The present study is based on oribatid mite materials (Acari, Oribatida) collected from Macao SAR, southern China during 2016 to 2018. A list of identified taxa, including 25 species from 23 genera and 14 families, is presented; of these, 11 species and 1 genus are recorded for the first time in the fauna of this country, and 1 species is recorded for the first time in the Oriental region. The taxonomic data on two species of the family Galumnidae is provided. A new species, Dimiodiogalumna ilhaverdeensis Ermilov sp. nov. is described; it differs from Dimidiogalumna comoroensis Mahunka, 1994 by the presence of four pairs of notogastral porose areas and one strium on genital plates, and the absence of ridges on anal plates. The systematic placement of Acrogalumna bipartita Aoki & Hu, 1993 is discussed, resulting in the following taxonomic proposal: Allogalumna bipartita (Aoki & Hu, 1993) comb. nov.; the supplementary description of this species is presented based on specimens from China.
Introduction
Oribatid mites (Acari, Oribatida) of China are moderately known (Chen et al. 2010). Our work is based on materials collected from Macau Peninsula, Taipa Island and Coloane Island, Macao SAR, China during 2016 to 2018. The primary goal of the paper is to present a list and new findings of identified oribatid taxa.
A second goal of the paper is to present taxonomic data on two species of the family Galumnidae: (a) to describe and illustrate a new species of Dimidiogalumna Engelbrecht, 1972. This genus was proposed by Engelbrecht (1972) with Dimidiogalumna villiersensis Engelbrecht, 1972 as type species. At present, it comprises five species, which are distributed in the Ethiopian, Oriental and southern Palaearctic regions (Ermilov & Klimov 2017). The main generic traits were summarized by Ermilov and Klimov (2017). An identification key to previously known species of Dimidiogalumna was given by Ermilov and Anichkin (2014); (b) to provide the supplementary description of Allogalumna bipartita (Aoki & Hu, 1993) comb. nov. based on specimens from China, adding new information about some morphological structures and their measurements, identification of leg setae and solenidia, and morphology of the gnathosoma, and to discuss the generic placement of this species.
Material and methods
Material examined
Material was collected by C.-M. Leong in secondary forests from the Macau Peninsula, Taipa Island and Coloane Island, Macao SAR, China during 2016 to 2018 (Fig. 1). The field surveys included two periods: July to August and February to March. Samples were extracted by the two liters of shifted leaf litter and soil, using Winkler extractor into 75% ethanol during seven days in the laboratory conditions. Localities of samples:
#1—Siu Tam Hill, 22.1622 °N, 113.5463 °E, 97 m a.s.l., 15.II.2018.
#2—Hac Sa Reservoir, 22.1263 °N, 113.5727 °E, 65 m a.s.l., 12.II.2018.
#3—Ilha Verde Hill, 22.2116 °N, 113.5380 °E, 30 m a.s.l., 09.II.2018.
#4—Guia Hill, 22.1967 °N, 113.5502 °E, 74 m a.s.l., 10.II.2018.
#5—Ka Ho Reservoir, 22.1341 °N, 113.5766 °E, 56 m a.s.l., 27.I.2018.
#6—Siu Tam Hill, 22.1613 °N, 113.5501 °E, 52 m a.s.l., 18.I.2018.
#7—Mong Ha Hill, 22.2085 °N, 113.5476 °E, 32 m a.s.l., 18.II.2018.
#8—Ecological Protection Area I, 22.1425 °N, 113.5538 °E, 0 m a.s.l, 14.II.2018.
#11—Hac Sa Reservoir, 22.1264 °N, 113.5733 °E, 71 m a.s.l., 29. VII.2017.
#12—Ilha Verde Hill, 22.2117 °N, 113.5375 °E, 5 m a.s.l., 13.VIII.2017.
#13—Siu Tam Hill, ca. 22.1608 °N, 113.5466 °E, ca. 75 m a.s.l., 26.VIII.2016.
#14—Hac Sa Reservoir, ca. 22.1344 °N, 113.5725 °E, ca. 65 a.s.l., 09.I.2017.
#15—Hac Sa Reservoir, ca. 22.1344 °N, 113.5725 °E, ca. 65 a.s.l., 20.VIII.2016.
#16—Hac Sa Reservoir, 22.1233 °N, 113.5700 °E, 38 m a.s.l., 02.VIII.2017.
#17—Guia Hill, 22.1983 °N, 113.5523 °E, 77 m a.s.l., 01. VII.2017.
