Phytophagous insects associated with the reproductive structures of mesquite (Prosopis spp.) in Argentina and their potential as biocontrol agents in South Africa

F. Mc Kay* & D. Gandolfo South American Biological Control Laboratory, USDA-ARS, Hurlingham, Argentina

Mesquites (Prosopis spp.) are thorny leguminous shrubs or trees native to Southwest Asia, Africa, and predominantly North and South America. Introduced as beneficial plants to India, Pakistan, South Africa, Egypt, Kuwait, Australia, U.S.A. (Hawaii), and Brazil, some Prosopis species have become invasive in some of these countries. Mechanical and chemical control methods implemented in the U.S.A., Australia and South Africa have been only marginally effective. In an attempt to preserve the beneficial attributes of mesquite while arresting its spread, biocontrol efforts in South Africa have only focused on seed-feeding agents. Between 1987 and 1993, three species of bruchids were introduced from the U.S.A., but their impact has been limited as ripe pods are eaten by livestock before the agents have had a chance to eat the seeds. In this paper we present a list of insects that attack the repro- ductive structures of Prosopis species in Argentina, focusing on those that feed on green, unripe pods and flower buds. We provide information on the biology, distribution and host range of nine beetles, four moths and one gall midge species. Their potential as biological control agents against mesquite in South Africa is discussed with special reference to that of a new weevil species, Coelocephalapion gandolfoi Kissinger which was selected for further studies. The adults feed and oviposit on green pods, while the larvae feed on the immature seeds, apparently without causing a substantial reduction in the value of pods as fodder. Key words: mesquite, weed biocontrol, natural enemies, Prosopis, Coelocephalapion.

INTRODUCTION

Prosopis species, commonly known as mesquite, Prosopis glandulosa Torrey (honey mesquite) and are thorny, usually xerophilous shrubs or trees in Prosopis velutina Wooton (velvet mesquite), native the family Fabaceae. About 44 species are found in to southwestern North America and Prosopis southwest Asia, Africa, and predominantly North ruscifolia Grisebach (vinal) native to northern and South America, with the centre of polymor- Argentina, constitute serious weeds within their phism in Argentina (Burkart 1976). A few species native range (Burkart 1976). of Prosopis have been introduced as useful plants Mechanical and chemical methods to control in many countries, notably India, Pakistan, South Prosopis spp. have been implemented for example Africa, Egypt, Kuwait, Australia, U.S.A. (Hawaii) in the U.S.A., South Africa, Australia, but have met and Brazil, for their wood, erosion control, the use with limited success because they are labour- of pods as fodder,and for the production of honey, intensive and expensive, with costs of application gum and charcoal. However, in several of these usually far exceeding the value of the land (De countries some Prosopis species have become Loach 1985; Zimmerman 1991; van Klinken & weeds (De Loach 1985). Campbell 2001). By contrast, biological control is In South Africa, where all Prosopis species are seen as a long-term, economical and non-polluting exotic, their value is lost when impenetrable alternative method to control this weed. thickets of densely packed, thin-stemmed plants A great number of phytophagous insects develop providing minimal shade, and producing associated with Prosopis have been reported both few flowers and pods (Zimmermann 1991). from the New World (Swenson 1969; Ward Mesquite also has a negative impact on scarce et al.1977; Cordo & De Loach 1987; Erb 1988; Silva water resources, especially in the drier parts of 1988), and the Old World (Gerling & Kugler 1973; South Africa (Impson et al. 1999). Moreover, Ward et al.1977). These studies constitute a valuable *To whom correspondence should be addressed. Present address: Agricultural Council, ARS Laboratory, U.S. Embassy Unit 4325, APO source of information when selecting natural AA 34034-0001, U.S.A. E-mail: [email protected] enemies for implementing biological control

