reliable.
specimens from local people. Large series were secured of many species. In fact' as can be seen in the tables below, coefficients ofvariation are unvaryingly low.
Lizards of the genus Cnemidophorøs are extremely common in and around towns in Three body proportions were studied: tail length (regression on body, or snout-to-vent, length), length
Amazonia; we obtained good collections both of bisexual and of parthenogenetic oftibia (regression on body length) and length ofhead (regression on trunk, or snout-to-vent minus head,
length).
populations of what is currently called the Cnemidophorus lemniscatas "complex"
(FROST & WRIGHT, 1988). The taxonomy of Amazonian Cnemidophorus is not yet
well understood (e.g., AVILA-PIRES, 1995: 499; COLE & DESSAUER, 1993), and I
Methods
have been appraising the materials at hand for a tentative review of geographical
variation. I routinely checked for homogeneity among samples from localities represent- This is essentially a statistical investigation. It is my beliefthat zoological systematics demands conceptu-
ed by more than one collection (with a view to combining them), and unexpectedly ally simple methods, close to the physical nature ofthe characters. The present issue consists ofcomparing
discovered temporal fluctuations of some meristic characters and body proportions. This meristic and morphometric characters of two or more samples from the same locality, taken at different
report summarizes these pattems. times. In the case of two samples I have used l-tests for the differences between means or regression
statistics' In the case of three or more samples I used analysis of variance, followed, when adequate, by
Tukey's test. The methods used are available in elementary texts, such as DIXON & MASSEY (1933),
Materials ZAR (1999) or VANZOLINI (1993).
The Map shows localities from which C. lemniscatus samples were collected, and Table I summarizes The following conventions have been adopted throughout:
collecting dates and sample sizes. a: regression constant (intercept) + its standard deviation
Oriximina (01"4ó'S, 55"51'W). A sizable town (about 5,000 inhabitants at the time of collecting), on b: regression coefficient (slope) t its standard deviation
the left bank, some 30 km above the mouth of the Rio Trombetas, a tributary of the Amazonas on its left df: degrees of freedom
(north) bank. This is the place where the first parthenogenetic Amazonian population of Cnemidophorus F: variance ratio
was discovered (VANZOLINI 1970). The parthenogenetic populations have not been formally named, but m: mean + its standard deviation
are known by their cytotypes; the Oriximiná cytotype is designated B (PECCININI-SEALE 1989). COLE n: individuals in sample
& DESSAUER ( 1993) identify this population as belonging to their C. cryptus. We collected there nine R, R (x), R (y): ranges
times between 1965 and 1973, and Prof. Miguel T. Rodrigues, University of São Paulo, made a further l: coefficient ofdetermination
collection in 1993. s: sample standard deviation
Alter do Chão (02"32'5,54'57'W). A small town on the right bank of the Rio Tapajós (a tributary of t: Student's , the quotient ofa statistic by its standard deviation
the Amazonas from the south), some l0 km above the mouth. We made two collections there. The V: coefficient of variation
population is bisexual, referrable to C. lemniscatus (L., 1758) (COLE & DESSAUER 1993). n*:s : not significant at the 5Yo level
Breves (01'41'5, 50"28'W). Another sizable town, about the size of Oriximiná, on the landward **: significant at the 5% level
(western) end of the large island of Marajó (that plugs the mouth of the Amazonas), on the strait named ***: significant at the l% level
after the town. The population is parthenogenetic; its cytopype is B, as in Orimixiná (PECCININI-SEALE significant at the 0.1% level
1989) and the liza¡d should thus be C. cryptus. Breves was visited twice.
Corcovado (01'39'S,50'32'W). A very small settlement some 8 km to the northwest of Breves, on In the case ofTukey's test, vertical lines on the left ofthe table encompass samples that do not differ
the same bank as the strait. The lizard population is ofparthenogenetic cytoptype B, and thus C. cryptus. significantly.
