Corresponding author: Makoto Kato (kato@zoo.zool.kyoto-u.ac.jp)
Academic editor: Owen Lonsdale
Kato M, Yamamori L, Imada Y (2022) Diversity underfoot of agromyzids (Agromyzidae, Diptera) mining thalli of liverworts and hornworts. ZooKeys 1133: 1–164.
In vascular plants, a leaf represents a flat lamina borne on a shoot with abaxial-adaxial polarity, and the adaxial side is oriented towards the sun for the primary function of photosynthesis. The leaves are highly nutritious organs and thus are often consumed by various arthropods with various feeding strategies. Some insects consume specific layers of foliage while dwelling inside the internal plant tissues; such a means of herbivory is known as leaf mining (
The bryophytes (non-vascular plants) consist of three major clades: hornworts, liverworts, and mosses (
The
We have conducted an extensive taxonomic and ecological assessment of phytophagous insects on bryophytes in the Japanese archipelago. We discovered an immense diversity of agromyzid fauna on thalloid liverworts and hornworts. The morphology of the imagoes of the collected agromyzid flies is consistent with that of
In total, 128 thalloid bryophyte species have been recorded in the Japanese Archipelago: 111 thalloid liverwort species (2 classes, 5 orders, 19 families, 31 genera) and 17 hornwort species (1 class, 3 orders, 3 families, 6 genera) (Table
Thalloid liverwort and hornwort genera, with number of species of each genus and phytophagous insect species associated with the genera in Japan.
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1 | 0 | |||
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5 | 2 | |||||
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1 | 1 | |||||||
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22 | 1 | |||||||
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1 | 1 | ||||||
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10 | 2 | |||||||
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3 | 2 | 5 | |||||
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1 | 1 | ||||||
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1 | 1 | ||||||
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3 | 1 | ||||||
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1 | 0 | ||||||
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2 | 0 | |||||||
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4 | 1 | |||||||
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1 | 1 | ||||
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1 | 1 | |||||||
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6 | 6 | 5 | ||||
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1 | 1 | |||||||
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1 | 1 | 2 | |||||
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2 | 2 | 3 | |||||
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7 | 3 | 2 | ||||||
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4 | 1 | |||||||
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1 | 1 | 4 | 6 | |||||
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1 | 1 | ||||||
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2 | 0 | |||||||
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3 | 0 | |||||||
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1 | 1 | 1 | 1 | ||||
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1 | 1 | ||||||
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1 | 0 | ||||||
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5 | 5 | 19 | 4 | 15 | |||
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18 | 8 | 3 | ||||
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1 | 1 | |||||||
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1 | 0 | ||||||
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5 | 1 | 1 | ||
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2 | 1 | 2 | ||||||
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3 | 2 | 1 | ||||
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4 | 1 | 1 | ||||||
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2 | 1 | |||||
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1 | 1 | 1 | ||||||
Total number of species | 130 | 54 | 19 | 14 | 43 (39)* |
*Because two agromyzid species are associated with plural bryophyte genera, total number of agromyzid species is 39.
At nearly 120 sites in Japan, we extensively sampled mined thalli of thalloid liverworts and hornworts during the 2000s. Collected thalli were placed in plastic cases (13.6 × 8.7 × 2.5 or 13.6 × 8.7 × 3.5 cm, “clean-cup NK”, Risu-pack, Gifu Plastic Industry Co.) and kept in incubators with temperature and light conditions maintained similar to their natural habitats. The thalli were prevented from desiccation by modestly spraying with water during incubation. We checked the plastic cases frequently for emergence of adult flies. For each emerged fly specimen, the date of thallus collection (as larva or puparium) and the date of adult emergence are recorded. Additional searches for agromyzid flies walking on thalli were also performed. Emerged and collected flies were pinned with minute pins and freeze-dried in the ice boxes of the refrigerators, dried, or fixed in 99% ethanol.
The morphology of adult agromyzid fly specimens was examined under a microscope (VHS-7000; Keyence); it was photographed by synthesizing virtual images from a sequence of corresponding depth images. The morphological characters that were important to discriminate species were color of antenna, frons, and maxillary palpus; color and pattern of scutum and scutellum; and color of haltere.
For observation of genitalia, the abdomens of fly specimens were dipped in 10% KOH for 1 day, then rinsed off with water, and dissected under a binocular microscope. Male and female genitalia of prepared specimens were photographed under a microscope (VHS-7000; Keyence) by synthesizing virtual images from a sequence of corresponding depth images. The genital morphological characters that were important to discriminate species were number and position of tubercle-like setae on male epandrium, along with the shape and sclerotization pattern of hypophallus, mesophallus, and distiphallus.
Terminology follows that outlined in
The type specimens (holotypes and paratypes) and other materials are deposited in the
With rearing of phytophagous insects on 47 thalloid liverwort and 7 hornwort species (Table
Agromyzid species associated with thalloid liverworts and hornworts in Japan, with their host plants and distributions.
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1 |
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1 | 1 | 1 | |||||||
2 |
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1 | 1 | 1 | 1 | ||||||||
3 |
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1 | 1 | 1 | |||||||||
4 |
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1 | 3 | 1 | 1 | 1 | 1 | |||||||
5 |
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1 | 3 | 1 | 1 | 1 | 1 | 1 | 1 | |||||
6 |
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1 | 1 | 1 | 1 | 1 | 1 | 1 | ||||
7 |
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1 | 1 | 1 | 1 | 1 | |||||||
8 |
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1 | 2 | 1 | |||||||||
9 |
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2 | 2 | 1 | |||||||||||
10 |
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1 | 1 | 1 | |||||||||
11 |
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1 | 1 | 1 | |||||||||
12 |
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1 | 1 | 1 | 1 | 1 | |||||||
13 |
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1 | 1 | 1 | 1 | 1 | 1 | ||||||
14 |
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1 | 1 | 1 | 1 | ||||||||
15 |
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1 | 1 | 1 | |||||||||
16 |
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1 | 1 | 1 | 1 | ||||||||
17 |
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1 | 1 | 1 | 1 | 1 | |||||||
18 |
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1 | 1 | 1 | ||||||||
19 |
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1 | 2 | 1 | 1 | 1 | |||||||
20 |
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1 | 3 | 1 | 1 | 1 | ||||||||
21 |
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1 | 2 | 1 | ||||||||||
22 |
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1 | 1 | 1 | 1 | 1 | |||||||
23 |
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1 | 1 | 1 | 1 | 1 | 1 | 1 | |||||
24 |
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1 | 2 | 1 | 1 | |||||||||
25 |
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1 | 3 | 1 | ||||||||||
26 |
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1 | 1 | 1 | 1 | 1 | |||||||
27 |
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1 | 3 | 1 | 1 | |||||||||
28 |
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1 | 2 | 1 | ||||||||||
29 |
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1 | 2 | 1 | 1 | 1 | 1 | |||||||
30 |
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1 | 2 | 1 | 1 | |||||||||
31 |
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1 | 1 | 1 | 1 | 1 | 1 | ||||||
32 |
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1 | 4 | 1 | 1 | 1 | ||||||||
33 |
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1 | 1 | 1 | 1 | ||||||||
34 |
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1 | 7 | 1 | 1 | 1 | 1 | ||||||
35 |
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1 | 3 | 1 | ||||||||||
36 |
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1 | 2 | 1 | ||||||||||
37 |
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1 | 1 | 1 | |||||||
38 |
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1 | 1 | 1 | 1 | 1 | ||||||
39 |
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3 | 3 | 1 | 1 | 1 | 1 | 1 | 1 | ||||||
Total number of species | 11 | 32 | 18 | 17 | 5 | 6 | 4 | 4 |
1 HK, Hokkaido; HN, Honshu; SK, Shikoku; KS, Kyushu; YK, Yakushima Is.; Am, Amami Is.; OK, Okinawa Is.; YY, Yaeyama Isls.
All these species are thallus-miners; pupation takes place within their mines, except for some species mining with small thin thalli, which pupate in soil outside of the thalli. Mines of most species are linear, at least in young instars; they often enter the midrib or thicker parts of thalli in the last instar. In some species, mines are obscure from the outside. Adult fly emergence of most species occurs once in spring, while at least a few species are multivoltine.