African Entomology 15(1): 121–131 (2007) 122 African Entomology Vol. 15, No. 1, 2007 programmes. In Australia and South Africa, flower buds, giving special attention to seed- current integrated management strategies for feeding insects damaging immature pods. Insects mesquite include implementation of biologi- were sampled by hand, by beating and collection cal control programmes. In South Africa, these onto sheets, and by collecting reproductive struc- programmes have been restricted to the use of tures for subsequent emergence of insects. seed-feeding insects to arrest the spread of the Voucher specimens of the plants and insects col- weed while preserving its beneficial attributes. To lected were maintained at the USDA-ARS-SABCL. date, three species of bruchid beetles (Algarobius prosopis (Le Conte), Algarobius bottimeri Kingsolver RESULTS AND DISCUSSION and Neltumius arizonensis Shaeffer) have been introduced from the U.S.A. into South Africa. Al- Fourteen insect species, nine Coleoptera, four though two of these beetles have established (A. Lepidoptera and one Diptera were found damaging prosopis and N. arizonensis), they failed to provide reproductive structures of 10 Prosopis species the desirable level of control, mainly because live- (Table 1). Although most of these insects have pre- stock consume the ripe pods before they are uti- viously been reported (Gerling & Kugler 1973; lized by the beetles (Impson et al. 1999). The Ward et al. 1977; Cordo & De Loach 1987; Erb realization that the problems caused by mesquite 1988; Di Iorio 1995), new information on their outweigh the benefits, has recently led to consid- distribution, host plants, biology and potential as eration of additional control agents (Impson et al. biocontrol agents is now provided. 1999). In August 2002, the ARC-Plant Protection Re- Insects attacking immature pods search Institute (ARC-PPRI), Pretoria, South Africa, Coelocephalapion gandolfoi Kissinger (Coleoptera: and the USDA-ARS South American Biological Apionidae). This seed-feeding weevil (Fig. 2) Control Laboratory (SABCL) in Argentina initi- was collected along a wide latitudinal range ated a cooperative research programme for the (c. 1500 km), on nine Prosopis species within the biological control of mesquite in South Africa. The Algarobia group, namely P.affinis (Series Pallidae), main objective was to survey for natural enemies P. alba, P. alpataco, P. caldenia, P. chilensis, P. flexuosa of Prosopis species in Argentina and select poten- and P. nigra (Series Chilenses), P. ruscifoliae and tial biocontrol agents for use against problem P. vinalillo (Series Ruscifoliae) (Fig. 1). mesquite trees in South Africa. In this paper we Overwintering, reproductively mature adults present a prioritized list of insects that attack repro- were first found at the beginning of spring, late ductive structures of Prosopis species in Argentina. September, when mesquite trees started to produce We provide information on their biology, distribu- leaves and flowers. Adults became more abundant, tion, host plants and discuss their potential as bio- or at least more noticeable, as young green pods logical control agents against mesquite in South appeared. Adults fed actively on immature pods Africa. and oviposited near the seeds, inside the tender pod tissues. Newly hatched larvae entered the MATERIAL AND METHODS one-seeded endocarp units and fed on the seed, pupating within it (Fig. 3). Adults emerged from Four extensive exploratory trips were made the ripening pods approximately 40 days after between October 2002 and February 2004, in the oviposition. Newly emerged adults appeared in main distribution area of Prosopis in Argentina, the field when most of the pods hanging from the which ranges from the semi-arid eastern slopes of trees were ripening or ripe, and consequently the Andes to the central and eastern plains of the unsuitable for oviposition or even feeding. Monte and Chaco phytogeographical regions Under laboratory conditions, when suitable (Fig. 1). Field surveys were mainly conducted on excised young green pods were offered to newly Prosopis species from Section Algarobia, Series emerged adults, they laid few eggs, suggesting Chilenses, where the most troublesome species, that newly emerged adults are reproductively P. glandulosa var. torreyana and P. velutina growing immature, and have to overwinter to become in South Africa are classified. Collection of natural sexually mature. According to this hypothesis, enemies was restricted to insects attacking repro- there would be one weevil generation per year. ductive structures, seeds, pods, flowers and In terms of biological control of mesquite in Mc Kay & Gandolfo: Phytophagous insects associated with reproductive structures of mesquite 123

Fig. 1. Survey area showing Prosopis species and collecting sites of Coelocephalapion gandolfoi in Argentina. (BA, Buenos Aires; C, Corrientes; Ca, Catamarca; Ch, Chaco; Co, Córdoba; ER, Entre Ríos; F, Formosa; J, Jujuy; LP, La Pampa;LR, La Rioja;M, Mendoza;Mi, Misiones;N, Neuquén;S, Salta;SE, Santiago del Estero;SF,Santa Fé;SJ, San Juan; SL, San Luis; T, Tucumán).

