Heredity 57 (1986) 149159

The Genetical Society of Great Britain

Received 28 August 1985

Chromosomal polymorphism, climatic factors, and variation in population size of Drosophila willistoni i n southern
Brazil
V. L. S. Valente and A. M. Arajo
Departamento de Gentica, Universidade Federal do Rio Grande do Sul, Cx. Postal 1953, 90.001-Porto Alegre, RS, Brazil.

Natural populations of Drosophila willistoni collected every three months over banana baits between September, 1978 and May, 1982 at Parque de Itapui and Parque do Turvo, Rio Grande do Sul, Brazil, localised in different regions in terms of climate, flora and fauna, were studied with respect to the variation of their chromosomal polymorphism associated with meteorological variables and population fluctuations.

Multiple regression applied to the variation in the population size (dependent variable), macroclimatic data and heterozygous frequency for inversions of chromosomes IlL and III showed that in

The flexible and adaptive nature of its chromo-

somal polymorphism has been shown by Da

Cunha et al. (1950, 1959), Da Cunha and Dobzhansky (1954), Battaglia and Birch (1956), Birch and Battaglia (1957), Da Cunha (1956a, b; 1957), Ayala et a!. (1971), Valente et aL (1981), Valente and Arajo (1985), who found shifts in the inver-

both places those parameters play very distinct roles regarding D. willistoni populations. At Parque do Turvo, where the climate is more constant, the minimum temperature of the month that pre-

sion frequencies associated with environmental

variation or ecological factors. However, there are few works on the association of this marker with

ceded each month of collection explains 90 per cent of the population variation of the species, whereas at Itapu, a region presenting greater meteorological instability, intrinsic factors, such
as the inversion frequencies together with two climatic variables can account for the same proportion of the variation in population size. Significant correlations were also found between inversions and climate in both places, as well as associations

climatic variability in well defined seasonal

regions, such as the State of Rio Grande do Sul,

Brazil, situated near the southern limit of the

species distribution. Our objective was to add new information on the chromosomal polymorphism variability of D. willistoni populations and try to establish relation-

among inversions of the same and different

chromosomes.

ships between the cytological data and demographic and environmental parameters, especially the effects of climatic oscillations.
MATERIAL AND METHODS

INTRODU CTIO N

Drosophila willistoni is an ecologically versatile species, dominant in most Neotropical forests,

Studies

were made in two forest parks separated

although it had also been found in Mexican forests,

by about 500 km, and in different climatic regions: Parque de Itapu (30 17' 5, 51 1' W) and Parque

in sympatry with species from temperate regions (Ehrman and Powell, 1982). Its geographic distribution ranges from central Mexico and Florida to northern Argentina. It was also found by A. R. Cordeiro (unpublished results) at the southernmost locality of Montes Tordillos, 300 km from Buenos Aires (Winge, 1971).

Florestal Estadual do Turvo (27, 21' S, 53 and 54 10' W). More details about both places are found in Valente and Morales (1985). Parque do Turvo is situated in a "hot winter" region, and Parque de Itapu is localised in a "cold winter" area (Plano Integrado para o Desenvolvimento do Litoral Norte do Rio Grande do Sul, 1976).

150

V. L. S. VALENTE AND A. M. ARAUJO

The natural populations of Drosophila willistoni were collected over conventional banana baits dur-

great oscillations are found. Furthermore, it is also

ing an average of 5 days (the time spent at each site was about 30 minutes) every three months
between September, 1978 and May, 1982. The baits measured about 50 cm in diameter and were made from a total of 3 kg of bananas. Nearly half of the

noticeable that, in the 19 samples studied, the frequencies of the overlapping inversions IlL D and IlL E exceed 50 per cent in 17 of them,
exhibiting extreme values, ranging from 42 to 80 per cent, being, therefore, in most cases heterotic.

In chromosome III this tendency also appears,

samples can be considered synchronic, since they had only an interval of a week between places.

