Morphological study of the Stridulatory Organ in two ... · Ann. soc. entomol. Fr. (n.s.), 2006, 42 (1) : 99-105 99 ARTICLE Morphological study of the Stridulatory Organ in two species

Ann. soc. entomol. Fr. (n.s.), 2006, 42 (1) : 99-105

99

ARTICLE

Morphological study of the Stridulatory Organ in two species of Crematogaster genus: Crematogaster scutellaris (Olivier 1792) and Crematogaster auberti (Emery 1869) (Hymenoptera: Formicidae)

Abstract. The stridulatory organ of the Crematogaster scutellaris (Olivier 1792) workers is being described, comparing their pars stridens present in six nests of this species, with one nest of Crematogaster auberti Emery 1869 and with the bibliographical data regarding other neighbouring species at our disposal. Both species and some Crematogaster scutellaris nests have shown significant differences. We propose several hypotheses which could explain these differences.

Résumé. Etude morphologique de l’organe stridulatoire chez deux espèces du genre Crematogaster : Crematogaster scutellaris (Olivier 1792) et Crematogaster auberti (Emery 1869) (Hymenoptera : Formicidae). L’organe de stridulation des ouvrières de Crematogaster scutellaris (Olivier 1792) est décrit. On compare les pars stridens issus de 6 nids de cette espèce avec ceux d’un nid de Crematogaster auberti Emery 1869 et avec les données de la littérature sur les espèces voisines disponibles. Les deux espèces et certains nids de Crematogaster scutellaris ont montré des différences significatives. Nous proposons plusieurs hypothèses pour expliquer ces différences.Keywords: Acoustic communication, stridulation, stridulatory organ, Crematogaster scutellaris, Crematogaster auberti, Iberian Peninsula.

Eduardo Ruiz, M. Hoz Martínez, M. Dolores Martínez & J. María Hernández Dpto. de Zoología y Antropología Física, Facultad de Biología, Universidad Complutense de Madrid, 28040-Madrid, Spain

E-mail: [email protected]é le 27 octobre 2005

Ant social organization is based on very effective systems of communication (Grasso et al. 1998).

Like most social insects, ants spend most of their time in the darkness of their nests, for this reason, vision has a limited use in communication. Pheromones and sounds are mainly used to exchange information (Kirchner 1997).

Acoustic communication plays a very important role in several behaviours such as alarm, recruitment, mating ending and modulation of other signals (Hölldobler & Wilson 1990; Kirchner 1997).

According to Markl (1973) acoustic communication can be accomplished by drumming or stridulating. The most extended method in ants that nest in the ground is stridulation, as drumming is for arboreal ants. Crematogaster auberti Emery 1869, that inhabits in the ground and Crematogaster scutellaris (Olivier 1792), that lives on trees, where chosen in order to establish differences between their stridulatory organs.

Within the subfamily Myrmicinae, the genus Crematogaster (Lund 1831) is distributed throughout the world and includes 427 species (Bolton 1995); in the Iberian Peninsula only four of them are present:

C. scutellaris (Olivier 1792), C. auberti Emery 1869, C. laestrygon Emery 1869 and C. sordidula (Nylander 1849).

Crematogaster scutellaris has a wide allocation in the Iberian Peninsula and a clearly Mediterranean distribution. It nests regularly in different trees (Soulié 1956); in the Iberian Peninsula it has preference for Quercus ilex and Quercus suber, although they can use other trees or big bushes (Casevitz- Weulersse 1972; Martínez et al. 1996). Crematogaster auberti generally nests in the ground (Soulié 1956) and have nocturnal habits (Bernard, 1968).

The establishment of the colonies presents slight differences between both species: in C. scutellaris the foundation is independent, whereas in C. auberti the regular method is fission (Soulié 1962a; Casevitz-Weulersse 1991).