All these flies had the following common morphological characteristics: costa extending to vein M1; orbital setulae minute and erect or proclinate; male epandrium each side with a row of fused long tubercle-like setae and/or one or a few well-developed tubercle-like setae (rarely lacking both); distiphallus comprising a pair of unfused sclerotized tubules; female tergite 10 trifurcate or cruciform, cercus with two stout, apical, trichoid sensilla. These characteristics suggest that all the 39 species belong to the genus
The morphological characteristics of the species associated with bryophytes coincided with the characteristics unique to the genus
All the species associated with bryophytes are distinguished by color of antenna, color of maxillary palpus, color pattern of scutum and morphology of male genitalia. Colors of 1st flagellomere of antenna and maxillary palpus are dark or yellow, and unique to each species. Scutum was yellow or dark, and the yellow scutum often had three pairs of dark longitudinal stripes: medial stripes on anterior 2/3 (always merged together except in one species), intra-alar stripes (inner lateral stripes) and supra-alar stripes (outer lateral stripes) on anterior 4/5. In some species, the intra-alar stripe and the supra-alar stripe are merged together to shape a wide band, and often merged with the medial stripe. The color pattern of scutum is unique to each species at least among the species sharing the same host bryophyte genus. Although some species are very similar in external morphology, they could be clearly distinguished by the number of tubercle-like setae in a comb on the male epandrium and other male genital morphological characters.
The 39 recorded
A synopsis of morphological characteristics of Japanese bryophyte-associated
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dark | dark | 1.2–1.4 | 2–3 | dark | dark | dark | 0-0-0 | dark | lobate | 6 | 0 | 6 | tapering out | short |
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yellow | yellow | 1.1–1.3 | 3–4 | dark | partly yellow | yellow | 0-0-0 | yellow | lobate | - | 0 | 0 | tapering out | long |
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dark | yellow | 1.2–1.5 | 0 | dark | dark | yellow | 0-0-0 | dark | lobate | - | 0 | 0 | tapering out | long | ||
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dark | yellow | 1.1–1.3 | 3–4 | dark | dark | dark | 0-0-0 | dark | elongated | - | 0 | 0 | tapering out | short |
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dark | dark | 1.0–1.3 | 0–2 | dark | dark | yellow | 1-1-1 | dark | elongated | (2) | 0 | 2 | tapering out | short |
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dark | yellow | 1.2–1.5 | 1–3 | dark | dark | dark | 0-0-0 | dark | elongated | (40) | 0 | 0 | tapering out | short |
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dark | yellow | 1.4–1.7 | 0 | dark | dark | dark | 0-0-0 | dark | elongated | (3) | 0 | 1 | tapering out | short |
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dark | dark | 1.6–1.7 | 6–7 | dark | yellow | dark | 0-0-0 | dark | lobate | 4 | 2 | 1 | truncated | short |
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dark | dark | 1.7–1.8 | 5–6 | dark | yellow | dark | 0-0-0 | dark | lobate | 6 | 0 | 1 | truncated | short |
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dark | dark | 1.6–1.8 | 5–6 | dark | yellow | dark | 0-0-0 | dark | lobate | 8 | 0 | 0 | truncated | short |
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dark | dark | 1.3–1.7 | 6–7 | dark | yellow | dark | 0-0-0 | dark | lobate | 7 | 0 | 0 | truncated | short |
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dark | dark | 1.3–1.7 | 5–6 | dark | yellow | dark | 0-0-0 | dark | lobate | 5–6 | 0 | 2 | truncated | short |
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dark | dark | 1.8–1.9 | 5–6 | dark | yellow | dark | 0-0-0 | dark | lobate | 7 | 0 | 1 | truncated | short |
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dark | dark | 1.3–1.5 | 5–6 | dark | dark | dark | 0-0-0 | dark | lobate | 6 | 0 | 1 | truncated | short |
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dark | dark | 1.9–2.2 | 7–8 | dark | yellow | dark | 0-0-0 | yellow | lobate | 2 | 0 | 1 | truncated | short |
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dark | dark | 2.0–2.3 | 7 | dark | yellow | dark | 0-0-0 | yellow | lobate | (1) | 0 | 2 | truncated | short |
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dark | yellow | 1.3–1.6 | 5 | dark | yellow | dark | 0-0-0 | yellow | lobate | 7 | 0 | 3 | truncated | short |
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dark | yellow | 2.1–2.7 | 10–12 | dark | dark | yellow | 0-0-0 | dark | lobate | 6 | 0 | 1 | truncated | short |
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dark | yellow | 1.9–2.0 | 8–9 | yellow | yellow | yellow | 0-0-0 | yellow | lobate | 3–4 | 0 | 1 | truncated | short |
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yellow | yellow | 1.8–2.1 | 5–6 | yellow | yellow | yellow | 0-0-0 | yellow | lobate | 3 | 0 | 1 | truncated | short |
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dark | yellow | 1.9–2.0 | 8–9 | yellow | yellow | yellow | 0-1-1 | yellow | lobate | 4–5 | 0 | 1 | truncated | short |
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yellow | yellow | 1.5–1.9 | 5 | yellow | yellow | yellow | 1-1-1 | yellow | lobate | 4–5 | 0 | 1 | truncated | short |
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yellow | yellow | 1.5–1.6 | 5 | yellow | yellow | yellow | 1-1-1 | yellow | lobate | 6 | 1 | 1 | truncated | short |
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yellow | yellow | 1.6–1.7 | 7 | yellow | yellow | yellow | 1-1-1 | yellow | lobate | 6 | 1 | 1 | truncated | short |
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dark | yellow | 1.9–2.2 | 7–8 | yellow | yellow | yellow | 1-1-1 | yellow | lobate | 5–7 | 0 | 1 | truncated | short |
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dark | yellow | 1.7–1.8 | 8–9 | yellow | yellow | yellow | 1-1-1 | yellow | lobate | 6 | 0 | 2 | truncated | short |
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yellow/dark | yellow | 2.2–2.3 | 8–10 | yellow | yellow | yellow | 1-1-1 | yellow | lobate | 5–7 | 0 | 1 | truncated | short |
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dark | yellow | 2.2–2.3 | 22–26* | yellow | yellow | yellow | 1-1-0 | yellow | lobate | 3–4 | 1 | 1 | truncated | short |
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dark | yellow | 2.0–2.2 | 8 | yellow | yellow | yellow | 1-1/1 | yellow | lobate | 3+3 | 2 | 1 | truncated | short |
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dark | yellow | 2.7–2.9 | 8–10 | yellow | yellow | yellow | 1-1/1 | yellow | lobate | 7–8 | 0 | 1 | truncated | short |
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dark | yellow | 2.4–2.5 | 7–9 | yellow | yellow | yellow | 1-1/1 | yellow | lobate | 9–12 | 0 | 1 | truncated | short |
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dark | yellow | 2.2–2.9 | 8–9 | yellow | yellow | yellow | 1/1/1 | yellow | lobate | 6 | 0 | 1 | truncated | short |
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dark | yellow | 2.1–2.3 | 8–9 | yellow | yellow | yellow | 1/1/1 | yellow | lobate | 8 | 0 | 1 | truncated | short |
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dark | yellow | 2.0–2.1 | 6–7 | yellow | yellow | yellow | 1/1/1 | yellow | lobate | 5 | 0 | 1 | truncated | short |
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dark | yellow | 1.4–1.8 | 5 | yellow | yellow | yellow | 1/1/1 | dark | lobate | 5 | 0 | 0 | truncated | short |
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yellow | yellow | 1.3–1.6 | 6 | yellow | yellow | yellow | 1/1/1 | yellow | elongated | (2) | 1 | 1 | truncated | short |
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yellow | yellow | 1.4–1.5 | 6 | yellow | yellow | yellow | 1/1/1 | yellow | elongated | (2) | 0 | 1 | truncated | short | ||
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yellow | yellow | 1.6–2.0 | 6–8 | yellow | yellow | yellow | 1/1/1 | yellow | elongated | (2) | 1 | 1 | truncated | short |
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yellow | yellow | 1.9–2.3 | 7–8 | yellow | yellow | yellow | 1/1/1 | yellow | elongated | (2) | 1 | 1 | truncated | short |
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1 1, present; 0, absent; 1-1, stripes unfused; 1/1, stripes fused.<br/> 2 Number of unfused tubercle-like setae forming a row is shown in parenthesis; -, lacking tubercle-like setae forming a row.<br/> * Number of setulae in four rows.