The egg samples from females captured in nature and placed individually into tubes with
culture medium were processed using Ashburner's technique (1967), during the late third larval stage. Macroclimatic data such as temperature (C), humidity, rainfall and insolation corresponding to the collecting month were obtained from meteoro-

although sporadically, for inversion B. Table 2 presents the relative frequencies of D. willistoni in the samples and its variations during

the whole collecting period. It is the dominant

frequency is 46 per cent and the second most

abundant species at Parque de Itapu (11 per cent),

species at Parque do Turvo, where its total

logical stations nearer the places of study: IraI Station (about 60 km from Parque do Turvo 27 il'S, 53 14' W) and Porto Alegre Station
(about 60 km from Itapu30 01' 5, 530 13' W). (Appendix 1).

where the introduced species D. simulans is the dominant one (77 per cent of total frequencies) (Valente and Arajo, submitted).
Fig. 1 represents the variation in the frequency of Drosophila willistoni (from table 2B) in parallel with total heterozygosity for chromosomes IlL and

The Pearson Correlation and Multiple

III, and the heterozygous frequencies for inver-

sions of the same chromosomes at Parque de

Itapu and Parque do Turvo (from table 1). It can be seen that the total heterozygosity of

Regression subprograms belonging to the Statistical Package for the Social Sciences (Nie et al., 1975) were utilised in order to obtain correlations between heterozygous frequencies for inversions
of Drosophila willistoni and the climatic parameters considered, as well as to evaluate their effect over

chromosome III for Itapu and IlL for Turvo

oscillates more or less in parallel with the population size of the species. In relation to the frequencies of each inversion it was verified that in Itapu IlL D, E and F present an oscillation apparently the inverse of that of the population in the first samples, tending to be syn-

population size oscillations, estimated by the

indirect method (No. of D. willistoni/total No. of flies per sample). The tests were carried out with the data transformed as follows: population size, temperature,

chronic with it in the succeeding ones. IhI and

rainfall and insolation in common logarithms, humidity and inversion frequencies of chromosomes IlL and III by angular transformation.
RESULTS

"B, particularly the latter, exhibit larger frequencies when the population is in expansion. Among the samples from Itapu there is also
and E (excepting those of January and April, 1981) and an inverse tendency in those of inversions IlL

a strong agreement in the frequencies of IlL D

A and B and hhI9 and hII. The inversion frequencies of IlL D and E at Parque do Turvo, as well as those of IILH and III seem to follow the largest relative frequencies of D. willistoni, whereas in those of III, it does not occur in all samples. Strong linkage was also observed here between IlL D and E inversions,
with practically identical frequencies. An analysis of the role of the inversions and

The

analysis of the associations among climate,

population size and inversion frequencies is related

only to chromosomes IlL and III, which are the most polymorphic in our samples among the five chromosome arms of the larval salivary glands (Dobzhansky, 1950). The chromosome arms XL, XR and hR are practically homozygous in our
samples. Table 1 shows the frequencies for the heterozy-

gotes of the most common inversions (minimum

value of 10 per cent, at least for one of the samples).

climatic factors on population size was made through the multiple regression technique. The

As can be seen in the columns regarding total

heterozygosity, the values are very high in most
samples for both chromosomes. When the frequen-

cies of each inversion are observed one by one,

results for Parque de Itapu and Parque do Turvo are shown in tables 3 and 4, respectively. There are remarkable differences for both localities when the contribution of each variable is compared. By examining the multiple coefficient of determina-

tion (r2) in table 3 it is possible to see that five

-u

0 0
-o

:i
C/)

Table 1 Heterozygotefrequencies (%) for the most common inversions* of chromosomes IlL and III of D. willistoni collected in the two areas of study Chromosome II L Chromosome III
No. Inversions

z
0
No.

No.

Inversions Total
heterozygosis

0 -o
Ca)

of
of A
B

of
B C

Place

Month! Year

females
F 21-0

D
E 67-0 H larvae 256 20
896

Total heterozygosis
75-0 62-0 79-0

J
150 380
240

larvae

Parque

de 800
15-0

Sept. 78

68

800
15-0

660
140
7-0 212
108 558

420
4-0 46-0 36-0

10

4-0

480
46-0

258

I-.