Crematogaster scutellaris diet is mainly based on sweet substances and C. auberti is omnivorous. C. scutellaris forms foraging columns while C. auberti randoms alone or in small groups (Soulié 1962b). Temperature may influence the collecting cycle; nevertheless other climatic elements such as light intensity and relative humidity do not seem to be very important factors (Villagrán et al. 1992, 1998a, 1998b).

Crematogaster is a unique genus due to the origin of its track pheromones, which are not specific of the

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E. Ruiz, M.H. Martínez, M.D. Martínez & J. M. Hernández

Figure 1Location of stridulatory organ.

species included in this paper. Espadaler & Martí (1994) explain that the reason for this is that the C. auberti and the C. scutellaris inhabit different ecological niches.

Crematogaster ants are aggressive. C. scutellaris is the most hostile of the species studied in this paper (Soulié 1960). Crematogaster ants are unique among Myrmicinae in the defence of their nests because they use the secretion of Dufour gland as a defensive contact secretion (Daloze et al. 1987; Pasteels et al. 1989).

Stridulation sounds have been detected in four subfamilies of ants: Ponerinae, Pseudomyrmecinae, Mymicinae and Nothomyrmecinae.

Two structures are involved in the stridulatory organ: the plectrum or scraper, located in the back edge of the third abdominal tergite and the pars stridens or file, placed in the fourth abdominal segment (Grasso et al. 1998; Hernandez et al. 2002).

Sound occurs when gaster moves up and down; causing rubbing of the scraper with the file (Spangler 1967).

Some descriptive studies of the stridulatory organ for ant species have been completed: Grasso et al. (1998) for Messor minor (André,1883) and Messor wasmanni Krausse,1910; Schilliger & Baroni Urbani (1985) for Messor capitatus (Latreille 1798) and Messor structor (Latreille 1798); Hernández et al. (2002) for Messor barbarus (L. 1767); Le Roux (1976) for Myrmica laevinodis Nylander 1846; Hickling & Brown (2000) for Solenopsis richteri Forel 1909; Kermarrec et al. (1976) for Acromyrmex octospinosus (Reich 1793); and Álvarez et al. (in press) for Aphaenogaster senilis Mayr 1853.

However, no detailed studies have been undertaken with regards to the Crematogaster genus, with the exception of Raignier in 1933.

All the reasons mentioned above have impulsed us to study the Crematogaster scutellaris and the Crematogaster auberti stridulatory organ.

Material and MethodsWorker ants were sampled during the months of September and October 2001.Crematogaster scutellaris specimens were collected at “Casa de campo” in Madrid, UTM: 30TVK 364 749, altitude: 645 m, nests were located in the trunks of oak trees. Six different nests were sampled, collecting 50 specimens in each. 50 workers of Crematogaster auberti were collected from a single nest, located in the ground under stone, at Majadahonda (Madrid), UTM: 30TVK 242 744, altitude: 712 m. For the study of the stridulatory organ, the gaster of each specimen was detached and fixed on the same cardboard to which the ant had been mounted previously. A mould of the first gaster tergite was obtained by means of covering it with transparent nail lacquer; it was dried for 24 hours before proceeding to separate the obtained layer. This was mounted on microscope slides, fixing them with a cover slide by means of two strips of sticky paper.Measurements of the stridulatory organ were obtained from the

Stridulatory Organ in Crematogaster

101

digitalized images of preparations. We used a Presentco CCD 3630V camera with microscope adapter and connected to a digitized card 3400 ASUS AGP TNT for PC.The studied specimens and preparations obtained are deposited in the collection of the “Department of Zoology” at “Complutense University of Madrid” (UCME Collection).Three variables were measured from each stridulatory organ: Maximum width, maximum length, and distance between ridges. This last variable was taken considering it as the sum of the width of a ridge and its corresponding inter-ridge. From each specimen twenty measurements were performed and the mean value was computed. The relation between Maximum length/Maximum width was also calculated for each specimen.In order to compare the size of the stridulatory organ with the size of the ant, the width of the head was also considered. According to Elmes (1976) it represents a good indicator of the full-size of the specimen. The measurements were made following Dubois’ criterion (1998). A binocular microscope with a micrometric eyeglass was used to measure the width of the head, right behind the eyes.Inter and intra-specific relationships between the nests were calculated using a one way ANOVA test, and a Tamhane T2 test of multiple comparisons.

