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1 This article was downloaded by: [Ingenta Content Distribution - Routledge] On: 8 March 2011 Access details: Access Details: [subscription number ] Publisher Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Journal of Natural History Publication details, including instructions for authors and subscription information: A new species of Josephiella (Hymenoptera: Agaonidae) forming leaf galls on Ficus microcarpa L. (Moraceae) John W. Beardsley a ; Jean-Yves Rasplus a a INRA, Laboratoire de Modélisation et Biologie Evolutive, 488 rue Croix de Lavit, F Montpellier, France, Online publication date: 03 December 2010 To cite this Article Beardsley, John W. and Rasplus, Jean-Yves(2001) 'A new species of Josephiella (Hymenoptera: Agaonidae) forming leaf galls on Ficus microcarpa L. (Moraceae)', Journal of Natural History, 35: 1, To link to this Article: DOI: / URL: PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: This article may be used for research, teaching and private study purposes. Any substantial or systematic reproduction, re-distribution, re-selling, loan or sub-licensing, systematic supply or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

2 Journal of Natural History, 2001, 35, A new species of Josephiella (Hymenoptera: Agaonidae) forming leaf galls on Ficus microcarpa L. (Moraceae) JOHN W. BEARDSLEY1 and JEAN-YVES RASPLUS* INRA, Laboratoire de Mode lisation et Biologie Evolutive, 488 rue Croix de Lavit, F Montpellier, France (Accepted 30 November 1999) A second species of Josephiella Narendran, J. microcarpae, is described and gured, based on specimens reared in Hawaii, California and the Canary Islands. This wasp develops in galls on the leaves of Ficus microcarpa L., a widely cultivated ornamental banyan tree that is native to South-East Asia and adjacent islands. It has not been found on any other hosts. Both sexes are described, and males, which are apparently all fully winged, are relatively rare. We postulate that this wasp is a recent introduction into Hawaii, California and the Canary Islands, from an unknown location in South-East Asia, where Ficus microcarpa is native. Keywords: Agaonidae, Epichrysomallinae, Josephiella, new species, pest, Ficus, Moraceae. Introduction In May 1989, the rst author noticed for the rst time the presence of small, pustule-like galls on the leaves of a common ornamental Ficus tree, F. microcarpa L. ( gure 1) on the campus of the University of Hawaii in Honolulu. Specimens of an Epichrysomallinae agaonid wasp that emerged from galled leaves held in the laboratory were identi ed as possibly a species of Camarothorax by E. Grissell at the US National Museum in Washington, DC (Beardsley, 1992). Specimens were sent also to Z. BoucÏ ek at the Natural History Museum in London. BoucÏ ek (1997: 119) referred brie y to the species stating `...one species of Acophila Ishii ( Epichrysomallinae), found recently in Hawaii, is believed to produce galls on the leaves of a Ficus. In September 1997, the same wasp was discovered forming leaf galls on F. microcarpa at Santa Ana, California by J. N. Nisson (Anon., 1998). In October 1997, the species had spread as far as the Canary Islands (Spain), where it was found causing heavy infestation on foliage of cultivated F. microcarpa in Tenerife 1Present address: 1026 Oak Dale Lane, Arcadia, CA 91006, USA. * To whom correspondence is addressed; rasplus@ensam.inra.fr. Journal of Natural History ISSN print/issn online Ñ 2001 Taylor & Francis Ltd

3 34 J. W. Beardsley and J.-Y. Rasplus Fig. 1. Galls of Josephiella microcarpae sp. n. on foliage of Ficus microcarpa L. (A) Twig with infested leaves. (B) Upper ( left) and lower surfaces of F. microcarpa leaves bearing fresh (unemerged) galls. (C) Enlarged view of J. microcarpae emergence holes on lower surface of old gall cluster. (J. L. Nieves-Aldrey, personal communication). Because F. microcarpa is widely cultivated as an ornamental shade tree throughout the tropics and subtropics of the world, and because the leaf galls of this wasp sometimes produce unsightly deformed foliage and occasionally defoliation, we believe that it will develop into a signi cant pest in many places. Generic placement Narendran (in Narendran and Sheela, 1993) described the genus Josephiella, with a single included species, J. malabarensis Narendran, reared from leaf galls of Ficus bengalensis in India. That wasp apparently was the rst species of Epichrysomallinae described which develops within leaf galls; all previously described species for which hosts are known inhabit g syconia. However another species, Leeuweniella Wcophila FerrieÁ re, 1929 which is an Epichrysomallinae, despite its description in Perilampidae, develops in galls on twigs of Ficus recurva Blanco (FerrieÁ re, 1929 ). The new species possesses most of the diagnostic characters cited by Narendran and de ning Josephiella: Female with subglobose head. Bilobed clypeus. Supraclypeal area well delimited, subquadrate. Antenna inserted above the ocular

