This species is similar to Psammoecus hiranoi Yoshida & Hirowatari, 2013, especially in darker specimens. However they can be distinguished by the ventral shape of the head. The temples of this species are immediately narrowed behind eyes, while those of Psammoecus hiranoi are widened behind the eyes and gradually narrowed toward the anterior margins.

Body length. 2.65–3.26 mm (n=48).

Coloration (Fig. 1A). Head and pronotum yellowish-brown to reddish-brown. Elytra black with yellowish-brown maculae; horizontal band at anterior 1/4, sometimes enlarged toward bases of elytra, round maculae at posterior 1/4, connected to lateral margins. Antennae yellowish-brown basally, 6th antennomere darker, 7th to 10th blackish-brown, 11th (apex) very bright.

Head (Fig. 3A, B, C). Broad, HW/HL 1.31–2.00; IE/HL 0.87–1.30. Temples immediately narrowed behind eyes, slightly incised at bases. Dorsal surface with comparatively coarse punctuation, ventral surface with very sparse punctures. Antennae 1.40–1.43 mm long, slender; covered relatively sparsely with long erect pubescence on each antennomere; approximate ratio of antennomeres of holotype as follows: 2.5: 1.1: 1.2: 1.3: 1.3: 1.3: 1.1: 1.2: 1.2: 1.0: 2.0 (Fig. 3A).

Pronotum (Figs 3B, C). Subquadrate, PW/PL 1.14–1.32. Punctuation on dorsal surface comparatively sparse, punctuation on ventral surface sparser than on dorsal surface. Pubescence composed of many short setae, and long setae on teeth on lateral margins and anterior and posterior angles. Anterior angles with a few very small protrusions, lateral margins with several small teeth, these teeth variable, slightly longer on lateral margins around anterior and posterior angles, and each posterior angle with very small teeth.

Elytra (Fig. 3E). Elongate-oval, EW/BL 0.38–0.51. Rows of punctures narrower than interstices. Pubescence composed of many semi-long medium length setae, and some long erect setae in a row around lateral margins, longer toward anterior portion.

9th abdominal sternite (Fig. 13A). Strut very long, narrow, cut at anterior 3/8, diverging gradually at posterior 1/8, branches comparatively small. Lateral sclerites elongate with small sclerites attached around posterior apical portion.

Aedeagus (Fig. 13B, C). Parameres broad, depressed at base, narrowed on inner margins in posterior half, incised around apex, punctuated very sparsely, a few long setae around apex, several short setae on lateral margins and posterior half of inner margins. Phallobase diverging in anterior half, each branch twisted at anterior 1/4. Penis relatively elongate, curved dorsally before apex.

Holotype: male, Kûra, Ishigaki Island, Okinawa Prefecture, Japan, 22–IX–2012, T. Yoshida leg. (EUMJ). Paratypes: [Kagoshima Pref.] 1 ex., Honcha, Amami-Ôshima Island, 23–VII–1962, N. Ohbayashi leg. (EUMJ); 1 ex., Nadakawa, Amami-Ôshima Island, 16–VII–1962, N. Ohbayashi leg. (EUMJ); 1 ex., Kamiya, Amami-Ôshima Island, 4–XI–1984, M. Tomokuni leg. (EUMJ). [Okinawa Pref.] 1 ex., Hiji, Kunigami Village, Okinawa Island, 15–20–VI–1994, K. Okada leg. (EUMJ); 1 male & 1 female, Takazato, Kunigami Village, Okinawa Island, 18–22–VI–2003, I. Tanaka leg. (ITC); 2 exs., Inamine, Nago City, Okinawa Island, 26–IX–2012, T. Yoshida leg. (ELKU); 1 ex., Shokubutsuen, Miyako Island, 3–III–1999, T. Mizoguchi leg. (EUMJ); 1 female, Kûra, Ishigaki Island, 22–IX–2012, T. Yoshida leg. (ELKU); 4 exs., Ôura Dam, Ishigaki Island, 7–X–2013, R. Itô leg. (ELKU); 2 exs., Mt. Yarabu-dake, Ishigaki Island, 26–III–2000, T. Kurihara leg. (EUMJ); 7 exs., same locality, 16–VI–2002, T. Watanabe leg. (ELKU); 5 exs., same locality, 8–X–2013, R. Itô leg. (ELKU); 1 male, same locality, 8–III–2006, I. Tanaka leg. (ITC); 1 female, Kabira, Ishigaki Island, 19–IV–2010, I. Tanaka leg. (ITC); 2 exs., Takeda Rindô, Ishigaki Island, 14–III–1999, T. Mizoguchi leg. (EUMJ); 2 exs., same locality, 16–VI–2002, T. Watanabe leg. (ELKU); 2 exs., same locality, 22–IX–2012, T. Yoshida leg. (ELKU); 6 exs., same locality, 23–IX–2012, T. Yoshida leg. (ELKU); 3 exs., Funaura, Iriomote Island, 18–III–2008, S. Yamamoto leg. (ELKU); 1 ex., Urauchi-gawa, Iriomote Island, 2–VII–1994, K. Okada leg. (EUMJ); 3 exs., Komi, Iriomote Island, 20–III–2008, S. Yamamoto leg. (ELKU); 1 ex., Ôtomi Rindô, Iriomote Island, 13–III–1999, T. Mizoguchi leg. (EUMJ); 1 ex., Mt. Inbi-dake, Yonaguni Island, 31–III–2009, S. Yamamoto leg. (ELKU).

JAPAN: Amami-Ôshima, Okinawa, Miyako, Ishigaki, Iriomote and Yonaguni Islands.

This species is found in piled dead leaves such as Japanese pampas grass Miscanthus sinensis Andersson, Banana Musa sp. and some other kinds of broad-leaved tree. This species is most common in the Nansei Islands.