In spite of the close geographicál proximity to Breves and of the coincidence of cytotypes, samples
collected at both localities on the same day (September 9,1969) differed significantly in the number of
femoral pores, fourth toe lamellae, and dorsal granules. Corcovado was visited twice, on the same Analysis
occasions as Breves.
ln all 674 specimens were used, and from practically all localities we have large samples (Table l). We have time series from four localities (Table 1): Oriximiná (10 collections), Alter do Chão and
In some cases subsamples of 20 or 50 specimens were chosen at random for taxonomic work, depending Corcovado 13 apiece) and Breves (2).
on the variability of the most variable character. Of course it has always been possible to increase the
subsamples, but no dubious results made that necessary: all statistical tests gave clear responses. Oriximiná
The samples are well suited to statistical analysis. They comprise (Table 6) specimens ofall sizes (= ages),
540 541
from hatchlings, 29-33 mm in body length, to large adults, over 70 mm. The statistics ofthe scale counts length on trunk length.
are shown on Table 2, those of the regressions on Table 6.
The coefflrcients ofvariation (V) ofthe scale counts are uniformly low (1.0 to 4.6) even in the samples
with very few individuals. Analysis of variance of the scale counts (Tables 3 and 4) indicates temporal Summary
heterogeneities in two characters, number of ventral scales and of dorsal granules. In Tukey's test the
statistics are sorted in ascending order. In this analysis (Table 5) I ananged the samples in temporal order Table 20 summarizes the changes in the characters studied, specifying the time of change, although this
and treated them as in Tukey's test: samples that do not differ significantly are encompassed by continuous was not ascertainable in the case of dorsal granules in Oriximiná.
lines on the left ofthe table. It is clear that in the case ofgranules the heterogeneities did not involve any The salient facts seem to be:
abrupt breaks, but that, in the case of ventrals, 1968b and 1972 arc excepcional samples, especially the l. Changes occuned in polygenically determined characters.
former. In the 136 days between January l3 and May 28 there was a shift in the mean of the ventrals, with 2. They occurred both in parthenogenetic and bisexual populations.
a retum to "normal" values within a further 92 days (Fig. l). 3. The geographical compass is ample: there are some 600 kilometers between Oriximiná and Breves. In
It is interesting to note that the sample means of ventrals and granules are conelated (F=22.301**,f= fact, the whole geographical space sampled was involved.
0.7360, I 8 df). No other pair of variables is so correlated, neither are any pair of variables within samples. 4. Several of the changes were practically synchronous in distant localities.
The regressions oflength oftail and length oftibia (Table 6) are, as usual in lizards, excellent, with
the exception of sample 1967b, composed of 5 very young individuals, which did not afford a significant
regression. The regressions ofhead length on trunk length are somewhat less good: the 1967b sample fails Comments
again to provide a signifìcant regression.
The study of regressions is divided in two phases, the first is tests the homogeneity of the slopes; if It must be avowed that the present findings, however interesting, are fortuitous. The
heteronogeneities are found the analysis stops. Otherwise the intercepts are tested and, in this case,
collections were made on general principles and analyzed much later. The analysis was
simultaneous comparison of significant regressions (Table 7) indicates signifìcant heterogeneity in the
made looking for changes in time, but in truth without much hope of finding any (not
cases ofhead x trunk (slope) and tail x body (intercept). Tukey's tests (Tables 5,8; Figs.2,3) reveal in
quite as a matter of mere form, but frankly of seemliness). Furthermore, in a field
both cases breaks inside the year 1967 and between 1967 and 1968.
where cytological and biochemical information are paramount, we are limited to a few
Breves and Corcovado scale counts and body measurements. Thus much cannot be made of the case, but some
These two localities are geographically very close (ca. 8 km), and are topographically and ecologically points deserve coÍrment, as contributing to the rich and difficult field of speciation in
identical. On two occasions they were visited on the same (September 19, 1969) or practically the same Amazonian Cnemidophorus.
