Alex. J. Agric. Sci. Vol. 64, No. 5, pp. 331-339, 2019

Parasitoid Dinarmus acutus (: ) Parasitizing on bruchid beetle, Acanthoscelides macrophthalmus on Leucaena Tree at Alexandria, Egypt Amany M.H.Abu-shall1and M.E.Tawfeek Department of Applied Entomology and Zoology, Faculty of Agriculture, Alexandria University, Egypt ABSTRACT Acanthoscelides macrophthalmus (Schaeffer) (Coleoptera: Chrysomelidae: Bruchinae) hold over high host specificity to Leucaena leucocephala (Lamark) deWit (Fabales: Fabaceae) at Alexandria (Egypt). The infestation rate of the seed beetle and its population fluctuation were determined under natural conditions. The rates were low between February and May 2017, While, it increased and reached a peak in August and decreased in the next January. The parasitoid Dinarmusacutus (Thompson) (Hymenoptera: Pteromalidae) emerged in August with high numbers and reached a peak in September, 2017. The current study recorded A.macrophthalmus as a new host of D. acutus at Alexandria. The average survival rate of A. macrophthalmus and D. acutus until the death of adults were 30±3 days and 21±3 days, respectively. This study presented some baseline information regarding the development of A. macrophthalmus that may be beneficial as a biological control agent of L. leucocephala at Alexandria, Egypt. In contrast, if L. leucocephala was considered as beneficial tree, the mass rearing of D. acutus should be done to control A. macrophthalmus.

Keywords: biological control, population fluctuation, Leucaena leucocephala, seed bruchid beetle, Acanthoscelides macrophthalmus, parasitoid, Dinarmus acutus.

INTRODUCTION (Bruchinae) and distributed for the fauna of the world (Gupta et al., 1997; Andriescu and Mitroiu, Acanthoscelides macrophthalmus (Schaeffer) 2004; Ghahari et al., 2010; Tselikh, 2012; Noyes, (Coleoptera: Chrysomelidae: Bruchinae) is a seed 2014; Dzhanokmen, 2015; Alrubeai, 2017; Tselikh predator for controlling the weedy shrub Leucaena and Kostjukov, 2017). leucocephala (Lamark) de Wit (Fabales: A. macrophthalmus was first recorded in Fabaceae). Therefore, it is considered to be both a Egypt by Mohammad (2017). But, there are no pre- and post-dispersal ‘seed predator’(Kergoat et studies about the efficacy of it as a biological al., 2005; Raghu et al., 2005; Vassiliou and control agent. This study aims to determine the Papadoulis, 2008; Tuda et al., 2009) studies were population fluctuation of A. macrophthalmus and carried out on this beetle in several countries such its parasitoid, D. acutuson L. leucocephala at as Australia (Raghu et al., 2005), in South-East Alexandria, Egypt. This information is necessary Asia (Wu et al., 2013) and in Ethiopia (Yirgu et to use A. macrophthalmus as a biological agent for al., 2015). Bruchinae beetles are highly host- control invasive weed L. leucocephala or a host for specific seed feeders during the larval stage rearing the parasitoid D. acutus in a mass (Shobam and Olckers, 2010; Wood et al., 2017). production for biological control. Leucaena leucocephala is a tree/shrub cultivated for fodder, green manure, reforestation, MATERIALS AND METHODS windbreak, fuel, pulp, erosion control and 1-Natural infestation of Acanthoscelides vegetable crop (Barrett, 1990). On the other hand, macrophthalmus on Leucaena leucocephala:- It eventually became one of the 100 worst invasive Infestation by A. macrophthalmus was alien species in more than 20 countries where it followed in five trees of Leucaena leucocephala was introduced (Lowe et al., 2000 and Walton, planted in different parts around the Faculty of 2003). Its seed production is heavy (up to 1700 Agriculture, University of Alexandria, ElShatby pods/tree) and each pod containing approximately district. Weekly random pick up of 50 ripe pods on 20 seeds (Raghu et al., 2005). 5 unlike trees (10 pods from each tree) of L. The species richness and evenness of leucocephala were collected throughout the year parasitoids attacking the bruchid beetle, A. 2017. The ripe pods were transferred inside paper macrophthalmus feeding on L. leucocephala seeds packages to the laboratory. In the laboratory, the were determined. A total of 1420 parasitoids collected pods and their seeds were examined (Hymenoptera) belonging to four families and five carefully under a dissecting microscope at 20 to 40 subfamilies were recorded (Wood et al., 2017). magnifications. The pods and seeds of Leucaena Dinarmus acutus (Pteromalidae) is a primary infested by A. macrophthalmus contained exit parasitoid of many species of bean weevils circular neat holes, but bigger in diameter than the