4 A new species of Josephiella 35 line. Antennal formula 11(1)53 (the annellus hardly visible in J. microcarpae). Scape short not reaching the median ocellus. Occipital carina absent (in dried specimens of J. microcarpae, the occiput may be partly collapsed and a consequent crease misinterpreted as a carina). Axillar grooves shallow or obliterated. Tarsi foursegmented. Stigmal vein longer than marginal vein. Stigmal vein at an angle of about 90ß. Marginal fringe present. Josephiella clearly diœers from Acophila in having four-segmente d tarsi (vs vesegmented), bilobed clypeus (vs truncate). Antennae inserted above the ocular line (vs inserted below the ocular line). Occipital carina absent (vs clearly visible). Scutellum without distinct punctures (vs with punctures). Marginal vein about as long as the stigmal (vs marginal clearly longer than the stigmal). Stigmal vein at an angle of about 90ß (vs stigmal vein at an angle less than 90ß ). Marginal fringe present (vs absent). The closest genus to Josephiella is Odontofroggatia Ishii. In the key to genera of Nearctic Agaonidae (BoucÏ ek, 1997), couplet 10(8) should be altered as follows to allow identi cation of these genera: 10(8) Stigmal vein at a right angle to wing margin and about as long as marginal vein, the postmarginal vein very short and stub-like; body brown to yellow, shiny; tarsi 4-segmented EPICHRYSOMALLINAE, 10¾ ± Stigmal vein more or less at an acute angle to wing margin and shorter, etc ¾ (10) Female: head transverse. Antennal formula 1163, scape long, reaching the median ocellus. Supraclypeal area not delimited. Scutellum bearing 2Ö 2 setae. Axillar grooves well marked. Male: mandible long and falcate. Petiole apparent, sometimes with a ventral hook or lateral teeth Odontofroggatia Ishii ± Female: head subglobose ( gure 3C ). Antennal area formula 11(1)53 ( gure 2A), the annellus hardly visible in J. microcarpae; scape short, not reaching the median ocellus. Supraclypeal area well delimited, subquadrate ( gure 2B). Scutellum bearing at least 2Ö 3 setae. Axillar grooves shallow or obliterated ( gure 2D). Male: mandible normal. Petiole short Josephiella Narendran In the key to the Oriental genera of Epichrysomallinae (Narendran and Sheela, 1996), the new species described here will run to the genus Josephiella only with di culty. In the rst couplet of their key these authors refer to the relative widths of the anterior margin of the scutellum and posterior margin of the midlobe of the mesoscutum. In the new species described below, the sutures that de ne these sclerites are weakly developed, and not easily seen; they are well de ned in the type species. However, in specimens of the new species in which the axillary sutures are discernible, they intersect the transscutal articulation close together near the midline, as in the type species of Josephiella. The species described here shares another important, apparently apomorphic character with the type species of Josephiella: the unusual habit of developing within leaf galls. Josephiella microcarpae, sp. n. ( gures 2± 4) Female ( Wgure 4B). Length of holotype 2.2 mm. Body uniformly dark brown; antennae and legs, including coxae, mostly pale yellow, femora slightly darkened, pretarsi dark; wings hyaline, veins pale yellow-brown, semi-transparent; mouthparts pale. Body weakly sclerotized, gaster and mesosoma tending to collapse and shrivel

5 36 J. W. Beardsley and J.-Y. Rasplus Fig. 2. Josephiella microcarpae sp. n. details. (A) Female antenna; (B) male antenna; (C) female mandible; (D) female hind tibia; (E) male genitalia. in air-dried specimens. Integumental sculpture consisting of weakly to moderately developed reticulation; head ( gure 3A± C ) reticulate in dorsal occipital area behind ocelli, genal areas and face lateral of antennal scrobes, from clypeal margin upwards, becoming evanescent near upper margin of parascrobal areas. Pronotum with moderately well-developed reticulation, similar to head in pattern; mesoscutum ( gure 3D) smooth centrally with weakly developed semi-diagonally oriented reticulation on lateral lobes; notauli weakly or not at all developed, not reaching posterior margin of scutum; scutellum ( gure 3D) smooth, with weak, longitudinally oriented reticulation; axillary sutures weakly indicated, intersecting transscutal articulation near midline. Propodeum ( gure 3D) weakly reticulate, lacking carinae except on margins