1 | Acrostichal setulae vestigial, at most 4 pairs in 2 rows (Fig. |
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– | Acrostichal setulae at least 5 pairs in 2 (or rarely 4) rows (Fig. |
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2 | Surstylus small, lobate, not elongated, not setose apically (Fig. |
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– | Surstylus large, elongated, setose apically (Fig. |
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3 | Legs dark brown (Fig. |
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– | Legs yellow or brownish yellow (Fig. |
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4 | 1st flagellomere and haltere yellow (Fig. |
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– | 1st flagellomere and haltere dark (Fig. |
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5 | Maxillary palpus dark-colored (Fig. |
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– | Maxillary palpus yellow (Fig. |
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6 | Scutum completely gray (Fig. |
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– | Scutum bluish gray with longitudinal dark stripes (Fig. |
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7 | Scutum with 3 pairs of longitudinal dark gray stripes (Fig. |
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– | Scutum with a medial stripe and a pair of lateral longitudinal dark gray stripes (Fig. |
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8 | Legs dark-colored (Fig. |
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– | Legs yellow (Fig. |
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9 | Haltere yellow (Fig. |
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– | Haltere dark-colored (Fig. |
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10 | Maxillary palpus yellow (Fig. |
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– | Maxillary palpus dark-colored (Fig. |
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11 | Pleuron with ventral half dark-colored (Fig. |
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– | Pleuron wholly yellow (Fig. |
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12 | Scutellum wholly gray (Fig. |
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– | Scutellum wholly yellow or at least partly yellow (Fig. |
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13 | Pedicel of antenna yellow (Fig. |
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– | Pedicel of antenna brown (Fig. |
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14 | Legs dark brown (Fig. |
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– | Legs yellowish brown (Fig. |
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15 | Male surstylus lacking tubercle-like seta (Fig. |
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– | Male surstylus with 1 or 2 tubercle-like setae (Fig. |
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16 | Male epandrium with a comb comprising 8 basally fused, long, tubercle-like setae (Fig. |
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– | Male epandrium with a comb comprising 7 basally fused, long, tubercle-like setae (Fig. |
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17 | Male epandrium with a comb comprising 5 or 6 basally fused, long, tubercle-like setae (Fig. |
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– | Male epandrium with a comb comprising 7 basally fused long tubercle-like setae (Fig. |
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18 | Scutum and scutellum gray (Fig. |
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– | Scutum yellow at least in part, and scutellum yellow (Fig. |
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19 | Scutum wholly yellow, lacking dark stripes (Fig. |
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– | Scutum yellow, with 1 or 2 pairs of dark longitudinal stripes (Fig. |
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20 | 1st flagellomere black (Fig. |
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– | 1st flagellomere yellow (Fig. |
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21 | Scutum lacking a medial dark stripe, but with brown unfused intra-alar and supra-alar stripes (Fig. |
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– | Scutum with a medial dark stripe (Fig. |
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22 | Intra-alar and supra-alar stripes of scutum separate and unfused (Fig. |
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– | Intra-alar and supra-alar stripes of scutum fused, forming a pair of wide bands (Fig. |
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23 | Scutum subglossy with pruinosity; intra-alar and supra-alar stipes with similar thickness (Fig. |
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– | Scutum glossy; supra-alar stipe is pale and indistinct (Fig. |
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24 | Intra-alar stripes not confluent with a pair of lateral presutural dark ovoid spots (Fig. |
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– | Intra-alar stripes adjoining and confluent with 1 pair of lateral presutural dark ovoid spots (Fig. |
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25 | 1st flagellomere yellow (Fig. |
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– | 1st flagellomere black (Fig. |
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26 | Intra-alar stripes and lateral presutural dark ovoid spots with silvery pruinosity (Fig. |
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– | Intra-alar stripes and lateral presutural dark ovoid spots without silvery pruinosity (Fig. |
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27 | Male epandrium with an extremely elongated arm, bearing 2, dark, ventrally curved, tubercle-like setae borne at wide angle (Fig. |
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– | Male epandrium with a basally enlarged, slightly flattened hypertrophied arm, which bears a dark laterally enlarged tubercle-like seta (Fig. |
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28 | Scutellum yellow with lateral margins dark-colored (Fig. |
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– | Scutellum wholly yellow (Fig. |
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29 | 1st flagellomere black (Fig. |
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– | 1st flagellomere yellow (Fig. |
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30 | Scutellum brownish yellow, tergite of abdomen brown (Fig. |
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– | Scutellum yellow, tergite of abdomen yellow (Fig. |
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31 | Lateral fused band of scutum reaching just before scutellum (Fig. |
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– | Lateral fused band of scutum ending at anterior 7/8 length of scutum before scutellum (Fig. |
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32 | Medial stripe of scutum not confluent with the lateral fused band, and the 2 bands are delimited by a yellow line (Fig. |
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– | Medial stripe of scutum confluent with the lateral fused band (Fig. |
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33 | Scutellum wholly yellow (Fig. |
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– | Scutellum yellow with lateral margins darkened (Fig. |
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34 | Abdomen dorsally with a pair of brown lateral semicircular patches on anterior half of the 2nd tergite (Fig. |
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– | Abdomen dorsally without dark patches on tergites (Fig. |
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35 | Male epandrium with a comb comprising 6 fused tubercle-like setae (Fig. |
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– | Male epandrium with a comb comprising 8 fused tubercle-like setae (Fig. |
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36 | Yellow patch of midposterior scutum darkened and ill-defined (Fig. |
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– | Yellow patch of midposterior scutum rectangular and well-defined (Fig. |
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37 | Male epandrium with a pair of elongated, hypertrophied arms, each bearing a tubercle-like seta apically (Fig. |
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– | Male epandrium without a pair of elongated, hypertrophied arms (Fig. |
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38 | Hypertrophied arm on male epandrium strongly curved outward (Fig. |
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– | Hypertrophied arm on male epandrium protruded forward (Fig. |
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We describe 39
Japan: 1♂1♀ (MK-AG-a410, a26), Namari-kawa, Yakumo, Futami, Hokkaido (
A medium-sized yellow species (wing length 1.7–1.8 mm) that has a pruinose yellow scutum with one medial and two pairs of gray lateral stripes, black 1st flagellomeres, yellow maxillary palpi, yellow halteres, and yellow legs. Male epandrium inner-laterally with a comb consisting of six fused long tubercle-like setae, and surstylus with two tubercle-like setae. Larva mines the thallus of
Usuobi-zenigoke-hamoguribae.
(Fig.
The host plants from which this species emerged grow on wet cliffs along roads in a cool temperate forest of
Thus far, recorded from Europe, North America, and Japan. The authors recorded only from southern Hokkaido (Fig.
Locality records of three
The morphology of the Japanese specimens closely coincided with the description of
This species resembles
Japan: 35♂37♀, same data as holotype, emerged on 8–17-V-2021; 9♂5♀, Iwaobetsu, Shari, Hokkaido, 1-V-2021 (as larva), emerged on 26–28-V-2021; 3♂1♀, Horoman-kyo, Samani, Hokkaido, 30-IV-2021 (as larva), emerged on 26–31-V-2021; 1♂2♀, Odarumi, Makioka, Yamanashi Pref., 27-VI-2014 (as larva), emerged on 15-VII-2021; 1♂1♀, Shirabiso-toge, Kamimura, Iida, Nagano Pref., 27-IV-2021 (as larva), emerged on 23–25-V-2021; 22♂20♀, Ikawa-toge, Aoi-ku, Shisuoka Pref., 26-V-2021 (as larva), emerged on 8–17-VI-2021.
A large yellow species (wing length 2.0–2.1 mm) having a pruinose dark gray scutum with a trapezoid yellow patch medially on posterior 1/3, a yellow scutellum, black 1st flagellomeres, yellow maxillary palpi, yellow halteres, and yellow legs. Male epandrium with a comb comprising five or six fused long tubercle-like setae. Larva mines the thallus of
The specific name (
Kitsunebi-zenigoke-hamoguribae.
Larvae construct linear-botch mines in the thallus, particularly in the midrib, and pupate in the mines (Fig.
Habitat of
The host plants from which this species emerged, grow on mesic soils along roads in cool temperate forests (beech forests dominated by
Japan: Hokkaido and Honshu (Fig.
This species is distinguished from all other species of
Japan: 1♂1♀, Ikawa-toge, Aoi-ku, Shizuoka Pref., 26-V-2021 (as larva), emerged on 31-V–2-VI-2021; 1♂, Tokuzo-ji, Yana, Kisarazu, Chiba Pref., I-XI-2021 (as larva), emerged on 3-XII-2021.
A medium-sized species (wing length 1.6–1.7 mm) having pruinose dark gray scutum, light yellow scutellum, black 1st flagellomere, dark maxillary palpus, dark halteres, and dark legs. Male epandrium inner-basally with a comb of four unfused tubercle-like setae, and inner-laterally with a comb of three or four fused, long, tubercle-like setae; surstylus bilobed and hypertrophied dorsal arm with one tubercle-like seta. Larva mines the thallus of
The specific name
Tsukuyomi-zenigoke-hamoguribae.
Larvae mine the thallus of the host plant and pupate in the mines. The mines are not apparent from the outside.
The host plants from which this species emerged, grow on mesic soils on levee of paddy fields and along roads in natural beech forests (Fig.
Japan: Honshu (Fig.
This species resembles
Japan: On
On
On
On
A medium-sized species (wing length 1.6–1.8 mm) having dark gray scutum, dark-cornered yellow scutellum, black 1st flagellomere, black maxillary palpus, gray halteres, and brown legs. Male epandrium with a comb comprising seven or eight fused long tubercle-like setae. Larva mines the thallus of
Female morphology and larval ecology of
The yellow patch on posterior scutum slightly varied from distinct to obscure ones among localities but not among host liverwort species/subspecies (Fig, 6F–K).