The specific name means ‘pretty’ and ‘beautiful’. This new species has vivid coloration.

Psammoecus scitus sp. n. has been misidentified as Psammoecus quadrimaculatus Reitter, 1874 for a long time. Psammoecus quadrimaculatus was originally described from ‘Japonia’ (=Japan) by Reitter (1874). Sato (1989) first gave the Japanese name ‘Yotsumon-semaru-hiratamushi’ to Psammoecus quadrimaculatus. Hirano (2009) and Hirano (2010) provided figures of this species as Psammoecus quadrimaculatus. However, examination of the holotype of Psammoecus quadrimaculatus deposited in BMNH revealed that external characters such as the shape of the teeth on the lateral margins of the pronotum and the male genital structure are distinctly different.

This species is similar to Psammoecus trimaculatus and Psammoecus boreas sp. n., but can be distinguished from the former by the shorter teeth on the lateral margins of the pronotum and from the latter by the shorter body length and the shorter antennae.

Body length. 2.25–2.83 mm (n=9).

Coloration (Fig. 1B). Head dark brown, gradually darker toward anterior. Pronotum yellowish-brown to reddish-brown. Elytra yellowish-brown with round dark maculae on posterior 1/3, elytral suture dark in posterior half. Elytra of lighter colored specimens with reduced dark maculae and dark areas of elytral suture. Antennae yellowish-brown basally, 9th, 10th and posterior end of 8th antennomeres blackish-brown, 11th (apex) yellowish-brown.

Head (Fig. 4A, B, C). Triangular, HW/HL 1.55–2.06; IE/HL 1.03–1.33. Dorsal surface with strong dense punctures, temples and posterior ventral surface also with irregularly dense punctures. Temples well enlarged, immediately narrowing thereafter. Eyes prominent, relatively small. Antennae 1.09–1.20 mm, comparatively thick, short; covered with medium length erect pubescence on each antennomere; approximate ratio of one of examined specimens as follows: 2.6: 1.1: 1.2: 1.1: 1.2: 1.2: 1.1: 1.0: 1.1: 1.0: 1.8 (Fig. 4A).

Pronotum (Fig. 4B, C). Roundly subquadrate, PW/PL 1.16–1.24. Dorsal surface with strong moderately dense punctuation, denser than that on ventral surface; some punctures on ventral surface in rows. Pubescence composed of many short setae on dorsal surface and several medium length setae on each lateral margin, distance between medium length setae approximately regular. Each anterior angle with several very small teeth, each lateral margin with four similarly sized small teeth, the distance between them irregular and variable; each posterior angle with a small tooth, similar in size to those on lateral margins.

Elytra (Fig. 4E). Elongate, EW/BL 0.36–0.47. Rows of punctures almost same width as interstices. Pubescence composed of numerous comparatively short setae, without long setae.

9th abdominal sternite (Fig. 13D). Strut Y-shaped, cut at anterior 1/3, diverging in posterior 1/4, branches relatively wide, ends of each branch pointed and curved inwards. Lateral sclerites elongate, membranous.

Aedeagus (Figs 13E and F). Parameres club-shaped, wide and thick at bases, each apical portion with a long seta, stick-shaped portions with sparse punctuation and several short setae, lateral portions and inner margins of bases with successions of dense punctures, inner margin of bases with numerous short setae. Phallobase enlarged toward posterior margin from posterior 1/3, anterior margin roundly incised, anterior portion extended. Penis flat, wide, narrowed gradually toward apex, apical portion pointed, apical portion of dorsal part with sparse punctures.

JAPAN: [Hokkaido Pref.] 1 male, Takkobu-numa Lake, Kushiro City, 1–IX–2006, T. Yoshida leg. (ELKU); 1 ex., Iwabokki, Kushiro City, 25–VIII–1990, M. Sakai leg. (EUMJ); 6 exs., same locality, 26–VIII–1990, M. Sakai leg. (EUMJ); 1 ex., same locality, 29–VIII–1990, M. Sakai leg. (EUMJ).

JAPAN: Hokkaido; Africa; Europe; Russia; Turkestan.

This species is known to inhabit marshland. Warne (1963) reported that large numbers of this species were found on Carex.

The habitat and distribution of this species are unusual in the genus Psammoecus. This species inhabits marshland and is distributed in comparatively high latitudes where species richness of Psammoecus is poor. Coloration is variable. In the past, four aberrations and two varieties (e.g. Lucas 1843; Gerhardt 1912) were described.

This species is closely similar to Psammoecus triguttatus and Psammoecus labyrinthicus sp. n. but it can be distinguished from the former by the larger basal parameres and the broader distance between the posterior margin of the phallobase and the deepest point of incision of the anterior margin of the phallobase, and from the latter by the larger bases of parameres and the wider protuberances on the inner margins of the branches of the anterior phallobase. The rows of punctures on the elytra of Psammoecus trimaculatus (Fig. 5E) are comparatively narrower than in these two closely similar species. However, their external characters, especially their coloration, are variable. These features may cause misidentification.

Body length. 2.32–2.96 mm (n=38).

9th abdominal sternite (Fig. 13G). Strut Y-shaped, elongate, cut at anterior 3/8, diverging in posterior 1/5, branches relatively wide, narrowed gradually toward apex, ends of each branch curved inward. Lateral sclerites elongate, membranous.

Aedeagus (Fig. 13H, I). Parameres hatchet-shaped, inner margin around bases pointed, punctuated sparsely on stick-shaped portions, densely on anterior portions, a long seta at each apex, numerous short setae on stick-shaped portions and around inner margins of anterior parts. Posterior margin of phallobase rounded, phallobase roundly hollowed, its anterior branches markedly protruding inward. Penis gradually narrowed toward apex, posterior 1/9 flattened, punctuated sparsely on posterior 1/9.

