Bruggmanniella cinnamomi Tokuda & Yukawa, 2006

Lin, Sheng-Feng, Yang, Man-Miao & Tokuda, Makoto, 2020, Molecular Phylogeny Revealing the Single Origin of - associated (Diptera: Cecidomyiidae) in Asia, with Descriptions of Three New and One Newly Recorded Species from Taiwan., Zoological Studies 59 (66), pp. 1-16 : 9-13

Bruggmanniella cinnamomi Tokuda & Yukawa ( Fig. 7 ; Table 5)

Description: See Tokuda and Yukawa (2006). Number of palpal segment is four in both sex ( Fig. 7A ) that was misidentified in Tokuda and Yukawa (2006). Additional descriptions are as follows: female tarsal claws thicker than male ( Fig. 7B–C ); Wing 2.6–2.9 mm long in male (n = 8, Fig. 7D ) and 3.1–3.6 mm long in female (n = 8); Frontoclypeal and thoracic setal counts as in table 5.

Specimen examined: Holotype. Male (on slide, ELKU), JAPAN: Nakagusuku , Okinawa, adult reared by M. Tokuda and emerged on 15.iii.2001 from stem galls that collected on 25.ii.2001, leg. J. Yukawa, S. Yamauchi. Paratypes. JAPAN: (Cecid. Nos. C7101– 7114; C7151–C7164; see Tokuda and Yukawa (2006) for detailed information).

Other specimens: TAIWAN: 8ò, 8ñ, 2 pupal exuviae (on slide, NCHU), Pingtung Co., Dahan forest road, adult emerged on 25–27.iv.2016 from galls that collected on 19.v.2016, S.F. Lin leg.; 4 pupal exuviae (on slide, NCHU) Pingtung Co., Dahan forest road, 28.iii.2014, S.F. Lin leg.; 5 larvae (on slide, NCHU) Nantou Co., Gaofeng Ln, 19.iv.2016, S.F. Lin leg.

Host and Distribution: The host species is C. yabunikkei (= C. japonica ) in south-western islands of Japan ( Tokuda and Yukawa 2006) and C. insularimontanum in Taiwan (new host and distribution records).

Gall: Swollen multi-chambered stem gall, which is 1 to 12 cm long and 1 cm wide.

Life history: Adults emerge in mid-March and early April in Japan ( Tokuda and Yukawa 2006) and in early May in Taiwan. Further details of the life history remain unclear.

Taxonomic remarks

As mentioned earlier, B. sanlianensis sp. nov. possesses a bidentate apical tooth of gonostylus, which does not fit the previous definition of the genus Bruggmanniella . However, other morphological features and current molecular phylogenetic works (see below) support close relationships with other Cinnamomum - associated Bruggmanniella species. Therefore, B. sanlianensis sp. nov. should be treated as a member of Bruggmanniella ( Fig. 8 ).

The two other new species, B. turoguei sp. nov. and B. shianguei sp. nov., both possess finger-like pupal antennal horns ( Fig. 6A–B ) similar to those of B. cinnamomi . Within these three species, the gonostylus tooth is well separated in B. shianguei sp. nov. ( Fig. 5B ), but closely situated in B. turoguei sp. nov. ( Fig. 5A ) and B. cinnamomi . Bruggmanniella turoguei sp. nov. and B. cinnamomi are distinguishable by the shape of the hypoproct, which is deeply emarginated in the former but only slightly in the latter. In addition, pupa and larva differ in size among them: pupae of B. turoguei sp. nov., B. shianguei sp. nov. and B. cinnamomi are 3.2–3.4, 1.8–2.4 and 3.4–4.3 mm long, respectively, and larvae of B. shianguei sp. nov. and B. cinnamomi are 1.6–1.8 mm and 2.7–3.5 mm ( Tokuda and Yukawa 2006), respectively.

