Daily activity patterns of two co-occurring tropical ... - Semantic Scholar

Journal of Insect Science: Vol. 9 | Article 54

Peixoto and Bensen

Daily activity patterns of two co-occurring tropical satyrine butterflies Paulo Enrique Cardoso Peixotoa and Woodruff W. Bensonb Institutional Affiliation: Department of Zoology, IB, State University of Campinas, São Paulo, Brazil, C.P. 6109, 13083-970

Abstract 5F=E>2=6D2?576>2=6D@7>2?J:?D64EDA64:6D2C66IA64E65E@25;FDEE96:C52:=J24E:G:EJA2EE6C?:?@C56CE@2G@:5 DEC6DD7F=4=:>2E:44@?5:E:@?D2?5:?4C62D6E96492?46DE@6?4@F?E6CD6IF2=A2CE?6CD-D:?8D42?D2>A=:?8>6E9@5D 2DD@4:2E65H:E97@42=:?5:G:5F2=@3D6CG2E:@?D:EH2D7@F?5E92EEH@D2EJC:?63FEE6C7=:6D@7D:>:=2CD:K62?5>@CA9@=@8J Hermeuptychia hermes 23C:4:FD$6AE:5@AE6C2&J>A92=:5262?5 Paryphthimoides phronius @52CED9@H4@>A=6E6=J 5:776C6?E52:=J24E:G:EJA2EE6C?D@?7@C6DE6586D:?D@FE962DE6C?C2K:= Hermeuptychia hermes AC6D6?ED@?623F?52?46A62< :?E96>@C?:?82?52?@E96C:?E96=2E627E6C?@@?H9:=6 P. phronius 23F?52?46A62:552JC6>2:?:?8DE23=6 F?E:=

9,9:D5:776C6?46:DAC@323=J5F6E@E96@44FCC6?46@7E6CC:E@C:2=3692G:@C:?E96=2E6CDA64:6D,96368:??:?8 @7E6CC:E@C:2=5676?D63J P. phronius >2=6D4@:?4:565H:E9E96E:>6@7?6H3@C?76>2=624E:G:EJ @H6G6C?6H=J92E4965 76>2=6DH6C6?@ED6IF2==JC646AE:G6,9627E6C?@@?E6CC:E@C:2=:EJ:?>2=6 P. phronius >2J36:?A2CEC6=2E65E@>2E6 24BF:D:E:@? @H6G6CH9JE9623F?52?46@7 H. hermes 564C62D6DH96?E9623F?52?46@7 P. phronius :?4C62D6D:D=6DD4=62C Keywords: Satyrinae, mating tactics, territoriality, daily activity pattern, mate location Correspondence: [email protected], [email protected] Received: 13 February 2008 | Accepted: 8 October 2008 | Published: 14 July 2009 Copyright: This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 9, Number 54 Cite this paper as: Peixoto P, Benson WW. 2009. Daily activity patterns of two co-occurring tropical satyrine butterflies. 14pp. Journal of Insect Science 9:54, available online: insectscience.org/9.54

Journal of Insect Science | www.insectscience.org

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Figure 1. Daily variations of temperature on the dry and the rainy seasons averaged across all areas and 5 sampling days per season. Bars indicate standard error.


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Peixoto and Bensen

Table 1. Behavioral ethogram of Paryphthimoides phronius males Behavior

Description

Perch

Male perched on the vegetation or on the ground

Flight

Horizontal shift inside the territory area or over greater distances

Forage

Adult feeding on grass inflorescence or decomposing fruit

Territorial patrol

Back and forth flight, inside and adjacent to the territory area

Territorial defense

Linear, circular or spiral chases between two P. phronius males

Territorial display Periods during the territorial defense when both males remain perched in opposite directions, flapping its wings at regular intervals Copulation

Male and female with its abdominal tips in contact

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@? 6249 9@FC 2?5 D2>A=6 52J H2D DEC@?8=J 4@CC6=2E65 H:E9 E6>A6C2EFC6 @? E96 5CJ $:?62C C68C6DD:@? 3 C A