India: [Andhra Pradesh State] 1 ex., Hyderabad, 18–X–1969, T. Ishihara leg. (EUMJ). JAPAN: [Hokkaido Pref.] 1 male, Suehiro, Asahikawa City, 18–IX–2013, T. Yoshida leg. (ELKU); 1 male, Aizankei, Kamikawa Town, 2–IX–1977, A. Oda leg. (EUMJ); 1 ex., Sôunkyo, Kamikawa Town, 4–IX–1977, A. Oda leg. (EUMJ); 1 ex., Chûbisei, Memuro Town, 25–VIII–1995, S. Hisamatsu leg. (EUMJ); 2 exs., Fushimi Marsh, Memuro Town, 25–VIII–1995, S. Hisamatsu leg. (EUMJ). [Aomori Pref.] 1 male, Namioka Ôaza Yoshinoda, Aomori City, 16–IX–2012, K. Ikeuchi leg. (ELKU). [Gifu Pref.] 1 male, Tentoku, Mizunami City, 31–VII–2009, K. Itô leg. (ELKU). [Hyogo Pref.] 1 male, Mt. Mikusa-yama, Inagawa Town, 27–VI–1991, K. Ikeuchi leg. (ELKU). [Wakayama Pref.] 2 males, 3 females & 7 exs., Biwadani, Kinokawa City, 25–IV–2012, T. Yoshida leg. (ELKU). [Kagawa Pref.] 1 male, Yoshima Island, Sakaide City, 10–12–IX–1973, S. Kinoshita leg. (EUMJ). [Tokushima Pref.] 2 males, 2 females & 4 exs., Kamojima-chô, Yoshinogawa City, 4–III–2012, T. Yoshida leg. (ELKU). [Kochi Pref.] 1 male, Godaisan-Park, Kochi City, 21–V–1983, K. Ishida leg. (EUMJ). [Fukuoka Pref.] 1 male, Kitano-chô, Kurume City, 1–VII–2013, T. Yoshida leg. (ELKU). [Kagoshima Pref.] 1 male, Kotoku-gawa, Amami-Ôshima Island, 14–III–1988, M. Satô leg. (EUMJ). [Okinawa Pref.] 1 male, Yona, Kunigami Village, Okinawa Island, 17–VII–1965, Y. Hori leg. (EUMJ); 1 male, same locality, 19–X–1987, M. Sakai leg. (EUMJ); 1 male & 1 ex., Inaba, Iriomote Island, 9–VIII–1962, M. Satô & Y. Arita leg. (EUMJ).

JAPAN: Hokkaido, Honshu, Shikoku, Kyushu, Amami-Ôshima, Okinawa, and Iriomote Islands. (Madagascar recorded by Karner (2012); Nepal, India and Bhutan recorded by Pal (1980).)

This species was redescribed by Pal (1980) with a description of the larva. It is distributed worldwide except for Europe and North America and common at least in Japan and India (Hirano 2009; Karner 2012; Pal 1980).

Hirano (2009) and Hirano (2010) illustrated this species as Psammoecus triguttatus and suggested that the distribution of Psammoecus trimaculatus in Japan is doubtful. In this paper, it is regarded as Psammoecus trimaculatus on the basis of comparison of the male genital structure of the species illustrated by Pal (1985).

Pal (1985) gave as the distribution of this species Madagascar, Sri Lanka, Nepal, India, Malaysia, Burma, New Guinea, Australia and Japan, synonymizing the following species with Psammoecus trimaculatus: Telephanus cruciger Waterhouse, 1876 and Psammoecus cephalotes Grouvelle, 1919 from New Guinea, Cucujus incommodus Walker, 1859 from Sri Lanka and Psammoecus ypsilon Blackburn, 1903 from Australia (Walker 1859; Waterhouse 1876; Blackburn 1903; Grouvelle 1919). In addition, Thomas and Yamamoto (2007) recorded this species from Brazil. However, in view of the evidence that at least two other closely similar species have been confused with this species, past records of this species should be reexamined. The specimen figured by Thomas and Yamamoto (2007) seems to be another species. In this paper, records of this species from outside Japan are recognised from label data described by Pal (1980) and Karner (2012), on the basis that they examined the male genital structure.

This species is closely similar to Psammoecus trimaculatus and Psammoecus labyrinthicus sp. n. Morphological differences among these species were stated in diagnosis of Psammoecus trimaculatus.

Description. Body length. 2.32–2.93 mm (n=23).

Coloration (Fig. 1D). Head and pronotum yellowish-brown to reddish-brown. Elytra yellowish-brown with variable dark maculae: round ones at half, oval ones on the posterior half to posterior 1/4 of elytral suture, sometimes connected with maculae at half, darkened around end of elytra. Antennae reddish-brown basally, blackish-brown from 8th to 10th antennomeres, 11th (apex) comparatively bright.

Head (Fig. 6A, B, C). Triangular, HW/HL 1.46–1.90; IE/HL 0.92–1.20. Temples slightly expanded behind eyes, narrowed toward posterior. Eyes large, prominent. Dorsal surface with relatively dense punctuation. Antennae 1.06–1.38 mm, moderately long; covered with considerable medium length or long semi-erect setae and short pubescence on each antennomere; approximate ratio of lectotype as follows: 2.7: 1.0: 1.1: 1.2: 1.3: 1.3: 1.3: 1.1: 1.1: 1.1: 2.0 (Fig. 6A).

Pronotum (Fig. 6B, C). Broad, PW/PL 1.31–1.50. Dorsal surface with relatively dense punctuation. Pubescence composed of relatively fine short or medium length setae, a long seta on each tooth on lateral margins and anterior and posterior angles. Each anterior angle with four or five small teeth; each lateral margin with four teeth, tooth I small, tooth II longer than tooth I, teeth III and IV relatively narrow and of almost identical size, longer than tooth II, tooth V smallest or absent; each posterior angle with a few very small protuberances of variable shape.