day (30 luly - 2 August, 2-4 August 1968). This opens an interesting possibility of study, as discussed An analogous case has been reported by CARSON (1985), in which abrupt changes
below. ',¡/ere noticed, with subsequent retum to norm. He followed a population of the Hawaiian
Table 9 summarizes the statistics of the distributions of frequencies of the scale counts for both Drosophila silvestris from 1976 to 1980; his paper presents measurements of the foreleg
localities. The two columns on the right ofthe table refer, respectively, t (B) to the differences between the tibia and counts of the cilia on three rows on the same tibia. It is possible from
1968 and 1969 Breves samples and t (B * C/ to the differences between Breves and Corcovado collected Carson's Table 2 and Figure 1, to recover the data necessary for an analysis of the type
during the same visit. It is seen that in three characters, pores, lamellae and granules, the two localities adopted here. I have done so and found out that, while tibial length and number of cilia
differed both in 1968 and 1969. Samples from Breves showed significant differences between the two
on row 6 remained constant in the time interval sampled, cilia on rows 5 and 5a showed
visits in ventrals and granules. Table l0 shows that only the distribution of granules in Corcovado varied parallel increases in 1979 and a return to previous values in 1980. It was Carson's
in the interval studied, and that 1968 was the outlying year.
conclusion that these findings indicated the presence ofconsiderable genetic variability
Table 12 presents the statistics ofthe regression analyses. Table 13 shows that in Breves the two years
differ in the three regressions: in tail and head length the difference is in intercept, in length of tibia it is in some quantitative traits and their sensitivity to stabilizing selection. I think the same
in slope. The analysis ofvariance for Corcovado is shown on Table 14, and Table 1l summarizes the explanation applies to the present case.
situation in this locality; there are significant heterogeneities in the number of granules, and 1968 is the Cnemidophon¿r is a speciose genus (ca. 50 species) with a high incidence of parthe-
distinctive year. In the case ofBreves it is possible only to say that the differences appeared between 1968 nogenetic lineages. This has stimulated a considerable amount of research on its genet-
and 1969. In Corcovado, howeve¡, in the case ofthe body proportions, there were heterogeneities both in ics and evolution (e.g., see papers in WRIGHT & VITT 1993; DAWLEY & BOGART
the regression of tail on body and of tibia on body; in both cases the break occurred between 1968 and 1989). One of the issues that has been considered in the literature is the contrast
1969. berween intra-clonal and inter-clonal variability. DESSAUER & COLE (1989), for
instance, state that "there is very little individual variation within lineages of unisexual
Alter do Chão species as compared with bisexual species". This is clearly not the case with Amazonian
Tables 15 and 17 show the statistics ofthe distributions ofscale counts and ofregressions ofthe only
Cnemidophorus: the parthenogenetic and bisexual populations behave in the same way,
bisexual locality. In both tables the columns headed ¡ refer to sexual differences for the character in the
and both reveal a considerable stock of genetic variability in some quantitative traits,
individual years. Tables 16 and l8 contain the respoctive analyses ofvariance and Table l9 the application
of Tukey's test. It is clear that in the seven comparisons made (grânules for females and the three which is reflected in phenotypic plasticity. This is also indicated by the significant
regressions, for both sexes) there was a break within the year 1968. In fact, within this year the break differences found between the neighboring localities of Breves and Corcovado in the
caused even a change in the sexual dimorphisms, in the number of ventrals and in the regression of head same day. I think these facts will have to be taken into account in studies of speciation
s42 543
in the group. MORITZ, C., BROWN, W.M., DENSMORE, L.D., WRIGHT, J.W., VYAS, D., DONNELLAN, S.,
A final issue is the simultaneous occurrence in 1968 of phenotypic shifts in localities ADAMS, M. & P. BAVERSTOCK (1989): Genetic diversity and the dynamics of hybrid parrhenogen-
as far apart as Oriximiná and the west coast of Marajó. There are two prima facie esis in Cnemidophorus (Teüdae) aîd Heteronotict (Gekkonidae). - In: R.M. DAWLEY & J.p. BO-
explanations for this synchronism. Either there was a powerful pan-Amazonian ecologi- GART (eds.): Evolution and ecology ofunisexual vertebrates: 87-112.
cal phenomenon operating at the time, or this type of occurrence is rather common, and PECCINININI-SEALE, D.M. (1989): Genetic studies onbisexual and unisexual populations of Amazonian
the coincidence just that, a coincidence. Cnemidophorus. - In: R.M. DAWLEY & J.P. BOGART (eds.): Evolution and ecology of unisexual
I was on site when all changes ocurred, and did not notice any ecological peculiari- vertebrates: 241-251.