The specific name refers to host plant genus,
Uzumibi-zenigoke-hamoguribae.
The main host plants are
Larvae construct linear mines in the thallus, particularly in the midrib, and pupate in the mined thalli (Fig.
Japan: Hokkaido, Honshu, Shikoku, Kyushu (Fig.
This species resembles
This species also resembles
Japan: On
On
On
A small black species (wing length 1.3–1.6 mm) having subshiny black scutum with an oval yellow pattern extending from mid-posterior margin to scutellum, black 1st flagellomere, yellow maxillary palpus, yellow halteres, and brownish legs. Male epandrium with a comb comprising seven fused tubercle-like setae, and surstylus with a comb comprising three fused tubercle-like setae. Larva mines the thallus of
Female morphology (
The specific name
Nubatama-tosazenigoke-hamoguribae.
Larvae construct linear mines in the thallus, and pupate in the midrib of the mined thalli (Fig.
The main habitats of this species are river banks where
Japan: Honshu, Shikoku, Kyushu, Amami, Okinawa and Yaeyama Islands (Fig.
This species resembles
Japan: 5♂4♀, Abe-toge, Aoi-ku, Shizuoka Pref., 5-I-2015 (as larva), emerged on 18-V-2016; 2♂3♀, Tango-kanzaki, Maizuru, Kyoto Pref., 19-V-2021 (as larva), emerged on 30-V–5-VI-2021; 1♂1♀, Tategasaki, Kumano, Mie Pref., 23-IV-2021 (as larva), emerged on 11-V-2021; 3♀, Gakuen-ji, Bessho, Izumo, Shimane Pref., 31-III-2015 (as larva), emerged on 9-V-2015; 3♂1♀, Tazukawa-keikoku, Katsuura, Tokushima Pref., 30-III-2021 (as larva), emerged on 11–20-V-2021; 3♂4♀, Narutaki, Ichiu, Tsurugi, Tokushima Pref., 12-VI-2017 (as larva), emerged on 17–21-VI-2017; 1♀, Tashiro, Kinko, Kimotsuki, Kagoshima Pref., 22-III-2015 (as larva), emerged on 1-V-2015; 1♂1♀, Isso, Yaku Is., Kumage, Kagoshima Pref., 11-III-2016 (as larva), emerged on 617–19-IV-2016; 11♂14♀, Higashinakama, Sumiyo, Amami, Kagoshima Pref., 21-II-2015 (as larva), emerged on 30-III–8-IV-2015.
A large dark species (wing length 1.9–2.2 mm) having subshiny dark brown scutum, brownish yellow scutellum, black 1st flagellomere, dark maxillary palpus, dark halteres, and dark gray legs. Male epandrium inner-laterally with an incomplete comb comprising three short, fused, tubercle-like setae medially. Larva mines the thallus of
Female morphology and larval ecology of
The color of frons and scutellum varies from yellow to brownish yellow in some localities.
The specific name refers to its host plant genus
Zangetsu-kezenigoke-hamoguribae.
(Fig.
The main habitats of this species are along streams in warm temperate evergreen forests (Fig.
Japan: Honshu, Shikoku, Kyushu, Yaku Island and Amami-oshima Island (Fig.
Locality records of three
This species resembles
Japan: 1♀, Sanekawa-keikoku, Iide, Aga, Higashikanbara, Niigata Pref., 3-V-2015 (as larva), emerged on 12-VI -2015; 1♀, Mt. Kiyosumi, Kamogawa, Chiba Pref., 24-I-2012 (as larva), emerged on 25-V–5-VI-2012; 17♂32♀, Inago, Shibakawa, Fujinomiya, Shizuoka Pref., 17-II-2002 (as larva), emerged on 23-IV–5-VI-2021; 1♂, Shogawa-kyo, Tonami, Toyama Pref., 3-V-2016 (as larva), emerged on 9-VI -2016; 1♀, Takeda-gawa, Maruoka, Sakai, Fukui Pref., 6-IV-2002 (as larva), emerged on 13-VI-2019; 2♂2♀, Mt. Nabejiri, Taga, Shiga Pref., 23-V-2015 (as larva), emerged on 26–15-VI-2015; 1♂1♀, Boumura, Kazuragawa, Otsu, Shiga Pref., 7-IV-2002 (as larva), emerged on 24-V-2002; 1♂1♀, Mitake, Kamitsushima, Nagasaki Pref., 9-IV-2009 (as larva), emerged on 6-VI-2009.
A medium-sized species (wing length 1.4–1.8 mm) having subshiny brown scutum with brownish yellow pattern extending from mid-posterior margin to scutellum, black 1st flagellomere, brown maxillary palpus, brown halteres, and brownish yellow legs. Male epandrium inner-laterally with a comb comprising five long fused tubercle-like setae. Larva mines the thallus of
The specific name (
Yuuzuki-kezenigoke-hamoguribae.
(Fig.
The main habitats of this species are along streams in warm temperate forests and cool temperate deciduous forests (Fig.
Japan: Honshu, Shikoku, Kyushu, Tsushima Island (Fig.
Although this species used the same host plant as
Japan: On
On
A small species (wing length 1.3–1.6 mm) having subshiny brown scutum with yellow pattern extending from mid-posterior margin to scutellum, yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-subdistally with hypertrophied, elongated, strongly curved arm bearing a dark tubercle-like seta. Larva mines the thallus of
Female morphology and larval/adult ecology of
The yellow posterior patch on the scutum varied from distinct to obscure ones among localities and even among individuals in some localities.
The specific name (
Yumihari-tsubozenigoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus and pupate in the mines (Fig.
The habitats of this species are limestone outcrops in temperate deciduous forests, where the host liverworts grow (Fig.
Japan: Honshu (Fig.
Locality records of three
This species resembles
Japan: 1♂3♀, same data as holotype, emerged on 19–25-V-2021; 3♀, Ozasu, Ogano, Chichibu, Saitama Pref., 28-XI-2014 (as larva on
A small species (wing length 1.4–1.5 mm) having subshiny brown scutum with an oval yellow pattern extending from mid-posterior margin to scutellum, yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium with an imperfect comb comprising two short tubercle-setae. Larva mines the thallus of
The specific name refers to the host plant genus
Tsukikage-tsubozenigoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus and pupate in the mines (Fig.
The habitats of this species are outcrops of lime stones in temperate deciduous forests, where the host liverworts grow (Fig.
Japan: Honshu (Fig.
This species resembles
Japan: 1♀, Narahara, Ueno, Tano, Gunnma Pref., 18-IV-2021 (as larva), emerged on 1-VI-2021; 1♀1♂, Ozasu, Ogano, Chichibu, Saitama Pref., 10-IX-2017 (as larva), emerged on 9–13-X-2017; 1♀, Irisawai, Oshika, Nagano Pref., 29-IV-2011 (as larva), emerged on 22-V-2011.
A medium-sized yellow species (wing length 1.6–1.7 mm) having subshiny yellow scutum with two pairs of lateral stripes, yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-subdistally with a hypertrophied arm which bears an enlarged tubercle-like seta; inner-basally with a hypertrophied arm which bears a comb comprising five or six fused tubercle-like setae. Larva mines the thallus of
Female morphology and larval/adult ecology of
The specific name (
Kurosuji-tsubozenigoke-hamoguribae.
Larvae construct radiate mines in the thallus and pupate in the mines (Fig.
The habitats of this species are limestone outcrops in temperate deciduous forests, where the host liverworts grow (Fig.
Japan: Honshu (Fig.
This species has several unique characteristics in male genitalia: a flattened stout tubercle-like seta on distal margin of epandrium; a comb of tubercle-like setae borne on enlarged projection of inner surface of epandrium; thin, bare surstylus; short distiphallus unpigmented at basal half. The unique characteristics suggest distant relation of this species from other liverwort-associated species. This species resembles
Japan: 1♀, Yutsun, Iriomote-Is. Yaeyama, Okinawa Pref., 8-XI-2021 (as larva), emerged on 17-XI-2021.
A small species (wing length 1.4–1.7 mm) having pruinose gray scutum and scutellum, brown 1st flagellomere, yellow maxillary palpus, dark halteres, and yellow legs. Scutum lacks acrostichal setulae. Male epandrium lacks tubercle-like seta; distiphallus of male genitalia tapering apically. Larva mines the thallus of
The specific name refers to the type locality, Iriomote Island.
Okinawasaihaigoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus in early instars, later entering the midrib, and pupating there (Fig.
The habitats of this species are rocky stream banks in subtropical evergreen forests (Fig.
Japan (Honshu). Recorded only from Iriomote Island in the Ryukyu Archipelago (Fig.
