Foraging strategies of two dominant native ant species associated with Toxoptera aurantii (Hemiptera, Aphididae) on cocoa trees in Centre Region of Cameroon

doi:10.21203/rs.3.rs-4707689/v1

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Foraging strategies of two dominant native ant species associated with Toxoptera aurantii (Hemiptera, Aphididae) on cocoa trees in Centre Region of Cameroon

https://doi.org/10.21203/rs.3.rs-4707689/v1

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Mutualism between ants and Hemiptera is well known on specific plant species. Cocoa is a tree-crop with a high economic value and harbour many ant-tending hemipteran species causing damage on shade trees. Herein, we examined the foraging strategies including frequency of ant activity, palpation and honeydew collection patterns used by local ants Myrmicaria opaciventris and Pheidole megacephala associated with Toxoptera aurantii. Field investigations were conducted from january to december 2022 in five locations in Centre Cameroon. At each location, one plot (0.25 ha) containing cocoa trees were monthly selected. Within each plot, only plants showing association between the target local ants and T. aurantii were investigated for our experiment. The overall number of local ants attending hempiterans was counted, and the palpation and honeydew collection duration were determinate following three time bands (8 a.m to 10 a.m; 12 a.m to 2 p.m; 4 p.m to 6 p.m). A total of 20,189 workers (13,015 individuals of M. opaciventris and 7,174 individuals of P. megacephala) was recorded associated with T. aurantii. Myrmicaria opaciventris foraged twice than P. megacephala at different time bands. In both ants, the highest number of workers was recorded between 8 a.m to 10 a.m and the lowest between 12 a.m to 2 p.m. All ants foraging on cocoa trees did not participate in palpation and honeydew collection activities. foraging strategies used by M. opaciventris appear more efficient and detrimental to cocoa growing. There is an urgent need to develop control strategies to reduce their populations and preserve cocoa-agrosystems.

Myrmicaria opaciventris

Pheidole megacephala

Toxoptera aurantii

trophobiotic

relationship

Foraging strategies

Cocoa (Theobroma cocoa L., 1753) is a cash crop of global importance. Cocoa trees grown under a dense canopy of various shade trees are a good model of agroforestry as they contribute more to the preservation of ecosystem integrity and diversity compared with more intensively managed cocoa systems (De Beenhouwer et al. 2013; Tscharntke et al. 2011). Further, cocoa’s global trade represented $8.6 billion in 2017 (Voora et al. 2019) with 68.4% of the world’s cacoa being produced in Central/West Africa nowadays (FAOSTAT 2020). In Cameroon, cocoa seedling and beans constitute an important source of incomes for many people (Ndoumbe et al. 2004). Although cocoa is considered the fastest expanding export-oriented crop across Central/West Africa (Ordway et al. 2017), cocoa productivity is decreasing in many producing countries due to problems like degraded soils, diseases and pests (Armengot et al. 2020; Blaser et al. 2018).

Cocoa pests such as the brown capsid Sahlbergella singularis Haglund, 1895 (Hemiptera: Miridae), is the major insect pest in the cocoa belt of Central/West Africa (Bagny Beilhe et al. 2018), and the mealybug (Hemiptera: Pseudococcidae spp.), responsible for spreading the Cocoa Swollen Shoot Virus Disease (Andres et al. 2018), are still among the most important factors limiting cocoa production (Wessel & Quist-Wessel, 2015). In some cases, these pests can cause annual crop losses of about 25–40% (Wessel and Quist-Wessel 2015). Beside S. singularis and other cocoa pests, Toxoptera aurantii is one of the most abundant aphid species involves in cocoa destruction as documented in some reports carried out in the centre region of Cameroon indeed, T. aurantii individuals have a high ecological adaptation and their abundance could enhance their harmfulness on cocoa tree. Sap-sucking insects (aphids) and ants have lived together on many ecosystems for a very long time and developed a trophobiotic relationship (Yede 2016; Ferreira et al. 2022).

Some hemipterans, especially those living in more or less large colonies, are often associated with one or several ant species (Steiner et al. 2004). Anyway, poor flying hemipterans such as aphids and coccids depend on their tending ants for dispersal. Ants might then play an important role in their dissemination and their spray in and within gardens (Delabie 2001). They can also amplify or reduce populations and protect them from natural enemies (Stadler and Dixon 2005; Oliver et al. 2007). In turn hemipterans provide considerable amounts of honeydew, a highly energetic food reward for various insect taxa, especially ants. Moreover, in the absence of ants, the honeydew could negatively affect the growth of hemipteran colonies as well as the fitness of the host-plants due to accumulation of honeydew that favour the development of sooty mold (Lehouck et al. 2004).

