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Diptera: Cecidomyiidae: Asphondyliini) from Palaearctic and Oriental Regions, with Descriptions of Two New Species That Induce Stem Galls on Lauraceae in Japan

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Diptera: Cecidomyiidae: Asphondyliini) from Palaearctic and Oriental Regions, with Descriptions of Two New Species That Induce Stem Galls on Lauraceae in Japan SYSTEMATICS First Records of Genus Bruggmanniella (Diptera: Cecidomyiidae: Asphondyliini) from Palaearctic and Oriental Regions, with Descriptions of Two New Species That Induce Stem Galls on Lauraceae in Japan 1, 2 3, 4 MAKOTO TOKUDA AND JUNICHI YUKAWA Downloaded from https://academic.oup.com/aesa/article/99/4/629/62315 by guest on 01 October 2021 Ann. Entomol. Soc. Am. 99(4): 629Ð637 (2006) ABSTRACT Two new species of the genus Bruggmanniella, Bruggmanniella actinodaphnes and Bruggmanniella cinnamomi (Diptera: Cecidomyiidae: Asphondyliini: Asphondyliina) are described from Japan. The former species induces stem galls on Actinodaphne lancifolia (Siebold et Zucc.) (Lauraceae) in the Palaearctic Region, and the latter species induces stem galls on Cinnamomum japonicum (Lauraceae) in the Oriental Region. Bruggmanniella, previously known to occur only in the Neotropical and southern Nearctic regions, is recorded for the Þrst time from the Palaearctic and Oriental regions. The number of genera, their components, and morphological features of Asphon- dyliina are compared between different zoogeographical regions. KEY WORDS Asphondyliini, Bruggmanniella, Cecidomyiidae, gall midge, Lauraceae THE TRIBE ASPHONDYLIINI IS a well circumscribed mono- Many species of the subtribe Asphondyliina have phyletic group sharing unique characteristics of the been known to induce various galls on broad-leaved adult postabdomen and is divided into two subtribes, evergreen trees of Lauraceae in the Oriental and Asphondyliina and Schizomyiina (Gagne´ 1994, 2004). eastern Palaearctic regions, such as India (Kieffer The tribe is a taxonomically and phylogenetically 1905), southwestern Japan (through which the well studied group compared with other tribes of boundary of the Palaearctic and Oriental regions ex- Cecidomyiidae (Mo¨hn 1961, Gagne´ 1994), but many tends; Fig. 1) (Yukawa 1974, Yukawa and Masuda species are still unnamed in various parts of the world. 1996), Taiwan (Yang et al. 1999), and Indonesia and In addition, Asphondyliini exhibit various unique eco- Thailand (Yukawa et al. 2005). Accordingly, we have logical and life history traits, such as host alternation been paying special attention to Lauraceae trees in our (Harris 1975; Orphanides 1975; Yukawa et al. 2003; search for galling species of Asphondyliina at various Uechi et al. 2004, 2005), polyphagy (Gagne´ and Woods localities in Japan. In recent Þeld surveys, we have 1988, Tokuda et al. 2005), prolonged diapause (Maeda collected several galls of Asphondyliina on Lauraceae et al. 1982, Takasu and Yukawa 1984, Tabuchi and from Japan. In this article, we describe two new spe- Amano 2003), and association with fungal symbionts cies of the genus Bruggmanniella that induce stem galls (Meyer 1987, Bissett and Borkent 1988, Yukawa and on trees of Lauraceae. We compare generic compo- Rohfritsch 2005). Therefore, further taxonomic, phy- nents and morphological features of Asphondyliina, logenetic, and ecological studies of Asphondyliini are including Bruggmanniella, between the Neotropical required to elucidate the evolution of these traits, in and Palaearctic regions. particular the processes of host range expansion, in- cluding host plant shift and galled organ shift (Tokuda and Yukawa 2005). Materials and Methods Collection of Galls and Gall Midges. Some of the collected galls were dissected under a stereoscopic 1 Entomological Laboratory, Graduate School of Bioresources and microscope to obtain larval and pupal specimens. Bioenvironmental Sciences, Kyushu University, Fukuoka 812-8581, When some of the dissected galls contained mature Japan. larvae or pupae, the rest of the collected galls were 2 Corresponding author, address: Institute for Biological Resources and Functions, National Institute of Advanced Industrial Science and maintained in plastic bags (350 by 250 mm) to obtain Technology, Tsukuba, Ibaraki 305-8566, Japan (e-mail: tokuda- adults and pupal exuviae. All specimens collected in [email protected]). this study were preserved in 75% ethanol or 99.5% 3 Entomological Laboratory, Faculty of Agriculture, Kyushu Uni- acetone. versity, Fukuoka 812-8581, Japan. 