Results and Discussion The two studied species present a typical stridulatory

organ for Formicidae, conformed by a scraper or plectrum in the third abdominal tergite and a file or pars stridens placed on the medial dorsal region of fourth abdominal pretergite (Fig. 1).

General pattern is similar in all the observed specimens. Plectrum is shaped by a thickening of the back edge on the third abdominal tergite. It has not any other modified structure (Fig. 1).

Pars stridens is an elliptical area that fills the medial region of the fourth abdominal pretergite. It reaches, almost, the anterior or posterior edges of the former. In this area, tegumentary sculpture adopts the structure of fine parallel ribs, showing a uniform striated aspect (Fig. 1).

The stridulatory organ is similar to the one present in other Myrmicinae species, both in its shape and in its structure and position, although remarkable differences may appear concerning the number of ridges of pars stridens, (Le Roux 1976; Schilliger & Baroni Urbani 1985; Hernández et al. 2002).

The shape and general structure is similar for all the nests under study and differences have only been found in the full size of pars stridens and in the distance

Table 1. One-way ANOVA results for pars stridens and head variables.

1-way ANOVA (BETWEEN SPECIES)Sum of squares df Mean

square F Sig.

MAXIMUM

WIDTH

Inter-groups 79.78 1 79.78 0.53 0.47

Intra-groups 25134.69 167 150.51

Total 25214.47 168

LENGTH

Inter-groups 53.00 1 53.00 0.49 0.48

Intra-groups 17968.33 167 107.59

Total 18021.34 168

SHAPE

Inter-groups 1.43E-03 1 1.43E-03 0.08 0.77

Intra-groups 2.82 165 1.71E-02

Total 2.82 166

RIDGES

Inter-groups 9.33E-02 1 9.33E-02 13.05 0


Total 1.29 168

HEAD

WIDTH

Inter-groups 0 1 0 0.01 0.91

Intra-groups 3.69 167 0.02

Total 3.69 168

HEAD

LENGTH

Inter-groups 0.04 1 0.04 2.05 0.15


Total 3.16 168

WIDTH/

LENGTH

Inter-groups 56.28 1 56.28 2.39 0.12

Intra-groups 3926.07 167 23.51

Total 3982.35 168

1-way ANOVA (BETWEEN NETS) SPECIES 1Sum of squares df Mean

square F Sig.

MAXIMUM

WIDTH

Inter-groups 5976.24 5 1195.25 10.84 0

Intra-groups 12566.71 114 110.23

Total 18542.95 119

LENGTH

Inter-groups 4330.98 5 866.20 11.23 0

Intra-groups 8792.46 114 77.13

Total 13123.44 119

SHAPE

Inter-groups 0.16 5 3.14E-02 1.70 0.14


Total 2.22 117

RIDGES

Inter-groups 8.11E-02 5 1.62E-02 2.54 0.03


Total 0.81 119

HEAD

WIDTH

Inter-groups 1.048 5 0.21 16.66 0


Total 2.48 119

HEAD

LENGTH

Inter-groups 0.47 5 0.09 6.98 0


Total 1.99 119

WIDTH/

LENGTH

Inter-groups 533.97 5 106.79 8.01 0

Intra-groups 1519.32 114 13.33

Total 2053.28 119

Significant difference between means at level 0.05

102


between ridges, although the two species have different nesting behaviours (edaphic and arboreal),their stridulatory structures are similar.

The first ANOVA results (Table 1) show significant differences only in one variable among the two species under study: the distance between ridges (p<0.05), but significant variations in the size of pars stridens were not detected.

Cephalic width does not confirm significant inter-species differences. This indicates a similar size on both groups of workers.