Dec/Jan

Iapu

600
300 110 280

140

12 224 48 81-0 84-0

70
100

135
261

5-0 3-0

24 910 260 110

(1)

40
20
85-0

450 380 370

52-0

219

78/79 Mar./Apr. 79 July/79 Oct./79

10-0

740 600

21 143

M.ar./Apr. 80 Jan/Feb. 81

77 80-0

870 920 790 300

24-0 30-0 1FO

66-0 75-0 78-0 79-0 43-0

4-0
509
205

13-0 19-0

50 370 290
15-0 40-0

32-0

108

80
1-0

460
500

546
137

Apr./81

116

60
260

90
420 700 630
63-0 70-0 24-0 20-0 19-0 17-0

10
4-0

48-0

260
511

July/81 78-0

820
0-4 15-0

100

320 45

60-0 68-0 75-0 77-0 79-0 50-0 60-0 70-0 9-0 9-0 8-0 6-0

340 390 430 550 460

79-0 44-0

460 490
240 220
23-0

206 37-0

123

40
710
8-0 12-0 7-0 16-0 16-0

Parque do Turvo

July/79 Oct./79 Jan./80

810 830
88-0 80-0

90
110

840 730 770 780 840 790 800

529

Mar/Apr. 80

69-0

270 265 97

7-0 2-0

800
220 200
17-0 17-0 17-0

71-0 72-0 80-0 69-0

220
220
22-0

450 460

102 10

790
170

30
670
78-0

670
780

70 90
220

460 360
28-0 76-0 78-0 72-0

39-0 24-0

1268 1402 228 467

920
33-0

88

30
750
79-0

840
160
17-0

440 410 820

76-0 46-0 9-0 4-0 50

127

85-0 86-0

10
0-4

6-0 20-0

July/80 Oct./80 Jan./Feb. 81 Apr.! May 81 Jan./82 May/82

264

810
130

750 790 730 730

519 1265 1369 228 467 36 383 254 563

280 170 190 430

19-0

46-0 46-0

36 386 259
564

* Minimum frequency of 10%

Absence

at least one sample

01

C,'

Table 2 Relative frequencies(%)

of D.
Month/year July
May
80 80 80 81 81

willistoni in each sample (A) and along the collection period (B) (from Valente and Araijo, submitted)

Jan.
July Oct.
80 80

Apr./ Jan.
81

Mar./ Apr.

Apr./ May

July

Jan.
82

May
82

willis-

Place
1479 158 37 1234
15 12
1

Aug. 78 79

Sept. 78

Dec./ Mar./ Jan. Apr. 78/79 79

Oct. 79 50
235
10

Total

toni

Parque de Itapu
5
25 3
1

N 5 20
8297
2815

%A
2
27450
141

111
1587 56 26

272

%B
3204 157 4 3552 6245 1797

8 2

9 5

100 21

4
650 26 11

5913

257 5
1

412 2299
1107 19

2509

Number of flies (all species) 1382 Parque do Turvo

3148

471

107 15
1

3938 79 51 5 2765 60 14

2254 47
11

53784
20523

11

N 4
50
17

85
30 7342
1 1

%A

4037 7151 4010 717

%B

Number of flies

(all species)

4885

4972

2191

4582

4750

44637

46

I-

rn
-I

0 >

L 0

POLYMORPHISM IN DROSOPHILA WILLISTONI

153

0/ I0
III

80
70

II

60
J 50

40 30 20 10
0

-'I

II L

II L

01

100

90 80

.v

70 60
50

iiL
A S DJAMJ

40 30 20 10

A SDJAMJ OJMAMJ OJFAMJ J M

iS

75

SO

SI

is

is

0 JMAMJ 0 JrAMJ so si

J N
$2

I TA PU

TURVO

Figure 1 Variation in population size of D. willistoni at Itapu and Turvo (thick line, below) and total frequency of heterozygotes inversions in chromosomes "L and III. Upper graphs: variation in the frequency of specific inversions in chromosomes "L and III.

variables (inversion B of the third chromosome, humidity at the collection month, inversions IlL E, III J and insolation at the collection month) can account for 90 per centof the variation in size.
This same quantitative effect is achieved at Parque do Turvo by a single variable, namely, minimum temperature in the month before collection (table

4). The contribution of inversions IlL E, IlL B, IlL F, and IlL H is negligible there. Table 5 shows the correlation matrix among 19 selected variables of the present study. Values for Parque de Itapu are placed above and those for Turvo below the diagonal. There are three types of significant associations that deserve attention