Many studies have showed that invasive ants, Linepithema humile and Wasmannia auropunctata are frequently associated with an increase in the abundance of Hemipterans which have a dramatic effect on plant growing (Helms and Vinson 2002; Holway et al. 2002; Abbott et al. 2007: Mbenoun et al. 2023). Numerous studies have investigated interaction between ants Tapinoma spp., Crematogaster spp., Tretamoruim spp., Paratrechina spp. associated with hemipteran T. aurantii on lemon trees (Dartigues 1988; Rosen 1967). In cameroonian’s fauna, recent studies have demonstrated that Myrmicaria opaciventris and Pheidole megacephala are two dominant local ants associated with high population of hemipterans including T. auranttii on Cocoa trees (Mbenoun et al. 2023; Yede et al. 2023). To date, little is known about the strategies used by these ants to attend hemipterans species on host plants. The main objective of this study was to investigate the foraging strategies of these two local ants with T. aurantii in cocoa plantations with the broader aim to develop pest management strategies to reduce cocoa pests and their negative effects on cocoa growing.

Study sites

Field studies were carried out from january to december 2022 in five locations in Centre Region of Cameroon namely: Minkoameyos in Mfoundi division (3°51’13.4’’N ; 11°25’15.8’’E ; 746 m a.s.l); Mve I in Mefou et Afamba division (3°57’33.0’’N ; 11° 43’31.9’’E ; 751 m a.s.l); Nkolbiyé: in Mefou et Akono division (3°40’15.1’’N ; 11°28’44.9’’E ; 660 m a.s.l); Libel-lingoi in Nyong et Kellé division (3°54’12.6’’N ; 10°55’12.6’’E ; 406 m a.s.l.) and Aboé in Nyong et Mfoumou division (4°04’23.5’’N ; 12°25’03.0’’ E ; 590 m a.s.l.) (Fig. 1). These locations were chosen along rural-urban gradient based on the presence of cocoa farms, distance between households and cocoa farms, density of human population, and level of vegetation disturbance by anthropogenic activities. These different locations were situated within the Guinean and equatorial transitional climate type. Rainforest pattern is characterized by a bimodal with the short wet season from March to June, followed by a short dry season from July to the end of August, and then a major wet season from September to mid-November. The short, wet season is followed by a long dry season from mid-November to February. The average annual rainfall ranging from 1400 to 1600 mm/year (Sighomnou 2004). The mean annual relative humidity is 79.5%. The mean air temperature ranges from 19.2 to 28.6°C.

Frequency of local ant activities associated with Toxoptera aurantii

Preliminary surveys were carried out early morning between 6 a.m to 8 a.m to investigate the presence of ant species and hemipterans in different locations. At each location, we have selected one plot (0.25 ha) containing cocoa trees. Within each plot, only plants showing association between the target local ants and T. aurantii were selected for our experiment (Fig. 2). For field observations, we divided the day in three parts (Morning, afternoon and evening) after dividing, three time bands were chosen (8 a.m to 10 a.m; 12 a.m to 2 p.m; 4 p.m to 6 p.m). During each time band, M. opaciventris and P. megacephala individuals associated with T. aurantii were counted on the chupon of the cocoa trees.

Duration of palpation and honeydew collection

Ant species are considered associated with T. aurantii if it was found on the same chupon infested with T. aurantii. During each time band, the total number of ant species was recorded, after that the palpating and honeydew collection individuals were recorded. We used both ant species to record the duration of palpation and honeydew collection during the sampling. Some Abiotic parameters such as Temperature and relative humidity were recorded at each time band.

