Pseudasphondylia elaeocarpi Tokuda & Yukawa, 2005

Hidaka, Chiho, Yang, Chien-Hui & Wakabayashi, Kaori, 2022, Scyllarides squammosus Nisto, Zoological Studies 61 (39), pp. 1-13 : 7-8

Pseudasphondylia elaeocarpi Tokuda & Yukawa

( Fig. 4 )

Diagnosis: Adult palpus three-segmented, first to sixth tergites with a single row of posterior setae, hypoproct basally rather wide and extremely narrowing at apex. Pupal antennal horn triangular, well developed ( Fig. 4A ). Pupal stigmatal tubercles present on second to fourth abdominal segments ( Fig. 4B ), reduced on fifth to eighth abdominal segments. Larval sternal spatula with four lobes and the inner pairs longer than the outer ( Fig. 4C ).

Material examined: Type materials: JAPAN: ò (on slide, Holotype, ELKU), Shiroyama, Kagoshima, Kyushu, galls collected 1.v.1976 and adult emerged on 2.v.1976., leg. J Yukawa; 10ò, 6ñ, 7 Larvae (on slides, Paratypes, ELKU), same data as holotype; 3ò (on slides), Shiroyama, Kagoshima, Kyushu, 15.iv.1971, J Yukawa; 2ñ (on slides) Shiroyama, Kagoshima, Kyushu, galls collected on 22.iv.1974 and adults emerged 26.iv.1974, leg. K Takahashi; 2ò, 5ñ, 2 Pupae (on slides) Kekura, Kagoshima, Kyushu, galls collected on 29.iv.1977 and adults emerged 4–11.v.1977, leg. T Sunose; 1ò, 1ñ, 2 Pupae (on slides), Yona, Kunigami, Okinawa Island, 24.ii.2001, leg. M Tokuda and J Yukawa.

Other materials examined: TAIWAN: [Taichung City] 2ò, 2ñ, 6 Pupae (on slides, NCHU), Mt. Tung-mao, galls collected on 25.iv.2017 and adults emerged on 3.v.2017, leg. SF Lin; 2ò, 7ñ (in EtOH, NCHU), Mt. Tung-mao, galls collected on 25.iv.2017 and adults emerged on 8–11.v.2017, SF Lin; [Pingtung Co.] 10ò, 7ñ, (in EtOH, NCHU), Dahan forest rd., galls collected on 4.v.2017 and adults emerged on 8–25.v.2017, leg. YM Chao and SF Lin.

Distribution: JAPAN: Honshu, Kyushu, Tanegashima, Nakanoshima, Amamiohshima, and Okinawajima ( Tokuda and Yukawa 2005; Yukawa et al. 2013; Tokuda 2018); and TAIWAN (New distribution records, see Table 3).

Genetic distance and species delimitation

Sequences of five P. kiwiphila sp. nov. (one Japanese and four Taiwanese individuals) and two Taiwanese P. elaeocarpi individuals were obtained for a partial COI gene (658–689 bp). Species delimitation of ABGD showed three presumptive species ( Fig. 5 ) corresponding to our morphological examinations: (1) P. matatabi : one Japanese sequence; (2) P. elaeocarpi : one Japanese and two Taiwanese sequences; and (3) P. kiwiphila sp. nov.: one Japanese and four Taiwanese sequences.

The genetic distance of P. elaeocarpi is 0.1% within Taiwanese individuals and 2.7% between Taiwanese and Japanese populations. As to Actinidia - associated species, the genetic distance is 0–2.6% among P. kiwiphila sp. nov. and 5.1–6.7% between P. kiwiphila sp. nov. and Japanese P. matatabi .

Phylogeny and divergence time

The phylogenetic trees constructed by Bayesian inference ( Fig. 5A ) and by Maximum likelihood shared a similar topology except for the phylogenetic relationship among individuals of P. kiwiphila sp. nov., which was resolved in the IQ tree ( Fig. 5B ) but not the BI tree. Leaf gallers of Pseudasphondylia , P. elaeocarpi , and P. saohimea , situated at the stem part of the tree and formed a paraphyletic group, whereas Pseudasphondylia species including all flower-bud and fruit gallers formed a monophyletic clade (PP: 0.51 and ML: 40%). This clade is further divided into Actinidia - associated species, P. kiwiphila sp. nov. and P. matatabi , (PP: 1 and ML: 97%) and Japanese species on other hosts (PP: 0.62 and ML: 95%).

In Actinidia -associated species, the divergence time for the two species was estimated to be 2.2–2.9 mya (COI genetic distance: 5.1–6.7%). At the interspecific level, Taiwanese individuals of P. kiwiphila sp. nov. were paraphyletic to the Japanese individual. Taiwanese taxa of P. elaeocarpi was reconfirmed as a sister group of Japanese P. elaeocarpi (PP: 1 and ML: 100%), and their divergence time was estimated to be around 1.2 mya (COI genetic distance: 2.7%).

Figures

Fig. 4. Pseudasphondylia elaeocarpi (A) Pupal face and antennal horns. (B) Pupal abdomen, arrows indicate stigmatal tubercles. (C) Larval sternal spatula. Scale bars: A and C = 0.3 mm; B = 0.5 mm.

Fig. 5. Phylogenetic tree of Pseudasphondylia species based on DNA partial COI region. (A) Bayesian tree of Pseudasphondylia species, supported values are shown beside nodes with the Bayesian posterior probability (lift) and ultrafast bootstrap of the IQ tree (right). Taxa with different colors refer to distinct species based on summarized results of species delimitation (Automatic Barcode Gap Discovery analysis). (B) Cladogram of Pseudasphondylia kiwiphila sp. nov. lineage in IQ tree.