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exit hole by parasitoid Dinarmus acutus (Fig. 1). at Alexandria [gardens of El-montaza (31o According to Sanders et al. (2013) 17’18.582” N, 30o57’488”E), Antoniads (31o A.macrophthalmus establish circular exit holes 13’18.004” N, 29o56’48.042”E), Faculty of middling 1.7 mm in diameter in both seed and pod. Agriculture El-Shatby (31o12’18.62”N, While, holes from the parasitoid were 29o55’8.86”E), Police hospital (31o 9’55.223.582” performed similar to the bruchid beetle exit hole N, 29o56’0.005”E), Alexandria Agricultural road but smaller mean 0.6 mm in diameter. (30°48.0316 N, 30°59.8407 E) and Smouha (31o Infested pods and percentage of natural 21‘29.17” N, 29o98‘78.06” E)], Egypt. Climatic infestation (%predation rate) of the pods and seeds mean values during the year of 2017, obtained by beetles of A. macrophthalmus were counted. from data reported by the weather station: 623180 From the counts, mean number of seed bruchid (HEAX), Ministry of Agriculture as showing in beetle per pod and percentage of unemerged Table (1). This ecological zone is characterized by beetles of A. macrophthalmus (the mummified monthly mean temperatures from 15 to 32°C remains as adults in exit hole) (Fig. 2) per pod were throughout the year and relative humidity determined. fluctuated between 65% and 87% during this 2- Population fluctuation and development of season at Alexandria, Egypt. A. macrophthalmus and its parasitoid wasp, Dinarmus acutus:- The present work was carried out under prevailing natural field conditions in six locations

(A) (B) Figure 1: (A) The exit circular holes of Acanthoscelides macrophthalmus on pods and seeds of Leucaena leucocephala. (B) The exit hole by parasitoid, Dinarmus acutus on seeds. (C)

Figure 2: The mummified remains of un-emerged Acanthoscelides macrophthalmus adults in exit hole on pods of Leucaena leucocephala.

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The population fluctuation of A. of A. macrophthalmus and D. acutus (Fig. 3). macrophthalmus and its parasitic wasp, Dinarmus Emerging bruchid beetles and the parasitoid in the acutus were estimated from January 2017 on L. glass tubes were picked up and counted daily. The leucocephala. Five of leucaena trees were selected removed adults of A. macrophthalmus and D. in each of the chosen locations with fix-space plant acutus from each tube were put separately in established. From each tree, 10 pods were collected labeled glass vial and covered with muslin cloth, monthly (50 pods per location). Samples of mature then observed daily until death of adults. Male and pods were taken and put in labeled paper packages. female adults of A. macrophthalmus and D. acutus Mature pods were placed in covered plastic jars in were identified by morphological traits (Fig. 4). the ambient conditions (28 ± 2° C and 75 ± 5% This information is essential to calculate the sexual RH) in the laboratory. Each plastic jar was fixed to ratio and survival rate of A. macrophthalmus and dark muslin cloth and covered in the top by plastic D. acutus in nature conditions at Alexandria, cover provided with glass tube for trapping adults Egypt. Table 1: Climatic mean values during 2017 at Alexandria, Egypt. Climatic mean values Season 2017 Temperature (oc) RH% January 17.3 + 1.9 70% February 18.5 + 2.6 68% March 21.1 + 2.01 65% April 23.6 + 2.6 65% May 27.8 + 3.9 67% June 29.5 + 2.5 81% July 30.8 + 2.7 80% August 31.9 + 0.6 78% September 29.2 + 1.2 87% October 27.7 + 5.6 77% November 24.1 + 4.7 80% December 19.2 +1.3 70%

(A) (B) Figure 3: (A) Plastic jar provided with glass tube for trapping adults of Acanthoscelides macrophthalmus and Dinarmus acutus. (B) Plastic jarscovered with dark muslin cloth.

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♀ ♂ ♀ ♂ (A) (B) Figure 4: (A) Adults (♀ and ♂) of Acanthoscelides macrophthalmus. (B)Adults (♀ and ♂) of Dinarmus acutus.