Elytra (Fig. 6E). Elongate-oval, EW/BL 0.39–0.45. Rows of punctures narrower than interstices. Pubescence composed of many semi-erect setae of medium length; several long setae in a row around anterolateral margins.

9th abdominal sternite (Fig. 13J). Strut Y-shaped, cut at anterior 2/5. Lateral sclerites curved inward, relatively elongate.

Aedeagus (Fig. 13K, L). Parameres club-shaped, wide around bases, posterior inner margins of wide portions a little prominent, punctuation on narrow portions relatively sparse, a little denser on wide portions, posterior half of inner margins of wide portions with many setae, narrow portions with several sparse setae, apex with a long seta. Posterior half of phallobase wider towards posterior margin, distance between posterior margin and deepest point of incision of margin of upper layer narrow, protuberances around anterior 1/4 pointed inwards, posterior margin of lower layer widely deeply incised. Penis relatively flat and thin, narrowed toward apex, punctuation around apex denser toward apex.

Lectotype: male, Nagasaki, Nagasaki Prefecture, Japan, 1869, G. Lewis leg. (BMNH). Paralectotype: 1 female, same locality, 1869, G. Lewis leg. (BMNH). (here designated)

JAPAM: [Tokyo Pref.] 1 male, Hatagaya, Shibuya-ku, 30–VII–1956, K. Tanaka leg. (EUMJ). [Gifu Pref.] 1 male, Tentoku, Toki-chô, Mizunami City, 6–VIII–2010, K. Itô leg. (ELKU); 1 male, Konokure, Toki-chô, Mizunami City, 16–VII–2011, K. Itô leg. (ELKU). [Kagoshima Pref.] 1 male, 4 females and 3exs., Sata Cape, Minamiôsumi Town, 6–VII–1968, K. Suga leg. (HUSE); 1 male, Nakanoshima Island, Toshima Village, 9–VII–1969, M. Satô leg. (EUMJ); 1 male, same locality, 26–VI–1965, S. Fukuda leg. (HUSE). [Okinawa Pref.] 1 male, Aha beach, Kunigami Village, Okinawa Island, 23–I–2008, Y. Hirano leg. (YHC); 2 males and 2 exs., Motobu Town, Okinawa Island, 19–VIII–2008, T. Yoshida leg. (ELKU); 1 male, Fusato, Nanjô City, Okinawa Island, 5–VI–1970, M. Takagi leg. (EUMJ); 1 male, Mt. Omoto-san, Ishigaki Island, 1–VII–1965, Y. Hori leg. (EUMJ); 1 male, Ushuku-no-mori, Iriomote Island, 26–VI–1965, Y. Hori leg. (EUMJ).

JAPAN: Honshu, Kyushu, Nakanoshima (Tokara Islands), Okinawa, Ishigaki and Iriomote Islands.

Psammoecus sp. 4 illustrated by Hirano (2009) and Hirano (2010) seems to be this species or Psammoecus labyrinthicus sp. n. However, identification is difficult, because the specimen is a female. The Japanese name of Psammoecus sp. 4 given by Hirano (2009) and Hirano (2010) “Nise-mitsumon-semaru-hiratamushi” is adopted as the Japanese name of this species in this paper.

Halstead et al. (2007) recorded distribution of this species from Russia (Far East), Korea, China and Japan. However, it is now clear that at least two closely similar species occur in Japan, so, past records of this species should be reconfirmed. In addition, two or more closely similar species were sometimes found in the same limited area such as Tentoku, Toki-cho, Mizunami City, Gifu Prefecture where Psammoecus triguttatus and Psammoecus trimaculatus were collected. Thus, these species should be identified carefully.

Syntypes of this species consist of two specimens, one male and one female. The male specimen is designated as a lectotype, and the female specimen is designated as a paralectotype.

This species is closely similar to Psammoecus trimaculatus and Psammoecus triguttatus. However, it can be distinguished by the male genital structure, especially the shape of the parameres, and the comparatively dense punctuation of the dorsal pronotum.

Body length: 2.66–3.38 mm (n=25).

Coloration (Fig. 1E). Head and pronotum reddish-brown. Elytra yellowish-brown with dark maculae; round ones at half, oblong ones on the posterior half to posterior 1/4 along elytral suture, darkened around end of elytra: these maculae sometimes connected each other. Antennae reddish-brown basally, 7th antennomere darkened, from 8th to 10th blackish-brown, 11th (apex) comparatively bright.

Head (Fig. 7A, B, C). Rounded-triangular, HW/HL 1.25–2.00; IE/HL 0.75–1.27. Temples slightly expanded behind eyes, gradually narrowed. Eyes large, prominent. Dorsal surface with dense and strong punctuation. Antennae 1.35–1.56 mm, moderately long; covered with medium length semi-erect pubescence and some long erect setae on each antennomere; approximate ratio of holotype as follows: 2.4: 1.0: 1.0: 1.2: 1.3: 1.1: 1.1: 1.1: 1.1: 1.1: 1.9 (Fig. 7A).

Pronotum (Fig. 7B, C). Roundly subquadrate, PW/PL 1.25–1.53. Relatively strong and dense punctuation on dorsal surface. Pubescence composed of short setae, a long seta on each tooth on lateral margins and anterior and posterior angles. Each anterior angle with several small teeth, each lateral margin with four short teeth; tooth I small, tooth II longer than tooth I, teeth III and IV almost the same size, longer than tooth II, teeth II, III and IV broadened at base, each posterior angle with a very small tooth, the shape of these teeth variable.

Elytra (Fig. 7E). Elongate-oval, EW/BL 0.41–0.47. Rows of punctures narrower than interstices. Pubescence composed of numerous medium length semi-erect setae and some long setae in a row around lateral margins.