#18—Ilha Verde Hill, 22.2117 °N, 113.5375 °E, 41 m a.s.l., 27.VII.2017.
#19—Siu Tam Hill, 22.1614 °N, 113.5502 °E, 52 m a.s.l., 15. VII.2017.
Except the holotype and two paratypes, specimens examined are deposited in TSUMZ.
Methods
Specimens were mounted in lactic acid on temporary cavity slides for identification of all taxa and for measurement and illustration of the new species. Body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the notogaster. Notogastral width refers to the maximum width of notogaster (behind pteromorphs) in dorsal view. Lengths of body setae were measured in lateral aspect. All body measurements are presented in micrometers. Formulas for leg setation are given in parentheses according to the sequence trochanter-femur-genu-tibia-tarsus (famulus included). Formulas for leg solenidia are given in square brackets according to the sequence genu-tibia-tarsus.
Drawings were made with a camera lucida using a Leica transmission light microscope “Leica DM 2500”. Microscope images were taken with an AxioCam ICc5 digital camera.
Morphological terminology follows that of F. Grandjean (see Ermilov & Klimov 2017 for review and application).
The following abbreviations are used: irt—inner rostral tooth; L—lamellar line; S—sublamellar line; N—prodorsal leg niche; E, T—lateral ridges of prodorsum; ro, le, in, bs—rostral, lamellar, interlamellar and bothridial setae, respectively; Ad—sejugal porose area; D—dorsophragma; c, la, lm, lp, h, p—notogastral setae; Aa, A1, A2, A3—notogastral porose areas; mp—median pore; ia, im, ip, ih, ips—notogastral lyrifissures; gla—opisthonotal gland opening; h, m, a—subcapitular setae; or—adoral seta; sac—axillary saccule; v, l, d, cm, acm, ul, sul, vt, lt—palp setae; ω—palp and leg solenidion; cha, chb—cheliceral setae; Tg—Trägårdh's organ; Pd I, Pd II—pedotecta I, II, respectively; 1b, 3a, 3b, 3c, 4a, 4b—epimeral setae; dis—discidium; cp—circumpedal carina; g, ag, an, ad—genital, aggenital, anal and adanal setae, respectively; iad—adanal lyrifissure; Ap— post anal porose area; p.o.—preanal organ; Tr, Fe, Ge, Ti, Ta—leg trochanter, femur, genu, tibia and tarsus, respectively; p.a.—leg porose area; σ, ϕ —leg solenidia; ε—tarsus I famulus; v, ev, bv, l, d, ft, tc, it, p, u, a, s, pv, pl—leg setae.
The following abbreviations of collections are used: NTU—National Taiwan University, Taipei, Taiwan; SMNH—Senckenberg Museum of Natural History, Görlitz, Germany; TSUMZ— Tyumen State University Museum of Zoology, Tyumen, Russia.
List of identified taxa1
The list of identified oribatid mites collected from Macao, China includes 25 species from 23 genera and 14 families. Of these, 11 species and 1 genus are recorded for the first time in the fauna of this country; and 1 species is recorded for the first time in the Oriental region.
Lohmanniidae
Lohmannia corallium Nakatamari, 1982. Locality: 19 (5 ex.). Distribution: Japan. New record of the species in the Oriental region.
Javacarus porosus Hammer, 1979. Locality: 11 (1 ex.). Distribution: Tropical and Subtropical regions.
Meristacarus pubescentius Ren, Yang, Liang & Xie, 2018. Locality: 13 (2 ex.). Distribution: China.
Meristolohmannia macaoensis Ermilov, 2018. Locality: 12 (3 ex.). Distribution: China.
Trhypochthoniidae
Archegozetes longisetosus Aoki, 1965. Locality: 19 (5 ex.). Distribution: Oriental and Neotropical regions.
Malaconothridae
Tyrphonothrus crassisetosus (Willmann, 1932). Locality: 4 (1 ex.). Distribution: Oriental, Australian and Ethiopian regions. New record of the species in China.
Nothridae
Nothrus anauniensis Canestrini & Fanzago, 1877. Localities: 4 (1 ex.), 18 (1 ex.). Distribution: Cosmopolitan.
Basilobelbidae
Basilobelba retiaria (Warburton, 1912). Locality: 19 (2 ex.). Distribution: Tropical and Subtropical regions, Japan. New record of the species in China.
Damaeidae
Tectodamaeus armatus Aoki, 1984. Locality: 18 (1 ex.). Distribution: Palaearctic and Oriental regions.