Data analysis

Descriptive statistics (abundances and means of ant species) were used, the Kruskal-Wallis H test and Mann Witney Test (U) for the comparison of means, the chi-square (χ2) test for the comparison of frequencies at 5%. Microsoft Office Excel 2013 and PAST 3.14 software were used for this purpose. The mean number of ant species interacting with T. aurantii counted during the different time bands was considered as optimal foraging for different ant species. In the same way, the palpating and honeydew collection individuals of each species observed during the different time bands were considered as the optimal foraging. The duration of palpation and honeydew collection of both species during the different time bands were considered as the optimal foraging. Different seasons have been use to show the correlation between ant species and abiotic factors

Overall number of foraging ants

A total of 20,189 workers (13,015 individuals of M. opaciventris (64.47%) and 7,174 individuals of P. megacephala (35.53%)) was recorded associated with T. aurantii in cocoa trees during the study period (Fig. 3). Individuals of M. opaciventris were higher than individuals of P. megacephala except in February and March during the survey. Two major peaks were recorded in both ant species during the year: October and December (M. opaciventris), February and December (P. megacephala). From January to April (long dry season), abundance was higher in P. megacephala than in M. opaciventris with a small peak (65.57 ± 23.49 workers) in February. Both species showed a decrease of individuals from April end to May (Short wet season). A number of individuals increase drastically in M. opaciventris from may (Short dry season) to August (Major wet season) while a slight increase in number was observed in P. megacephala during the same period. We found not significant correlation between M. opaciventris and P. megacephala individuals (U = 41; p = 0.07).

Daily foraging activity

Myrmicaria opaciventris ants foraged twice as much as P. megacephala in the morning, midday and afternoon. The maximum mean number of workers 107.25 ± 12.93 individuals was recorded in the morning between 8 a.m to 10 a.m and the minimum mean number of 91.52 ± 11.02 workers was collected between 12 a.m to 2 p.m in M. opaciventris. The same pattern was observed in P. megacephala within time bands (Table 1). There was not a significant difference of individuals of different time bands of both ant species; but abundance between both species differed significantly between 8 a.m to 10 a.m (U = 567: p = 0.001), 12 a.m to 2 p.m (U = 624; p = 0.004), and 4 p.m to 6 p.m (U = 612: p = 0.003) (Table 1). M. opaciventris is most associated in T. aurantii than P. megacephala in cocoa tree.

Table 1

Foraging patterns of *M. opaciventris* and *P. megacephala* recorded associated with *T. aurantii* in cocoa trees according to time bands. (Lowercase letters = comparison between columns).
	Time bands
Species	8 a.m- 10 a.m	12 a.m-2 p.m	4 p.m-6 p.m
Myrmicaria opaciventris	107.25 ± 12.93^a	91.52 ± 11.02 ^a		97.02 ± 12.13 ^a
Pheidole megacephala	55.86 ±11.26^b	52.31 ± 10.45 ^b	55.31 ± 11.05 ^b
Mann Witney Test (U) p-value	567 0.001***	624 0.004***	612 0.003***
^*** = more significant

Correlation between foraging activities and abiotic factors

Foraging individuals of M. opaciventris workers was not significantly correlated with temperature and relative humidity in short dry season according to time bands (Table 2). A positive correlation was found between M. opaciventris workers and temperature in short rainy season and long dry season (4 p.m to 6 p.m). The same pattern of positive correlation was found between M. opaciventris workers and relative humidity between 8a.m to 10 a.m (short rainy season), 12a.m to 2p.m (long rainy season and long dry season), and 4p.m to 6p.m (major and short wet seasons. In P. megacephala workers, we found not significant correlation neither with temperature nor relative humidity during the long dry season (Table 2). A negative correlation was found between P. megacephala workers and temperature in short wet season and short dry season in all the time bands. In contrary, a positive correlation was found between P. megacephala workers and relative humidity between 8a.m to 10 a.m ; 4p.m to 6p.m (in short wet season) and during all the time bands (short dry season).

Table 2

Correlation between dominant ants (*M. opaciventris* and *P. megacephala*), and temperature (T°) and relative humidity (RH) according to different wet and dry seasons (Legend: MWS = major wet season; LDS = long dry season; SWS = short wet season ; SDS = short dry season
Foraging ants	Season	T°			RH
Foraging ants	Season	8 a.m-10 a.m	12 a.m-2 p.m	4 p.m-6 p.m	8 a.m-10 a.m	12 a.m-2 p.m	4 p.m-6 p.m
M. opaciventris	MWS	-0.36	-0.93*	-0.14	0.01	0.78*	0.7*
	LDS	0.02	-0.64*	0.98*	-0.97*	0.97*	-0.1
	SWS	-0.69*	-0.73*	0.97*	0.81*	0.27	0.82*
	SDS	-0.37	-0.03	0.44	0.11	-0.29	-0.4
P. megacephala	MWS	-0.7*	-0.99*	-0.77*	-0.96*	0.95*	0.88*
	LDS	0.48	0.46	0.16	-0.21	-0.19	-0.17
	SWS	-0.84*	-0.51*	-0.72*	0.83*	0.18	0.7*
	SDS	-0.73*	-0.94*	-0.99*	0.88*	0.99*	0.99*
*₌ significant difference