South Africa, we consider this weevil to be suitable Anypsipyla sp. nr. univitella Dyar (Lepidoptera: in that it oviposits exclusively on immature green Pyralidae) and Cydia sp. (Lepidoptera: Tortricidae). pods thereby destroying the seeds before livestock Anypsipyla nr. univitella was previously reported can gain access to them. In addition, although by Cordo & De Loach (1987) on P. alpataco, P. larvae destroy the seeds, pods still develop, appar- chilensis and P. flexuosa and on P. alba. During our ently without losing much of their value as fodder, surveys, this moth was also found damaging pods an important benefit of mesquite trees. In terms of of P. caldenia, P. nigra, P. ruscifolia and P. vinalillo, its specificity, studies conducted in Argentina on which constituted new host plant records (Table 1). 27 leguminous plants, indicate that the host range Females laid eggs in the mesocarp of the young of C. gandolfoi is restricted to nine Prosopis species green pods. The bluish-red larvae tunneled inside in the Section Algarobia, the same botanical group the immature pods feeding on the mesocarp and to which the weed species growing in South Africa the seeds. Larval feeding activity was recognized belong (Mc Kay & Gandolfo, in prep.). Additional by the presence of frass on the surface of the fruits host-specificity studies conducted in South Africa, (Fig. 4). specifically on Acacia, Prosopis species and hybrids Cydia sp. was more common on Prosopis species growing in South Africa, have confirmed the occurring in the west-central provinces of Argen- specificity of C. gandolfoi to Prosopis (A. Witt, pers. tina (Table 1). Eggs were laid on the pods and as in comm.). A.nr.univitella, the larvae fed on the mesocarp and 124 African Entomology Vol. 15, No. 1, 2007

Continued on p. 125 M; N; SSE; SL; SJ species in Argentina.

Prosopis or modeSeed feeders (unripe pods) Beating/emerged from ripening pods method ER; CA; CH; CO; F; J; LR; LP; Common/locally abundant Flower budsSeed feeders (ripe pods) Emerged from ripening pods Beating/emerged from flower buds ER LPSeed feeder (unripe/ripe pods) Emerged from ripening podsSeed feeder (ripe pods) Locally abundant Locally abundant CH; LP; LR; SL Emerged from ripening pods CH; CO; LP; LR; Occasional SE Abundant Seed feeders (unripe/ripe pods) Emerged from ripening podsSeed feeder (unripe/ripe pods) Emerged from ripening pods CH; F; LP; LR CH; CO; LP; LR; SL Occasional Common/abundant spp. Attack site Collection Distribution* Abundance Prosopis P.caldenia P.alpataco P.chilensis P.flexuosa P.nigra P.alba P.alpataco P.caldenia P.chilensis P.flexuosa P.nigra P.ruscifolia P.vinalillo P.affinis P.caldenia P.flexuosa P.flexuosa P.ruscifolia P.caldenia P.chilensis P.flexuosa P.nigra P.alba P.caldenia P.flexuosa P.alba (Fahr.) ) spp. S. vinalicola ; . Natural enemies found attacking reproductive structures of spp. S. terani Coleoptera Apionidae Coelocephalapion gandolfoi P.affinis Bruchidae Pectinibruchus longiscutus P.alpataco ( Table 1 Natural enemies Apion Rhipibruchus atratusRhipibruchus psephenopygus P.alba P.alba Rhipibruchus picturatus Scutobruchus Mc Kay & Gandolfo: Phytophagous insects associated with reproductive structures of mesquite 125

Continued on p. 126 S; SE; SJ; SL; T SE; SJ; SL; T Emerged from ovary galls CA; CH; CO; F; LP; LR; Common/abundant or modeRipe pods (mesocarp and seeds) Emerged from ripe pods method Flower (gall maker) CH; SE; SLUnripe pods (gall Occasional maker) Emerged from gallsUnripe pods(mesocarp and seeds) CA; CH; CO, F; LP; LR; S; Common/locally abundant Emerged from ripenig pods CA; CH; CO; F; LP; LR; S; Common/abundant SE; SJ; SL spp. Attack site Collection Distribution* Abundance Prosopis P.nigra P.chilensis P.caldenia P. chilensis P.flexuosa P.nigra P.kuntzei P.ruscifolia P.alba P.caldenia P.chilensis P.flexuosa P.kuntzei P.nigra P. alpataco P. caldenia P. flexuosa P.chilensis P. nigra P.ruscifolia P.vinalillo prosopidis P.alba prosopidis P.affinis ) univitella P.alba sp. nr. sp. nr.