Locality records of three
This species resembles
In Japan, six
Japan: 6♂7♀, Izu-oshima Is. Tokyo Pref., 22-III-2009 (as larva), emerged on 15–20-IV-2009; 2♀, Yugashima, Izu, Shizuoka Pref., 7-III-2012 (as larva), emerged on 12–14-IV-2012; 4♂8♀, Sinjo, Mihama, Mikata, Fukui Pref., 11-III-2012 (as larva), emerged on 18–23-IV-2012; 3♂2♀, Shimaji-gawa, Ujitachi, Ise, Mie Pref., 3-IV-2010 (as larva), emerged on 22-IV–2-V-2010; 2♂1♀, Urashima-jinja, Honjo-hama, Ine, Kyoto Pref., 31-VII-2011 (as larva), emerged on 22-III-2011; 2♂5♀, Ukawa, Tango, Kyotango, Kyoto Pref., 5-III-2021 (as larva), emerged on 2–8-IV-2021; 1♀, Sannoko, Kawakami, Higashi-yoshino, Nara Pref., 26-II-2016 (as larva), emerged on 11-IV-2016; 1♀, Tazukawa-keikoku, Katsuura, Tokushima Pref., 11-X-2016 (as larva), emerged on 19-IV-2011.
A large yellow species (wing length 2.0–2.2 mm) having a pruinose dark gray scutum with an oval yellow pattern extending from the mid-posterior margin to the scutellum, a black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a hand-like comb comprising
Four or five basally fused, long, tubercle-like setae. Larva mines the thallus of
Female morphology and larval ecology of
The specific name (
Suisei-jingasagoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus, and pupate in the mines (Fig.
The habitats of this species are rocky cliffs in warm temperate evergreen forests (Fig.
Japan (Honshu, Shikoku).
This species resembles
Japan: 5♂3♀, Takasuka, Joso, Ibaragi Pref., 2-XI-2021 (as larva), emerged on 11–15-XII-2021; 1♀, Kuchisakamoto, Aoi-ku, Shizuoka Pref., 20-IX-1998 (as larva), emerged on 18–23-IX-1998; 1♀, Mt. Gozaisho, Komono, Mie Pref., 1-V-2001 (as larva), emerged on 25-V-2001; 3♂6♀, Ukawa, Tango, Kyotango, Kyoto Pref., 5-III-2021 (as larva), emerged on 2–8-IV-2021; 16♂10♀, Kibune, Sakyo-ku, Kyoto Pref., 24-VI-2011 (as larva), emerged on 12-VII-2011; 16♂25♀, Wadagawa-kyo, Kumanogawa, Shingu, Wakayama Pref., 7-VI-2021 (as larva), emerged on 17-VII–8-VIII-2021; 1♂1♀, Kibune, Sakyo-ku, Kyoto Pref., 23-IV-2021 (as larva), emerged on 2–6-V-2021; 6♂8♀, Ryugakyo, Yamashiro, Miyoshi, Tokushima Pref., 1-II-2014 (as larva), emerged on 24-IV–3-V-2014; 1♂1♀, Tazukawa-keikoku, Katsuura, Tokushima Pref., 30-III-2021 (as larva), emerged on 26–29-IV-2021; 3♂4♀, Sui, Anan, Tokushima Pref., 30-III-2021 (as larva), emerged on 14–17-IV-2021; 1♂2♀, Kurase-keikoku, Tanbara, Saijo, Ehime Pref., 2-II-2014 (as larva), emerged on 20–26-IV-2014; 1♀, Yasui-keikoku, Niyodogawa, Agawa, Kochi Pref., 27-II-2011 (as larva), emerged on 17-IV-2011; 1♂, Kinsakubaru, Amami, Kagoshima Pref., 4-VII-1999 (as larva), emerged on 25-VII-1999; 5♂11♀, Tachijami, Kume Is. Okinawa Pref., 20-III-2020 (as larva), emerged on 15-IV–8-V-2020.
A medium-sized species (wing length 1.5–1.9 mm) having subshiny yellow scutum with a medial and two pairs of lateral dark stripes; inner stripes with silvery reflection. Adults with yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a hand-like comb comprising four or five basally fused, long, tubercle-like setae. Larva mines the thallus of
Female morphology and larval ecology of
Background color of scutum and scutellum varies from yellow to grayish yellow, and the blackening is obvious in the Yaku Islands.
The specific name (
Ginsuji-jingasagoke-hamoguribae.
Larvae construct digitate mines in the thallus, and pupate in the mine (Fig.
The habitats of this species are rocky cliffs in warm temperate evergreen forests (Fig.
Japan: Honshu, Shikoku, Yaku Island, Amami-Oshima Island, Kume Island (Fig.
Locality records of three
This species resembles
Japan: 4♂3♀, Kamihirayama, Tatsuyama, Tenryu, Hamamatsu, Shizuoka Pref., 7-XI-2010 (as larva), emerged on 18–28-IV-2011; 1♂1♀, Chiromo, Toyotama, Tsushima, Nagasaki Pref., 28-XI-2011 (as larva), emerged on 1-V-2011; 2♀, Kibune, Sakyo-ku, Kyoto Pref., 20-VI-2016 (as larva), emerged on 14-VII-2021.
A medium-sized species (wing length 1.5–1.6 mm) having a subshiny yellow scutum with a medial and two pairs of lateral black stripes, yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-basally with a comb comprising six fused long tubercle-like setae, and inner-subdistally with an extremely elongated arm, which apically bears two dark, diverging, ventrally curved, tubercle-like setae. Larva mines the thallus of
Female morphology and larval/adult ecology of
The specific name (
Sasumata-jingasagoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus, and pupate in the mines (Fig.
The habitats of this species are rocky cliffs in warm temperate evergreen forests (Fig.
Japan: Honshu, Shikoku (Fig.
This species resembles
Japan: 17♂22♀, Kanna-gawa, Nakatsugawa, Chichibu, Saitama Pref., 14-XI-2010 (as larva), emerged on 14-IV-2011; 6♂11♀, Oochi-gawa, Chichibu, Saitama Pref., 13-III-2017 (as larva), emerged on 25–28-IV-2011; 1♂2♀, Ukawa, Tango, Kyotango, Kyoto Pref., 5-III-2021 (as larva), emerged on 9–17-IV-2021; 1♀, Doro-kyo, Totsugawa-mura, Nara Pref., 29-III-2019 (as larva), emerged on 3-IV-2019; 1♂4♀, Tazukawa-keikoku, Katsuura, Tokushima Pref., 30-III-2021 (as larva), emerged on 23–30-IV-2021; 2♂1♀, Kurase-keikoku, Tanbara, Saijo, Ehime Pref., 2-II-2014 (as larva), emerged on 20–26-IV-2014.
A medium-sized yellow species (wing length 1.6–2.0 mm) having subshiny brown scutum with an oval yellow pattern extending from the mid-posterior margin to the scutellum, a yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a long hypertrophied arm which apically bears a dark, long, apically flattened, obliquely truncated tubercle-like seta. Larva mines the thallus of
Female morphology and larval ecology of
The specific name (
Naginata-jingasagoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus, and pupate in the mines (Fig.
The habitats of this species are rocky cliffs in warm temperate evergreen forests. This species is sympatric with
Japan: Honshu, Shikoku, Tsushima Island (Fig.
Locality records of three
This species resembles
Japan: 1♂, Ryugakyo, Yamashiro, Miyoshi, Tokushima Pref., 21-IV-2014 (as larva), emerged on 2-V-2014.
A medium-sized yellow species (wing length 1.9–2.3 mm) having a subshiny brown scutum with an obscure oval yellow pattern extending from the mid-posterior margin to the scutellum, a yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a long hypertrophied, ventrally curved arm that apically bears a dark, apically bifid tubercle-like seta. Larva mines the thallus of
The specific name (
Karasuki-jingasagoke-hamoguribae.
Larva constructs linear mine in the thallus, and pupate in the mine (Fig.
The habitats of this species are rocky cliffs in warm temperate evergreen forests (Fig.
Japan: Honshu, Shikoku, Tsushima Island (Fig.
This species resembles
Japan: 1♀, Izu-oshima Is. Tokyo Pref., 22-III-2009 (as larva), emerged on 19-IV-2009; 2♀, Ashiu, Nantan, Kyoto Pref., 12-III-2018 (as larva), emerged on ?-IV-2018; 2♂6♀, Sannoko, Kawakami, Higashi-yoshino, Nara Pref., 26-II-2016 (as larva), emerged on 18–27-IV-2016; 19♂13♀, WD, 7-VII-2021 (as larva), emerged on 24-VII–6-VIII-2021; 2♂, Yajiemon-jinja, Sakurae, Gotsu, Shimane Pref., 24-VI-2012 (as larva), emerged on 19-VII–23-VIII-2012; 2♂2♀, Tazukawa-keikoku, Katsuura, Tokushima Pref., 11-X-2016 (as larva), emerged on 1-V-2016; 1♂3♀, Yasui-keikoku, Niyodogawa, Agawa, Kochi Pref., 27-II-2011 (as larva), emerged on 18-IV-2011; 1♂3♀, YSI, 27-II-2011 (as larva), emerged on 18-IV-2011; 1♀, Shiibaru, Izumi, Yatsushiro, Kumamoto Pref., 23-III-2015 (as larva), emerged on 28-IV-2015; 1♀, Okujisso, Isa, Kagoshima Pref., 17-XII-2012 (as larva), emerged on 12-IV-2013; 2♂1♀, Sumiyo, Amami, Kagoshima Pref., 17-II-1999 (as larva), emerged on 25–28-II-1999.