Duration of foraging activities

The palpation duration of each dominant ant species is presented in the Table 3. According to the observation made, higher palpation activities of both species were recorded between 12 a.m to 2 p.m and lower between 4 p.m to 6 p.m out of a total of 65 individuals. Pheidole megacephala performed more palpation (3.29 ± 0.07 min) than M. opaciventris (2.99 ± 0.11 min) on T. aurantii before honey collection. We only notice a significant different between palpation time in P. megacephala according to time bands (Table 3). Controversy, the honeydew collection time was higher in M. opaciventris (3.57 ± 0.08 min) and lower in P. megacephala (3.49 ± 0.78 min) at all-time bands except between 4 p.m to 6 p.m. There is a significant different between honey collection time in both species according to time bands (Table 4).

Table 3

Duration of foraging activities (time in minutes) of *M. opaciventris* and *P. megacephala* recorded associated with *T. aurantii* in cocoa trees according to time bands. (Lowercase letters = comparison between lines).
		Time bands
Duration of foraging activities	Species	8 a.m-10 a.m	12 a.m-2 p.m	4 p.m -6 p.m	H	p-value
Palpation	Myrmicaria opaciventris	2.61 ± 0.26^a	2.99 ± 0.11^a	2.57 ± 0.15^a	H = 3.14	p = 0.2
Palpation	Pheidole megacephala	2.29 ± 0.18^a	3.29 ± 0.07^b	2.32 ± 0.12^a	H = 20.38	p = 0.0001
Honeydew collection	Myrmicaria opaciventris	2.63 ± 0.25^a	3.57 ± 0.08^b	2.76 ± 0.18^a	H = 11.84	p = 0.002
Honeydew collection	Pheidole megacephala	2.49 ± 0.22^a	3.49 ± 0.78^b	2.86 ± 0.20^a	H = 11.82	p = 0.003

Difference between number of ants per trees and foraging patterns

All ants foraging on cocoa trees did not participate in palpation and honeydew collection activities. Significant different was found between the mean number of both local ants and foraging patterns (p < 0.001) (Fig. 3A and B). In M. opaciventris, the mean number of individuals (13.73 ± 1.98) was higher between 8 a.m to 10 a.m and lower (11.70 ± 1.73) between 12 a.m to 2 p.m. There was a significant difference between palpation and honeydew collection at the first and second time bands (8 a.m-10 a.m and 12 a.m -2 p.m) whereas no significant difference was found between 4p.m and 6p.m (Fig. 4). In Pheidole megacephala, the same pattern of difference between palpation and honeydew collection was found at the second time bands (12a.m-2p.m) and no difference was observed at the first (8 a.m-10 a.m) and the third time bands (4p.m and 6p.m) (Fig. 5).

Palpation and honeydew collections between local ants

The overall number of local ants performing palpation and honeydew collection was compared according to the time bands (Table 5). Individuals of both local ants involved in palpation varied between 800 and 2740. There is a significant different in Palpation between Pheidole megacephala and M. opaciventris according to the time bands (p < 0.001). Myrmicaria opaciventris individuals performed more palpation than P. megacephala ones. Individuals of both local ants involved in honeydew collection varied between 300 and 1060. There is a significant different in honeydew collection between both local ants according to the time bands (p < 0.05). Myrmicaria opaciventris individuals collected more honeydew than P. megacephala.

Table 5

Comparison of foraging patterns between *M. opaciventris* and *P. megacephala* in association with *T. aurantii* according to the time bands
Times bands	Palpation				Honeydew collection
Times bands	M. opaciventris	P. megacephala	Test (X²)	p-value	M. opaciventris	P. megacephala	Test (X²)	p-value
8 a.m-10 a.m	1600 (66.67%)	800 (33.33%)	137.14	< 0.001**	380 (55.88%)	300 (44.12%)	4.72	0.02*
12 a.m-2 p.m	2740 (58.80%)	1920 (41.20%)	72.70	< 0.001**	1010 (58.80%)	800 (41.20%)	1.22	0.004**
4 p.m-6 p.m	1660 (62.88%)	980 (37.12%)	89.03	< 0.001**	1060 (77.94%)	300 (22.06%)	230.34	< 0.001**