ty. Additionally, having in mind that the explanation might be linked to abrupt massive PRICE' 4.W., LAPOINTE, J.L. & J.W. ATMAR (1993): The ecology and evolutionary implications of
immigration, I checked the records of the level of the Rio Negro at Manaus, and found competition and parthenogenesis in Cnemidophorus. - In: J.W. WRIGHT & L.J. VITT (eds.): Biology
no signs ofunusual floods. I think, as does CARSON (1985) that such sudden pheno- VANoZf OwLhiIpNtaI,i lP l.iEza. r(d1s9 7(g0e):n uUsn iCsenxeumaidl oCpnheomruísd)o: ph3o7r uI s-4 Il e0m.niscatus in the Amazonas valley: a preliminary
typic shifts are, as said, due to high genetic variability and sensitivity to selection, and note (Sauria, Teiidae). - Papéis Avulsos Zool., S. Paulo, 23(7): 63-68.
that they are not more often noticed because nobody looks for them. VANZOLINI, P.E. (1993): Métodos estatísticos elementares em sistemática zoológica. Hicitec, S. Paulo:
A final issue, and not a trivial one is one practical import of this find: what consti- 130 pp.
tutes a proper geographical sample? Should the concept of time (the moment of collec- VRIJENHOEK, R.C. (1989): Genetical and ecological constraints on the origin and establishment of
tion) be introduced into alpha systematics? Different occasions may mean different unisexual ve¡tebrates. - In: R.M. DAWLEY & J.P. BOGART (eds.): Evolution and ecology of unisexu-
phenotypes - the differences possibly reaching a magnitudejudged signifrcant in studies al vertebrates:24-31.
of geographical differentiation. This is an unavoidable issue. The presence of stabilizing WRIGHT, J.W. (1993): Evolution of the lizards of the genus Cnemidophorus. - In: J.W. WRIGHT & L.J.
selection, i.e., the impermanence of the morphological changes, suggests that it would VITT (eds.): Biology of whiptail lizards (genus Cnemidophorus): 27-77.
take much bad luck to hit upon a significantly contaminating shift in characters across WRIGHT, J.W. & C.H. LOWE (1968): Weeds, polyploids, pathenogenesis and ecological distribution of
a geographically meaningful area, but the admonition stands. all-female species ofCrernidophorus. - Copeia 1968(l): 128-138.
' WRIGHT, J.W. & L.J. VITT (1993): Biology of whiptail lizards (genus Cnemídophorus). - Oklahoma
Mus. Nat. Hist. Norman, Oklahoma: 417 pp.
ZAR, J.H. (1999): Biostatistical analysis. - Prentice-Hall, Upper Saddle River: 663 pp.
Acknowledgments
This paper has not been funded by any granting agency. W.R. Heyer (Smithsonian Institution), C.W.
Myers (American Museum of Natural History) and H.L. Carson (University of Hawaii at Manoa) have
kindly criticized the manuscript. Francisca Carolina do Val helped with matters genetical.
It is an honor and a pleasure to participate in this homage to Harald Sioli, whose pioneer work on
Brasilian limnology is outstanding both for its scientific quality and for its importance to the country.
References
AVILA-PIRES, T.C.S. (1995): Lizards of Brazilian Amazonia (Reptilia: Squamata). - Zool. Verhandl.
Leiden 299: l-706.
CARSON, H.L. (1985): Genetic variation in a courtship-related male character in Drosophila silvestris
from a single Hawaiian locality. - Evolution 39(3): 678-686.
COLE, C.J. & H.C. DESSAUER (1993): Unisexual and bisexual whiptail lizards of the Cnemidophorus
lemniscatus complex (Squamata: Teiidae) of the Guiana region, south Americam, with the description
of new species. - Amer. Mus. Novit. 3081: l-30.
DAWLEY, R.M. & J.P. BOGART (1989): Evolution and ecology of unisexual vertebrates. - Bull. New
York State Mus. 46ó: 302 pp.
