1025

Cuticular Hydrocarbon Variation of Castes and Sex in the 
Weaver Ant Camponotus textor (Hymenoptera: Formicidae)

by

Maria Claudia G. Campos1, Maria Lucia G. Campos1, Izabel C. Turatti2                                        
& Fabio S. Nascimento1*

AbSTrACT

Cuticular hydrocarbons play important roles as chemical signatures of 
individuals, castes, sex and brood. They also can mediate the regulation of 
egg laying in ants, by informing directly or indirectly the reproductive status 
of queens. In this study we asked whether cuticular hydrocarbon profiles are 
correlated with castes and sex of Camponotus textor. Cuticular hydrocarbons 
were extracted from part of a mature colony (80 workers, 27 major workers, 
27 queens, 27 virgin queens and 27 males). results showed that cuticular hy-
drocarbons varied quantitatively and qualitatively among the groups and this 
variation was sufficiently strong to allow separation of castes and genders. We 
discuss the specificity of some compounds as possible regulatory compounds 
of worker tasks and reproduction in C. textor.

Key-words: cuticular hydrocarbons, Camponotus textor, castes and sex

INTroduCTIoN

The integrity of social insect colonies is possible due to the ability of 
individuals to recognize and to discriminate nestmates, non-nestmates and 
heterospecific individuals (Wilson 1971; Hölldobler & Michener 1980; 
breed & bennett 1987; Nunes et al. 2008). In a nestmate recognition system, 
individuals detect and perceive the odor present in the cuticle of another 
individual and compare it with an internal template, thus distinguishing 
nestmate cues from non-nestmate cues (Sturgis & Gordon 2012; Leonhardt 
et al. 2007). Several studies have shown that cuticular hydrocarbons are the 
main compounds playing roles as chemical signatures of individuals, castes, 

1departamento de biologia, Faculdade de Filosofia Ciências e Letras de ribeirão Preto
universidade de São Paulo, Av. bandeirantes 3900, 14040-901, ribeirão Preto, São Paulo, brazil
2 departamento de Física e Química, Faculdade de Ciências Farmacêuticas de ribeirão Preto, 
universidade de São Paulo
* author for correspondence: fsnascim@usp.br



1026  Sociobiolog y Vol. 59,  No. 3, 2012

brood and colonies (Vander Meer & Morel 1998; Tannure-Nascimento et 
al. 2009; reviewed in blomquist & bagneres 2010).

In ants, for example, workers that perform distinct tasks present variable 
hydrocarbon profiles (Wagner et al. 1998). Patrollers and foragers of Pogono-
myrmex barbatus yield a higher amount of more complex compounds than nest 
maintenance workers (Wagner et al. 2001). In this same species, Greene and 
Gordon (2003) showed that a worker can access the task status of a nestmate 
from its cuticular hydrocarbon profile. In addition, Kaib et al. (2000) showed 
that, irrespective to their age, Myrmicaria eumenoides workers performing 
nest tasks retain their hydrocarbon profiles unaltered. They stated that the 
variation of the cuticular blend is not only relatively correlated with age, but 
also strongly correlated with task allocation of each individual.

Moreover, cuticular hydrocarbons can mediate the reproduction and 
regulation of egg laying in ants (Peeters et al. 1999; Liebig et al. 2000) and the 
reproductive status is related to the cuticular hydrocarbons of queens, mated 
workers and gamergates (Monnin et al. 1998; Liebig et al. 2000; Cuvillier-Hot 
et al. 2001). reproductive queens have differences in their cuticular profiles 
when compared with infertile workers (Heinze et al. 2002). In Camponotus 
floridanus the presence and the fertility of queen may be indicated by presence 
of hydrocarbons directly from the queen or her eggs (Endler et al. 2004). 

Camponotus textor is a weaver ant that inhabits forests of Central and 
South America (Schremmer 1979a, 1979b). Their nests are built with larval 
silk and present a round shape with several galleries around the branches 
and dead leaves (Santos 2002; Longino 2006). behavioral observations and 
natural history of this species has previously been studied (cited as C. senex: 
Santos et al. 2005a, 2005b; Santos & del-Claro 2009).