As differences in the size of pars stridens are not present, the variation in the distance between ridges may determine a different number of ridges, which in turn could mean certain specificity in the sound produced.

A second ANOVA for the six C. scutellaris nests shows significant differences (p<0.01) in the maximum

length and width, whilst the index relating both variables does not show significant differences (Table 2). This implies that, while the size seems to be significantly variable between nests, the general proportions of the stridulatory organ remain the same.

The Tamhane’s multiple comparisons test places the length and width differences in nests 4 and 6. Table 3 shows a gradation in shape and length in workers from different nests, being those from nest 4 the smallest and those from nest 6 the largest. This explains the reason why these nests show Tamhane test significant differences when compared to other nests. Several hypotheses that could be verified later, may show that probably the smaller size of the stridulatory organ of nest 4 workers, is due to the fact that the individuals collected conformed the first generation of the colony, which is usually smaller in size (Gray 1971; Passera 1984).

Table 2. Tamhane T2 test results for pars stridens variables.

DISTANCE BETWEEN RIDGES

(I) Nest

(J)Nest

DifferenceMean (I–J)

Standarderror Significa.

1 2 2.53E-02 2.52E-02 0.99

3 5.39E-02 2.52E-02 0.50

4 3.73E-02 2.52E-02 0.64

5 -9.90E-03 2.52E-02 1

6 -1.80E-02 2.52E-02 1

2 1 -2.53E-02 2.52E-02 0.99

3 2.86E-02 2.52E-02 1

4 1.20E-02 2.52E-02 1

5 -3.52E-02 2.52E-02 0.96

6 -4.34E-02 2.52E-02 0.82

3 1 -5.39E-02 2.52E-02 0.50

2 -2.86E-02 2.52E-02 0.99

4 -1.66E-02 2.52E-02 1

5 -6.38E-02 2.52E-02 0.40

6 -7.19E-02 2.52E-02 0.22

4 1 -3.73E-02 2.52E-02 0.64

2 -1.20E-02 2.52E-02 1

3 1.66E-02 2.52E-02 1

5 -4.72E-02 2.52E-02 0.55

6 -5.54E-02 2.52E-02 0.28

5 1 9.90E-03 2.52E-02 1

2 3.52E-02 2.52E-02 0.96

3 6.38E-02 2.52E-02 0.40

4 4.72E-02 2.52E-02 0.55

6 -8.13E-03 2.52E-02 1

6 1 1.80E-02 2.52E-02 1

2 4.33E-02 2.52E-02 0.82

3 7.19E-02 2.52E-02 0.22

4 5.54E-02 2.52E-02 0.29

5 8.13E-03 2.52E-02 1

PARS STRIDENS MAXIMUN WIDTH

(I) Nest

(J)Nest



1 2 -0.69 3.32 1

3 -9.32 3.32 0.19

4 10.59 3.32 0.05

5 -0.51 3.32 1

6 -10.88 3.32 0.06

2 1 0.69 3.32 1

3 -8.63 3.32 0.29

4 11.28 3.32 0.03

6 0.18 3.32 1

7 -10.19 3.32 0.09

3 1 9.32 3.32 0.19

2 8.63 3.32 0.29

4 19.91 3.32 0

6 8.81 3.32 0.01

7 -1.56 3.32 1

4 1 -10.59 3.32 0.05

2 -11.28 3.32 0.03

3 -19.91 3.32 0

6 -11.10 3.32 0.004

7 -21.47 3.32 0

5 1 0.51 3.32 1

2 -0.18 3.32 1

3 -8.81 3.32 0.10

4 11.10 3.32 0.004

7 -10.37 3.32 0.02

6 1 10.88 3.32 0.06

2 10.19 3.32 0.09

3 1.56 3.32 1

4 21.47 3.32 0

6 10.37 3.32 0.02


103

PARS STRIDENS LENGTH

(I) Nest

(J)Nest



1 2 2.60 2.78 0.10

3 -2.25 2.78 1

4 9.13 2.78 0.06

5 2.60 2.78 1

6 -10.72 2.78 0.01

2 1 -2.60 2.78 1

3 -4.85 2.78 0.59

4 6.52 2.78 0.30

5 7.5E–04 2.78 1

6 -13.32 2.78 0

3 1 2.25 2.78 1

2 4.85 2.78 0.60