3- Morphometeric measurements of the (Table 2). The present study showed that hymenopterous parasitoid Dinarmus uninfested pods contained (7.9 ± 0.2) intact seeds, acutus:- while pods infested by this beetles included Measurements of ten individuals from each significantly (P < 0.05) reduced number of intact males and females of the parasitoid Dinarmus seeds (2.6 ± 0.2). Previous studies indicated that acutus were conducted using research microscope between 8 and 30 seeds are created per uninfested and micrometer lens. The means of ten pod (Walton, 2003; ISSG, 2006; Sankaran, 2007). measurements in mm were obtained with Standard An average of 14.67 seeds per pod was collected deviation. (Sanders et al., 2013). 4- Statistical analyses:- The present data shoewd that A. The mean numbers of the distinct parameters macrophthalmus reduce seed production in L. of the experiments were calculated using the leucocephala. Green and Palmbald (1975) analysis of variance test (ANOVA), with the mean established that the seed predation by Acanthosce separation at 5% level of significance using iides fraterculus (Horn) overstep 60% in computer program Costat according to the method Astragalus cibarius Sheld. And Astragalus of Snedecor and Cochran (1967). utahensis (Tom) populations, and reach nearly 100% (Rogers and Garrison, 1975) in natural plant RESULTS AND DISCUSSION communities, and this variation related to years and 1-Natural infestation of Leucaena leucocephala locations (Baskin and Baskin, 1977). Seed by Acanthoscelides macrophthalmus:- predation of Glycyrrhiza lepidota Pursh vary The mature pods of Leucaena leucocephala between 7 and 71% with an general average of 41 infested by A. macrophthalmus were 72 ± 7.8% ± 2% (Boe and Wynia, 1985). So, Legume seed from all the collected pods and the in general predation by bruchid beetles may overstep 50%. average percentage of seed loss due to infestation While, Boe et al. (1989) found 36% of the pods by this beetle were 80 ± 7.7%. Emerged percent of included Acanthosce iides perforatus (Horn) A. macrophthalmus adults were 69 ± 7.3% from (Coleoptera: Bruchinae) in both lobules and larvae the pods on both sides, which contained the exit of the seed predator taken place in 77% of the circular holes to the bruchid beetle. But, the mature pods. mummified remains of un-emerged A. macrophthalmus adults in exit hole on pods of L. leucocephala were 11 ± 7.6% from both sides Table 2: The infested pods and percentage of predation rate by beetles of Acanthoscelides macrophthalmus on Leucaena leucocephala during 2017 at Alexandria, Egypt. %Infested Mean number of % Predation rate by beetles % unemerged beetles pods beetles / pod ±S.E pods seeds from pods 72 ± 7.8 13.25 ± 2.1 69 ± 7.3 80 ± 7.7 11 ± 7.6 -Means significantly different at the 0.05 level. - Means of 5 different trees of L. leucocephala; 10 pods per tree (50 sample).