In the present study we evaluated and compared the chemical profile of 
cuticles of C. textor workers, queens, virgin queens and males.

MATErIAL ANd METHodS
A mature colony (27 queens, approximately 24,600 workers and major 

workers, approximately 500 virgin queens and 500 males) of C. textor obtained 
from the university of São Paulo (uSP-ribeirão Preto, São Paulo, brazil), 
was collected and freezer-killed (-20°C). The cuticular hydrocarbons of 80 



1027 Campos, M.C.G et al. — Cuticular Hydrocarbon Profiles of Camponotus textor

workers, 27 major workers, 27 queens, 27 virgin queens and 27 males were 
extracted. 

Cuticular compounds were extracted from individuals in 0.5 ml hexane for 
2 min. After evaporating the solvent, the apolar extract was resuspended in 
160 µl of hexane. The samples were analyzed in a GC-MS (Shimadzu, model 
QP2010 plus) equipped with silica capillary column of 30m and helium as 
carrier gas at 1 ml/min. The oven temperature was initially set to 150°C, 
and ramped up 3°C/min until it reached 280°C. Analyses were performed 
in splitless mode. data were analyzed to characterize the mass spectrum. 
Equivalent chain lengths were determined using n-alkane standards (Sigma 
Chemical Co.) and quantification was based on the peak areas obtained from 
the chromatograms. 

The data were analyzed with GCMS solutions for Windows (Shimadzu 
Corporation) and the chemical compounds were identified based on their 
mass spectra by comparison with Wiley Library data and with a standard 
solution of different synthetic hydrocarbons.

Principal components analysis (PCA) was used to define the main com-
pound peaks to be compared. Compounds missing in the most individuals 
of an analyzed group as well as compounds contributing less than 5% to the 
first two factors, as indicated by the PCA, were excluded from the statistical 
analysis. Following this, a stepwise discriminant function analysis was used to 
observe if combinations of variables could be useful in the predicting group. 
In this method, variables are successively added to the model based on the 
higher F to enter values, adding no more variables when the F ratio is no longer 
significant. Wilks’ λ values were used to verify the individual contribution of 
each variable to the model. The Wilks’ λ statistic for the overall discrimination 
is computed as the ratio of the determinant of the within-groups variance/
covariance matrix to the determinant of the total variance/covariance ma-
trix. Mahalanobis distance (D) was used as parameter to evaluate chemical 
distance between groups.

To avoid errors in compositional sample data, the area under each peak 
was transformed according to the following formula: Z = ln[Ap/g(Ap)], 
where Ap is the area under the peak, g(Ap) is the geometric mean for each 
individual compound group and Z is the transformed peak area (Aitchison 



1028  Sociobiolog y Vol. 59,  No. 3, 2012

1986). The statistical analyses were performed using the software Statistica 
7.0 (Statsoft, Inc.).

rESuLTS

Analysis of the worker cuticular waxes identified 90 hydrocarbons (Table 
1). These compounds were classified as linear alkanes, monomethylalkanes, 
and dimethylalkanes. The groups presented many qualitative differences in 
their respective cuticular profiles (Table 1).

of ninety compounds found, eighty-nine compounds were present in 
queens. Apart from these, eight compounds were exclusively found in the 
queens and six compounds were only found in queens and virgin queens. 
Workers, major workers and males had no exclusive compounds, but major 
workers and queens had four compounds in common, and one compound 
was only found in workers and queens.

Queens, workers and major workers presented nine compounds and these 
compounds were not present in virgin queens and males. one compound was 
present only in major workers.

Some compounds were more representative in the relative proportion 
of cuticular hydrocarbon profile of studied groups: 17-, 15- and 13-Me C32 
(>10%), n-C

28
 and n-C

30
 (> 5%), 9-, 11- and 13-, 15-diMe C

32
 (>6%), and 

13-, 15-diMe C
29

 (>4%) (Table 2).
Stepwise discriminant analyses of these compounds significantly separated 

castes and sex (Table 2: Global Model: Wilks’ λ= 0.001; F = 52,668; p< 
0.001) and the most important hydrocarbons for this separation (F to enter 
>1) were 9-, 11-diMe C