9th abdominal sternite (Fig. 14A). Strut relatively short, cut at anterior 1/3. Lateral sclerites fused with strut, curved inwards.

Aedeagus (Fig. 14B, C). Parameres stout, club-shaped, punctuated on anterior half of inner margins, anterolateral portions well punctuated, anterior half of inner margins with many setae of variable size, apex with a long seta. Phallobase broadened toward posterior margin, posterior margin widely incised, protuberances around 1/3 of inner margins of branches, especially right protuberance thin. Penis flat, narrowed toward apex, apex pointed, punctuated around anterior 1/8, apex densely punctuated.

Holotype: male, Hachijô Island, Hachijô Town, Tokyo Prefecture, Japan, 21–VII–1957, S. Hisamatsu leg. (EUMJ). Paratypes: [Tokyo Pref.] 3 females, same data as holotype. (EUMJ). [Mie Pref.] 4 males, Ujitachi-chô, Ise City, 29–VIII–2009, N. Narukawa leg. (ELKU). [Nagasaki Pref.] 1 male and 11 exs., Ta, Toyotama-chô, Tsushima Island, 7–VII–2013, T. Yoshida leg. (ELKU). [Kagoshima Pref.] 2 males and 3 exs., Takarajima Island, Toshima Village, 2–VII–1960, M. Satô leg. (EUMJ); 1 ex., same locality, 3–VII–1960, M. Satô leg. (EUMJ); 1 ex., same locality, 1–VI–1962, M. Satô leg. (EUMJ).

JAPAN: Honshu, Hachijô (Izu Islands), Tsushima, Takarajima Islands (Tokara Islands).

The first author (Yoshida) collected many individuals of this new species from dead leaves of evergreen trees in Tsushima Island, Nagasaki Prefecture (Fig. 15A).

The specific name means ‘labyrinthine’. Identification of Psammoecus trimaculatus and closely similar species is very difficult. The addition of this new species to this group of closely similar species made their identification more difficult like the labyrinth.

Psammoecus trimaculatus is closely related to this species, however the former has been often collected from dead leaves of monocotyledon plant. These two species may have each distinct ecological trait.

This species is similar to Psammoecus trimaculatus and other species closely similar to Psammoecus trimaculatus. Nakane (1963) provided a figure of this species as Psammoecus triguttatus. It differs from the aforementioned species by the shorter lateral teeth of the pronotum. It is also very similar to Psammoecus harmandi Grouvelle, 1912 both in external characters and male genital structure as illustrated by Pal (1980), but can be distinguished from it by the longer antennae and the comparatively oblong 10th antennomere.

Body length. 2.74–3.27 mm (n=19).

Coloration (Fig. 2A). Head and pronotum yellowish-brown. Elytra yellowish-brown with dark brown maculae at half, oval ones at center of each elytron and dark oblique bands toward posterior portion, sometimes connected at elytral suture, forming a V-shaped band. Elytra of lighter color specimens with reduced maculae, oval ones and bands separated. Antennae yellowish-brown basally, posterior ends of 8th to 10th antennomeres darkened, or all antennomeres yellowish-brown in lighter colored specimens.

Head (Fig. 8A, B, C). Broad, HW/HL 1.35–1.65; IE/HL 0.96–1.13. Temples narrowed around each base. Eyes small, moderately rounded. Dorsal surface with moderately dense punctuation, ventral surface punctuated sparsely. Antennae 1.54–1.71 mm, thin, very long; covered with considerable long semi-erect pubescence on each antennomere; approximate ratio of holotype as follows: 2.6: 1.0: 1.1: 1.2: 1.2: 1.3: 1.2: 1.0: 1.0: 1.1: 1.7 (Fig. 8A).

Pronotum (Fig. 8B, C). Subquadrate, PW/PL 1.18–1.30. Dorsolateral portions lightly impressed. Dorsal surface with relatively dense punctuation, no punctures around posterior margin, comparatively sparse punctuation on ventral surface. Pubescence composed of many short setae and fine long setae on teeth on lateral margins and anterior and posterior angles. Each anterior angle with a distinct group of a few very small teeth, each lateral margin with four small teeth of almost the same size, each posterior angle with a small tooth, almost the same size as those on lateral margins.

Elytra (Fig. 8E). Oval, EW/BL 0.32–0.45. Rows of punctures wider than interstices. Pubescence composed of many short setae, no long setae.

9th abdominal sternite (Fig. 14D). Strut Y-shaped, cut deeply at anterior 1/5, diverging for posterior 1/4. Lateral sclerites elongate.

Aedeagus (Fig. 14E, F). Parameres cone-shaped with almost even sparse punctuation, sparser on bases, a few long setae around apical portions, a few short setae distributed sparsely. Phallobase tube-like, consisting of two layers, anterior margin rounded, dorsal surface around anterior margin thin, protuberances of upper layer directed towards anterior portion, small protuberances at beginning of divergence of upper layer. Penis stout, punctuated on posterior 1/9, rather coarsely on ventral surface.

Holotype: male, Yoshin, Tanzawa, Kanagawa Prefecture, Japan, 26–V–1989, Y. Hirano leg. (EUMJ). Paratypes: [Hokkaido Pref.] 2 exs., Chûbisei, Memuro Town, 25–VIII–1995, S. Hisamatsu leg. (EUMJ); 1 ex., Mt. Sapporo-dake, Sapporo City, 5–VIII–1970, S. Kinoshita leg. (EUMJ). [Niigata Pref.] 1 ex., Mikuni Touge, 1–VII–1967, K. Baba leg. (HUSE). [Kanagawa Pref.] 1 ex., Yoshin, Tanzawa, 26–V–1989, Y. Hirano leg. (YHC). [Nagano Pref.] 1 ex., Tokugo Touge, 29–VII–1955, T. Nakane leg. (HUSE); 5 exs., Ôbora, Ueda City, 24–VII–2013, T. Yoshida leg. (ELKU). [Oita Pref.] 6 ex., Mt. Sobo-san, 7–VI–2009, S. Yamamoto leg. (ELKU).