154

V. L. S. VALENTE AND A. M. ARAUJO

Table 3 Results of the stepwise multiple regression among population size (dependent variable), climatic variables and inversions (independent variables) at Parque de Itapu (B = partial regression coefficient; 13 = standardized regression coefficient; r2 = coefficient of determination; F = variance ratio)
Variable

B
1104656
8698744

/3

r
145937

F
913468891: 1258309721: 1912210541: 94382751: 98898 0721: 445489921: 35848471:

Inversion B (chromosome III) Humidity at the collection month Inversion E (chromosome II L) Inversion J (chromosome III) Insolation at the collection month Inversion H (chromosome III) Maximum temperature at the month before the collection Intercept

275940

1533116 739525
10-74973 10-27072

1i6495 032712
192376 082853

025618 043503 071285 081765 090443 099359

10000

075096
1265875

0-11588

1:=P<0-00l
Table 4 Results of the stepwise multiple regression among population size (dependent variable), climatic variables and inversions (independent variables) at Parque do Turvo. (B = partial regression coefficient; /3 = standardized regression coefficient; r2 = coefficient of determination; F = variance ratio)
Variable

/3

r2

F
174579171: 7979311: 51062791: 25498491: 18786341: 10929761:
39-4581:

Minimum temperature at the month before the collection Humidity at the month before the collection Rainfall at the collection month Inversion E (chromosome II L) Inversion B (chromosome II L) Inversion F (chromosome II L) Inversion H (chromosome II L) Intercept

469060
795321 127748 17-49238 292460

118305
028224
0-68421 0-69452

090201
0-95448 0-97186

098155
0-98970

1399988 0-48770 2120593

026644 0-21793 0-02355

099968 099999

1:=P<0-001

here: those between chromosome inversions and climatic variables, those between inversions in the same chromosome and those between different
ones.

When correlation among inversions in the same chromosome are examined, four significant values
_0.80* and rA,B=_0.85**). At Parque do Turvo inversion A, D and E from the 2nd chromosome

At Itapu the two climatic variables that corre-

are found for Itapu involving only the 2nd chromosome (rD,E = o.94**; rA,D = _0.67*; rA,E =
are related (rD,E=0.99*; rA,B _O.65* and rA,H=

late with inversion (humidity at the collection

month = UmiCol, and insolation at the collection month = InsCol) are both highly negatively corre-

lated (r=_0.94**), as would be expected; each

inversion correlates significantly with InsCol and

0.63*), while only inversions B and C from the third chromosome are associated (rB,c = O.82**).

UmiCol by a similar correlation coefficient,

although with a reverse sign (the exception being inversion III B). At Turvo, inversions of the third chromosome are not correlated with climatic variables with only one exception: inversion J with InsAnt (insolation

As for association of inversions belonging to different chromosomes, at Itapu both "L D,E
correlates with II C,H,J together with "L A, III C and "L H, III B; for Parque do Turvo these are "L A with II B,C and "L H, 11 C. Tables 6 and 7 show the estimated mean num-

in the month before the collection; r=0.81**). Inversion IlL D and E are correlated, of course,
with the same climatic variables (TAntMed, TCo1-

Med, InsAnt and PCo1); it is interesting to note, however, their very different behaviour at Parque de Itapu (above diagonal). IILF is another inversion significantly correlated with climatic variables (UmiCol and UmiAnt).

ber of inversions per chromosome arm and per female in the samples from Itapu and Turvo, respectively. The left arm of chromosome II and chromosome III are the most polymorphic in the two localities. Considering the total mean per
female there are no differences between Itapu and Turvo, although the latter showed a smaller stan-

dard deviation, suggesting a more stable distribution.