continued a sp. nr. ( Cerambycidae Lophopoeum bruchi P.alba Table 1 Natural enemies Diptera Cecidomyiidae Asphondylia Asphondylia Lepidoptera Pyralidae Anypsipyl 126 African Entomology Vol. 15, No. 1, 2007

on the seeds. In the field, future adult emergence holes, previ- ously formed by the final larval instar, were distinguished as round marks on the exocarp of the pods. Although pupation in the laboratory took place outside the pods, in the field, pupal skins were commonly found attached to the emergence holes.

ndoza; N, Neuquén; S, Salta; Both Cydia sp. and A. nr. univitella larval feeding cause a mix of frass and fruit juices that seems to facilitate secondary pathogenic infections. The com- bined effect of larval feeding and fungal activity may make these agents potentially more damag- ing than C. gandolfoi. Conse- quently, if the host range of these moths is narrow enough for their safe introduction in South Africa, the use of these insects as biocontrol agents will depend on the necessity for insects more injurious to mesquite. Asphondylia sp. nr. prosopidis (Diptera: Cecidomyiidae) (R.J. Gagné, pers. comm.) This gall midge induced galls on the mar- gin of immature green pods and in flower buds (Figs 5, 6). Erb (1988) previously reported the existence of this midge as the most frequent and abundant cecidomyiid infesting the flow- ers of P. ruscifolia in northwest- Unripe pods(mesocarp and seeds) Flowers Emerged fromripening pods CH; CO; LP; LR; SLPeduncles (gall Emerged maker) from flowers Locally abundant Emerged from peduncle galls CA; CH; CO; F; LR; S,SE; T CH; COR; ER; F; LP; Common/abundant SE; SL Occasional or mode method ern Argentina. According to Gagné (pers. comm.), this spe- spp. Attack site Collection Distribution* Abundance cies is closely related to A. prosopidis, a gall midge that pro- P.alba P. caldenia P.flexuosa P.alba P.chilensis P.flexuosa P.nigra P.kuntzei P.ruscifolia P.alba P.caldenia P.chilensis P.nigra P.ruscifolia Prosopis duces ovary galls on Prosopis glandulosa in southwestern North America. The only dis- tinction between these two spe- cies is a slight difference in the

) shape of the larval sternal spat- ula, which is distinctively sp.

sp. reduced in size in comparison to

continued (

sp. other Asphondylia species. Fruit margin galls were re- SE, Santiago del Estero; SJ, San Juan; SL, San Luis; T, Tucumán. Gelechiidae Polyhymno Gracillaridae Parectopa Tortricidae Cydia Table 1 Natural enemies *Provinces of Argentina: CA, Catamarca; CH, Chaco; CO, Córdoba, COR, Corrientes; ER, Entre Ríos; F, Formosa;corded J, Jujuy; LP, La Pampa; LR, La Rioja; M, Me on eight Prosopis species, Mc Kay & Gandolfo: Phytophagous insects associated with reproductive structures of mesquite 127