2?5 E96 C2:?J $:?62C C68C6DD:@? 3 C A

D62D@?D :8FC6 642FD6 @7 E96 DEC@?8 2DD@4: 2E:@? 36EH66? E96 EH@ G2C:23=6D 2?5 E96 ;F58>6?E E92E 9F>:5:EJ H2D A@E6?E:2==J =6DD :>A@CE2?E E92? E6>A6C2EFC6 2D2DEC6DD724E@C9F>:5:EJH2D6I4=F5657C@>2?2=JK6D Daily behavioral pattern of P. phronius males 6EH66? DF446DD:G6 4@F?E EC2?D64ED C2?5@>=J 49@D6? P. phronius >2=6D H6C6 @3D6CG65 7@C >:? H:E9 E96 2:5 @7 2 >:?: 42DD6EE6C64@C56C 2?5 E96 E:>6 :E DA6?E :? 6249 24E:G :EJ H2D 42=4F=2E65 ,23=6 @42= @3D6CG2E:@?D H6C6 C6 DEC:4E65 E@ >2=6D 3642FD6 >2E6 D62C49 :D 4@?46?EC2E65 :? 6IA@D65 A=246D H96C6 @3D6CG2E:@? :D C6=2E:G6=J 62DJ H96C62D 76>2=6D H6C6 ?@E 4@>>@? @? E96 DEF5J 2C62 2?5 H96? AC6D6?E @44FC @? E96 F?56C3CFD9 H96C6 E96J 2C65:77:4F=EE@:?FE6D ,96 6I4=FD:@? @7 E96D6 @3D6CG2 E:@?D 7C@> 2?2=JD6D 4@F=5 DF3DE2?E:2==J C65F46 E96 ?F> 36C @7 D2>A=65 >2=6D 2?5 H@F=5 :?EC@5F46 2 3:2D 3642FD6 >@C6 24E:G6 >2=6D H6C6 AC@323=J >@C6 5:77:4F=E E@ 7@==@H 7@C E96 E@E2= @3D6CG2E:@? A6C:@5 ,9FD H6 @AE65 E@ :? 4=F56 :? E96 2?2=JD6D :?5:G:5F2=D E92E H6C6 7@==@H65 7@C =6DD E92? >:? 2?5 :? @C56C E@ C65F46 E96 5:776C6?46D :? E96 @3D6CG2E:@? E:>6 2>@?8 >2=6D E96 AC@A@CE:@? @7 E:>6DA6?E:?62493692G:@CH2DFD65 Behavior of sexually receptive P. phronius females !? @C56C E@ :?G6DE:82E6 E96 :?7=F6?46 @7 D6IF2==J C646AE:G6 76>2=6D @? E96 24E:G:EJ A2EE6C? @7 >2=6 P. phronius 76>2=6D H6C6 C62C65 7C@> 688D :? E96 =23@C2E@CJ 2?5 C6=62D65 :? E96 7:6=5 88D H6C6 @3E2:?65 7C@> 7:6=542F89E 8C2G:5 76 >2=6D :?5F465 E@ @G:A@D:E 3J 9@=5:?8 E96> :? A=2DE:4 328D F?56C 2 56D< =2>A $2CG26 H6C6 C2:D65 D6A2C2E6=J :?

>= A=2DE:4 3@EE=6D >2:?E2:?65 ?62C 2 H:?5@H 2E C@@> E6>A6C2EFC6 ,96 3@EE=6D H6C6 4=62?65 6G6CJ 52J 2?5 =2CG26 H6C6 765 EH:46 2 52J H:E9 7C6D9 4FEE:?8D 7C@> E96 9@DEA=2?E Axonopus compressus +H2CED 62FG (@2=6D(@24626

4


Peixoto and Bensen

Table 2. Repeated measures ANOVA comparing total counts of Paryphthimoides phronius and Hermeuptychia hermes over the day in the area 1. The analysis was conducted separately for each season.

Season

Effect

SS

df MS

Dry

Species

6.25

1

6.25

Dry

Residual

59.44

8

7.43

Dry

Hour

35.89

9

3.98

Dry

F

P

0.84

0.386

2.47

0.016

Hour*species 34.05 9 3.78 2.34 0.022

Dry

Residual

116.16 72 1.61

Rainy

Species

10.24

1 10.24 3.11

Rainy

Residual

26.32

8

3.29

Rainy

Hour

44.36

9

4.93

Rainy Rainy

3.24

0.116 0.002

Hour*species 88.16 9 9.57 6.28 :==:8C2> @? E96 52J 7@==@H:?8 AFA2E:@? 6>2=6D H6C6 4=62C=J D6A2C 23=6 7C@> >2=6D 32D65 @? E96 AFA2= H6:89E 367@C6 6>6C 86?46 C@> 2 E@E2= @7 D2>A=6D :? 2 A:=@E DEF5J 2== >2=6D ? 6>6C865 7C@> AFA26 H:E9 E@ >8 2?5 2== 76>2=6D 6>6C865 H:E9 E@