0 0
-D
(1)

I
of heterozygo te inversi ons in chromosomes IlL and III

Table 5 LII Umi D

B

Correlation matrix among macroclimaticvariables, population size and freque ncy (above diagonal) and Parque do Turvo (below diagonal)

t
N

0-941: 0-771:

Parque de Itapu

IlL H
C
PCol 0-18
0-851: 0-28

0
UmiCol Ant
0-01
0-921: 0-03 0-01 0-841: 0.64* 0-851: 0.70*

F
J
PAnt InsCol InsAnt

TCol Med

TAnt Med
0-21

0-39 0-50 0.72* 0-36

0-34 0-45 0-33 0-37 0-34 0-43 0-18

0-35

027
0-51

032

036

0-03 0-20

0-02 0-32 0-23

0-25

0-39 0-35 0-22

0-09 0-24 0-31 0-09

0-30 0-26 0-30 0-28 0-01 0.72*

0-12 0-49 0-15 0-09

041
0.73* 0-45 0-08
0-27

0-40

0.67* 0-55
0-32

0-29

0-09 0-06 0-801: 0-09

TAntMed 0-871: TColMed 0-20 0-53 0-48 InsAnt 0.65* 0.73* 0-781: InsCol 0-37 0-11 UmiAnt 0-57 0-53 UmiCol
0-01
0-38 0-811:

0-06

0.79*
0-41 0-02

0.66*
0-57

0-48 0-37 0-40

0-32
0-37
0-17

0.72* PAnt PCoI 0-45 J 0-04 0-14 0-07

0-003 0-09 B

0-56

0-20 0-43 0.73* 0-38

0-36 0-37

0-54

0-31

0-05 0-44 0-20 0-53 0-14 0.71* 0-47 0-19 0-26 0-14 0.67* 0-811: 0.66* 0-10 0-13 0-11 0-01 0-53 0-10 055 0-43 0-02 0-64 0-62 0-50 0-821: 0-14

021 0-09 0-01 0-47 0-10 011 0-51 0-38 0-14 0-17 0-53 0-29 0-05 0-45 0-36 0.73* 0-17 0-40 094t 0-03 0-42 0-11 0-24 0-21 0-28 054 0-09 0-20 0-58 0-37 0-56 0-40 0-11 0-07 0-03 0-28 0-43 0-18 0-54 0-11 0-06 0-26

005 C
H 0-59 0-61 0.63* 0-20
0-18 0-48 0-41 0-10 0-22

0-09 0-09

0-10 041 0.57 009 0-04 0-11 0-03 0-26 0.71* 0-53 028 0-17 031 0-03 0-40 0-15 0-38 0-43 0-50 0-28 0-26 0-38 0-27 0-43 0.71* 0-16 057 015 0.79* 0.71* 0-40 0-941: 0-48 0.99*

0.74*
0-03

0.71* 0-881: 0.74* 0-891:

0-56 0-36

083t D
0-841: 0-23

E
0-20
0-26
0-18

014 0-18
0-10 0-871: 0-43

0-34 0-35 0-23

026

0-11 0-05

0.64* 0.64* 0-04 0-01 0-10

A
H
B

0-61 0-08 0-40 029 0-63 0-18 0-14 0-13 0-28 0-49 0-58 0-10 0-04 0-04 0-851: 0-17 002 0.63* 0-01 0-34 0-34 0-24 0-06 0-39 0-39 0.64* 030 0-05 0-05 045 0-46 0-50 049 0-54 0-50 0-29 0.68* 0-27 0-37 0-20 0-33 0-35 0-55

0-39 0-32

0-12 0-41 0-18

0-22 0-36 0-10

PCol=rainfall in the collection month; PAnt=rainfall in the month before the collection;UmiCol=humidity in the collection month; UmiAnt=humidityin the month before the collection; InsCol=insolation in the collection month; InsAnt=insolation in the month before the collection; TColMed=mean temperature in the collection month; TAntMed= mean temperaturein the month before the collection; N=population size.

1:

=P<0-01.