6 A new species of Josephiella 37 Fig. 3. Josephiella microcarpae sp. n. female. (A± C ) Head. (A) Dorsal; ( B) frontal (antennae removed); (C) lateral; (D) dorsum of mesosoma and base of gaster. of anteriorly directed spiracular foveae. Hind coxae ( gure 3D) with dorsal surfaces reticulate. Gaster smooth and shining. Setae of head and mesosoma sparse, distributed as in J. malabarensis, but apparently more elongate, those of face averaging about 45± 55 mm long ( gure 3A± C). Longer setae of mesosoma mostly 60± 75 mm long; sparsely scattered, and in a continuous row (ca 10) on posterior margin, on pronotum; less numerous on mesonotum. Mesonotum with one conspicuous pair of forward directed setae near posterior margin and three or four laterally on each side; axillae each with one such seta; scutellum with two pairs of elongate setae sublaterally, one near posterior margin and one near midlength; a few short inconspicuous short setae (12± 15 mm) present on scutellum and axillae, particularly laterally. Gaster largely bare, with a few ne setae in transverse line on each tergite. Head globular, in dorsal view ( gure 3A) wider than long (ca 7:4), ocelli in broadly obtuse triangle, area between them slightly raised and attened, distance

7 38 J. W. Beardsley and J.-Y. Rasplus Fig. 4. Josephiella microcarpae, sp. n. female. (A) Detail of distal part of venation; ( B) habitus. between posterior ocelli more than two times distance from ocellus to ocular margin, posterior ocellus (po)± anterior ocellus (ao) distance greater than distance between po and ocular margin; in frontal view ( gure 3B) slightly wider than high (7:6), face broad, the interocular distance equal to more than three times width of compound eye; clypeal margin bilobed, with relatively deep, narrow mesal notch. Occipital carina absent, although in dried specimens a crease sometimes present, often apparently nearly encircling foramen magnum below. Antennae inserted close together near middle of face, scrobes shallow ( gure 3B); antenna ( gures 2A, 3C) total length about 570 mm, scape slightly more than two times as long as wide, pedicel constricted basally, shorter than scape, longer than individual funicle segments, funicle with ve segments, plus very small basal anellus, hardly visible in pointmounted specimens, funicle segments subequal in length, becoming progressively slightly broader toward apex, club about as long as last two funicle segments combined, apparently unsegmented, basic three segments indicated by marginal constrictions and sensory ridges. Mandibles identical, with three teeth, lower tooth more strongly developed and somewhat separated from upper teeth ( gure 2C). Mesosoma ( gure 3D) with scutellum broadly rounded behind; propodeum ca four times as wide as long, spiracles at ends of relatively deep, anteriorly curved fovea.