>8 ? ,96C6 H6C6 ?@ AFA26 H:E9 E@ >8 2?5 E96C67@C6 ?@ H6:89E @G6C=2A 36EH66? E96 D6I6D %@C6@G6C :E H2D A@D D:3=6 E@ AC65:4E E96 52J @7 6>6C86?46 3642FD6 2== AFA26 492?865 4@=@C H96? 25F=ED H6C6 C625J E@ 6>6C86 6?46 AFA26 H6C6 A=2465 :? E96 7:6=5 @? E96 52J @7 6>6C86?46 E@ 2G@:5 A@DD:3=6 76>2=6 3692G:@C2= 492?86D 42FD65 3J 92?5>2?:AF=2E:@?@725F=ED ,96 9@FC @7 76>2=6 6>6C86?46 H2D >62DFC65 2D E96 AFA2= D2=6 AFA26 H6C6 2==@H65 E@ 6>6C86 U?2EFC2==JV 2 D9@CE 5:DE2?46 E@ > 7C@> A=246D 7C6 BF6?E65 3J 25F=E >2=6D == 3692G:@C2= @C >@CA9@=@8:42= 492?86D H6C6 E:>65 27E6C AFA2= D2=6D H6C6 @3D6CG65 7@C 9 27E6C E96:C 7:CDE DA@?E2?6@FD 7=:89E@CF?E:=E96JH6C6=@DE7C@>D:89E !? 255:E:@? :? @C56C E@ :?G6DE:82E6 A@DD:3=6 56=2JD :? E96 76>2=6 D6IF2= C646AE:G:EJ @E96C =23@C2E@CJC2:D65 76 >2=6D @?6 E@ 7@FC 52JD @7 286 H6C6 @3D6CG65 :? E96 7:6=5 3J 42C67F==J C6=62D:?8 E96> @? =@H G686E2E:@? ?62C E6CC:E @C:2= >2=6D -D:?8 2 3=24< >2C2=6D H6C6 >2C:E E96:C G:DF2= :56?E:7:42E:@? H:E9@FE C642AEFC6 7E6C C6=62D:?8 E96 36 92G:@C @7 E96D6 76>2=6D H2D @3D6CG65 7@==@H:?8 E96 D2>6 AC@E@4@=FD657@C?6H3@C?76>2=6D


Results Daily abundance pattern Hermeuptychia hermes H2D 7@F?5 @?=J :? 2C62 2?5 2= E9@F89 ?F>6C:42==J D:>:=2C E@ P. phronius D66>65 E@ 5:776C :? E96 =6G6=D @7 24E:G:EJ 5FC:?8 E96 52J :? 5CJ 2?5 E96 C2:?J D62D@?D ,23=6 Hermeuptychia hermes 92D EH@ 52:=J 23F?52?46 A62

E@

9 2?5 E96 @E96C :? E96 27E6C?@@? 2E

9 :8FC6 FC:?8 E96 5CJ D62D@? E96 A62:552J

T

9 2?5

T

9 2?5 H6C6 D6A2C2E65 3J 23C:673FE>2C E92E @7 H. hermes 3FE H2D BF2=:E2E:G6=J D:>:=2C 36EH66? E96 EH@ 2C62D :8FC6 !? 2C62 >2=6 23F?52?46 D9@H65 2 D>2== :?4C62D6 2C@F?5 >:552J :? 3@E9 D62D@?D !? E96 5CJ D62D@? >2=6 23F?52?46 C6 >2:?65 DE23=6 F?E:= =2E6 27E6C?@@? 27E6C H9:49 6?4@F?E6C C2E6 564C62D65 DF556?=J ,96 C2:?J D62D@? A2EE6C? H2D D:>:=2C 2=E9@F89 >2=6D H6C6 24E:G6 6G6? =2E6C F?E:= 27E6C

9 !? 2C62 P. phronius >2=6D A626 >2=6D H6C6 DE:== AC6D6?E 27E6C E9:D E:>6 5FC:?8E96C2:?JD62D@? !? E96 2C62 E96 23F?52?46 @7 P. phronius >2=6D D9@H65 D:>:=2C :?4C62D:?8 C2E6D H:E9 E6>A6C2EFC6 :? 3@E9 D62D@?D ,23=6 :8FC6 !? E96 2C62 E96 A2EE6C? H2D D:>:= 2C 2=E9@F89 2? :?E6C24E:@? 36EH66? D62D@? 2?5 E6>A6C 2EFC6 4@F=5 36 D66? H:E9 2 9:896C :?4C62D6 :? >2=6 23F?52?46 7@C 6249 6=D:FD 568C66 5FC:?8 E96 C2:?J D62 D@? H96? 4@>A2C65 E@ E96 5CJ D62D@?

5


Peixoto and Bensen

Figure 2. Variation in the abundance of Hermeuptychia hermes along forest edge transects taken hourly over the day. Bars represent the 95% confidence interval for means of total count based on samples from 5 days.

&'.E6>A6C2EFC6D62D@? A

:8FC6 Hermeuptychia hermes @? E96 @E96C 92?5 AC6D6?E65 2 G6CJ 5:776C6?E A2EE6C? !? 2C62 :ED 23F?52?46 H2D ?@E C6=2E65 E@ E6>A6C2EFC6 G2C:2E:@? :? E96 5CJ D62D@? 5FC:?8 E96 52J 3FE :? E96 H6E D62D@? :ED 23F?52?46 564=:?65 H:E9 :?4C62D:?8 E6>A6C2EFC6D &'.E6>A6C2EFC6D62D@? A

:8FC6 Table 3. Covariance analyses showing the relationship between mean male abundance of Paryphthimoides phronius and temperature along the day and season in the area 1. Effect

ANCOVA

P

Season

F(1,18) = 0.22

0.643

Temperature

F(1,18) = 9.27

0.008

Season*temperature

F(1,18) = 0.10

0.754

Daily behavioral pattern of P. phronius males ,96 52:=J 3692G:@C2= A2EE6C? 42=4F=2E65 7C@> 7@42= @3D6C G2E:@?D @? >2=6 P. phronius H6C6 G2C:23=6 2?5 E96 D>2== D2>A=6 D:K6 >256 :E A@DD:3=6 E@ C64@8?:K6 @?=J DEC@?86C E6?56?4:6D ,23=6 :8FC6 FEE6C7=J 7665:?8 @? Journal of Insect Science | www.insectscience.org