JAPAN: Hokkaido, Honshu, Kyushu.

The specific name is from the god of the north wind of ancient Greek mythology. Most Psammoecus species are distributed in tropical or subtropical zones, however, this new species is exceptionally distributed in Hokkaido or on mountains of high altitude located in Honshu and Kyushu.

Psammoecus sp. 3 illustrated by Hirano (2009) and Hirano (2010) was conspecific with this species and named same Japanese name proposed by him.

This species is similar to Psammoecus trimaculatus and other species closely similar to Psammoecus trimaculatus. However, it can be distinguished from them by the male genital structure, especially the parameres which are fused to the phallobase. The distinguishing external characters of this species are tooth IV of the lateral margins of pronotum extended in a posterolateral direction and the long antennae, especially the 1st antennomere.

Body length. 2.71–2.77 mm (n=2).

Coloration (Fig. 2B). Head and pronotum yellowish-brown to reddish-brown. Elytra yellowish-brown with dark maculae, round ones at half of each elytron, black ones on posterior half of elytral suture narrower toward posterior elytral suture, a thin short black band between these maculae. Antennae yellowish-brown, almost unicolorous, 11th (apical) antennomere comparatively bright.

Head (Fig. 9A, B, C). Transverse, HW/HL 1.59–1.63; IE/HL 1.07. Temples well enlarged behind eyes, narrowed at bases. Eyes small, moderately rounded. Frequent pubescence and very sparse punctuation on ventral surface, moderately dense punctuation on dorsal surface. Antennae 1.44–1.49 mm long; covered with frequent medium length semi-erect fine setae and some long erect setae on each antennomere; approximate ratio of holotype as follows: 2.6: 1.0: 1.2: 1.4: 1.4: 1.4: 1.3: 1.1: 1.2: 1.0: 1.9 (Fig. 9A).

Pronotum (Fig. 9B, C). Transverse, PW/PL 1.30–1.35. Punctuation on dorsal surface coarse. Pubescence composed of short fine setae, a long seta on each tooth on lateral margins and anterior and posterior angles. Each anterior angle with several very small prominences and two small teeth, each lateral margin with four teeth; tooth I comparatively small, teeth II and III almost the same size, longer than tooth I, tooth IV extended in posterolateral direction, longer than tooth II and III, each posterior angle with a very small protuberance.

Elytra (Fig. 9E). Oval, EW/BL 0.37–0.44. Around half of lateral margins relatively expanded. Rows of punctures wider than interstices. Pubescence composed of medium length semi-erect setae and a few long setae around anterolateral margins.

9th abdominal sternite (Fig. 14G) Strut Y-shaped, anterior 2/7 slightly thickened, cut from anterior 2/7, diverging in posterior 1/3, branches long, ends of each branch curved inwards. Lateral sclerites elongate, membranous.

Aedeagus (Figs 14H and I) Parameres fused with phallobase, incised shallowly at inner and lateral margins of bases, relatively stout, gradually curved inwards, three long setae around posterior 1/3, some short setae on posterior half, well punctuated on anterior half of inner margins. Phallobase broad and somewhat flat, anterior margin widely but shallowly incised, lateral margins slightly depressed around anterior 1/4. Penis wide, thinner toward apex, narrowed around apex, punctuated around apex, densely punctuated at apex.

Holotype: male, Mt. Omoto-san, Ishigaki Island, Okinawa Prefecture, Japan, 23–III–2000, T. Kurihara leg. (EUMJ). Paratype: [Okinawa Pref.] 1 male, same data as holotype, T. Kurihara leg. (EUMJ).

JAPAN: Ishigaki Island.

The specific name is derived from the type locality of the new species, Mt. omoto-san, Ishigaki Island, Okinawa Prefecture.

This species is distinguished from other Japanese Psammoecus species by the black elytra with yellow maculae and the posterior teeth on lateral margins of pronotum longer than those of anterior margins.

Body length. 2.24–2.56 mm (n=8).

9th abdominal sternite (Fig. 14J). Strut comparatively short, cut at anterior 2/5, diverging gradually, branches comparatively long and wide. Lateral sclerites elongate, slightly curved inwards, apical sclerites pointed.

Aedeagus (Fig. 14K, L). Parameres fused with phallobase, wide, short, posterior and lateral portions punctuated sparsely, densely punctuated triangular areas in anterior portions, a few long setae and short setae on posterior half, several short folds around beginning of divergence of parameres, a few lines on upper densely punctuated areas. Phallobase stout, broad in lateral aspect, anterior margin straight, a few subparallel lines on posterior dorsal portion. Penis comparatively broad, flattened around apex, apical margin rounded, punctuated sparsely on apex.

JAPAN: [Okinawa Pref.] 1 male, 4 females & 2 exs., Inamine, Nago City, Okinawa Island, 26–IX–2012, T. Yoshida leg. (ELKU); 1 ex., Mt. Yarabu-dake, Ishigaki Island, 16–VI–2002, T. Watanabe leg. (ELKU).

JAPAN: Okinawa, Hateruma and Ishigaki Islands; Madagascar; India; Sri Lanka; Malaysia; Indonesia; Philippines.

This species is found in dead leaves to which fungi are attached and occurs sympatrically with other Silvanid species such as Psammoecus trimaculatus, Psammoecus scitus sp. n., Cryptamorpha desjardinsi (Guérin-Méneville, 1844) and Monanus concinnulus (Walker, 1858) (Fig. 15B).

Pal (1980) and Karner (2012) redescribed this species. Grouvelle (1892) described it as ‘simonis’. However, Grouvelle (1908) referred to it with the specific name ‘simoni’. Pal (1980), Hirano (2010) and Karner (2012) used the latter spelling, which we adopt in the present study.