32
, 4-Me C

30
, n-C

32
, 13-, 15-diMe C

30
, n-C

30
, n-C

28
 

and n-C
29

. Chemical distances (D) between castes and sex were extremely 
high (Fig. 1): queens vs workers: 102.57, queens vs majors: 72.92, queens 
vs virgin queens: 127.87, queens vs males 184.58, workers vs majors: 41.95, 
workers vs virgin queens: 54.16, workers vs males: 58.20, majors vs virgin 
queens: 98.91, majors vs males: 121.06 and virgin queens vs males: 44.85 
(all values were highly significant p<0.001). 

dISCuSSIoN

In social insects, physiological variation of individuals is well reflected in 
their cuticular hydrocarbons (reviewed in blomquist & bagneres 2010). In 



1029 Campos, M.C.G et al. — Cuticular Hydrocarbon Profiles of Camponotus textor

Table 1: Cuticular hydrocarbons present in the Camponotus textor workers (W), major workers (LW), 
queens (Q), virgin queens (VQ) and males (M). rT = retention time in minutes. different letters 
indicate significant differences (p<0.001).

CoMPouNd rT
W LW Q VQ M

Mean Sd Mean Sd Mean Sd Mean Sd Mean Sd

n-C
26

28.841 0.54 ±0.36a 0.28 ±0.17 a 0.16 ±0.06 b 0.03 ±0.03 b 0.10 ±0.09 b

12-Me C
26

10-Me C
26

29.669 - - 0.12 ±0.09 0.09 ±0.04 -

3-Me C
26

30.661 0.40 ±0.21 a 0.44 ±0.16 b 0.35 ±0.17 a 0.31 ±0.05 a 0.14 ±0.08 c

n-C
27

31.359 1.32 ±1.56 d 0.45 ±0.25 c 0.24 ±0.11ac 0.05 ±0.02 b 0.09 ±0.05 ab

13-Me C
27

11-Me C
27

9-Me C
27

32.124 0.15 ±0.15 0.18 ±0.05 0.15 ±0.06 - -

5-Me C
27

32.725 0.12 ±0.08 0.07 ±0.02 0.18 ±0.48 0.05 ±0.02 -

11-.13-diMe C
27

13-.15-diMe C
27

32.879 - 0.09 ±0.03 0.16 ±0.08 0.05 ±0.02 0.01 ±0.01

3-Me C
27

33.122 - 0.21 ±0.09 0.11 ±0.05 - -

n-C
28

33.828 9.63 ±4.28 d 4.60 ±2.68 a 2.75 ±1.28 ab 0.80 ±0.47 c 1.89 ±0.75bc

14-Me C
28

13-Me C
28

12-Me C
28

34.561 4.13 ±1.72ac 3.62 ±0.98 ab 2.46 ±1.20 c 4.26 ±0.39 b 3.16 ±0.91 c

7-Me C
28

34.745 0.43 ±0.53 b 1.33 ±0.34 a 0.28 ±0.13 b 1.47 ±0.17 a 1.02 ±0.22 a

6-Me C
28

34.950 - 0.21 ±0.07 0.27 ±0.30 - -

4-Me C
28