564@>A@D:?8 7CF:E 2?5 8C2DD H2D @3D6CG65 2=>@DE 6I4=FD :G6=J :? E96 62C=J >@C?:?8 2?5 =2E6 27E6C?@@? ,9:D A2E E6C? :D :? 244@C52?46 H:E9 @442D:@?2= @3D6CG2E:@?D >256 5FC:?8 E96 DEF5J %2=6D D9@H65 E6CC:E@C:2= 3692G:@C AC6 5@>:?2?E=J 7C@>

E@

9 ,9:D 3692G:@C H2D 6G:56?E 3@E9 5FC:?8 E96 D42?D 2?5 E96 7@42= @3D6CG2E:@?D @? >2=6D =:89ED H6C6 @3D6CG65 2== 2=@?8 E96 52J 2E 2 C2E6 @7 2AAC@I:>2E6=J T ,96 >@DE 7C6BF6?E 24E:G:EJ H2D A6C49:?8 H9:49 @44FA:65 @? 2G6C286 + @7E96>2=624E:G:EJ3F586E ,96 E6CC:E@C:6D @7 >2=6D P. phronius E92E 6?82865 :? E6CC:E @C:2= A2EC@= 2?5 5676?D6 H6C6 86?6C2==J =@42E65 :? DF??J 4=62C:?8D ?62C EC66D @? E96 7@C6DE 6586 2=E9@F89 2 76H 42D6D @7 >2=6D 5676?5:?8 2C62D H:E9 564@>A@D:?8 72==6? 7CF:ED H6C6 @3D6CG65 ,96 D2>6 E6CC:E@C:6D E6?565 E@ 36 FD65 2E E96 D2>6 D:E6D @? 5:776C6?E DFCG6J 52JD 2?5 3J 5:D E:?4E E6CC:E@C:2= >2=6D E92E 86?6C2==J C6>2:?65 A6C4965 @? E96=627=:EE6C@C@?=62G6DFAE@ 4>7C@>E968C@F?5 @42= >2=6D E92E 5676?565 E6CC:E@C:6D EJA:42==J 7=6H E@ H2C5D 2?J 3FEE6C7=J E92E A2DD65 ?62C3J 6G6? >F49 =2C86C DA64:6D @H6G6C E96 5:DAFE6D :?G@=G:?8 4@?DA64:7:4 >2=6D H6C6 =@?86C >62? D + D E92? 6


Peixoto and Bensen

Figure 3. Variation in the abundance of Paryphthimoides phronius along forest edge transects taken hourly over the day on area 1 (A) and area 2 (B). Bars represent the 95% confidence interval for means of total count based on five sample days.


7


Peixoto and Bensen

Figure 4. Relationship between mean male abundance of Paryphthimoides phronius and temperature in area 1 (A) and area 2 (B). Each point represents the temperature and male total counts averaged per hour across five sampling days.


8


Peixoto and Bensen

Figure 5. Relationship between mean abundance of Hermeuptychia hermes and temperature. Each point represents the temperature and butterfly total count averaged per hour across five sampling days.

5:DAFE6D 36EH66? P. phronius >2=6D 2?5 @E96C 3FEE6C7=J DA64:6D >62? D + D %2??/9:E?6J E6DE - ? ? A

!?E6C24E:@?D 36EH66? P. phronius >2=6D FDF2==J 4@?D:DE65 @7 2 D6C:6D @7 C6=2E:G6=J =6?8E9J 4:C4F=2C @C DA:C2= 7=:89ED E92E 6?565 H96? @?6 @7 E96 3FEE6C7=:6D =67E E96 2C62 ,96D6 AFCDF:ED D@>6E:>6D >@G65 72C 2H2J 7C@> E96 2C62 E92E H2D 4@> >@?=J :?DA64E65 3J E96 E6CC:E@C:2= >2=6 7E6C 2 492D6 3@E9 C6D:56?E 2?5 :?ECF56C >2=6D 4@F=5 C6EFC? 2?5 A6C49 282:? :? E96 E6CC:E@CJ 367@C6 C6:?:E:2E:?8 E96 5:DAFE6 FC:?8 E96 4@?E6IE 3@E9 C:G2=D D@>6E:>6D :?E6CCFAE65 7=:89E 2?5 H9:=6 A6C49:?8 @? =62G6D :?D:56 E96 E6CC:E@CJ @7E6? @A6?65 2?5 4=@D65 E96:C H:?8D 2E C68F=2C :?E6CG2=D 6C:2= 5:DAFE6D H6C6 EJA:42==J 4@?5F4E65 2E 9:89 DA665D D@ E92E :E H2D FDF2==J ?@E A@DD:3=6 E@ D66 56E2:=D @7 E96 4@? E6DEE24E:4D@7E964@?E6?56CD