DESSAUER, H.C. & C.J. COLE (1989): Diversitybetween andwithin nominal forms of unisexual teiid
lizards. - In: R.M. DAWLEY & J.P. BOGART (eds.): Evolution and ecology of unisexual vertebrates:
49-7 t.
DIXON, W.J. & F.J. MASSEY, jr. (1983): Introduction to statistical analysis. - Fourth ed. McGraw-Hill,
Inc., New York: 678 pp.
FROST, D.R. & J.W. WRIGHT (1988)l The taxonomy of uniparental species, with special reference to
parthenogenetic Cnemidophorus (Squamata: Teiidae). - Syst. Zool. 37:200-208.
544 545
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Cnemidophorus, Oriximiná, regression of tail length on body length.
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Cnemidophorus, Oriximiná, changes in the number of rows of ventral scales with time.
Fig. 3:
Cnemidophorus, Oriximiná, regression of length of tibia on body length.
546
547
Table I : Cnentidophorus, materials. Table 2: Cnemidophorus, Oriximiná, Statistics of the distributions of lrequencies of the scale counts.
Locality Dâte Code dd 9C Sample N R m s V
Oriximiná l9 September 1965 l 965 5 Ventrals
I - 24 January 1967 1967 a 70 1965 5 28-30 28.40 r 0.400 0.89 3.2
ll February 1967 t967b 5 196',7 a 70 28-3t 29.04 0.080 0.67 2.3
12 - l3 January 1968 I 968a 1967b 5 29-30 29.40 0.245 0.5 5 1.9
28 May 1968 l 968b l0 I 968a 47 27 -30 29.00 0. l l4 0.78 2.7
28 August 1968 I 968c 67 l 968b l0 29-31 29.90 0.233 0.74 2.5
l8 November 1968 I 968d 39 I 968c 67 27 -3t 29.25 0.101 0.82 2.8
4-8January1972 1972 20 I 968d 39 29 ,31 29.s | 0.089 0.56 1.9
3-7November1973 1973 20 1972 20 21 -30 28.80 0.156 0.70 2.4
2l - 3l January 1993 l 993 20 1973 20 27 -29 28.30 0.128 0.67 2.0
I 993 20 27 -30 28.3s 0.166 0.75 ¿.ft
Breves 2-4August1968 I 968 50 Pores
9 September 1969 1969 50 I 965 5 44 - 4'l 44.60 0.600 1.34 3.0
1967a 70 43-48 45.67 0.156 1.30 2.9
Corcovado l8 - 19 October 1965 I 965 79 1967b 5 46-47 46.80 0.200 0.45 1.0
30 July - 3 August 1968 I 968 50 I 968a 42-48 45.47 0.201 L38 3.0
9 September 1969 1969 50 I 968b 10 42-48 45.20 0.554 1.7 5 3.9
I 968c 65 43-48 45.34 0.163 1.3 l 2.9
Alter do Chão 2l - 22 February 1968 I 968a 30 30 1 968d 37 42-49 45.89 0.247 I .51 3.3
17 - 2l July 1968 1 968b 30 30 t972 20 43-49 45.60 0.400 t.79 3.9
l9 September 1969 1969 30 30 1973 20 43-49 45.7 5 0.3 l5 t.41 3.1
))
1993 t9 44-48 45.26 0.227 0.99
Lamellae
I 965 5 2t-23 22.00 0.316 0.71 3.2
1967 a 70 2t-25 22.34 0.083 0.70 3.t
1967b 5 22 22.00
I 968a 47 22.19 0.078 0.54 2.4
I 968b l0 22-23 22.10 0.100 0.32 1.4
I 968c 66 20-24 22.20 0.070 0.57 2.6
l 968d 39 22-24 22.28 0.090 0.56 2.5
1972 20 22-23 22.29 0.105 0.47 2.t
t973 20 22-23 22.10 0.069 0.31 t.4
))_)\
1993 t9 22.16 0.086 0.3'7 1.7
Granules
I 965 5 94- 106 l0l .20 2.059 4.60 4.6
1967a 70 95- 105 99.36 0.290 2.43 2.4
t967b 5 97- 100 98.20 0.583 1.30 1.3
I 968a 47 93- 102 98.96 0.284 1.94 2.0
I 968b t0 96- 100 98.00 0.577 1.83 1.9
I 968c 66 96- 106 99.94 0.280 2.27 2.3
I 968d 38 96- 102 99.37 0.265 1.63 1.6
1972 20 97- 104 100.50 0.495 2.21 2.2
1973 20 94- 108 100.45 0.7 45 J.JJ 3).)3
I 993 20 97- 105 100.60 0.499 2.23
s48 s49
Table 3: Cnemidophorus, Oriximiná, analyses ofvariance, scale counts. Table 5: Cnemidophorus, Oriximiná, Tukey's test in temporal sequence.