35.162 0.28 ±0.26 a 0.22 ±0.06 a 0.19 ±0.13 a 0.23 ±0.07 a 0.04 ±0.03 b

2-Me C
28

35.582 5.91 ±1.81 b 5.54 ±1.35 a 6.05 ±2.76 abc 4.15 ±0.86 c 8.79 ±1.75 d

3-Me C
28

35.660 - 0.15 ±0.06 0.06 ±0.09 0.09 ±0.03 -

n-C
29

36.175 3.24 ±1.68 d 1.16 ±0.84ac 1.78 ±0.70 c 0.22 ±0.15 b 0.46 ±0.10 ab

15-Me C
29

13-Me C
29

11-Me C
29

36.845 0.32 ±0.35 d 0.81 ±0.41bc 0.30 ±0.16 ad 0.92 ±0.15 c 0.54 ±0.23 ab

9-Me C
29

36.970 - 0.11 ±0.05 0.16 ±0.11 - -

11-.15-diMe C
29

37.454 - 0.09 ±0.12 0.43 ±0.85 - -

9-.13-diMe C
29

37.593 0.21 ±0.12 0.23 ±0.08 0.37 ±0.18 - -

3-Me C
29

37.824 0.13 ±0.11 a 0.17 ±0.07 b 0.38 ±0.62 b 0.08 ±0.03 a 0.07 ±0.03 a

n-C
30

38.491 7.22 ±3.11 b 3.91 ±2.40 a 6.56 ±2.73 b 1.00 ±0.68 a 2.77 ±0.95 a

13-. 15-diMe C
29

38.941 5.12 ±1.63 - 1.74 ±2.34 9.29 ±1.28 11.15 ±2.87

16-Me C
30

14-Me C
30

12-Me C
30

39.111 0.58 ±0.42 ab 7.71 ±1.89 d 1.00 ±0.86 a 0.30 ±0.09bc 0.19 ±0.09 c



1030  Sociobiolog y Vol. 59,  No. 3, 2012

10-Me C
30

39.205 1.11 ±0.57 d 0.26 ±0.10 c 1.61 ±0.78 a 1.96 ±0.25 b 1.89 ±0.40 ab

? 39.321 0.76 ±0.63 1.86 ±0.42 1.11 ±0.39 - -

17-Me C
30

15-Me C
30

13-Me C
30

39.556 1.24 ±0.52 - 0.82 ±0.84 1.24 ±0.13 0.21 ±0.40

11-.15-diMe C
30

39.683 1.17 ±0.56 ab 0.79 ±0.30 c 0.65 ±0.37bd 0.46 ±0.08acd 1.36 ±1.21 d

9-. 17-di Me C
30

39.756 1.29 ±0.69 a 0.65 ±0.48 ab 0.46 ±0.28 b 2.72 ±1.42a 8.86 ±2.84 c

13-.15 diMe C
30

39.939 2.61 ±1.53 a 1.46 ±0.53 a 0.45 ±0.33 b 1.71 ±0.80 a 0.65 ±0.38 b

4-Me C
30

40.151 4.04 ±1.23 b 3.92 ±1.77 b 7.70 ±3.24 c 2.05 ±0.75 a 1.04 ±0.19 a

n-C
31

40.680 1.41 ±0.59 d 0.71 ±0.39 ab 3.82 ±3.26 c 0.73 ±0.28 b 0.47 ±0.13 a

14-Me C
31

15-Me C
31

41.279 1.19 ±0.45 b 2.15 ±0.68 a 1.17 ±0.58 b 2.27 ±0.60 a 1.59 ±0.28 c

9-Me C
31

41.435 - - 0.14 ±0.07 - -

5-Me C
31

41.579 - - 0.45 ±1.00 - -

11-.15-diMe C
31

9-.13-diMe C
31

41.898 0.52 ±0.21 b 0.96 ±0.30 a 1.50 ±0.90 a 0.43 ±0.19 b 0.25 ±0.13 c

7-. 15-diMe C
31 

7-. 17-diMe C
31

41.998 - - 0.59 ±0.41 - -

5-. 13-diMe C
31.