Table 4. Number of observed males and total observation time for each hour during the daily activity pattern sampling (for wet and dry seasons) of Paryphthimoides phronius. Hour N 8:30

6

Expected time (min)

Total time observed (min)

10800

4869.3 (45%)

9:30

5

9000

4724.8 (52%)

10:30

9

16200

9897.8 (61%)

11:30

8

14400

11602.7 (81%)

12:30

9

16200

9886.9 (61%)

13:30

8

14400

11114.4 (77%)

14:30

7

12600

11713.1 (93%)

15:30 10

18000

11486.4 (64%)

16:30

16200

8574.0 (53%)

9

The expected time was calculated as the total time that would have been expended if we had accompanied all males for complete 30 min. Values between parentheses represent the proportion of the observations done in relation to the total expected time.


9


Peixoto and Bensen

Figure 6. Daily activity budget of male Paryphthimoides phronius (based on 71 behavioral observations, 48 during the rainy season and 23 during the dry season). Each hourly estimate is based on means of five butterflies observed each on separate days. Bars represent standard error. Time spent perching is not shown.

=E9@F89 7@42= @3D6CG2E:@?D @? H. hermes H6C6 ?@E 4@? 5F4E65 6IE6?D:G6 @3D6CG2E:@?D @? E9:D 3FEE6C7=J 2E E96 DEF5J 2C62 AC@G:56 ?@ 6G:56?46 @7 E6CC:E@C:2= 3692G:@C FC:@FD=J H. hermes 5676?5 E6CC:E@C:6D :? >:5 2?5 =2E6 27 E6C?@@? :? @E96C C6D6CG6D A6CD@?2= @3D6CG2E:@?D FD:?8 3692G:@CDC6D6>3=:?8E9@D6@7 P. phronius Behavior of sexually receptive P. phronius females 6>2=6D C2:D65 :? E96 =23@C2E@CJ 6>6C865 36EH66?

9 2?5 9 ? :8FC6 ,96 >2;@C:EJ 6>6C865 36EH66?

2?5

9 2?5 @?=J @?6 6>6C865 367@C6

9 ,96 6>6C86?46 E:>6 @7 >2=6D H2D ?@E >62DFC65 5FC:?8 E96 DE2?52C5 @3D6CG2E:@?D @H6G6C D:I >2=6D @3 D6CG65 :? 2 A:=@E DEF5J 6>6C865 36EH66? 2?5

9 ,96 76>2=6D E92E 6>6C865 7C@> AFA2 2?5 H6C6 EC2?D A@CE65 E@ E96 7:6=5 @? E96 52J @7 6>6C86?46 D9@H65 G6CJ D:>:=2C 3692G:@CD C6D9=J 6>6C865 76>2=6D EJA:42==J C6 >2:?65 A6C4965 @? @C ?62C E96 AFA2= DA=6E6 H:?8 6IA2?D:@? >62? >:? C2?86 T 2?5 D@>6E:>6D H2=2=6D H2D 7@==@H65 F? E:= E96J 7=6H D6G6? @7 E96> 7@C 2? 255:E:@?2= 9@FC 27E6C E96 7:CDE 7=:89E &@?6 @7 E96D6 76>2=6D @C:6?E65 E@ ?62C3J >2=6D @C E@ 2E65 76>2=6D H6C6 DF3D6BF6?E=J C6D:89E65 ?62C E96:C C6=62D:?8 A@:?ED @?6 @7 E96> H2D C6A62E65=J @3D6CG65 2E 2 A=246 ?62C 564@>A@D :?8 7CF:ED @AF=2E:@?D :?G@=G:?8 EH@ @E96C 76>2=6D H6C6 @3D6CG65 2>@?8 E96 FD65 :? E96 C6=62D6 6IA6C:>6?E '?6 :?G@=G65 E96 76>2=6 E92E H2D C6=62D65 H96? :E H2D 7@FC52JD@=5 ,9:D 3FEE6C7=J 7=6H E@ E96 42?@AJ 27E6C :ED C6=62D6 2?5 2AAC@I:>2E6=J >:? =2EE6C C6EFC?65 6?E6C65 2 >2=6 E6CC:E@CJ ?@ 5642J:?8 7CF:E H2D ?62C3J 2?5 H2D 4@FCE65 2?5 >2E65 ,@ DE2CE E96 4@AF=2E:@? E96 >2=6 7=6H :? 5:C64E:@? @7 E96 76>2=6 F?E:= E96J A6C4965 2?5 4@?E24E65 E96:C 235@>6?D 7E6C E96J C6>2:?65 :? 4@? E24E 7@C >:? E96 >2E:?8 A2:C D6A2C2E65 2?5 E96 76>2=6 7=6H @77 ,96 @E96C 4@AF=2E:@? :?G@=G65 2 76>2=6 E92E H2D C6=62D65 H96? :E H2D E9C6652J@=5 3FE H2D @3D6CG65 >2E:?8 @? E96 52J 7@==@H:?8 :ED C6=62D6 5FC:?8 E96 7@42= >2=6 @3D6CG2E:@? A6C:@5 36EH66? >2=6 D42?D ,96 4@AF =2E:@? DE2CE65 2E