Source Sum of squâres df mean square F Sample Ventrals (m) Granules (m) Tail x body (a) Head x trunk (b)
Ventrâls 1965 28.40 101.20 13.27 0.2380
Between samples 43.4964 9 4.8329 9.406**,¡ 1967a 29.04 99.36 13.7 6 0.2896
Error r 50.5505 288 0.5138 t967b 29.40 98.20 7.34 ns
Total 194.0469 302 I 968a 29.00 98.96 12.46 0.0912
Pores 1 968b 29.90 98.00 10. l6 0. 1913
Between samples 24.s625 9 2.7292 I .431ns 1 968c 29.25 99.94 13.09 0.2268
I
Error 549.09 I 288 1.9066 l 968d 29.51 99.37 15.85 0.2303
Total 573.6543 297 1972 28.80 100.50 14.22 0.2234
Lamellae t9't3 28.30 100.45 12.86 0.2228
Between samples 2.3125 9 0.2569 0.8l8ns I 993 28.35 100.60 14.95 0.1 829
Error 91.4084 291 0.3141
Total 93.7209 300
Granules
Between samples l3 r.9097 9 14.6566 2.'179**
Error t534.9142 291 s.2746
Total 1666.8239 300
Table 4: Cnemidophorus, Oriximiná, scale counts, Tukey's test.
Ventrals Granules
Sample m N Sample m N
1.1973 28.30 20 l. 1968b 98.00 l0
2. 1993 28.35 20 2. t967b 98.20 5
3. l96s 28.40 5 3.1968b 98.96 4'.7
4. 1972 28.80 20 4. 1967a 99.36 70
5.1968a 29.00 47 5.1968d 99.37 38
6. 196'la 29.04 70 6. 1968c 99.94 66
7.1968c 29.25 67 '7. 1973 100.4s 20
8. r967b 29.40 5 8. 1972 I 00.50 20
9. r968d 29.51 39 9. 1993 100.60 20
10. 1968b 29.90 l0 10. l9ó5 l0l .20 5
550 551
12 0.9502 0.9779 0.9304 0.9948 0.9'744 0.9882 0.9s67 0.983ó 0.9809 0.8467 0.9173 0.8615 0.9069 0.9573 0.9363 0.931l 0.9489 0.9775 0.9627 0.8968 0.4276l0.887 0.9494 0.8948 0.9301 0.9699 0.7567 *ì o{a\ :'\ô \o ot 3 Þoo ootsÞ o0ao Þ o Þ Þoo
ôûeão o râìo (u o z@Þ Ê I
57.230*+ 1324.793*** 334.162*** 379.043*** 180.752***I 1174.279+** 397.719*** 1081.015*** 924.892**+ 16.568*** 742.734*** 85ns0. I 279.936*** 77 -914+** 1412.892*** 543.623++* 243.077******333.98 I 783.374*** 7'7.404++* 582.3.73*** (ns)indet. l9***33.6 62.839*** 1219.290*** 306.326*** 239.617+*+ 590.107*** 55.992+*+ Þ .- 6 Sa?gqBãä€EEã* -'oã0a oq -S rr!¡ÈèNæaæN5¡.JU 19ÈL^b\o6*:'O\ O\*Ø*
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body proportion b * 1.986 0.26252.131 0.05862.005 0.10972.1t7 0.10872.014 0.05862.349 0.06851.957 0.09811.983 0.0603t.999 0.06s7 0.04620.188 0.1600.0059 0.0870.2023 0.1620.0097 0.1540.0175 0.1630.0043 0.1810.0078 0.1590.0102 0.0087590.1 0.1560.0056 0.02700.238 0.2200.0091 0 0.01570.091l9l0.02410. 0.00650.227 0.2300.0132 0.2230.0144 0.00920.2230.r830.0244 ÈÞdo 99c\ß\ oxãp- Þ ô o @0ao o -lx'o of