In this paper, we re-combine two species, B. cinnamomi comb. rev. and B. actinodaphnes comb. rev., with Bruggmanniella because of morphological and molecular similarites ( Fig. 8 ). Although these two species were transferred to Pseudasphondylia based on an unstable phylogenetic analysis in Garcia et al. (2020), they are definitely the relatives of Lauraceae-associated Asian Bruggmanniella ( Tokuda and Yukawa 2005 2006). We recognize that further studies are needed to determine the relationships between Asian and Neotropical (plus Nearctic) Bruggmanniella , and the species might be divided into several genera in future. However, we believe that more comprehensive studies are needed to conclusively determine the detailed relationship and, at least for the present, we prefer to retain all of them as members of Bruggmanniella .

Molecular phylogeny and genetic distances

The phylogenetic trees created from the three methods had similar topologies. Asian Bruggmanniella and genus Pseudasphondylia were monophyletic in the NJ ( Fig. 8 ), ML and BI trees, with more than 50% bootstrap support in the ML and NJ trees and 0.5 support in the BI topology. Among Bruggmanniella species, the basal lineage is a Litsea -associated species, B. actinodaphnes , which is a sister group to Neolitsea - associated species B. brevipes plus Cinnamomum - associated species. Within Cinnamomum -associated taxa, three stem gallers belong to a single clade whose monophyly is highly supported by bootstrap values (99%, 94% and 1 in NJ, ML and BI trees, respectively). The leaf gall midge, B. sanlianensis sp. nov., is situated at the basal part of stem gallers’ clade. Among stem gallers, B. shianguei sp. nov. occupies the basal position and received high supported value in three phylogenetic methods. However, the relationship between B. turoguei sp. nov. and B. cinnamomi was discordant in three phylogenetic methods. The monophyly of B. cinnamomi was supported in both NJ and ML trees, but undetermined in the BI tree due to an unsolved relationship with B. turoguei sp. nov. In B. cinnamomi, Taiwanese taxa are monophyletic and sister to the Japanese taxa based on the ML method, but become a paraphyletic group with respect to Japanese taxa in the NJ tree ( Fig. 8 ).

The genetic distances within species are 0.4%, 1.9%, 1.9% and 2.0% in B. cinnamomi , B. turoguei sp. nov., B. shianguei sp. nov. and B. brevipes , respectively. At the interspecific level within Cinnamomum - associated Bruggmanniella , the genetic distance of most species pairs do no exceed 7.0%. The lowest genetic distance is between B. cinnamomi (Taiwanese population) and B. turoguei sp. nov. (1.8%) ( Table 6). At the interspecific level among B. actinodaphnes , B. brevipes and Cinnamomum -associated species, the genetic distance are 12.8–16.4%.

Figures

Fig. 7. Bruggmanniella cinnamomi (A) Male head (ventral view). (B) Male 5th tarsomere. (C) Female 5th tarsomere. (D) Male wing. Scale bars:A = 0.03 mm; B–C = 0.1 mm; D = 1 mm.

Fig. 8. Cladogram of East Asian Bruggmanniella based on the Neighbor-joining method based on the DNA COI region. Support values at the nodes are Neighbor-joining (left)/ Maximum-likelihood (media)/ Bayesian inference (right). “–” indicates support values less than 50% in ML and NJ, and 0.5 in BI inference.

Fig. 6. Taiwanese Cinnamomum–associated Bruggmanniella. Bruggmanniella turoguei sp. nov. (A, D), B. shianguei sp. nov. (B, E), B. sanlianensis sp. nov. (C, F). Scale bars: pupal head (A–C) = 0.5 mm and for larval sternal spatula (D–F) = 0.1 mm. Larval lateral papillae shown in dotted blue circles.

Fig. 5. Male genitalia of Taiwanese Cinnamomum–associated Bruggmanniella. (A) Bruggmanniella turoguei sp. nov., (B) B. shianguei sp. nov., and (C) B. sanlianensis sp. nov. Scale bar = 0.1 mm.