Figs 2–6.2–3: Coelocephalapion gandolfoi; 2, adult; 3, empty one-seeded endocarp unit with pupa. 4, Anypsipyla sp. nr. univitella larval damage on green pod. 5–6: Asphondylia sp. nr prosopidis; 5, galls on a peduncle of Prosopis nigra. 6, galls on ovary and margin. a = ovary gall, b = intermediate gall, c = margin gall. over a wide distribution area (Table 1). In some lo- between 6 and 10 cm (mean ± S.D. = 5.84 ± 2.24; cations, 90 % of pods on specific trees were galled. n = 50) and, in most cases pods adopted a ‘C’ Situated on the edge and mainly on the proximal shape, with the gall as the inflection point (Fig. 6). third of pods, ovoid galls were conspicuous (3.1 ± Only one larva developed within the central cav- 0.52 mm long and 1.4 ± 0.38 mm wide) (mean ± ity of each gall, with adults emerging sometime S.D.; n = 20). Length of infested pods varied later. The presence of the gall on the young pods 128 African Entomology Vol. 15, No. 1, 2007 was consistently associated with a reddish color- alternate host in other plant families (R.J. Gagné, ation of the pods when ripening, and arrested pod pers. comm.). If this is the case for A. sp. nr. development followed by premature abscission prosopidis, field surveys on mesquite growing from the trees. sympatrically with related species should be con- The flower bud galls (3.4 ± 0.46 mm long by 1.5 ± ducted in order to identify the plants that may act 0.34 mm in diameter) (mean ± S.D.; n = 20) were as alternate hosts for this gall midge. red to green, thin-walled, ovoid-shaped structures Considering that this insect attacks not only the that consisted of the aborted and enlarged ovary flowers but also the young green fruits of Prosopis of the flowers. As in the fruit galls, inside each spp., it could be a promising biocontrol agent to flower gall, a single yellowish larva developed and arrest the spread of mesquite in South Africa, if pupated, after which adults emerged, leaving the suitably specific. pupal skin protruding from the gall. In the field, it was common to find inflorescences with almost all Insects attacking flowers of the flowers affected (Fig. 5). Polyhymno sp. (Lepidoptera: Gelechiidae). This Besides the existence of well-defined ovary and flower-feeding moth was collected at several margin fruit galls, we also found some intermedi- locations, frequently coexisting with A. sp. n. ate gall forms, which consisted of enlarged ovary prosopidis (Table 1). Larvae fed on the flowers, and galls that bore a little portion of a very young pod pupated inside a cocoon formed by silk and dry (2–3 cm long), distally to the gall (Figs 5, 6). Gerling flower material. Infested inflorescences contained & Kugler (1973) as part of a survey of natural two to three cocoons and became brown. After enemies of P. farcta conducted in Israel, found A. adults emerged, the silk net and dry flower material prosopidis responsible for the formation of ovary disintegrated and only a bare peduncle remained. galls, but indicated that in some cases, the gall, Generally no pods developed from an infested which starts developing as a hypertrophied ovary, inflorescence, but occasionally one or two pods eventually attains the shape of a young fruit. were observed. In addition, A. prosopidis, was reported to be In some areas, the combined action of both, responsible for producing two different types of A. sp. nr. prosopidis and Polyhymno sp., accounted bud galls on P. glandulosa var. glandulosa (Painter for 90 % of the damage to inflorescences, resulting in 1935; Felt 1935; Rogers 1973; Gagné 1989). Rogers substantial reduction in fruit and seed production. (1973) designated these bud galls as either ‘perse- Depending on its specificity and on whether the verant’ or ‘abortive’ bud galls, and indicated that use of biocontrol agents other than seed-feeders is gall types might be influenced by the stage of bud considered in the future, this flower-feeding moth development at the time of oviposition. Abortive would constitute a promising biocontrol candidate bud galls begin formation prior to fertilization of against mesquite in South Africa. the bud ovary, while perseverant galls avoid abortion by beginning their formation after the Insects attacking ripening pods ovary is fertilized. These observations support Bruchid beetles. Species of the genus Rhipibruchus the idea that A. sp. nr. prosopidis responsible for (3), Pectinibruchus (1) and Scutobruchus (2) emerged inducing the whole range of galls forms in Prosopis from pod samples of seven Prosopis species (Table 1). species in Argentina. Some species in these genera are known to be Our knowledge on the life cycle of A. sp. nr. restricted to feeding on seeds of Prosopis species prosopidis infesting mesquite in Argentina is (Johnson 1983). incomplete. In Texas, P. glandulosa produces flow- At least two species, Rhipibruchus pesephenopygus ers throughout the year, so A. prosopidis does not Kingsolver and R. atratus Kingsolver (Fig. 7) were require an alternate host. However, mesquite found to lay eggs on both green and ripe pods. plants in Argentina do not produce flowers and Dissection of young green pods revealed the pods during the winter months (June–August). It presence of burrowing larvae reaching the imma- seems unlikely that the fragile Asphondylia adults ture seeds (Fig. 8). In laboratory conditions, newly would be able to survive the winter months and emerged adults readily laid eggs on both imma- there are no examples of such long-lived adults in ture and mature pods. On the mature pods, white, the Cecidomyiidae. Also, some Japanese Asphondylia ovoid eggs (0.58 ± 0.02 mm long; 0.03 ± 0.02 mm species are known to have an occasional, facultative width) (mean ± S.D.; n = 20) were randomly laid Mc Kay & Gandolfo: Phytophagous insects associated with reproductive structures of mesquite 129