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ofthe atistics regressi R(y) 83-t4382-r6076-156 -14581 't7 t49- 76-129 80-1641)-15879-t57 t26l36 67 613 t26 t25 t26 t26 t25 t26 9- 159-t7 99-149- l58-179- 169-168-169-16 Ø= =oÞ c'. 6 Þ oD.< z Øçn otl 6-@DÈ z
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Oriximiná, midophorus, NR(x) dy35-645-'tl35 32- 33 277133-664 - 6'l32 33l633-5535-732029-722033-7020 ody35-64535-746934-38533-724736-771032-736532-673935-7320-1229 2033-6920 unk26-49526-576925-29524-624727-62t023-546723-513826-572020-562024-5420 F:"9.:l9:¡èIl|.l:.^\o\o\o\Õe€99€;ía{6o\o,!o.o\o\;aN{qæuææ{o Þ ã !¡|Èlr!r!"l..rl.r9Jæ\ol.J{ÞÕæÀ-aqqúo\l\ooôoè iÞÞaaþÞôôôNUAô{ÀA 990J9:^¡P!.J:-\o\o\o\o\o\o\o€€O\66{{O\6þO\qææNa{æaæÀoÞdÞ 'pi'riJieieþLLbÞaNñl..J-€æþæc)ôaN€-h-J-oaæsæôa9ì..) aO\NJ196É19èuæ{oô€oo{
6: Table Cne Sample Tail x bo t965 1967a 1968a 1968b 1968c 1968d 1972 1973 1993x Tibia b 1965 1967a 1967b 1968a 1968b 1968c 1968d 1972 1973 1993 x trHead 965l 1967a 1967b 1968a 1968b 1968c 1968d t972 19'13 r993
(t l.') qJ
Table 9: Cnemidophorus, Breves and Corcovado, statistics ofthe distribution offrequencies ofthe scale Table l0: Cnemidophorus, Corcovado, scale counts, analyses ofvariance.
counts.
Source Sum of squares df Mean square
v (B)
Sample N R m t(C x B)
Ventrals
Ventrals Between samples 1.2344 ) 0.6172 2.299ns
Breves 1968 50 27 -29 28.08 + 0.090 0.63 2.t 2.109* Error 46.1836 t72 0.2685
1969 50 27 -30 28.36 0.100 0.ó9 2.4 Total 47.4319 l't 4
Corcovado 1965 76 27 -29 28.30 0.056 0.49 t;7
1968 49 2'7 - 30 28.12 0.080 0.56 2.0 0.3 l2ns Pores
t969 50 27 -29 28.32 0.070 0.51 1.8 0.328ns Between samples 5.9063 ') 2.9531 l.978ns
Error 250.8359 168 1.4931
Po¡es Total 256.7422 170
Breves r 968 45 4t-48 43.73 0.230 1.54 3.5 0.644ns
1969 46 40 - 4't 43.52 0.234 1.59 3.7 Lamellae
Corcovado 1965 72 40-4s 42.65 0.147 t.25 2.9 Between samples 0.2t09 2 0. I 055 1 565ns
l 9ó8 49 40 - 4'l 43. l0 0.1 83 1.28 3.0 2.166* Error I1.7871 175 0.0674
1969 50 4l-46 42.86 0.159 1. 13 2.6 3. I 73** Total l 2.0000 t77
Lamellae Granules
Breves 1 968 48 2t-24 22.21 0.079 0.54 1< 0. I 57ns Between samples 124.6250 2 62.3t25 13.819*.**
1969 49 22-24 22.22 0.067 0.47 2.1 Error 775.0000 172 4.5058
Corcovado I 965 79 21-23 22.04 0.028 0.25 t.1 Total 899.62s0 174
1 968 49 21-22 21.98 0.020 0.l4 0.7 2.845f*
1969 50 21-23 21.96 0.049 0.35 1.6 3.195**
Granules
Breves I 968 50 96 - 106 101. l4 0.305 2.16 2.1 2.616**
1969 50 94 - 103 100.02 0.300 ', 1) 2.1
Corcovado r 965 75 94 - 106 100.29 0.24'.1 2.14 2.1 Table I l: Cnemidophorus, Corcovado, Tukey's test.