A small dark species (wing length 1.3–1.7 mm) having pruinose dark gray scutum with a small oval yellow pattern extending from the mid-posterior margin to the scutellum, a black 1st flagellomere, dark maxillary palpus, gray halteres, and brown legs. Male epandrium inner-basally with a comb comprising seven long fused tubercle-like setae. Larva mines the thallus of
Female morphology and larval ecology of
The specific name refers to the larval life within
Kainohi-jingasagoke-hamoguribae.
Larvae at first construct linear mines in the thallus at first, then enter the midrib, and pupate in the mine (Fig.
The habitats of this species are rocky cliffs in warm temperate evergreen forests (Fig.
Japan: Honshu, Shikoku, Tsushima Island (Fig.
This species resembles
Japan: 1♂1♀, Sendan-todoro, Izumi, Yatsushiro, Kumamoto Pref. (
A large dark species (wing length 2.0–2.3 mm) having subshiny dark gray scutum, yellow scutellum, black 1st flagellomere, dark maxillary palpus, gray halteres, and dark brown legs. Male epandrium inner-basally with a protruding, plate-like arm bearing one strong, tubercle-like seta apically. Larva mines the thallus of
Female morphology and larval ecology of
The specific name refers to the host plant genus,
Azumazenigoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later expanding their mines, and pupate in the mines (Fig.
The habitats of this species are mesic slopes in warm temperate evergreen forests. Our rearing records suggest that it is univoltine, with adults emerging from overwintered pupae in spring.
Japan: Kyushu (Fig.
This species is superficially very similar to
Japan: On
On
A large yellow species (wing length 2.7–2.9 mm) having pruinose yellow scutum with a medial and a pair of dark brown lateral stripes, entirely yellow scutellum, black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-distally with a long tubercle-like seta, and inner-basally with a comb consisting of 7–9 long fused tubercle-like setae. Larva mines the thallus of
The number of tubercle-like setae in a comb in the male genitalia varies from 7 to 9, but the variation did not involve in a geographical cline.
The specific name refers to the moon; a clear, rounded, yellow pattern on the scutum was likened to a full moon.
Meigetsu-jagoke-hamoguribae.
(Fig.
Larval ecology of
The habitats of this species are stream banks and mesic slopes in subalpine coniferous forests dominated by
Japan: Hokkaido, Honshu, Shikoku (Fig.
Locality records of three
The characteristics of this species and the following three related species (
This species also resembles
This species resembles
Among Japanese species, this species resembles
Japan: On
On
On
A large yellow species (wing length 2.4–2.5 mm) having a pruinose yellow scutum with a medial and a pair of dark brown lateral stripes, a yellow scutellum with dark lateral corners, black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-distally with a long tubercle-like seta, and inner-basally with a comb consisting of 9–12 long fused tubercle-like setae. Larva mines the thallus of
.
Female morphology and larval ecology of
The number of tubercle-like setae in a comb in the male genitalia varied from 9 to 12. Although the number varied among individuals within a population and even between left and right sides of the epandrium in an individual, and the number was generally greater in western Honshu and Shikoku than in northern regions.
The specific name
Izayoi-jagoke-hamoguribae.
(Fig.
The habitats of this species are stream banks and mesic slopes in temperate deciduous forests dominated by
Japan: Hokkaido, Honshu, Shikoku (Fig.
This species resembles
Among the Japanese species,
Japan: 1♂, Mt. Futago, Ogano, Chichibu-gun, Saitama Pref., 26-III-2021 (as larva), emerged on 21-IV-2021.
A large yellow species (wing length 2.2–2.9 mm) having a pruinose dark brown scutum with an obscure oval yellow pattern extending from the mid-posterior margin to the scutellum, a black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a long tubercle-like seta, and inner-basally with a comb consisting of six long, fused, tubercle-like setae. Larva mines the thallus of
The specific name (
Shungetsu-jagoke-hamoguribae.
(Fig.
The habitats of this species are stream banks and cliffs in temperate deciduous forests dominated by
Japan: Honshu, around Chichibu mountains in the Kanto Region (Fig.
This species resembles
Japan: 1♂1♀, Mt. Nokogiri, Kyonan, Awa, Chiba Pref., 24-I-2012 (as larva), emerged on 24-IV–8-V-2012 1♂1♀, Asahi-daki, Shuzenji, Izu, Shizuoka Pref., 7-III-2012 (as larva), emerged on 28-III–1-IV-2012; 3♂3♀, Tamaki-gawa, Totsugawa, Yoshino, Nara Pref., 9-III-2014 (as larva), emerged on 26-III–9-IV-2014; 2♂1♀, Wabuka, Kushimoto, Wakayama Pref., 4-III-2012 (as larva), emerged on 26-III–12-IV-2012; 1♂2♀, Kibune, Sakyo-ku, Kyoto Pref., 1-III-2011 (as larva), emerged on 25-III–3-IV-2011; 2♂, Yasukawa-keikoku, Tanabe, Wakayama Pref., 9-VII-2012 (as larva), emerged on 16–21-IV-2002; 1♂1♀, Takinohai, Kozagawa, Wakayama Pref., 13-IV-2014 (as larva), emerged on 2–15-V-2014; 1♀, Yoshiwa, Hatsukaichi, Hiroshima Pref., 30-V-2014 (as larva), emerged on ?-V-2014; 2♀, Mt. Ryuso, Aoi, Shizuoka Pref., 13-X-2015 (as larva), emerged on 14-IV-2015; 1♂, Shinkawa-keikoku, Kirishima, Kagoshima Pref., 31-III-2017 (as larva), emerged on 16-IV-2017.
A large yellow species (wing length 2.1–2.3 mm) having pruinose dark brown scutum with an obscure oval yellow pattern extending from the mid-posterior margin to the scutellum, a black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a long tubercle-like seta and inner-basally with a comb comprising eight or nine long fused tubercle-like setae. Larva mines the thallus of
The number of tubercle-like setae in a comb in the male genitalia varies from 8 to 9, but the variation does not involve a geographic cline.
The specific name (
Oborozuki-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in warm temperate evergreen forests dominated by
Japan: Honshu, Shikoku, Kyushu (Fig.
This species resembles
Japan: 6♂6♀, Mikisato, Owase, Mie Pref., 30-XII-2020 (as larva), emerged on 20-II–15-III-2021; 3♂5♀, Tategasaki, Kumano, Mie Pref., 23-IV-2021 (as larva), emerged on 26-IV–18-V-2021; 1♀, Mt. Kosho, Asakura, Fukuoka Pref., 11-IV-2010 (as larva), emerged on 18-IV-2010; 12♂15♀, Mt. Kora, Kurume, Fukuoka Pref., 29-IV-2008 (as larva), emerged on 1–3-V-2008; 1♂2♀, Mt. Osuzu, Tsuno, Miyazaki Pref., 15-XII-2020 (as larva), emerged on 23–28-II-2013; 1♂2♀, Shinkawa-keikoku, Kirishima, Kagoshima Pref., 31-III-2071 (as larva), emerged on 13–27-IV-2017; 1♀, Tashiro, Kinko, Kimotsuki, Kagoshima Pref., 18-V-2013 collected on thallus of
A large dark species (wing length 2.1–2.7 mm) having pruinose dark brown scutum, black 1st flagellomere with yellow arista, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-basally with a comb comprising six long fused tubercle-like setae, but lacking an inner-lateral tubercle-like seta. Larva mines the thallus of
The number of tubercle-like setae in a comb of the male epandrium varies from 6 to 8 within the same population and among localities.
The specific name
Ugetsu-jagoke-hamoguribae.
(Fig.
The habitats of this species are stream banks and mesic slopes in warm temperate evergreen forests dominated by
Japan: Honshu, Shikoku, Kyushu (Fig.