5-. 15-diMe C
31

5-. 17-diMe C
31

42.282 - 0.36 ±0.14 0.52 ±0.24 0.58 ±0.20 -

3-Me C
31

42.465 - - 0.39 ±0.41 0.12 ±0.05 -

n-C
32

42.843 1.47 ±0.79 a 0.74 ±0.45 a 5.36 ±2.06 c 0.16 ±0.13 b 0.33 ±0.11 b

13-Me C
32

15-Me C
32

17-Me C
32

43.415 13.76 ±3.67 d 12.73 ±2.99 a 7.14 ±3.72 b 15.21 ±1.02 c 15.12 ±2.48 a

9-Me C
32

43.533 1.95 ±0.50 a 0.37 ±0.33 c 2.84 ±1.10 b 2.04 ±0.19 b 2.24 ±0.74 ab

7-Me C
32

43.645 - 2.41 ±0.56 2.02 ±0.71 - -

10-. 14-diMe C
32

5- Me C
32

43.859 5.00 ±1.55 c 2.00 ±0.66 a 4.46 ±2.61bc 2.87 ±0.32 ab 4.53 ±2.35bc

12- 16-diMe C
32

43.957 1.12 ±0.60 c 1.98 ±0.60 a 2.00 ±2.24bc 1.27 ±0.18 ab 3.17 ±1.99 a

9-. 11-diMe C
32

13-.15-diMe C
32

44.055 4.15 ±1.12 c 0.95 ±0.29 b 1.54 ±1.11 b 9.45 ±0.99 a 7.27 ±1.42 a

6-. 10-diMe C
32

6-. 14-diMe C
32

6-. 16-diMe C
32

44.222 2.52 ±1.40 7.66 ±2.08 1.50 ±0.79 - -

Table 1 (continued): Cuticular hydrocarbons present in the Camponotus textor workers (W), major 
workers (LW), queens (Q), virgin queens (VQ) and males (M). rT = retention time in minutes. dif-
ferent letters indicate significant differences (p<0.001).

CoMPouNd rT W LW Q VQ M

Mean Sd Mean Sd Mean Sd Mean Sd Mean Sd



1031 Campos, M.C.G et al. — Cuticular Hydrocarbon Profiles of Camponotus textor

n- C
33

44.519 2.28 ±0.61 c 1.30 ±0.34 ab 10.91 ±4.25 d 1.62 ±0.23bc 0.91 ±0.28 a

13-Me C
33.

15-Me C
33

17-Me C
33

45.015 - - 1.23 ±0.54 1.54 ±0.22 -

9-Me C
33

45.153 0.06 ±0.10 - 0.78 ±0.49 0.78 ±0.19 -

7-Me C
33

45.684 0.64 ±0.51 0.31 ±0.12 0.93 ±0.48 - -

9-. 11-di Me C
33

9-.13-diMe C
33

45.759 - - 0.59 ±0.38 - -

6-Me C
33

46.070 0.10 ±0.24 - 0.29 ±0.19 3.93 ±0.76 -

5-Me C
33

46.300 0.21 ±0.26 1.22 ±0.41 0.53 ±0.74 - -

3-Me C
33

46.426 - - 0.93 ±1.62 - -

n-C
34

47.648 - - 0.96 ±0.90 3.70 ±0.42 -

13-Me C
34

48.531 2.41 ±2.23 ad 3.04 ±0.97 d 2.62 ±2.36 b 3.97 ±0.48 abc 2.86 ±0.55 c

11-Me C
34

48.620 1.62 ±1.55 c 2.51 ±0.74 b 1.09 ±1.13 c 3.53 ±0.73 a 3.11 ±0.93 ab

5-Me C
34

49.174 1.76 ±0.64 - 0.89 ±0.70 2.95 ±0.54 1.95 ±0.43

7-. 11-diMe C
34

49.340 - 1.76 ±0.67 - - -

9-. 11-diMe C
34

49.413 2.00 ±0.74 b 0.84 ±0.27 a 1.28 ±0.65 a 1.54 ±0.46 b 1.11 ±0.34 a

5-. 7-diMe C
34

49.635 - 1.19 ±0.42 1.12 ±0.57 - -

n-C
35

49.971 0.12 ±0.17 - 3.41 ±1.68 - -

8-Me C
35

51.046 - - 1.29 ±0.76 - -

Table 1 (continued): Cuticular hydrocarbons present in the Camponotus textor workers (W), major 
workers (LW), queens (Q), virgin queens (VQ) and males (M). rT = retention time in minutes. dif-
ferent letters indicate significant differences (p<0.001).

CoMPouNd rT W LW Q VQ M

Mean Sd Mean Sd Mean Sd Mean Sd Mean Sd

Table 2: Classification matrix for group comparison of predicted functional group using discriminant 
analysis.

% Correct W LW Q VQ M

W 98.76 80 1 0 0 0
LW 100 0 27 0 0 0
Q 100 0 0 27 0 0
VQ 88.88 0 0 0 24 3
M 100 0 0 0 0 27
Total 97.88 80 28 27 24 30



1032  Sociobiolog y Vol. 59,  No. 3, 2012

this study, castes and sex had qualitative differences in their respective cuticular 
profiles, mainly related to monomethylalkanes, and dimethylalkanes.