Figs 7–8. Rhipibruchus atratus. 7, Adult; 8, a dissected young green pod of Prosopis nigra showing an egg (a), larval tunnel (b), larva (c) and immature seed (d). on the pod surface, while on the immature pods, insects. Therefore the survey of natural enemies of eggs were preferentially cemented on the pod rim. Prosopis species in Argentina has been restricted to This pattern was also observed in the field. insects attacking reproductive structures, paying Infestation levels of these two species were special attention to seed-feeding insects. Among estimated on immature pods of Prosopis flexuosa in these, a new species, C. gandolfoi, was selected for northwestern Argentina. From a sample of 400 further studies. Based on the results presented pods taken from 100 trees (four pods per tree), 87 above, a motivation to release C. gandolfoi in South trees were infested, and 52 % (208 of 400) of the Africa is expected to be submitted to the regulatory pods had at least one egg. authorities in 2007. The available information strongly suggests that There are several additional agents that could be these beetles are specific to the genus Prosopis and considered for introduction into South Africa but the fact that R. psephenopygus and R. atratus both these are dependent on the efficacy of C. gandolfoi, lay eggs on immature green pods indicates that once released, and the removal of current restric- they may be more efficient agents than the tions to introduce agents other than seed feeders. bruchids established in South Africa. Although no formal host tests have been conducted Lophopoeum bruchi Monné & Martins (Coleoptera: with the bruchids, R. psephenopygus and R. atratus, Cerambyciidae). Adults of the pod-boring beetle the current information strongly suggests a were collected from ripening pods of Prosopis alba, suitably narrow host range. The bruchid, N. P.nigra and P.chilensis. According to Di Iorio (1995), arizonensis introduced in South Africa was thought L. bruchi also attacks P. affinis, P. caldenia, P. hassleri, to attack immature pods, but studies in South P. hassleri var nigroides, P. kuntzei and Acacia aroma. Africa showed that females strongly prefer mature The larvae feed on the mesocarp and seeds of all pods (Impson et al. 1999). Field observations these species, except P.kuntzei, where the seeds are reported here on the two Rhipibruchus species not attacked. However, the host record on A. aroma, conclusively showed that they attack green pods which has many native relatives in southern African and are probably better prospects than the three countries (Smit 1999), suggests an unacceptably species already introduced. A similar habit was broad host range for this long-horned beetle. reported for the North American bruchid, Mimosestes protractus (Horn), introduced into Prospects for the biological control of quarantine, but not released (Johnson 1983; mesquite in South Africa Moran et al. 1993) as attempts to establish a quaran- To preserve the benefits of Prosopis species, tine culture failed (Impson et al. 1999). biological control of mesquite in South Africa In Australia, where some North American should be limited to the use of seed-feeding Prosopis species have also become serious weeds, 130 African Entomology Vol. 15, No. 1, 2007 biological control programmes include the use of Africa is now available and most farmers do not insects that attack not only seeds but also the vege- regard mesquite plants as valuable (Harding 1987; tative structures. Two bruchids, A. prosopis and Impson et al. 1999). In such a situation, Evippe sp. A. bottimeri, previously introduced into South no. 1 seems an ideal candidate as only minor addi- Africa were released in Queensland and Western tional host tests will be needed. The flower feeders Australia between 1996 and 1997 (van Klinken & reported here, Polyhymno sp. and Asphondylia sp. Campbell 2001). However, their impact is likely to nr. prosopidis, also hold great potential, although be limited by herbivores which consume the pods they might substantially reduce pod production, before the bruchids have the chance to damage affecting the use of mesquite plants as fodder. the seeds (van Klinken & Campbell 2001). A renewed biological control programme against ACKNOWLEDGEMENTS mesquite, initiated in 1994, was re-directed to surveys of insects which targeted foliage and We thank H.A. Cordo for sharing his expertise reproductive structures (buds, flowers and pods) and knowledge on Prosopis species in Argentina, prior to their consumption by herbivores (van S. Neser (ARC-PPRI) for sharing his vast experi- Klinken & Campbell 2001). Two new agents from ence as field collector in some of the exploratory Argentina, the sap-sucking psyllid Prosopidopsylla trips, R.J. Gagné for identification, comments and flava Burckhardt, and the leaf-tying gelechiid observations made on Cecidomyiidae, F. Navarro moth, Evippe sp. no. 1, have subsequently been (Instituto Superior de Entomología ‘Dr. Abraham released since 1998. The latter is widely established Willink’, Argentina), O. Di Iorio (Universidad de and having a significant impact in the warmest Buenos Aires) for the identification of Lepidoptera regions of Western Australia (van Klinken et al. and Cerambyciidae, respectively, D.G. Kissinger 2003). for the identification and description of Coelo- It seems likely that present restrictions could be cephalapion gandolfoi and M.P.Hill (Rhodes Univer- removed in the near future as more information sity, South Africa) for comments on the manu- on the detrimental impact of mesquite in South script.

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Accepted 6 February 2007