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Johnson, 1981 stated that seeds of L. bruchind beetles, and the hymenopterous parasitoid leucocephala are small (about 1.5 mg/seed), and is important to comprehending their ecology and mean number beetles of A. macrophthalmus population fluctuation. The effect of bruchid devoured seeds about 13.25± 2.1 per pod, one or beetles on long-term seed production of native more bruchid larvae feed and develop inside a legumes and the efficacy of hymenopterous single seed in large-seeded legumes. While, the parasitoids for reducing seed losses can be decided larvae of A. perforatus devoured from 2 to 12 seeds after more extend evaluation. (Boe et al., 1989). 2-1.Temporal variation of A. macrophthalmus A. macrophthalmus has been invaded West and parasitic wasp, D. acutus density Africa, feeding on the host plant, L. leucocephala In the laboratory, collected L. leucocephala (Delobel and Johnson, 1998). Since 1999 this pods from six locations showed that A. bruchinae species has been introduced in South macrophthalmus is present all the year. The Africa for biological control of L. leucocephala number of emerged A. macrophthalmus adults was (ARC-PPRI, 2003; Olckers, 2004). Only, A. low during the first half of the season 2017, macrophthalmus was used as bio-agents control of between February and May. Number of adult’s weedy plants and emerge from the seeds of L. emergence increased from June and arrived a peak leucocephala (Tuda et al., 2009). A. in August, and starting to decrease up to January. macrophthalmus emerged from L. leucocephala On the other hand, adults of parasitoid began to was able to place eggs and hatching larvae were emerge in August with high numbers and reached a able to feed on L. leucocephala seeds, after that peak in September, but only a few adults of A. bruchid beetles emerged from these seeds. Our macrophthalmus emerged during this time of 2017 review of the issued work indicated that A. (Fig. 5). macrophthalmus had high host specificity to genus The present results agree with that of Elder Leucaena not in native regions only but also in (2002) and Effow et al. (2010) who mentioned that introduced regions (Pfaffenberger and Johnson, the females of bruchid beetle deposit their eggs 1976; Johnson, 1979; Jones, 1996; Hughes and throughout the year on ripe pods. The infestation Johnson, 1996; Delobel and Johnson 1998; Tuda et rates of this beetle were low between February and al., 2009; Olckers and Welsford, 2013; English and March, corresponding to the majority of L. Olckers, 2014). leucocephala trees partially loses their foliage and 2- Population fluctuation of A. macrophthalmus dry pods. These old pods did not prefer to be and its parasitoid, Dinarmus acutus during selected by the females for oviposition. But, the 2017 at Alexandria, Egypt:- rates of infestation increased in April and a This study identified Dinarmus acutus as a maximum rate of assault taken place between parasitoid of A. macrophthalmus and showed its August and December (64 and 72% of the pods direct effect on the reproductive capacity of the were charged). This period corresponds to the good plant L. Leucocephala and population fluctuation flowering and new pods that are preferentially of A. macrophthalmus. These findings propose that chosen by females for oviposition explanation of the interactions between host plant,

Figure 5: Population fluctuation of Acanthoscelides macrophthalmus and its parasitoid, Dinarmus acutus during 2017 at Alexandria, Egypt.

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The parasitoid Dinarmus acutus was parasitoid, D. acutus emerged from collected pods, parasitizing Acanthoscelides macrophthalmus as a respectively, during this season of 2017. Under new host recorded throughout the present study in laboratory conditions, a female of D. Alexandria, Egypt. This parasitoid has been acutus produced 35 offspring in a sex ratio of 1:1. breeded from Bruchidius ater (Marsham) and Females preferred mature pods infested with Bruchus brachialis Fahr. (Krombein et al., 1979) bruchid and 4th larval instars. The egg was and registered from many of seed legumes deposited inside the cavity of seed on the body including Acanthoscelides fraterculus, A. griseolus surface of the bruchid host. The larval stage of this (Fall) (Johnson, 1970) and A. perforatus (Boe et external parasite consists of 3 larval instars and al., 1989). D. acutus had a straight effect on L. overwintered as a 3rd instar inside seeds. The life leucocephala by decreasing the number of seeds cycle required for completion from egg to adult devoured by larvae of A. perforatus. Since larvae about 24 days and 10 months in non-overwintering of A. perforatus consumed more food of seeds 85% and overwintering brood, respectively (Leong and than those parasitized by D. acutus 62% (Boe et Dickason, 1975). Also, Effowe et al. (2010) found al., 1989). Fabres and Reymonet (1991) found that that the sex-ratio of A. macrophthalmus on pods the ectoparasite D. Acutus emerge in small and seeds were 50.3% and 50.9% females in numbers in autumn of the same year and then southern Togo (West Africa), respectively. abundantly in spring of the following year after a 3- Morphometeric measurements of the diapause in the last larval stage. While, few of the hymenopterous parasitoid Dinarmus D. acutus adults were found during July or early acutus:- August, and more between mid- August to During this study, observation showed that September (Leong and Dickason, 1975). the size of A. macrophthalmus adults was different 2-2. Adult survival of A. macrophthalmus and its between beginning and the end of the season. Haga parasitoid, D. acutus in the laboratory and Rossi (2016) reported that the L. leucocephala In laboratory, the survival rate of A. seed characteristics influence on the body size and macrophthalmus and its parasitoid, D. acutus until of A. macrophthalmus. According to that variation the death of adults were 30±3 days and 21±3 days, in measurements of body size of the parasitoid D. respectively. The adult female of both A. acutus adults was appeared. So, the measurements macrophthalmus and D. acutus lived from two to of males and females of the parasitoid D. acutus three weeks. adults were accomplished and the mean ± Standard During the development of A. deviation in mm were given in Table (3). macrophthalmus and the parasitic wasp, D. basalis Future studies are required to determine the in laboratory, the total developmental period (egg relation between the size of A. macrophthalmus to adult) continued 33.8 ± 2.9 days and 33.4 ± 2 larvae and the size of its parasitoid D. acutus. days on pods and seeds, respectively. These means were different according to substrate of CONCLUSION oviposition; which was longer on pods than on The bruchid beetle, Acanthoscelides seeds. The development was low at the first stages macrophthalmus (Schaeffer) (Coleoptera: because of very hardness of the tegument of Chrysomelidae: Bruchinae) oviposit on seed pods L. leucocephala seeds that can create a barrier for The the perception of the first instars inside the seed of the tree Leucana leucocephala (Lam.). (Mondedji et al., 2002). This is the main reason to beetles lay their eggs on seed pods and on loose describe the long development of A. seeds of its host. Hatching larvae burrow into the macrophthalmus when the eggs are laid on pods as seeds to consume the contents, emerging as adults. compared to on seeds (Effowe et al., 2010). It was released as biological control agent to During their lifetime, the sex-ratio (♂: ♀) was reduce the invasive Neotropical tree Leucaena approximately 1: 1.5 and 1: 2.7, when calculated leucocephala in certain countries. from the adults of A. macrophthalmus and its Table 3: Mean ± S.D. measurements (mm) of males and females of the parasitoid Dinarmus acutus adults on its host Acanthoscelides macrophthalmus at Alexandria, Egypt. Mean± S.D. of the parasitoid D. acutus adults Male Female L. Body L. Ant. L. Wi. L. Body L. Ant. L. Wi. 0.2153±6.7 0.54±3.4 0.129±2.9 0.371±16.6 0.106±1.2 0.214±2.1 *Abbreviations: L. Body= The length of Body, L. Ant. = The length of Antenna, L. Wi. = The length of Wing. *data from 10 individuals.