Locality records of three
This species is unique in the wholly dark brown color of both scutum and scutellum, and easily distinguished from all other
Japan: 7♂12♀, Soun-kyo, Kamikawa, Hokkaido, 31-V-2021 (as larva), emerged on 11–22-VII-2021; 2♂4♀, Aizankei, Kamikawa, Hokkaido, 4-X-2011 (as larva), emerged on 26-V–2-VI-2011; 2♂6♀, Yuni-ishikari-gawa, Soun-kyo, Kamikawa, Hokkaido, 1-VI-2020 (as larva), emerged on 5–14-VII-2020; 10♂4♀, Nozuka-toge, Urakawa, Hokkaido, 30-IV-2011 (as larva), emerged on 5–18-VI-2021; 1♂2♀, Mt. Tengu, Jozan-kei, Minami-ku, Sapporo, Hokkaido, 2-V-2021 (as larva), emerged on 7–10-VI-2021; 2♂1♀, Jozan-kei, Minami-ku, Sapporo, Hokkaido, 2-V-2021 (as larva), emerged on 7–19-VI-2021; 1♂, Horoman-kyo, Samani, Hokkaido, 30-IV-2021 (as larva), emerged on 19-VI-2021.
A medium-sized yellow species (wing length 2.2–2.3 mm) uniquely having sexual dimorphism in color of the 1st flagellomere: male yellow, female black. The adult has a pruinose yellow scutum with a medial and two pairs of black stripes, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a long tubercle-like seta, and inner-basally with a comb comprising 6–8 long fused tubercle-like setae. Larva mines the thallus of
Female morphology and larval ecology of
The color of the lateral stripes on the scutum varies among populations, with specimens in the southern population having darker stripes. The number of tubercle-like setae in a comb in male genitalia varies from 5 to 7 among localities.
The specific name (
Murasame-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are mesic slopes in subalpine coniferous forests dominated by
Japan: Hokkaido, Honshu (Fig.
This species is unique in that male and female respectively has yellow and black 1st flagellomere of antenna; intersexual color dimorphism in 1st flagellomere was observed only in this species among the studied species. This species resembles
Japan: On
On
On
A medium-sized yellow species (wing length 1.9–2.2 mm) having pruinose yellow scutum with a medial and two pairs of gray stripes, a black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a long tubercle-like seta, and inner-basally with a comb comprising 5–7 long fused tubercle-like setae. Larva mines the thallus of
The specific name (
Harusame-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in cool temperate deciduous forests dominated by
Japan: Hokkaido, Honshu (Fig.
This species resembles
Japan: 3♂1♀, Momiki, Izumi, Yatsushiro, Kumamoto Pref., 23-III-2015 (as larva), emerged on ?-VI-2015; 2♂, Yoro-keikoku, Otaki, Isumi, Chiba Pref., 17-III-2016 (as larva), emerged on 18–20-IV-2016; 1♂2♀, Amagi-toge, Izu, Shizuoka Pref., 19-IV-2012 (as larva), emerged on 8-V–3-VI-2021; 2♂2♀, Kuki, Owase, Mie Pref., 29-III-2019 (as larva), emerged on 9–30-IV-2019; 1♀, Takinohai, Kozagawa, Wakayama Pref., 13-IV-2014 (as larva), emerged on 19-IV-2014; 2♀, Wabuka, Kushimoto, Wakayama Pref., 4-V-2012 (as larva), emerged on 9-IV-2012; 7♂8♀, Narutaki, Ichiu, Tsurugi, Tokushima Pref., 31-III-2021 (as larva), emerged on 28-IV–20-V-2021; 1♂2♀, Yasui-keikoku, Niyodogawa, Agawa, Kochi Pref., 27-II-2011 (as larva), emerged on 25-IV-2011; 3♀, Mt. Kosho, Asakura, Fukuoka Pref., 11-IV-2010 (as larva), emerged on 1–13-V-2016; 1♂, Amagi-toge, Izu, Kaeda-keikoku, Kagamisu, Miyazaki, Miyazaki Pref. Pref., 11-IV-2021 (as larva), emerged on 19-IV-2021.
A medium-sized yellow species (wing length 1.9–2.0 mm) having pruinose yellow scutum with two pairs of pale brown stripes, a black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with an extended, apically flattened tubercle-like seta, and inner-basally with a comb comprising 3–5 long fused tubercle-like setae. Larva mines the thallus of
The color of the lateral stripes on the scutum varied among populations, but a geographical cline was not observed. The number of tubercle-like setae in a comb of the male epandrium varied from 5 to 6 among localities.
The specific name (
Kirisame-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in warm temperate evergreen forests dominated by
Japan: Honshu, Shikoku and Kyushu (Fig.
Locality records of three
This species resembles
Japan: On
On
On
A medium-sized yellow species (wing length 1.9–2.0 mm) having pruinose, entirely yellow scutum and scutellum, a black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with an extended, apically flattened tubercle-like seta, and inner-basally with a comb comprising 3–5 long fused tubercle-like setae. Larva mines the thallus of
The number of tubercle-like setae in a comb of the male epandrium varied from 3 to 5 among localities, with the specimens from northern populations and at high altitudes having fewer tubercle-like setae.
The specific name (
Kiiro-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are stream banks, mesic slopes and stone wall in cool temperate deciduous forests dominated by
Japan: Hokkaido, Honshu, Shikoku, Kyushu (Fig.
This species resembles
Japan: On
On
A medium-sized yellow species (wing length 1.8–2.1 mm) having a pruinose light yellow scutum and scutellum, a yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with an extended, apically flattened tubercle-like seta, and inner-basally with a comb comprising three or four long fused tubercle-like setae. Larva mines the thallus of
Female morphology and larval/adult ecology of
The number of tubercle-like setae in a comb of the male epandrium varied from 3 to 5 among localities, with the individuals from northern localities having fewer tubercle-like setae. Pigmentation pattern in distiphallus and morphology of paraphallus also differed between Hokkaido and Honshu populations.
The specific name (
Usuki-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in cool temperate deciduous forests dominated by
Japan: Hokkaido, Honshu (Fig.
This species resembles
Japan: On
On
On
A large yellow species (wing length 2.2–2.3 mm) having a shiny yellow scutum with a medial and two pairs of black stripes, a black 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with a hypertrophied tubercle-like seta, and inner-basally with a comb comprising three or four long fused tubercle-like setae. Larva mines the thallus of
The specific name (
Tsuyasuji-jagoke-hamoguribae.
Larvae construct linear mines in the midrib of the thallus, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in cool temperate deciduous forests dominated by
Japan: Hokkaido, Honshu, Shikoku and Kyushu (Fig.
Locality records of three
This species resembles
Japan: On
On
On
A medium-sized dark species (wing length 1.7–1.8 mm) having pruinose dark gray scutum, yellow scutellum, a black 1st flagellomere, dark maxillary palpus, dark halteres, and dark gray legs. Male epandrium inner-subdistally with a hypertrophied tubercle-like seta, and inner-basally with a comb comprising six or seven long fused tubercle-like setae. Larva mines the thallus of
Female morphology and larval ecology of
The morphology of the tubercle-like seta on the inner-distal margin of the male epandrium varied among localities from a simple seta to a flattened, basally enlarged acute spine. The relative position of a comb of tubercle-like setae and the separate tubercle-like seta neighboring the comb also varied among localities.
Mihikari-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in cool temperate deciduous forests dominated by
Japan: Hokkaido, Honshu (Fig.
This species was reported from Scotland by
Japan: On
A medium-sized dark species (wing length 1.8–1.9 mm) having pruinose dark gray scutum with mid-posterior yellow margin, yellow scutellum with dark lateral corners, black 1st flagellomere, dark maxillary palpus, dark halteres, and yellowish brown legs. Male epandrium inner-laterally with a long ventrally directed tubercle-like seta, and inner-basally with a siku-shaped comb comprising seven fused tubercle-like setae.
Larva mines the thallus of
Color pattern of scutum and subscutellum varied among localities. In specimens from Hachijo Island, the subscutellum had a large lateral dark corner.
The specific name (
Tomoshibi-jagoke-hamoguribae.
Larvae construct linear mines in the thallus in early instars, later entering the midrib, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in warm temperate evergreen forests dominated by
Japan: Hokkaido, Honshu, Hachijo Island, and Yaku Island (Fig.
This species resembles
Japan: On
On
On
On
A small dark species (wing length 1.3–1.7 mm) having a pruinose dark gray scutum with a mid-posterior yellow margin, a yellow scutellum with dark lateral corners, a black 1st flagellomere, dark maxillary palpus, dark halteres, and yellowish brown legs. Male epandrium inner-subdistally with a long ventrally directed tubercle-like seta, inner-laterally with a tubercle like seta, and inner-basally with a comb comprising five fused tubercle-like setae. Larva mines the thallus of all Japanese
Female morphology and larval/adult ecology of
The color pattern of the scutellum varied among localities; individuals at some localities had a larger dark corner on the scutum. The number of tubercle-like setae in a comb of the male epandrium varied from 5 to 6 among localities, among individuals in the same locality and even between right and left combs of an individual. The number of tubercle-like setae on the surstylus was consistently two, but the direction of each varied among localities.