The compounds that have distinct and variable isomers (double bonds 
or methyl groups) are those associated with colonial recognition. on the 
other hand, n-alkanes are considered important for physical protection of 
the individual (Howard & blomquist 2005).

In C. textor, some compounds were only found in queens and virgin queens, 
which suggest that these hydrocarbons are characteristics of reproductive 
females. In Camponotus floridanus, the prime distinction of cuticular hydro-
carbon profiles is observed between queens and workers, in which eighteen 
compounds are only found in fertile queens (Endler et al. 2004, 2006). 
In Harpegnathos saltator the hydrocarbon profiles were similar between 
gamergates and queens, but they were different between workers and young 
queens (Peeters & Hölldobler 1995), and in Pachycondyla inversa many 
compounds were different in profiles of queens and workers (Heinze et al. 
2002). In the polyg ynous ant Linepithema humile the cuticular hydrocarbon 
profiles of queens, young queens and workers were different (de biseau et al. 
2004). dietemann et al. (2003) showed that Myrmecia gulosa had differences 
between queen and worker hydrocarbon profiles. dahbi & Lenoir (1998) 
showed the similarity of chemical compound profiles in queens and their 

Fig. 1: discriminant analysis of cuticular hydrocarbons present in the castes. W (workers), LW (major 
workers), Q (queens), VQ (Virgin queens) and M (males).



1033 Campos, M.C.G et al. — Cuticular Hydrocarbon Profiles of Camponotus textor

respective daughter-workers, indicating a significant correlation between 
cuticular profiles and fertility.

In C. textor, no alkenes were present in the cuticular profiles. The same 
results were obtained for C. floridanus (Endler et al. 2004) and C. fellah (La-
lzar et al. 2010).This pattern is different from that found in Pogonomyrmex 
barbatus (Wagner et al. 1998), Harpegnathos saltator (Liebig et al. 2000) and 
other 37 ant species (van Wilgenburg et al. 2011).

discriminant analysis of compounds of all studied groups showed that 
of  90 total compounds, 42 were common to all castes (queens and workers) 
and genders (females and males). 4-Me C

30
, n-C

32
 and n-C

33
 were found 

in larger quantities in queens than in other groups, major workers had the 
compounds 16-,14- and 12-Me C

30 
in more representative quantities than any 

other group. Males had the compound 9-, 17-diMe C
30 

in larger quantities 
than other groups and virgin queens and males presented the compound 9-, 
11- and 13-, 15-diMe C

32 
in larger quantities than other groups. boulay et 

al. (2007) observed quantitative differences in the profile of Aphaenogaster 
senile, where reproductive queens had five compounds in larger quantities and 
workers had five other compounds in larger quantities. Moreover, Endler et 
al. (2006) showed that queens had a higher proportion of compounds com-
pared with workers. Also in Crematogaster smithii, workers, virgin queens, 
intermorphs and queens had qualitative differences in their compounds 
(oettler et al. 2008).

Colonial blend or Gestalt colony odor seems to be more important for 
individual recognition than a specific group of hydrocarbons. However, recent 
revision of cuticular hydrocarbon profiles in ants suggests that dimethylal-
kanes are good candidates for species and nestmate discrimination (reviewed 
in Martin & drijfhout 2009a). In C. textor, ten of ninety compounds were 
dimethylalkanes, and discriminant analysis showed that variation between 
groups was mostly yielded by these hydrocarbons. However, further behav-
ioral bioassays are needed to understand the role of these compounds within 
the colonies.

In summary, our study demonstrates that cuticular hydrocarbons varied 
sufficiently among castes and sex to separate individuals according to their 
functional roles. by using this variation, individuals may discriminate among 
workers, major workers, fertile queens, virgin queens and males like in other 



1034  Sociobiolog y Vol. 59,  No. 3, 2012

ant species. Variation in cuticular hydrocarbons might therefore play an im-
portant role in the organization of colonies of polyg ynous ant species.

ACKNoWLEdGMENTS

The authors thank Cnpq for a graduate research grant to M.C.G.C. and 
Fapesp (Proc. 10/10027-5 to F.S.N.).

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