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The data in this study introduce, some Dzhanokmen, K. A. (2015). Pteromalids information regarding the development of A. (Hymenoptera, Chalcidoidea: Pteromalidae) of macrophthalmus that may be beneficial for its the Sajram-Ugam State National Nature Park promotion as a biological control agent, if L. in Western Tien Shan. Entomological Review, leucocephala was considered as invasive weed in 95(4): 456–471. Egypt. The parasitoid, Dinarmus acutus Effowe, T. Q., Amevoin, K., Nuto, Y., Mondedji, (Thompson) (Hymenoptera: Pteromalidae) was D. and Glitho, I. A. (2010). Reproductive recorded for the first time on A.macrophthalmus at capacities and development of a seed bruchid Alexandria, Egypt. If the Neotropical tree L. beetle, Acanthoscelides macrophthalmus, a leucocephala consider as beneficial agroforestry, potential host for the mass rearing of the A. macrophthalmus could be used as a host for parasitoid, Dinarmus basalis. Journal of rearing the parasitoid D. acutus in a mass Science. 10: 129. ISSN: 1536-2442.Online at: productionto biocontrol A. macrophthalmus. insectscience.org/10.129. Elder, R. (2002). Leucaena seed bruchid in AKNOWLEDGMENT Leucaena. DPI Note 3181, Available online: Authors deeply thank Dr. Nevein Gad Allah http://www2.dpi.qld.gov.au/beef/3181.html. professor of Entomology - faculty of Science - English, K. F. and Olckers, T. (2014). Does the Cairo University who identify the parasitoid, size of the seeds and seed pods of the invasive D. acutus tree Leucaena leucocephala (Fabaceae) affect their utilization by the biological control agent REFERENCES Acanthoscelides macrophthalmus Alrubeai, H. F. (2017). Biological Control of Insect (Chrysomelidae: Bruchinae). African Pests in Iraq: 1) an Overview of Parasitoids Entomology. 22(4): 872–879. and Predators Research Development. Fabres, G. and Reymonet, C. (1991). L'induction Academic Journal of Entomology. 10 (2): 10- maternelle de la diapause larvaire chez 18. Dinarmus acutus [Hym.: Pteromalidae]. Andriescu, J. and Mitroiu, M. D. (2004). “Notes on Entomophaga. 36(1): 121-129. the Pteromalid Fauna (Hymenoptera: Green, T. W. and Palmbald, I. G. (1975). Effects of Chalcidoidea, Pteromalidae) of Dobrogea, insect seed predators on Astragalus cibarius Romania (II),” An. Şti. Univ. “AL. I. CUZA” and Astragalus utahensis (Leguminosae). laşi, serie Biologie . 50, 89–96. Ecology. 56: 1435-1440. [ARC-PPRI] Agricultural Research Council-Plant Gupta, H. C., Gupta, I. J. and Sharma, S. N. Protection Research Institute (2003). Releases (1997). Parasitic , Dinarmus acutus of biological control agents against weeds in (Thompson) and Dinarmus basalis (Rondani) South Africa. (Hymenoptera: Pteromalidae) parasitizing http://155.240.199.39/institutes/ppri/main/ pulse beetle divisions/weedsdiv/releases.htm. (Fabricius). Journal of Biological Control. 11 Barrett, R. P. (1990). Legume species as leaf (1/2): 77-78. vegetables. In: Janick J, Simon JE (eds) Haga, E. B. and Rossi, M. N. (2016). The effect of Advances in New Crops. Timber Press, seed traits on geographic variation in body size Portland, OR. 391–396. and sexual size dimorphism of the seed- Baskin, J. M. and Baskin, C. C. (1977). Predation feeding beetle Acanthoscelides of Cassia marilandica seeds by Sennius macrophthalmus. Ecology and Evolution. 6 abbreviatus (Co1eoptera: Bruchidae). Bull. (19): 6892-6905. Torr. Bot. Club. 104: 61-64. Hughes, C. E. and Johnson, C. D. (1996). New host Boe, A. and Wynia, R. (1985). Seed predation, records and notes on Bruchidae (Coleoptera) seedling emergence, and rhizome from Leucaena Benth. (Leguminosae, characteristics of American licorice. Journal of Mimosoideae) from Mexico, Central and Range Management. 38: 400-402. South America. Journal of Applied Boe, A., MCDniel, B., and Robbins, K. (1989). Entomology. 120: 137–141. Direct effect of parasitism by Dinarmus acutus ISSG.(2006). Ecology of Leucaena leucocephala. Thomson on seed predation by In: Invasive Species Specialist Group (ISSG) Acanthoscelides perforatus (Horn) in Canada (Eds) Global Invasive Species Database. milk-vetch. J. Range Manage. 42(6): 514-515. http://www.issg.org/database/species/ecology. Delobel, A. and Johnson, C. D. (1998). First record asp?fr=1&si=23 (accessed 11 April 2011). of a seed-beetle on Leucaena leucocephala in West Africa. Leucnet News. 5: 25–26.