01 01

156 Table 6

V. L. S. VALENTE AND A. M. ARAUJO

Mean number of inversions per chromosome and per female of Drosophila willistoni from Parque de Itapu Chromosome Mean no. Invs/9

Samples

Month/Year
Sept./78

Season

XL

XR 001
258

II L
186
256

II R

III
114
258

Spring
Summer

N N

O0O 258

001
259

300 273 298

3.22

Jan./79 Apr./79

000
23

014
21

170
20

009
23

088
24 125 900 125
219

Autumn
N

002
929

001
929

174
896

003
926

July/79
Oct./79 Mar./80 Jan./81
Apr./81 July/81

Winter
N

001
221

002
220

195
212

005
221

Spring
Autumn

000
N N
107

000
107

191
108

005
107

080
108

278
3.34

000
547

000
547

202
558

004
549

125
546

Summer
N

003
136

000
136

147
135

005
136

132
137

288
5.28

Autumn
N

000
139

001
139

292
139

012
139

221
138

Winter

000
511

000
511

183
509

002
511

108
511
124

292
324 063

Total

0006

002

193

005

Table 7 Mean number of inversions per chromosome and per female of Drosophila willistoni from Parque do Turvo

Chromosome
Samples

Month/Year
July/79
Oct./79

Season Season

XL

XR

II L

II R
005
205

Mean no.

III
117
206

Inv/
296 319 246 333

Winter

0004

N
Spring

001
205

172
205

205

002
N N
516

002
516

185
519

009
527

120
529

Jan./80
Apr./80

Summer

001
1289
0'OO

0004
1289

119
1265

004
1289

124
1268

Autumn
N

000
1406

211
1369

004
1413

122
1402

1407

July/80
Oct./80
Feb./81

Winter
N

000
228

000
228
003

182
228

006
228

111
228

298 293
344
337

Spring
Summer

000
N N
470

179
467

004
471

108
467

470 36 355

000
36

000 000
000

231
36

014
36

Apr./81

Autumn
N

000
355

182
383

003
384

100 36 142
386

Jan./82 May/82
Total

Summer
N

000
257

198
254

004
251

115
259

318

257

Autumn
N

0004
566

000
566

183
563

005
568

113
564

300

0004

0006

184

0'06

117

308008

DISCUSSION

The simultaneous occurrence of such a large num-

ber of inversions in populations of Drosophila willistoni in Rio Grande do Sul, geographically near

the southern border of the species distribution,

raises the following question: how does this high degree of polymorphism contribute to the adapta-

tion of the populations to the environmental variations? In this State, due to its climatic characteristics, the effects of the meteorological conditions over flora, fauna and genetics of populations are probably more effective than in typically tropical Brazilian regions.

Although the frequencies of inversions are

similar in both places, the correlation matrix (table

POLYMORPHISM IN DROSOPHILA WILLISTONI

157

5) shows clearly that they play different roles. As far as populations size is concerned, the differences found in multiple regression for Itapu and Turvo

P<0.05) from Turvo, since they are sufficiently distant to allow free recombination. Asthis is not

occurring their association probably is advantageous for individuals possessing them. The same

show how much biotic and abiotic variables may act differently in both places. Thus, the populations from Parque do Turvo seem to depend much more on the macroclimatic conditions, especially on the previous minimum temperature, than on any other of the variables

explanation can be offered to those inversions

belonging to the different chromosomes, as "L D,E

and III C,H,J; IlL A and III C, "L H and III B

for Itapu, and "L A with III B,C, and IlL H with III C for Turvo (table 5).

considered. On the contrary, at Itapu, the

Interchromosomal associations have been

chromosomal constitution of the species (especially in regard to the inversions of chromosome III)
seems to be the most effective factor with humidity playing a minor role. Such results were surprising at first sight, since

found in several species of Drosophila as for inst-

ance, Da Cunha et a!. (1950) for D. willistoni,

Levitan and Salzano (1959) forD. guaramunu (see also the review by Levitan (1958), Brncic (1961) for D. pavani, Sene (1981) for D. mercatorum. Any

Itapu was considered a marginal environment when compared to Turvo as far as lepidoptera populations are concerned (Pansera and Arauijo,
1983). It would be expected then, that biotic factors

attempt of interpretation of such relationships should involve experimental work as well as a

deeper knowledge of the physiological role played by inversions.

should be more relevant at Turvo than at Itapu; this was not the case. In view of the circumstances we conjecture that at Turvo the climatic factors

Finally, the constant mean number of inversions per female (about 3) over the years and in both places leads us to suggest that probably it
reflects the action of stabilising selection, thus sup-

that regulate the cycle of plants on which the

populations of D. willistoni depend are preponderant, whereas at Itapu the unpredictability of the environment makes chromosome polymorphism a useful way to overcome that difficulty.

porting Carson's hypothesis of homoselection at the limit of the species distribution. Similar values
were reported by Da Cunha et al. (1950), Da Cunha

and Dobzhansky (1954), Cordeiro (1961), Cordeiro et aL (1960) in samples from Rio Grande do
Sul.