The optimal foraging of ant species have varied according to the time band. Overall, the activity period of the different ant species was high during the 8 a.m–10a.m time band, and low during the 12 p.m–2 p.m time band. Our study show that the survival and foraging activity of both ant species are strongly influenced by abiotic conditions. A significant difference was observed between the ant numbers of the two species. The ability of ants to group together on the aphids estimated by the survival rate of these, depends on the accessibility of the insects, the distance between them and the ant colony, the number in the group visited by ants, their ability to group together, to produce honeydew, as well as the diversity of food sources available to ants (Way 1963). Indeed, the main benefit of the Ant-Homoptera association is the supply of nutrients, in particular sugars (honeydew), and proteins which are later consumed by ants (Way 1963) or harvested after their death (Buckley, 1987).

Several factors are at the origin of the Ant-Homopteran association; in particular the accessibility of the Homoptera, the distance between them and the ant colony, the density or the numerical importance of the group of Homoptera, their ability to group together and produce honeydew in a qualitative and quantitative way; and finally a protein deficiency in the food system of ants. (Way, 1963; Gullan 1997).

The highest activity of ants was located at the time band of 8a.m to 10a.m could be related to the period honeydew is highly produced by T. aurantii populations; the high abundance of workers of these different species of ants is related to intense sap-harvesting activity of this aphid. These result is similar with those of Fotso et al (2008) who obtained a peak of activity of the ant Anoplolepis tenella between 8 a.m. and 10 a.m. Indeed ants are very active when the temperature is low and the humidity is high; insect performed in warm or sunny days activity, the enhanced temperature positively influence the insect activity (Kasper et al. 2008). As with other ant species, daily and seasonal variations in foraging activity are linked to variations in temperature and humidity (Lévieux 1983; Torres 1984). In addition, foraging activity can be affected by many environmental factors such as predation, interspecific competition, density and turnover rate of the food resource, humidity and temperature (Torres 1984; Hölldobler and Wilson 1990; Cerdá et al. 1998; Holway 1998; Kaspari and Weiser 2000; Hahn and Wheeler 2002; Menke and Holway 2006).

Taking into account the different activities carried out by the two species of ants on T. aurantii; only palpation and honeydew sampling were observed. M. opaciventris and P. megacephala showed through palpation activity that they are more active during the 12a.m to 2 p.m. time band, and less active from 8 a.m. to 10 a.m. On the other hand, the activity of honeydew collection of these two species was greater during the time band from 4 p.m. to 6 p.m., and less from 12 a.m. to 2 p.m. A number of aphid species considered myrmecophilous interact with ants. For ants, the main advantage of this interaction is the source of food: either sugar (honeydew); or proteins if they are consumed by the latter (Way 1963; Buckley 1987). Palpation is the main activity carried out by ants to collect honeydew; this activity is almost systematic because it stimulates the aphids to produce more honeydew than usual in quality and quantity; the species concerned respond to the palpation of their abdominal extremity by ants by secreting honeydew which constitutes for the ants a food supplement rich in sugar (Way 1963; Hölldobler and Wilson, 1990). Our results are not in line with those of Fotso Kuate et al (2008) this could be explained by the fact that in their work, they used abundance to assess the period of activity; in the context of this study, palpation was used for this assessment.

The African big-headed ant, Pheidole megacephala performed more palpation duration than M. opaciventris on T. aurantii from 12 a.m to 2 p.m ; Our results are in line with those of Kingha et al in 2012 who obtained a peak activity of Xylocopa olivacea on Phaseolus vulgaris flowers between 10 a.m and 1p.m. The ecological success of a species depends on its ability to adjust the foraging strategies to maximize food intake with the least amount of energy possible (Stephens and Krebs 1971). Daily foraging adjustments are influenced mainly by limitations such as availability and space-time distribution of the food resource, as well as nutritional needs (Manly et al. 1993, Giraldeau and Caraco 2000). The investment in foraging of each ant colony depends on the ability and efficiency of each worker, on environmental changes, and on factors such as distance to the food source and demographic density (Gordon 1999; Silva and Noda 2000). The investment in foraging of each ant colony depends on the ability and efficiency of each worker, on environmental changes, and on factors such as distance to the food source and demographic density (Gordon 1999; Silva and Noda 2000). Therefore, a difference in the investment and efficiency of each colony is expected. For the social insects, efficient foraging depends on own actions as well as on those of other individuals of the colony. The social foraging efficiency of social insects such as ants is not easy to measure because of the difficulty in identifying the efficiency of each individual (Giraldeau and Caraco 2000). Different foraging strategies are used by ants based on environmental changes and food density (Fowler 1985, Herbers and Choiniere 1996). Foraging varies widely in the Ponerinae species, from solitary foraging to social foraging and from generalist to specialist predators (Peeters and Crewe 1987; Hölldobler and Wilson 1990; Leal and Oliveira 1995; Fowler 1997).