4 Current address: Matsuzaki 1-5-12, Higashi-ku, Fukuoka 813-0035, Preservation of Specimens Examined. All speci- Japan. mens examined, including holotypes and paratypes, 0013-8746/06/0629Ð0637$04.00/0 ᭧ 2006 Entomological Society of America 630 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 99, no. 4 Table 1. B. actinodaphnes sp. n.: frontoclypeal and thoracic (for each sex 7 ؍ setal counts (n Male Female Mean SD Range Mean SD Range Frontoclypeal setae 25.0 2.94 21Ð28 21.3 4.72 18Ð28 ADL setaea 73.0 19.76 61Ð108 69.6 5.13 65Ð76 PDL setaeb 96.2 12.03 83Ð112 91.8 17.63 81Ð118 Mesopleural setae 56.0 8.00 45Ð65 55.0 12.37 46Ð76 Mesepimeral setae 57.0 4.98 50Ð63 60.4 7.16 52Ð71 a Anterior dorsolateral setae. b Posterior dorsolateral setae. Downloaded from https://academic.oup.com/aesa/article/99/4/629/62315 by guest on 01 October 2021 Hemibruggmanniella Mo¨hn, 1961: 6. Type species Bruggmanniella oblita Tavares 1920 The genus Bruggmanniella belongs to the subtribe Asphondyliina and comprises six known species in the world (Gagne´ 2004, Gagne´ et al. 2004). Among them, Þve species occur in the Neotropical Region and one in the southern parts of the Nearctic Region. Because Fig. 1. Collection records of B. actinodaphnes and the morphological features of Bruggmanniella were B. cinnamomi based on Yukawa (1976), Usuba (1977), Yama- summarized and discussed in Gagne´ (1994) and uchi et al. (1982), Yukawa (1988), Yukawa and Masuda Gagne´ et al. (2004), we refer in this article only to (1996), and the present results. The broken line illustrates morphological differences between Bruggmanniella the boundary between Oriental and Palaearctic zoogeo- and allied genera in the Palaearctic Region as fol- graphical regions. lows: Bruggmanniella is similar to Pseudasphondylia Monzen, 1955 in most male morphological features are preserved in the collection of the Entomological such as the presence of parameres and the two sep- Laboratory, Kyushu University, Fukuoka, Japan. arate teeth of the gonostylus. However, Bruggman- Morphological Studies and Terminology. For mi- niella can be distinguished from Pseudasphondylia by croscopic study, some of the ethanol-preserved spec- the presence of setae on the larval cervical papillae imens were mounted on slides in Canada balsam by and on all papillae of the ventral surface (Tokuda and using the techniques outlined in Gagne´ (1989). Draw- Yukawa 2005). Bruggmanniella, with its two separate ings were made with the aid of a drawing tube. Some teeth on the gonostylus, is distinguishable from Pro- important structures of the pupae were examined with bruggmanniella Mo¨hn, 1961, which has a single tooth. a scanning electron microscope (S-3000N, Hitachi, It differs from Illiciomyia Tokuda, 2004 by having Tokyo, Japan) by using the acetone-preserved spec- several morphological features such as less constricted imens. male ßagellomeres, convolute male circumÞla, pres- Adult morphological terminology, except for that of ence of an apical spur on the Þrst tarsomeres, and thoracic plates, follows use in McAlpine (1981) and absence of asetose lateral papillae on larval thoracic that of thoracic plates follows use in Tokuda (2004) segments (Tokuda 2004). and Tokuda et al. (2004b). Counts of setae and scales were based on the setal and scale insertions because Bruggmanniella actinodaphnes many setae and scales become lost through the pro- Tokuda & Yukawa sp. n. cesses of collection, preservation, and preparation. (Figs. 2A and B, 3A and B; Tables 1 and 2) Morphological terminology of the immature stages follows use in Mo¨ hn (1955, 1961), which was orig- Male. Eye bridge four to six facets long. Frontocly- inally written in German and later translated into peal setal count as in Table 1. Palpus two-segmented; English in Yukawa (1971), but terminology of the Þrst palpal segment Ϸ38 ␮m long, Ϸ1.3 times as long pupal antennal horn follows that in Gagne´ (1994). as basal width; second Ϸ2.0 times as long as Þrst. In addition, the term “anterior dorsal papillae” (To- Antenna with 12 ßagellomeres; scape and pedicel with kuda et al. 2004a,b) is applied to pupal abdominal rather dense setae; Þrst and second ßagellomeres not papillae that are situated in the anterior fourth of fused; Þrst ßagellomere Ϸ215 ␮m long, 4.3 times as the dorsal surface and distinctly anterior to the row long as wide, 1.2 times as long as second, Þfth ßag- of “dorsal papillae.” ellomere Ϸ160 ␮m long, 3.3 times as long as wide. Thoracic setal counts as in Table 1. All legs covered with many blackish scales; length of respective seg- Taxonomy ments as in Table 2; Þrst tarsomeres of all legs each Genus Bruggmanniella Tavares with a short apical spur; claw simple on all legs, bent Bruggmanniella Tavares, 1909: 19. Type species: Brugg- nearly at right angle; empodium well developed, as manniella braziliensis Tavares 1909 long as claw; pulvillus much shorter than empodium. July 2006 TOKUDA AND YUKAWA: Bruggmanniella,NEW TO PALAEARCTIC/ORIENTAL REGIONS 631 (for each sex 7 ؍ Table 2. B. actinodaphnes sp. n.: measurements of legs (micrometers, n Male Female Mean SD Range Mean SD Range Foreleg Femur 1,272 53.8 1,197Ð1,344 1,118 91.2 985Ð1,241 Tibia 1,154 94.1 1,083Ð1,348 1,016 113.5 837Ð1,154 Tarsomere I 174 13.7 154Ð192 178 30.4 141Ð210 Tarsomere II 970 157.8 763Ð1123 889 156.8 641Ð1,053 Tarsomere III 441 25.6 415Ð475 366 46.4 314Ð410 Tarsomere IV 320 40.6
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