Unfortunately, we have not found in the

literature similar data in order to compare them with our data. However, it is also important to
point out that the coexistence and competition with other sympatric species (whose frequencies did not take part in the regression) must certainly play an important role in the regulation of Drosophila wil-

Findings from smaller latitudes where the mean number of inversions per female is around 9 (Da Cunha et aL, 1950) suggest that such populations

are close to the centre of origin of Drosophila

willistoni.

listoni populations. Pavan et al. (1957) have already suggested that when D. willistoni, D.
tropicalis and D. paulistorum are sympatric, the dominant one shows heterotic inversions. In our samples, however, sibling species of D. willistoni were not detected (in function of their own geographic distribution). A similar relationship of dominance between D. willistoni and D. simulans has been observed in the present study, although without any evident change in the frequencies of the heterotic inversions "L D, E. Heterotic inversions in Drosophila willistoni were recorded previously by many authors such
Acknowledgements Thanks are due to directors of Parque do Turvo and Parque de Itapu for allowing us to work in those

places. We also express our gratitude to 80 Distrito de

Meteorologia of Ministrio da Agricultura, Centro de Processamento de Dados of Universidade Federal do Rio Grande do Sul and to Conseiho Nacional de Desenvolvimento CientIfico e Tencolgico (CNPq), V Cmara (UFRGS), Fundao
(FAPERGS) and PNUDUNESCO (RLA 78024) for the financial support.

de Amparo a Pesquisa do Estado do Rio Grande do Sul

as Da Cunha (1953), Da Cunha et al. (1959),

Dobzhansky and Pavlovsky (1958) and Cordeiro et al. (1960), which seems to be advantageous for
the species in some extension, despite of increasing the homozygous mortality. Among the correlations involving inversions of the same chromosome arm we believe are the most

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0 0
-U

(I,

z
0 (I) 0
Insolation (hours) Preceding Collection month month 1321

Appendix 1

Meteorologicaldata of regions near Parque de Itapu and Parque do Turvo Temperature (C) Preceding month Temperature (C) Collection month Rainfa ii (mm) Preceding Collection month month Preceding Collection month month
Relative humidity (%)

()
Max.
Med. Mm.

()
907
875

Place

Month Year
Mm. Med.

Max. 147 175 111 169 207

Parque

198

N N

de

Itapu

228 289 259

138 193 183

235 216
1073

80
183 586 158

259 302 186 210 208 244 124 614 1344 907 243 231
135 191 253

1792 1334 914 584 1186

2002 2073
1575 1697 73
2468

1859 2541

C')

2050
1396 1322 1844 1980

293

2l65
146 1239 1082 810 2199 3346

78 70 75 79 82 78

2599 2577
1686 1618 1523 1553

282 278
22.5

186

116 207 193 185

234 299 299 247 245 201

146 140 195

97 110
195 239

95 159 220 206 164

Parque do Turvo

210 237 297

74
166 209 270

133 128

31i
297
221

21i
1875

1380 498 1079 3362 965

287 1036 1715

1556 1629 1658 1574

71

321 137 159

249

111 9.4 166 175 164

219 232 257 209

249 235 235

90

321

240 220 143 210 256

596 1216
1738

78 78 82 79 69 75 78

2122
1633

2824 2130 874

77
71

316 300

202 83 100 210 176

Sept./78 Dec./Jan. 78/79 Mar./Apr. 79 July/79 Oct./79 Mar./Apr. 80 Jan./81 Apr./81 July/81 July/79 Oct./79 Jan./80 Apr./80 July/80 Oct./80 Feb./81 Apr./81 Jan./82 May/82

18i
145

275 322 284 328

158 211 155 185 118

1493 1556 1785 1915

507 2038 357

1962 259 1326

79 74 73 80 74 73 76 75 81 84 71 79 79 80 71 74 71 72 73

61 77

2886
2075

609 2278 2583 2459 2472

01
Co