The activities of ants are the most important parameters to take into account when rating their activity. The foraging strategies of M. opaciventris and P. megacephala in association with T. aurantii was evaluated according to their foraging pattern. The period of activity of the different species of ants according to the abundance is from 8a.m to10a.m. The. The palpation activity showed a period of activity of ants from 12a.m to 2p.m while the honeydew collection activity took place during the time band of 4 p.m. to 6 p.m. Myrmicaria opaciventris forage more than P. megacephala. The data presented here are useful for managing populations of T. aurantii, using baits, which would be placed selectively during periods of high activity on these different ant species in order to reduce the impact of T. aurantii on cocoa.

Acknowledgements

We would like to thank Patrick Atagana and Alain Wandji for providing logistics during the study. We are grateful to Michel Dongmo, Christian Djuideu and Clinton Factheu for useful comments. We thank Paul Ntjam, Julienne Ngo Ntjam and Ebozoa for providing cocoa farms.

Conflict of interest statement

The authors declare no conflict of interest.

Abbott KL, Green PT (2007) Collapse of an ant–scale mutualism in a rainforest on Christmas Island. Oikos 116:1238–246. https://doi.org/10.1111/j2007-0030-1299-15629
Andres C, Blaser WJ, Dzahini-Obiatey HK, Ameyaw GA, Domfeh OK, Awiagah MA, Gattinger A, Schneider M, Offei SK, Six J (2018) Agroforestry systems can mitigate the severity of cocoa swollen shoot virus disease. Agric Ecosyst Environ 252:83-92. https://doi.org/10.1016/j2017-09-031
Armengot L, Ferrari L, Milz J, Velasquez F, Hohmann P, Schneider M (2020) Cacao agroforestry systems do not increase pest and disease incidence compared with monocultures under good cultural management practices. Crop Prot 130:105047. https://doi.org/10.1016/j2019-105047
Bagny BL, Babin R, Ten Hoopen GM (2018) Insect pests affecting cacao. In:Umaharan P (Ed) Achieving Sustainable Cultivation of Cocoa. Burleigh Dodds Science Publishing, Cambridge, pp 303–326
Blaser WJ, Oppong J, Hart SP, Landolt J, Yeboah E, Six J (2018) Climate-smart sustainable agriculture in low-to-intermediate shade agroforests. Nat Sustain 1:234-239. https://doi.org/10.1038/s41893-020-0594-6
Buckley RC (1987) Interactions involving plants, Homoptera, and ants. Annu Rev Ecol Evol Syst 8:111-135. https://doi.org/10.1146/a18-110187-000551
Dartigues D (1988) Influence de la fourmi Tapinoma simrothi Krausse sur les pucerons de l’oranger, Toxoptera aurantii Boyer, Aphis citricola V.D. Goot, et sur le puceron de la fève, Aphis fabae Scop. Ann Inst Nat Agroe El-harrach, Alger, 12:89-100.
Delabie JHC (2001). Trophobiosis between Formicidae and Hemiptera (Sternorrhyncha and Auchenorrhyncha): An Overview. Neotrop Entomol 30:501-516. https://doi.org/10.1590/s1519-566X2001000400001
Cerda X, Retana J, Manzaneda A (1998) The role of competition by dominants and temperature in foraging of subordinate species in Mediterranean ant communities. Oecologia 117:404-412.
De Beenhouwer M, Aerts R, Honnay O (2013). A global meta-analysis of the biodiversity and ecosystem service benefits of coffee and cacao agroforestry. Agric Ecosyst Environ 175:1-7. https://doi.org/10.1016/j2013-05-003
FAOSTAT (2020) Food and Agriculture Organization of the United Nations. https://www.fao.org/faostat/en/#data/QCL
Ferreira DF, Jarrett C, Wandji A C, Atagana JP, Hugo RB, Powell LL (2022) Birds and bats enhance yields in Afrotropical cacao agroforests only under high tree-level shade cover. Agric Ecosyst Environ 345:108325. https://doi.org/10.1016/j2022-108325