9 27E6C E96 2CC:G2= @7 E6CC:E@C:2= >2=6D :? E96 2C62 ?62C 3FE ?@E :?D:56 2 E6CC:E@CJ 4@> >@?=J @44FA:65 3J >2=6D @H6G6C E96 >2E:?8 >2=6 H2D ?@E >2C2E:?8 E24E:4 E6CC:E@C:2= @C ?@E 4@F=5 ?@E 36 :56?E:7:65 367@C6 E96 4@AF=2E:@? DE2CE65 ,96 >2E:?8 H2D @3D6CG65 7@C >:?FE6D F?E:= G:DF2= 4@?E24E H2D=@DE


Discussion ,96D6 C6DF=ED D9@H E92E EH@ >@CA9@=@8:42==J D:>:=2C D2EJC:?26 3FEE6C7=:6D E92E @44FC :? E96 D2>6 2C62 AC6D6?E 5:776C6?E 52:=J 23F?52?46 A2EE6C?D Hermeuptychia hermes AC6D6?E65 EH@ 23F?52?46 A62@C?:?8 2?5 C6 >2:?65 DE23=6 5FC:?8 E96 27E6C?@@? Paryphthimoides phronius 23F?52?46 :?4C62D65 H:E9 E6>A6C2EFC6 5FC:?8 3@E9 D62D@?D H9:=6 H. hermes 23F?52?46 H2D ?@E 27764E65 5FC :?8 E96 5CJ D62D@? 3FE 564C62D65 5FC:?8 9@E A6C:@5D :? E96 C2:?J D62D@? %2=6D @7 P. phronius DE2CE65 E@ 5676?5 E6CC:E@C:6D :? E96 =2E6 >@C?:?8 H9:=6 >2=6D @7 H. hermes H6C6 ?@E D66? 25@AE:?8 E9:D 3692G:@C =E9@F89 76>2=6D 5:5 ?@E D66> E@ 36 D6IF2==J C646AE:G6 @? E96 52J E96J 6>6C865 E96 E:>6 E92E >2=6D @7 P. phronius 3682? E@ 56 76?5 >2E:?8 2C62D 4@:?4:565 H:E9 E96 E:>6 E92E ?6H=J 6>6C865 76>2=6D 3682? E@ 7=J @H6G6C D:?46 E96 =@H ?F>36CD @7 ?6H=J64=@D65 76>2=6D C6=62D65 :? E96 7:6=5 H2D 2DD@4:2E65 H:E9 5:77:4F=E:6D :? 7:6=5 @3D6CG2E:@? DEC@?8 :?76C6?46D @? E96 A@DE64=@D:@? 76>2=6 C646AE:G:EJ A6C:@5 42??@E 36 >256 2?5 E96D6 4@?4=FD:@?D >FDE 36 EC62E65 H:E9 42FE:@? !? E9:D D6?D6 :E D66>D :?E6C6DE:?8 E@ ?@E6 E92E 2?@E96C EC@A:42= 3FEE6C7=J +2EJC:?26 DA64:6D

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Journal of Insect Science: Vol. 9 | Article 54 Hypolimnas bolina 2=D@ ?665D 2AAC@I:>2E6=J 7@FC 52JD E@ 364@>6 C646AE:G6 #6>A