This species is closely similar to Psammoecus hiranoi. It can be distinguished by the longer parameres, the shallower incision of the phallobase on the anterior 1/4 and the shorter teeth on the lateral margins of the pronotum.

Body length. 2.75–3.50 mm (n=49).

Coloration (Fig. 2D). Head and pronotum yellowish-brown to reddish-brown. Elytra with a variable horizontal dark band at middle, this band in lighter color specimens thinner towards elytral suture, lateral margins, apical portion and posterior elytral suture dark. Antennae brown or reddish-brown basally, 7th and 8th antennomeres darker, 9th and 10th almost black, 11th (apex) lighter than basal ones. Antennae of some specimens almost unicolorous.

Head (Fig. 11A, B, C). Broad, HW/HL 1.40–2.08; IE/HL 0.94–1.38. Temples enlarged behind eyes, narrowed gradually toward posterior margin. Eyes large, slightly prominent. Ventral surface with rough irregular punctuation, sparser punctuation on dorsal surface. Antennae 1.38–1.57 mm, 2nd antennomere short; covered with thick medium length pubescence and some long erect pubescence on each antennomere; approximate ratio of lectotype as follows: 2.5: 1.0: 1.3: 1.3: 1.3: 1.3: 1.3: 1.3: 1.3: 1.2: 1.8 (Fig. 11A).

Pronotum (Fig. 11B, C). Broad, PW/PL 0.69–1.52. Punctuation on dorsal surface very dense, but no punctures around posterior margin, punctuation on ventral surface sparser than dorsal surface, some of them in rows especially on posterior half. Pubescence composed of frequent short setae on dorsal surface and a long seta on each tooth on lateral margins and at anterior and posterior angles. Each anterior angle with a distinct group of a few very small teeth, each lateral margin with four teeth; tooth I comparatively small, teeth II, III and IV almost the same moderate size, each posterior angle with a comparatively small tooth.

Elytra (Fig. 11E). Elongate-oval, EW/BL 0.41–0.47. Punctures wider than interstices. Pubescence composed of semi-erect medium length setae and some long erect setae in a row around lateral margins, longer toward anterior portion.

9th abdominal sternite (Fig. 14M). Strut cut at around half, diverging widely around the posterior base, ends of each branch curved inwards. Lateral sclerites triangular, flat.

Aedeagus (Figs 14N and O). Parameres simple stick-shaped, elongate, slightly curved inwards, punctuated sparsely except base, a few semi-long setae around apical portion. Phallobase diverging gradually around anterior 1/4, a little bulging along inner margins of each branch, lateral margins broadening. Penis also elongate, punctuated on posterior 1/6, more densely on ventral portion, especially densely on apical portion.

Lectotype: male, Mt. Maya-san, Kobe City, Hyogo Prefecture, Japan, 1871, G. Lewis leg. (BMNH). Paralectotype: 1 female, same locality, 1871, G. Lewis leg. (BMNH). (here designated)

Russia: [Primorsky Krai] 1 ex., Chuguevka Village, 27–VI–1999, Y. Notsu leg. (EUMJ). JAPAN: [Niigata Pref.] 1 male & 1 ex., Renge-Onsen, Itoigawa City, 14–VIII–2003, I. Tanaka leg. (ITC). [Tokyo Pref.] 7 exs., Mikura Island, Mikurajima Village, 11–VI–1977, T. Nakane leg. (HUSE). [Nagano Pref.] 1 ex., Togakushi-Kôgen, Nagano City, 5–6–VII–2008, K. Mizuno leg. (KMC); 2 ex., Ôbora, Ueda City, 24–VII–2013, T. Yoshida leg. (ELKU). [Kyoto Pref.] 1 ex., Kifune, Kyoto City, 6–IV–1988, I. Tanaka leg. (ITC). [Hyogo Pref.] 1 ex., Itoi, Asago City, 26–IX–2004, I. Tanaka leg. (ITC); 5 exs., Sakanotani, Shisô City, 19–VII–2009, K. Itô leg. (KMC); 1 ex., Onzui, Shisô City, 19–VII–2009, K. Utô leg. (KMC); 1 ex., Mt. Yuzuruha-san, Nantan Town, Awaji Island, 18–X–2003, I. Tanaka leg. (ITC). [Nara Pref.] 1 ex., Mt. Kasuga-yama, Nara City, 5–V–1985, I. Tanaka leg. (ITC); 2 exs., same locality, 5–V–2012, K. Ikeuchi leg. (ELKU); 1 ex., same locality, 31–VII–2012, K. Matsuda leg. (ELKU). [Tokushima Pref.] 1 ex., Mt. Shibakoya-yama, Kamiyama Town, 31–VII–1975, M. Yoshida leg. (EUMJ). [Ehime Pref.] 1 ex., Nishidani, Yanadani Village, 15–16–VII–1994, K. Aita leg. (EUMJ). [Kochi Pref.] 1 ex., Muroto Cape, Muroto City, 7–VI–1959, S. Hisamatsu leg. (EUMJ). [Nagasaki Pref.] 1 ex., Mt. Tatera-san, Tsushima Island 16–IX–1995, N. Narukawa leg. (ELKU); 3 exs., Midake, 12–X–1977, Y. Notsu leg. (EUMJ). [Kagoshima Pref.] 1 male & 4 exs., Kirishima, 29–VII–1971, T. Nakane leg. (HUSE); 2 exs., Shiratani-unsuikyô, Yakushima Island, 15–VIII–2005, J. Ogawa leg. (EUMJ); 1 male & 1 ex., Onoaida Hodô, Yakushima Island, 14–15–IX–2002, N. Ohbayashi leg. (EUMJ); 2 exs., Kosugidani, Yakushima Island, 2–V–1984, K. Mizuno leg. (OMNH); 4 exs., Anbô, Yakushima Island, 16–VII–1989, N. Narukawa leg. (ELKU); 5 exs., Nogi, Nishino-omote, Tanegashima Island, 10–VII–1974, S. Hisamatsu leg. (EUMJ).