1 968 50 94 - t07 102.32 0.286 2.02 2.0 2.981*
1969 50 95 - i05 100.90 0.309 2.19 2.2 2.133x
Sample m N Sample bN Sample bN
Granules Tail x body Tibia x body
ll
I 965 100.29 75 1969 l .913 t965 0.1 60 78
1969 100.90 50 I 965 2.146 2l 1968 0.1 69 47
I 968 r02.32 I 968 2.282 22 1969 0.1 89 50
554 555
Table l2: Cnemidophorus, Breves and Corcovado, statistics ofthe regressions ofbody proportions. Table l4: Cnemidophorus, Corcovado, simultaneous comparisons ofregressions, analyses ofvariance.
Sample N R(x) R(y) b a FR2 Regression Source Sum of squares df Mean square F
Tail x body Tail x body general regression 23473.3s54 I
Breves 1968 20 32 74 '70 - 163 2.121 + 0.0682 7.81 t 5.632ns 968.096*** 0.9817 el1or 385.789 I 48 8.0373
1969 24 32 69 75- 146 2.090 0.087s 9.98 4.507* 570.535*'r.,t 0.9629 slope 82.5349 1 41.2674 5. I 35**
Corcovado 1965 2l 32 60 1)_ 132 2.146 0.0'703 5.01 4.5lOns 932.923*** 0.9800 Tibia x body general regression 657.8848 I
1968 22 3t 66 73- 152 2.282 0.0563 0.04 4.89lns I 649.538xrx 0.9880 error 45.4580 69 0.2690
1969 l l 35 65 85- t43 I .913 0. 1054 t7 .79 5.418** 328.902*** 0.9734 slope 2.8506 2 1.4253 5.299**
Tibia x body Head x trunk general regression 850.4820 I
Breves 1968 50 32-74 6-t2 0.162 0.0042 8.48 0.290:r'*x I 435.6!3*** 0.9687 efTor 45.57 46 t7t 0.2665
1969 50 32-73 6-13 0.1 83 0.0056 9.58 0.2 I 6*** I 085.894*** 0.9577 slope 0.6636 2 0.33 l8 l.245ns
Corcovado 1965 78 30-75 6-t2 0.1 60 0.0063 7.13 0. I 86*** 584.708*** 0.8850 intercept 0.1612 2 0.0806 0.302ns
1968 47 3t-69 5-t2 0.1 69 0.0053 8. 17 0.293*** I 0l 5.490*** 0.9576
1969 50 29-72 6-16 0.1 89 0.0068 8.62 0.327*** '166.154'þx* 0.9410
Head x trunk
Breves 1968 48 23-57 8-17 0.230 0.0070 l t.s6 0.345*** I 088.97 I *** 0.9595
1969 50 23-60 9-t7 0.239 0.0074 12.66 0.307*** I 056.9 I 7*** 0.9566
Corcovado 1965 78 2l-58 8 - 17 0.236 0.0089 9.94 0.217*** 707.'196*** 0.9030
1968 s0 23-s3 8 - 16 0.245 0.0068 n.64 0.340*** 1291.109*** 0.9642
1969 49 2t-56 8-16 0.228 0.0069 tt.49 0.3 l7*** I 101.633*** 0.9591
Table l3: Cnemidophorus, Breves, differences between regressions, 1968 and 1969
Slope Intercept
Regression difference (b) dft difference (a) dfr
Tail x body 0.0308 + 0.09091 40 0.339ns 12.97 ! 3.9t1 4l 3.316**
Tibia x body 0.0209 0.00699 96 2.991*
Head x trunk 0.0093 0.01110 94 0.845ns 0.97 0.466 95 2.082*
556 557