Fotso KA, Tindo M, Hanna R, Kenne M, Goergen G (2008) Foraging activity and diet of the ant, Anoplolepis tenella Santschi (Hymenoptera: Formicidae), in southern Cameroon. Afr Entomol 16:107-114.
Fowler HG (1985) Population foraging and territoriality in Dinoponera australis (Hymenoptera, Formicidae). Rev Bras Entomol 16:443-447.
Fowler HG (1997) Foraging, diet and community structure in an epigaeic ant (Hymenoptera: Formicidae) assemblage: The role of recruitment. Cien Cult 49:199-202.
Giraldeau LA, Caraco T (2000) Social foraging theory. Princeton, Princeton University Press
Gordon DM (1999) Ants at work: How an insect society is organized. The free press, New York
Gullan P J (1997) Relationships with ants. World Crop Pests 7:351-377. https://doi.org/10.1016/s1572-4379-80065-6
Hahn DA, Wheeler DE (2002) Seasonal foraging activity and bait preferences of ant on Barro Colorado Island, Panama. Biotropica. 34:348-356. https://doi.org/10.1111/j1744-7429-2002
Helms KR, Vinson SB (2002) Widespread association of the invasive ant Solenopsis invicta with an invasive mealybug. Ecology 83:2425-2438. https://doi.org/10.1890/0012-9658-083
Herbers JM, Choiniere E (1996) Foraging behavior and colony structure in ants. Anim Behav 51:141-153. https://doi.org/10.1006/a1996-0012
Hölldobler B, Wilson EO (1990) The Ants. Cambridge, Massachusetts
Holway DA (1998) Factors governing the rate of invasion:a natural experiment using Argentine ants. Oecologia 115:206-212
Holway D Lach L, Suarez AV, Tsutui ND, Case T (2002) The Causes and Consequences of Ant Invasions. Annurev Ecolsys 33:181-233. https://doi.org/10.1146/a33-10802-150444
Kaspari M, Weiser MD (2000) Ant activity along moisture gradients in a Neotropical forest. Biotropica 32:703-711. https://doi.org/10.1111/j1744-7429-2000-00518
Kasper M.L., Reeson AF, Mackay DA, Austin AD (2008) Environmental factors influencing daily foraging activity of Vespula germanica (Hymenoptera, Vespidae) in Mediterranean Australia. Insect Soc 55:288-296. https://doi.org/10.1007/s00040-008-1004-7
Kingha TBM, Tchuenguem FFN, Ngakou A, Brückner D (2012) Foraging and pollination activities of Xylocopa olivacea (Hymenoptera, Apidae) on Phaseolus vulgaris (Fabaceae) flowers at Dang (Ngaoundere-Cameroon). J Agric Ext Rural Dev 6:330-339.. https://doi.org/10.5897/j11-151
Leal IR, Oliveira PS (1995) Interactions between fungus-growing ants (Attine), fruits and seeds in cerrado vegetation in Southeast Brazil. Biotropica 30:170-178. https://doi.org/10.1111/j1744-7429-1998
Lehouck VS, Bonte DB, Dekoninck W, Maelfait, JPE (2004). Trophobiotic relationships berween ants (Hymenoptera: Formicidae) and Tettigometridae (Hemiptera: Fulgoromorpha) in the grey dunes of Belgium. Eur J Entomol 101:547-553.
Lévieux J (1983) Modes d’exploitation des ressources alimentaires épigées de savanes africaines par la fourmi Myrmicaria eumenoïdes Gerstaecker. Insectes Sociaux 30:165–176.
Manly B, McDonald L, Thomas D (1993) Resource selection by animals. Chapman, London
Mbenoun MPS, Yede, Mballa NPA, Mendoua E, Ebangue T (2023) Interaction and resource exploitations between invasive and native ants in cocoa and surronding farmlands in southern Cameroon rainforest. Acta Entomol. Zool 4:126-132. https://doi.org/10.33545/27080013-2023
Menke SB, Holway DA (2006) Abiotic factors control invasion by Argentine ants at the community scale. J Anim Ecol 75:368-376. https://doi.org/10.1111/j1365-2656-2006-01056
Ndoumbe-Nkeng M, Cilas C, Nyemb E, Nyasse S, Bieysse D, Flori A, Sache I (2004) Impact of Removing Diseased Pods on Cacao Black Pod Caused by Phytophthora megakarya and on Cacao Production in Cameroon. Crop Prot 23:415-424. https://doi.org/10.1016/j2003-09-010