=E9@F89 3FEE6C7=:6D 2D 2 CF=6 5:D2AA62C 7C@> D:89E :? E96 27E6C?@@? E96 EH@ D>2== D2EJC:?6D :?G6DE:82E65 96C6 H6C6 24E:G6 @G6C 6DD6?E:2==J E96 6?E:C652J=:89EA6C:@5 '?6 >:89E 6IA64E E92E 4=@D6=J C6=2E65 DA64:6D @7 D:>:=2C 2AA62C2?46 2?5 923:ED D9@F=5 C6DA@?5 D:>:=2C=J E@ 6?G:C @?>6?E2= 4@?5:E:@?D 2?5 252AE :? D:>:=2C H2JD H96? @4 4FCC:?8 E@86E96C &6G6CE96=6DD E96 5:776C6?46D :? E96 86? 6C2= 23F?52?46 A2EE6C?D @G6C E96 52J 36EH66? H. hermes 2?5 P. phronius DF886DE 6:E96C E92E E96J 92G6 5:776C6?E=J 252AE65 E@ 4@>>@? A9JD:42= 4@?5:E:@?D @C E92E @E96C 3:@=@8:42= 5:776C6?46D :? E96 EH@ DA64:6D E2:=2C 5:776C6?46D :? E96 52:=J 24E:G:EJ A2EE6C? 92G6 366? C6A@CE65 7@C D>2== D2EJC:?6 3FEE6C7=:6D :? EC@A:42= FDEC2=:2 C23J 44@C5:?8 E@ E9:D DEF5J E9C66 DJ>A2EC:4 DA64:6D @7 Micalesis +2EJC:?26 DF3EC:36 %J42=6D:?2 E6?5 E@ D9:7E 24E:G:E:6D DJ>>6EC:42==J 2C@F?5 >:552J H:E9 @?6 DA64:6D >@C6 24E:G6 2C@F?5 >:552J 2?5 E96 @E96C EH@ H:E9 3:>@52= 24E:G:EJ A6C:@5D E92E A62@C?:?8 2?5 282:? :? E96 27E6C?@@? D@>6H92E D:>:=2C A2EE6C? @44FCC65 H:E9 EH@ DJ>A2EC:4 Hypocysta DA64:6D +2EJC:?26 DF3EC:36 @6?@?J>A9:?2 :? H9:49 @?6 DA64:6D A62:552J 2?5 E96 @E96C :? E96 62C=J27E6C?@@? ,96 5:776C6?46D :? E96 24E:G:EJ A6C:@5D 36EH66? P. phronius 2?5 H. hermes AC@323=J H6C6 2 4@?D6BF6?46 @7 E96 4@? DA:4F@FD E6CC:E@C:2= 3692G:@C @7 E96 7@C>6C DA64:6D >22=6D G:D:3=6 2=@?8 A2E9D 2E E96 E:>6 >2=6D @7 H. hermes H6C6 5:DA6CD65 :? E96 3CFD9 !E D66>D C6=6G2?E E92E E96 E9C66 DA64:6D @7 Micalesis DEF5:65 3J C23J 2=D@ D6A 2C2E65 :?E@ AC6DF>23=J ?@?E6CC:E@C:2= DA64:6D E92E H6C6 24E:G6 62C=J 2?5 =2E6 M. perseus 2?5 M. sirius H96? E96 DF? :D =@H :? E96 D6C:42 >2J 36 2?2=@8@FD @H6G6C :E :D ?@E 4=62C H9J >2=6D 5676?5 E6CC:E@C:6D @? DF??J 2C62D H96? E6>A6C2EFC6 :D 9:89 ,96 E6CC:E@C:2= 56 76?D6 @7 2C62D H:E9 9:89 =:89E :?4:56?46 DF49 2D DF?DA@ED @C DF??J 4=62C:?8D :D 4@>>@? 2>@?8 3FEE6C7=J DA64:6D #?2AE@? C23J 6??:D 2?5 +A2C2? 6E 2=

!? E96D6 D:EF2E:@?D :7 76>2=6D 36 4@>6 C646AE:G6 @?=J :? E96 27E6C?@@? 2?5 FD6 E96D6 D:E6D 2D =2?5>2C< A@:?ED E@ 6?4@F?E6C D6IF2= A2CE?6CD >2=6D >2J 36 7@C465 E@ @44FAJ DF??J 2C62D 6G6? 5FC:?8 9@E A6C:@5D '? E96 @E96C 92?5 E96 E6>A@C2= 2?5 DA2E:2= D6A2C2E:@? D66? :? E96D6 D2EJC:?6D :D DE:== =:EE=6 F?56CDE@@5 2?5 E96 A@DD:3:=:EJ E92E 24E:G:EJ 5:776C6?46D 2C6 5F6 E@ :? E6CDA64:7:4:?E6C76C6?4642??@E366I4=F565 ,96 E6CC:E@C:2= 3692G:@C E92E H6 5@4F>6?E65 :? P. phronius :D D:>:=2C E@ E92E C6A@CE65 7@C @E96C E6CC:E@C:2= 3FEE6C 7=:6D #6>A 2?5 /:2E:?8 E6CC:E@CJ D:E6D 2C6 5676?565 3J C6D:56?E >2=6D E92E C6EFC? @? DF4 46DD:G6 52JD A6CD@?2= @3D6CG2E:@? *6D:56?E >2=6D 7=J 2E Journal of Insect Science | www.insectscience.org