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Vassiliou, V. A. and Papadoulis, G. (2008). First Wu, L. H., Wang, C. P. and Wu, W. J. (2013). record of Acanthoscelides macrophthalmus Effects of temperature and adult nutrition on (Schaeffer) (Coleoptera: Bruchidae) in Cyprus. the development of Acanthoscelides Entomologia Hellenica. 17: 52 – 55. macrophthalmus, a natural enemy of an Walton, C. S. (2003). Leucaena leucocephala in invasive tree, Leucaena leucocephala. Queensland. Pest Status Review Series –Land Biological Control. 65: 322–329. Protection, Department of Natural Resources Yirgu, A., Gezahgne, A. and Tsega, M. (2015). and Mines, Queensland.50 pp. First report of Acanthoscelides Wood, A., Haga, E. B., Costa, V. A. and Rossi, M. macrophthalmus (Schaeffer) on Leucaena N. (2017). Geographic distribution, large-scale leucocephala (Lam.) de wit in Ethiopia and a spatial structure and diversity of parasitoids of preliminary investigation into its impacts. the seed-feeding beetle Acanthoscelides African Entomology. 23 (2): 280-285. macrophthalmus. Bulletin of Entomological Research. 107 (3): 322-331.

Dinarmus acutus(Hymenoptera: Pteromalidae) Acanthoscelides macrophthalmus

(Coleoptera: Chrysomelidae: Bruchinae) Acanthoscelides macrophthalmus(Schaeffer) (Fabales: Fabaceae) Leucaena leucocephala (Lamark) deWit

Dinarmus acutus (Thompson)

(Hymenoptera: Pteromalidae)

D. acutus A. macrophthalmus

D. acutus A. macrophthalmus

A. macrophthalmus

D. acutus

A. macrophthalmus

339