8 A new species of Josephiella 39 Fore wing ( gure 4A, B) hyaline, with venation very similar to J. malabarensis, marginal vein shorter than stigmal, postmarginal ca one-half as long as marginal; with ca six large setae on submarginal vein, two on marginal and one at apex of postmarginal; marginal and postmarginal with several smaller setae; discal setae of wing sparse, as in J. malabarensis, mostly small (ca 6 mm, long) and somewhat spiniform; apical fringe setae short (ca 15± 18 mm long), moderately dense. Legs similar to J. malabarensis; hind tibia ( gure 2D) with a comb of ve slightly curved spines on inner face near apex, a series of ca eight spines and several long setae on outer margin. Gaster globular, dorsally humped and nearly as long as head plus mesosoma combined when distended with ova, nearly sessile, the petiolar segment much broader than long ( gure 3D), hypopygium not reaching apex. Male. Fully winged, similar to female in colour and form, except for slightly smaller average size ( length of allotype 1.1 mm), and details as follows: Antenna ( gure 2B) shorter (ca 480 mm long) with three funicle segments plus very narrow anellus (as in female); club distinctly two-segmented, basic three-segmented condition indicated by marginal constriction and sensory plates. Gaster less strongly developed, relatively at, elongate-oval in dorsal outline, about as long as mesosoma; genitalia ( gure 2E ) well developed, with conspicuous digiti and digital spines. Holotype female and male allotype. Hawaiian Is.: Oahu I.: Honolulu, University of Hawaii, nanoa, July 1994 (J. W. Beardsley) ex leaf galls on Ficus microcarpa. Paratype: Hawaiian Is.: Kauai I.: Alexander Dam, 2 February 1992 (A. Asquith), galls on Ficus leaves. Molokai I.: Kualapuu in coœee eld, 30 September to 14 October 1994, W. D. Perreira coll., blue sticky board trap (1 female); napulehu nr. Ililiopae Heiau, el. 60 ft, J. W. Beardsley and W. D. Perreira colls., sweeping (1 male); 22 July 1994, W. D. Perreira coll., reared ex leaves of Ficus microcarpa (3 females, 2 males); naunaloa, el. ca 1200 ft, 24 June to 8 July 1994, W. D. Perreira and M. Fukuda colls., yellow sticky board trap (2 females). Oahu I.: Honolulu, U. H. Manoa Campus, el ca 60 ft, 30 May 1989, J. W. Beardsley coll., reared ex leaf galls on Ficus microcarpa (30 females); same data as holotype and allotype (1 female, 2 males); Honolulu, U. H. Manoa, 28 June 1989, W. Nagamine coll., ex Ficus retusa ( 5 F. microcarpa) (4 females, USNM ); Honolulu, McCully St. overpass, el. ca 60 ft, 8 October 1994, J. W. Beardsley coll., reared ex leaf galls on Ficus microcarpa (15 females); Konahuanui, el ft, 14 February 1993, W. D. Perreira coll. (2 females). California: Orange County: Santa Ana, 3 October 1997, J. N. Nisson coll., reared from leaf galls on Ficus microcarpa (5 females); Orange, 12 November 1997, J. N. Nisson and B. Sanford colls., reared ex leaf galls on Ficus microcarpa (3 females, 2 males). Canary Islands: Tenerife: 1997, IIE 23716, in galls on Ficus nitida (5 F. microcarpa) (3 females). Holotype and allotype deposited in B. P. Bishop Museum, Honolulu, Hawaii. Paratypes in Bishop Museum and Hawaii Department of Agriculture, Honolulu, in California Department of Food and Agriculture, Sacramento; USNM Washington; National History Museum in London; INRA collection Montpellier. The new species can be easily separated from J. malabarensis Narendran by the very small annellus, hardly visible (vs well developed); the notauli obliterated (vs visible) and the gaster globular, not laterally compressed. All rearings of these wasps in Hawaii yielded predominantly females, and in

9 40 A new species of Josephiella some instances several hundred females were obtained without any males. Many of these specimens were preserved dry or in uid, were not designated as paratypes, and are not listed in the enumeration of specimens studied above. In the collections where some males were reared, these constituted around 2% of the total number obtained. No ightless male forms were ever found, although these were searched for both by visual scrutiny of emerged wasps and by dissecting leaf galls. These nding suggest that this species may reproduce normally by parthenogenesis, but further research is needed to clarify its mode of reproduction. Josephiella microcarpae is believed to have been introduced, apparently accidentally, into Hawaii shortly before its discovery there in 1989, and into California, probably from Hawaii, in We believe that it probably originated somewhere within the native range of Ficus microcarpa, in South-East Asia. Acknowledgements We would like to thank E. E. Grissell, J. M. Heraty, M. Hoddle, J. LaSalle and J. L. Nieves-Aldrey who help us to clarify the status of this wasp, and E. Barrau for preparing the illustration of the habitus. References Anonymous, 1998, Ficus leaf gall wasp, Josephiella sp., California Plant Pest and Disease Report, 16, 46. Beardsley, J. W., 1992, Note on gall-forming Epichrysomalline wasp from Ficus microcarpa leaves, Proceedings of the Hawaiian Entomolical Society, 31, 8. Boucª ek, Z., 1997, Agaonidae, in G. A. P. Gibson, J. T. Huber and J. B. Woolley (eds) Annotated Keys to the Genera of Nearctic Chalcidoidea (Hymenoptera) (Ottawa, NRC Research Press), Chap. 4, pp. 117± 133. Ferrière, C., 1929, Chalcidiens gallicoles de Java, Annales de la SocieÂte Entomologiqu e de France, 98, 143± 161. Narendran, T. C. and Sheela, S., 1993, Description of an interesting new genus and a new species of Epichrysomallinae ( Hymenoptera: Agaonidae) from India, Journal of the Zoological Society of Kerala, 3, 7± 12.