JAPAN: Hokkaido, Honshu, Shikoku, Kyushu, Mikura (Izu Islands), Awaji, Yakushima and Tanegashima Islands; Burma; Korea; Russia.

The first author (Yoshida) collected this species from various kinds of dead leaves from evergreen trees, deciduous trees and bamboo.

The syntypes of this species consist of three specimens: two of them belong to this species, but the remaining one represents Psammoecus trimaculatus or Psammoecus triguttatus. We designate a male specimen as lectotype, and a female specimen as paralectotype. The type specimens were collected from Mt. Maya-san, Kobe City, Hyogo.

This species is closely similar to Psammoecus fasciatus and morphological differences between these two species were stated in diagnosis of Psammoecus fasciatus.

Yoshida and Hirowatari (2013) described this species from the Nansei Islands, Japan including Nakanoshima (Tokara Islands), Amami-Ôshima, Tokunoshima, Okinawa, Ishigaki and the Iriomote Islands. It is closely similar to Psammoecus fasciatus, and these two species occur allopatrically across the Watase Line, which is one of the biogeographic borders proposed between the Palaearctic and Oriental regions passing through the Tokara Straits (Yoshida and Hirowatari 2013).

Yoshida and Hirowatari (2013) described this species and regarded the undetermined species, Psammoecus sp. 2 of Hirano (2009) and Hirano (2010), as conspecific. In addition, in the present study, Psammoecus sp. 1 illustrated by Hirano (2009) and Hirano (2010), which was represented by only one female, is also found to be conspecific with this species. We had the opportunity to examine some specimens possessing features of Psammoecus sp. 1 (Fig. 2E) and were able to conclude that morphological characters including the male genital structure of these specimens belonged to the range of morphological variation of Psammoecus hiranoi.

The following specimens were found after description of this species:

JAPAN: [Okinawa Pref.] 1 male & 4 exs., Mt. Yonaha-dake, Kunigami Village, Okinawa Island, 3–IV–1974, T. Kinoshita leg. (EUMJ); 10 exs., Ôkuni-rindô, Kuigami Village, Okinawa Island, 4–XI–2013, T. Yoshida leg. (ELKU).

This species is similar to Psammoecus trimaculatus, Psammoecus triguttatus and Psammoecus labyrinthicus sp. n., but can be distinguished from Psammoecus trimaculatus and Psammoecus triguttatus by the wide triangular basal portion of the parameres and the apically narrow portion of the penis, and from Psammoecus labyrinthicus sp. n. by the longer parameres, the apically narrow portion of the penis and the shape of the phallobase.

Body length. 2.50 mm (n=1).

Coloration (Fig. 2F). Head reddish-brown, pronotum somewhat light reddish-brown. Elytra blackish-brown with four large yellowish-brown maculae; macula around anterior 1/4 of each elytron almost quadrate, macula on posterior half longer than wide. Antennae yellowish-brown basally, 6th to 10th antennomeres black, 6th slightly brighter, 11th (apex) bright.

Head (Figs 12A, B and C). Rounded-triangular, HW/HL 1.68; IE/HL 1.09. Temples slightly expanded behind eyes, narrowed at base. Eyes large, prominent, diameter about half of length of head. Punctuation of dorsal surface moderately dense, on ventral surface sparse, and absent on center portion of ventral surface. Antennae 1.44 mm; covered with medium length pubescence and some relatively long erect setae on each antennomere; approximate ratio of holotype as follows: 2.4: 1.0: 1.0: 1.1: 1.1: 1.3: 1.0: 1.1: 1.2: 1.1: 2.2 (Fig. 12A).

Pronotum (Fig. 12B, C). Roundly subquadrate, PW/PL 1.34. Punctuation on dorsal surface relatively strong and moderately sparse. Pubescence composed of medium length setae, a long seta on each tooth on lateral margins and anterior angles, a relatively long seta on each posterior angle. Each anterior angle with several small teeth, each lateral margin with four short teeth; tooth I small, tooth II longer than tooth I, teeth III and IV almost same size, longer than tooth II, teeth II, III and IV relatively widened around base, each posterior angle with a few very small teeth.

Elytra (Fig. 12E). Elongate-oval, EW/BL 0.46. Rows of punctures wider than interstices. Pubescence composed of many medium length semi-erect setae, long erect setae in a row on lateral margins.

9th abdominal sternite (Fig. 14P). Strut cut at anterior 1/3, diverging deeply around posterior 1/3. Lateral sclerites rhomboid, comparatively large, curved inwardly.

Aedeagus (Fig. 14Q, R). Parameres club-shaped; narrow portions relatively broad, punctuated sparsely, with several sparse setae, apex with a long seta; wide portions punctuated densely on posterior half of inner margins and anterolateral portions of outer margins, posterior half of inner margins with many setae. Phallobase consisting of two layers, posterior margin incised roundly, distance between posterior margin and deepest point of incision of margin of upper layer narrow, anterior margin of lower layer relatively narrowly incised, protuberances around anterior 1/4 narrow, projecting inwards, posterior margin of lower layer broadly incised. Penis relatively elongate and flat, with relatively dense punctuation on posterior 1/8.

Holotype: male, Nagasaki, Nagasaki Prefecture, Japan, 1869, G. Lewis leg. (BMNH).

JAPAN: Nagasaki?.

Type specimen was mounted with a label reading ‘Nagasaki | 1869 | ? imported in Rice –’. We have not been able to find any specimen of this species from Japan other than holotype. Hence, occurrence of this species in Japan seems to be questionable.