Oliver TH, Mashanova A, Leather SR, Cook JM, Jasen VAA (2007) Ant semiochemicals limit apterous aphid dispersal. Proc R Soc 274:3127–3131. https://doi.org/10.1098/r2007-1251
Ordway EM, Asner GP, Lambin EF (2017) Deforestation risk due to commodity crop expansion in sub-Saharan Africa. Environ Res Let. 12:044015. https://doi.org/10.1088/1748-9326-6509
Peeters C, Crewe RM (1987) Foraging and recruitment in Ponerine ants: solitary hunting in queenless Ophtalmopone berthoudi (Hymenoptera: Formicidae). J Entomol 94:201-214. https://doi.org/10.1155/1987-74592
Rosen D (1967) On the relationships between ants and parasites of coccids and aphids on citrus. Contrib Entomol 17:281-286.
Sighomnou D (2004) Analyse et redéfinition des régimes climatiques et hydrologique du Cameroun: perspectives d’évolution des ressources en eau. Dissertation, Université of Yaoundé I
Silva ER, Noda SCM (2000) Aspectos da atividade forrageadora de Mischocyttarus cerberus styx Richards, 1940. (Hymenoptera, Vespidae): Duração das viagens, especialização individual e ritmos diário e sazonal. Rev Bras Zooc 2:7-20.
Steiner FM, Schlick-Steiner BC, Holzinger WE, Komposch C, Pazoutova S, Sanetra M, Christian E (2004) A novel relationship between ants and a leafhopper (Hymenoptera:Formicidae; Hemiptera: Cicadellidae). Eur J Entomol 101:689-692
Stephens DW, Krebs JR (1971) Foraging theory, Cambridge University Press, Princeton
Stadler B, Dixon AFG (2005) Ecology and evolution of Aphid-Ant Interactions. Annu Rev Ecol Evol Syst 36:345-372. https://doi.org/10.1146/a36-091704-175531
Torres JA (1984) Niches and coexistence of ant communities in Puerto Rico: repeated pattern. Biotropica 16:284-295.
Tscharntke T, Clough Y, Bhagwat SA, Buchori D, Faust H, Hertel D, H¨olscher D, Juhrbandt J, Kessler M, Perfecto I, Scherber C, Schroth G, Veldkamp E, Wanger TC (2011) Multifunctional shade-tree management in tropical agroforestry landscapes. J Appl Ecol 48:619-629. https://doi.org/10.1111/j1365-2664-2010-01939
Voora V, Bermúdez S, Larrea C (2019) Global Market Report:Cocoa. International Institute for Sustainable Development. http://www.jstor.com/stable/resrep22025 Accessed 1 January 2019
Way MJ (1963) Mutualism between ants and honeydew-producing Homoptera. Annu Rev Entomol 8:307-344.
Wessel M, Quist-Wessel PMF (2015) Cocoa production in West Africa, a review and analysis of recent developments. J Life Sci 75:1-7 https://doi.org/10.1016/j2015-09-001
Yede (2016) Diversité des peuplements des hémiptères dans les cacaoyères de la région du Centre Cameroun:impact économique et essai de lutte biologique. Dissertation, Université of Yaoundé I
Yede Mballa NPA, Tadu Z, Bilong BCF (2023) Diversity and structure of ant communities associated with Toxoptera aurantii (Boyer, 1941) (Hemiptera, Aphididae) in cocoa farms in the Centre Region of Cameroon: Role of anthropogenic activities. J Bio Env Sci 23:139-151

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Foraging strategies of two dominant native ant species associated with Toxoptera aurantii (Hemiptera, Aphididae) on cocoa trees in Centre Region of Cameroon

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Abstract

Figures

Introduction

Material and methods

Study sites

Frequency of local ant activities associated with Toxoptera aurantii

Duration of palpation and honeydew collection

Data analysis

Results

Overall number of foraging ants

Daily foraging activity

Correlation between foraging activities and abiotic factors

Duration of foraging activities

Difference between number of ants per trees and foraging patterns

Discussion

Conclusion

Declarations

References

Table

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