Peixoto and Bensen 2 H:56 C2?86 @7 @E96C :?D64ED 3FE =6?8E9J 4@>32ED @?=J @44FC H96? E96 :?ECF56C :D 2 4@?DA64:7:4 >2=6 (FCDF:ED :?4=F56 =:?62C DA:??:?8 2?5 @C DA:C2=:?8 7=:89ED 2?5 4@? E6DED 6?5 H:E9 E96 56A2CEFC6 @7 @?6 :?5:G:5F2= '? E96 @E96C 92?5 E96 E6CC:E@C:2= 5676?D6 @3D6CG65 :? P. phronius 5FC:?8 3@E9 5CJ 2?5 C2:?J D62D@?D :D :? 4@?EC2DE E@ @E96C +2EJC:?26 DA64:6D :? H9:49 E96 @44FCC6?46 @7 E6CC:E@C:2=:EJ :D D62D@?2= /:42? 2?5 /::E65 @3D6CG2E:@?D @? >2E:?8 6G6?ED DF886DE E92E E96 52:=J E6CC:E@C:2= E6?FC6 @7 P. phronius 4@CC6DA@?5D E@ E96 A6C:@5 :? H9:49 C646AE:G6 76>2=6D 364@>6 2G2:=23=6 7@C >2E:?8 2=@?8 7@C6DE 6586D ,9:D EJA6 @7 C6=2E:@? >265 E@ 36 252AE:G6 3FE 76H DEF5:6D 92G6 24EF2==J D@F89E 2 4@CC6DA@?56?46 36EH66? E96 >2E6 =@42E:@? D4965F=6 :? >2=6D 2?5 76>2=6 2G2:=23:=:EJ #6>A 2?5 *FE@HD:?8 @7 76>2=6 6>6C86?46 7C@> AFA26 :D ?@E 42FD2==J C6=2E65 E@ E96:C 2CC:G2= 2E E6CC:E@C:6D *2E96C 76>2=6D D66> E@ C6BF:C6 D6G6C2= 52JD 27E6C 6>6C86?46 E@ 364@>6 D6IF2==J C646AE:G6 23@FE 52JD :7 E96 52E2 2C6 C6AC6D6?E2E:G6 3FE H96? 2EE2:?:?8 >2EFC:EJ E96D6 @=56C C646AE:G6 76>2=6D >2J D9@H FA 2E E6CC:E@C:6D 2C@F?5 >:5 52J !E >2J 36 A@DD:3=6 E92E E@ 96=A 8F2C2?E66 >2E:?8 H:E9 2 BF2=:EJ >2=6 76>2=6D D9@F=5 H:E99@=5 >2E:?8 H:E9 C@G:?8 >2=6D DF49 2D >:89E @44FC :? E96 >@C?:?8 2?5 D66< >2E:?8 6I4=FD:G6=J :? E96 27E6C?@@? H:E9 E6CC:E@C:2= >2=6D 2E WC6?56KG@FDX D:E6D #6>A

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Journal of Insect Science: Vol. 9 | Article 54 E6CC:E@C:2=:EJ D9@F=5 @44FC :? P. phronius 2?5 ?@E :? H. hermes :D=6DD4=62C ,6CC:E@C:2=:EJ :? 3FEE6C7=:6D :D 4@>>@?=J 25@AE65 2D >2E6 24BF:D:E:@? E24E:4 #6>A 2?5 /:A 2?5 *FE@HD2E:4 G2C:2E:@?D :?7=F6?46D E96 >2=6 52:=J 24E:G:EJ A2EE6C? E96 ?665 E@ 7:?5 D6IF2==J C646AE:G6 A2CE?6CD >2J C6AC6D6?E 2 DEC@?86C 56 E6C>:?2?E @7 >2=6 3692G:@CD :? E96 EC@A:4D !? E9:D 4@? E6IE :7 76>2=6D FD6 DA64:7:4 6?4@F?E6C D:E6D E@ >2E6 >2=6D >2J 36 7@C465 E@ @44FAJ DF3@AE:>2= A=246D :6 DF??J @C 9@E 2C62D H96? E6>A6C2EFC6D 2C6 ?@?@AE:>2= '? E96 @E96C 92?5 :? DA64:6D E92E 5@ ?@E ?665 E@ FD6 DA64:7:4 >2E:?8 2C62D >2=6D 2?5 76>2=6D >2J C6DA@?5 E@ E6>A6C 2EFC6 G2C:2E:@?D :? 2 >@C6 AC65:4E23=6 H2J @44FAJ:?8 DF??J 2C62D H96? E6>A6C2EFC6 :D =@H 2?5 D92565 2C62D 5FC:?8 6IEC6>6=J 9@E A6C:@5D @H6G6C >@C6 52E2 @? EC@A:42= DA64:6D 2?5 6DA64:2==J DEF5:6D 56D:8?65 E@ D:>F= E2?6@FD=J E6DE E96 :?7=F6?46 @7 76>2=6 2G2:=23:=:EJ 2?5 4=: >2E:4 4@?5:E:@?D @? >2=6 3692G:@C 5FC:?8 E96 52J 2C6 ?646DD2CJE@E6DEE9:DA@DD:3:=:EJ

Acknowledgments /6 E92?< =2F4@ %24925@ 2?5 9C:DE:?: 2D6==: 7@C E96:C 96=A7F= 4@>>6?ED @? E96 >2?FD4C:AE 2?5 F?52ON@ "@DP (65C@ 56 '=:G6:C2 7@C A6C>:EE:?8 7:6=5 DEF5:6D 2E E96 +2?E2 6?63C2 %F?:4:A2= *6D6CG6 (2F=@ ?C:BF6 2C 5@D@ (6:I@E@ H2D DFAA@CE65 3J 8C25F2E6 76==@HD9:AD 7C@> E96 @@C56?2ON@ 56 A6C76:O@2>6?E@ 56 (6DD@2= 56 &QG6=+FA6C:@C(+

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Journal of Insect Science: Vol. 9 | Article 54 %@EE2 (

FEE6C7=:6D @7 E96 -36C=M?5:2 *68:@? 6?EC2= C2K:= +A64:6D =:DE 2?5 3:@=@8:42= 4@>>6?ED Brazilian Journal of Biology (6R2 /29=36C8 & /6:?82CE?6C #@52?52C2>2:29 - &J=:? + C6:E2D .$ C@H6C .1

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