4 11.38 2.78 0.005

5 4.85 2.78 0.63

6 -8.47 2.78 0.08

4 1 -9.13 2.78 0.06

2 -6.52 2.78 0.292

3 -11.38 2.78 0.005

5 -6.52 2.78 0.32

6 -19.85 2.78 0

5 1 -2.60 2.78 1

2 -7.5E–04 2.78 1

3 -4.85 2.78 0.63

4 6.52 2.78 0.32

6 -13.32 2.78 0

6 1 10.72 2.78 0.01

2 13.32 2.78 0

3 8.47 2.78 0.08

4 19.85 2.78 0

5 13.32 2.78 0

PARS STRIDENS - SHAPE (Length/Width)

(I) Nest

(J)Nest



1 2 3.38E-02 4.30E-02 1

3 9.60E-02 4.35E-02 0.26

4 -2.2197E-02 4.30E-02 1

5 3.84E-02 4.35E-02 1

6 2.34E-02 4.30E-02 1

2 1 -3.38E-02 4.30E-02 1

3 6.23E-02 4.35E-02 0.86

4 -5.60E-02 4.30E-02 0.97

5 4.62E-03 4.35E-02 1

6 -1.04E-02 4.30E-02 1

3 1 -9.60E-02 4.35E-02 0.26

2 -6.23E-02 4.35E-02 0.86

4 -0.12 4.35E-02 0.02

5 -5.76E-02 4.40E-02 0.95

6 -7.26E-02 4.35E-02 0.39

4 1 2.22E-02 4.30E-02 1

2 5.60E-02 4.30E-02 0.97

3 0.12 4.35E-02 0.02

5 6.06E-02 4.35E-02 0.97

6 4.56E-02 4.30E-02 0.98

5 1 -3.84E-02 4.35E-02 1

2 -4.62E-03 4.35E-02 1

3 5.76E-02 4.40E-02 0.95

4 -6.06E-02 4.35E-02 0.97

6 -1.50E-02 4.35E-02 1

6 1 -2.34E-02 4.30E-02 1

2 1.04E-02 4.30E-02 1

3 7.26E-02 4.35E-02 0.39

4 -4.56E-02 4.30E-02 0.98

5 1.50E-02 4.35E-02 1

The results of the cephalic size analysis are similar to the ones obtained for the measurements of the pars stridens. Significant differences have been observed in nests 4 and 6. When comparing them with the rest of the nests, workers of nest 4 are smaller in size and workers of nest 6 are bigger.

For that reason, the differences in size of the pars stridens in the Crematogaster scutellaris could be associated with the general size of ant. This could be explained by several variables, like the worker’s age for each studied nest or the food availability.

The ANOVA for C. scutellaris nests also shows a difference in distance between ridges (p>0.01). This difference is not explained by the Tamhane’s multiple comparisons test, as the two by two associations do not show a significant difference.

Although the presence of a developed stridulatory organ does not constitute evidence in itself, it could

be a sign that sound communication in these species exists, even at short range

Finally, the stridulatory organ structure is similar to the Messor species ants (Hernández et al. 2002) and other groups of insects with functional stridulation, like some Coleoptera families (Jing-Qiu 1991; Hernández et al. 1997). In beetles, structures like the ones described above, appear in other corporal segments. This convergence with other species that also have a well known sound communication is another sign of the stridulatory organ functionality in Crematogaster. More studies in the analysis of emitted acoustic signals may confirm all these aspects.

104


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105

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Morphological study of the Stridulatory Organ in two ... · Ann. soc. entomol. Fr. (n.s.), 2006, 42 (1) : 99-105 99 ARTICLE Morphological study of the Stridulatory Organ in two species