The specific name refers to the larval feeding on
Komorebi-jagoke-hamoguribae.
Larvae construct linear mines in the midrib of the thallus, and pupate in the mines (Fig.
The habitats of this species are stream banks and mesic slopes in warm temperate evergreen forests dominated by
Japan: Hokkaido, Honshu, Shikoku, Kyushu, and Yaku Island (Fig.
Locality records of three
This species is the second smallest (next to
This species also resembles
Japan: 3♂3♀, Arakawa, higashi-son, Okinawa Pref., 10-XI-2021 (as larva on
A small dark species (wing length 1.3–1.5 mm) having a pruinose dark gray scutum and scutellum, a black 1st flagellomere, dark maxillary palpus, dark halteres, and brown legs. Male epandrium inner-laterally with a tubercle like seta, and inner-basally with a comb comprising six long fused tubercle-like setae. Larva mines the thallus of
The specific name honors a botanist, Dr. Kenji Suetsugu, who collected thalli of
Yanbaru-jagoke-hamoguribae.
(Fig.
The habitats of this species are stream banks in warm temperate evergreen forests dominated by
Japan: Amami and Okinawa Islands (Fig.
This species resembles
Japan: On
On
On
On
On
On
On
A small species (wing length 1.0–1.3 mm) having a pruinose grayish yellow scutum with a medial and two pairs of lateral dark gray stripes, a gray scutellum, yellow pleuron, black 1st flagellomere, dark maxillary palpus, yellowish gray halteres, and yellow legs. Male epandrium with dorso-ventrally bilobed surstylus; dorsal arm with two short tubercle-like setae. Male epandrium with bilobed, dorsoventrally elongated surstyli. Distiphalli tapering toward apex and bilaterally asymmetrical. Larva mines the thallus of
Female morphology and larval ecology of
Geographical variation was found in the presence of an acrostichal seta on the scutum; the seta was almost absent in individuals at most localities but present in those from the Okinawa Island.
The specific name refers to the host plant genus
Yosame-hatakegoke-hamoguribae.
(Fig.
The host liverwort species,
Japan: Honshu, Shikoku, Kyushu, Okinawa Is. and Iriomote Is. (Fig.
Locality records of three
This species resembles an European
This species is also closely related to another European species,
Among the Japanese species,
Japan; On
On
On
A small species (wing length 1.2–1.5 mm) having a pruinose grayish scutum with six longitudinal dark gray bands, a gray scutellum, brown 1st flagellomere, brown maxillary palpus, yellowish gray halteres, and yellow legs. Male epandrium inner-distally with a strong tubercle-like seta and inner-basally with a cluster of 29–35 dense tubercle-like setae. Distiphalli bilaterally asymmetrical, with left one tapering toward apex. Larva mines the thallus of
Adult male.
Female morphology and larval morphology/ecology of
The specific name (
Mutsusuji-hatakegoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus, and pupate in or out of the mines (Fig.
The host liverwort species,
Japan: Honshu (Fig.
This species is unique in that the scutum is six-banded (medial bands on the scutum are confluent in other species), and in the male epandrium having a clump (not a comb) of 25–30 short tubercle-like setae on the basal margin. This species resembles
Japan: On
On
A small species (wing length 1.1–1.3 mm) having a pruinose gray scutum and scutellum, brown 1st flagellomere, yellow maxillary palpus, gray halteres, and yellowish brown legs. Male scutum uniquely with a pair of bluish bands. Male epandrium inner-laterally with two strong tubercle-like setae, and with ventrally elongated surstylus. Distiphalli bilaterally asymmetrical and tapering toward apex. Larva mines the thallus of
Female morphology and larval morphology/ecology of
The specific name (
Aosuji-hatakegoke-hamoguribae.
(Fig.
The two host liverwort species grow on levees of paddy fields in subtropical islands (Fig.
Japan: Ishigaki and Iriomote Islands (Fig.
This species is unique in that the female has blue lateral bands on the scutum. It resembles
Japan: 2♂1♀, Mt. Osuzu, Tsuno, Miyazaki Pref., 14-VII-2021 (as larva), emerged on 31-VII–18-VIII-2021.
A small species (wing length 1.1–1.3 mm) having a subshiny black scutum, black scutellum with small yellow spot centrally, yellow 1st flagellomere, yellow maxillary palpus, yellow halteres, and yellow legs. Male epandrium inner-laterally with one strong tubercle-like seta. Distiphalli tapering toward apex, fused after meeting, elongated over the length of phallapodeme. Larva mines the thallus of a hornwort,
The specific name refers to the host plant genus
Miyabetsunogoke-hamoguribae.
Mines are extremely inapparent because the thalli are thick and often overlapping (Fig.
The habitat of this species is a cliff along a river bank in warm temperate evergreen forests dominated by
Japan: Kyushu (Fig.
Locality records of three
This species resembles
The three agromyzid species recorded from hornworts all had a dark scutum, but varied among species in color of antenna, color of maxillary palpus, and comb of tubercle-like setae on male epandrium. They also had common characteristics in the male genitalia; the distiphallus is little sclerotized, elongated, and tapering toward the apex. These characteristics of the male genitalia in hornwort-associated species suggest their monophyletic origin.
Japan: 8♂9♀, Kotonotaki, Susami, Wakayama Pref., 24-III-2020 (as larva), emerged on 31-I–2-V-2020; 16♂28♀, Yasukawa-keikoku, Tanabe, Wakayama Pref., 31-VII-2015 (as larva), emerged on 25-VIII–2-IX-2020; 1♀, Wadagawa-kyo, Kumanogawa, Shingu, Wakayama Pref., 7-VII-2021 (as larva), emerged on 18-VIII-2021; 9♂25♀, Tashiro, Kinko, Kimotsuki, Kagoshima Pref., 22-III-2015 (as larva), emerged on 30-IV–30-IV–10-V-2020; 1♀, Isso, Yaku Is., Kumage, Kagoshima Pref., 29-III-2017 (as larva), emerged on 12-V-2021.
A small species (wing length 1.2–1.4 mm) having a pruinose gray scutum and scutellum, black 1st flagellomere, black maxillary palpus, dark gray halteres, and dark gray legs. Male epandrium inner-basally with a comb comprising five or six fused long tubercle-like setae; surstylus with a comb comprising five or six fused long tubercle-like setae. Larva mines the thallus of a riparian hornwort,
Larval ecology of
The specific name refers to the host plant genus
Ananashitsunogoke-hamoguribae.
Larvae construct linear-blotch mines in the thallus, and pupate in and rarely out of the mines (Fig.
The habitats of this species are watersides along river banks or near water fall in warm temperate evergreen forests dominated by
Japan: Honshu, Kyushu and Yaku Island (Fig.
This species resembles
Japan: On
On
On
On
A small species (wing length 1.2–1.5 mm) that has a pruinose gray scutum and scutellum, brown 1st flagellomere, yellow maxillary palpus, brown halteres, and yellow legs. Male epandrium inner-laterally with a short tubercle-like seta. Distiphalli elongated, tapering toward apex, more than 2 × longer than the phallapodeme. the larvae mine thalli of hornworts belonging to the following genera:
Female morphology and larval morphology/ecology of
The specific name refers to its host plant genus
Niwatsunogoke-hamoguribae.
Four hornwort species belonging to 3 families were recorded to be host plants:
Larvae construct linear-blotch mines in the thallus, and pupate in the mines (Fig.
The main habitats of this species are the paddy fields that have not experienced land improvement projects or spraying with herbicides, as mentioned for
Japan: Honshu, Shikoku, Kyushu, the Ryukyu Archipelago (Fig.
This species resembles
Immature stages, but not adults, were reported from a hornwort,
Our extensive rearing of phytophagous insects on bryophytes has revealed that thalloid liverworts and hornworts in the Japanese Archipelago harbor 39 bryophyte-associated agromyzid species. This diversity was unexpectedly diverse because previously just one species was known as a liverwort thallus-miner. Our taxonomical study based on a vast collection of reared specimens has elucidated a great cryptic diversity of bryophyte-associated agromyzid species in the world.
The monophyly of the thallus-mining
The larvae of all of these species are thallus-miners; they pupate within mines unless the thalli are particularly thin and minute (e.g.,
Among the 39
Although we explored the species diversity of
Among the Japanese
Some species (e.g.,
The liverwort-associated
In addition to male genitalia, these
We thank A. Kawakita, K. Suetsugu, T. Nishioka, T. Ohgue, M. Igawa, A. Wong Sato, S. Sakurai, Y. Yamane, and T. Kato for their help on collection of bryophytes. We also thank O. Lonsdale, C. Eiseman and anonymous referees for helpful comments and invaluable advice to improve our manuscripts. One of the authors (MK) is indebted to late Prof. M. Sasakawa for his kind guidance on biology of