Eremobelbidae
Eremobelba japonica Aoki, 1959. Locality: 19 (1 ex.). Distribution: Palaearctic and Oriental regions.
Dameolidae
Fosseremus laciniatus (Berlese, 1904). Locality: 12 (1 ex.). Distribution: Cosmopolitan.
Otocepheidae
Dolicheremaeus baloghi Aoki, 1967. Localities: 1 (1 ex.), 6 (1 ex.), 11 (1 ex.), 15 (1 ex.).
Distribution: Palaearctic and Oriental regions. New record of the species in China.
Tectocepheidae
Tegeozetes tunicatus Berlese, 1913. Locality: 12 (2 ex.). Distribution: Tropical and Subtropical regions, Hungary.
Mochlozetidae
Gephyrazetes fasciatus Hirauchi, 1999. Localities: 1 (1 ex.), 2 (1 ex.), 6 (1 ex.). Distribution: Japan, Taiwan. New record of the genus and species in China.
Unguizetes keralensis (Balakrishnan, 1989). Locality: 2 (2 ex.). Distribution: Oriental region. New record of the species in China.
Haplozetidae
Peloribates kaszabi Mahunka, 1988. Localities: 4 (3 ex.), 19 (4 ex.). Distribution: Oriental region. New record of the species in China.
Protoribates dentatus (Berlese, 1883). Localities: 8 (2 ex.), 13 (1 ex.). Distribution: Holarctic and Oriental regions, Fiji. New record of the species in China.
Rostrozetes ovulum (Berlese, 1908). Locality: 12 (1 ex.). Distribution: Tropical and Subtropical regions.
Scheloribatidae
Scheloribates praeincisus (Berlese, 1910). Localities: 12 (1 ex.), 18 (3 ex.), 19 (3 ex.). Distribution: Tropical and Subtropical regions.
Galumnidae
Allogalumna bipartita (Aoki & Hu, 1993). Localities: 5 (2 ex.), 7 (3 ex.). Distribution: Oriental region.
Dimidiogalumna ilhaverdeensis Ermilov sp. nov. Localities: 12 (4 ex.), 18 (5 ex.). Distribution: China.
Galumna acutirostrum Ermilov & Anichkin, 2010. Locality: 19 (3 ex.). Distribution: Vietnam. New record of the species in China.
Pergalumna cattienica Ermilov & Anichkin, 2011. Localities: 2 (1 ex.), 5 (1 ex.), 11 (1 ex.), 13 (3 ex.), 16 (2 ex.). Distribution: Vietnam. New record of the species in China.
Pergalumna kunsti Ermilov & Starý, 2017. Localities: 3 (1 ex.), 4 (1 ex.), 5 (1 ex.), 6 (3 ex.), 7 (2 ex.), 12 (1 ex.), 14 (1 ex.), 17 (1 ex.), 18 (2 ex.), 19 (1 ex.). Distribution: Vietnam. New record of the species in China.
Pergalumna paraelongata Ermilov & Anichkin, 2012. Localities: 7 (3 ex.), 11 (4 ex.). Distribution: Vietnam, Cameroon. New record of the species in China.
Taxonomy
Dimiodiogalumna ilhaverdeensis Ermilov sp. nov.
(Figs 2–13)
Diagnosis
Body size: 315–348 × 249–265. Rostral setae long, setiform, barbed, lamellar and interlamellar setae short, setiform, thin, smooth; ro thickest, in shortest. Bothridial setae clavate, barbed distally. Anterior parts of pteromorphs striate. With 10 pairs of notogastral microsetae and four pairs of porose areas, Aa transversely elongate, slightly triangular, A1, A2 and A3 rounded. Median pore present. Anterior margin of ventral plate dentate. Circumpedal carinae long, directed to pedotecta I. Genital plates with one longitudinal strium in medial parts. Epimeral and anogenital setae short, setiform, thin, smooth. Anterior edges of genital plates with one seta. Postanal porose area elongate oval. Solenidion of tibiae IV located in middle of the segments.
Description
Measurements. Body length: 315 (holotype, male), 315–348 (eight paratypes, four females and four males); notogaster width: 249 (holotype), 249–265 (eight paratypes). No clear differences between females and males in body size.
Integument (Figs 3, 13, 17). Body color brown. Body surface microfoveolate (visible in dissected specimens under high magnification, × 1000). Anterior parts of pteromorphs striate. Genital plates with one longitudinal strium in medial parts. All leg femora and trochanters III, IV with elongate tubercles antiaxially.
Prodorsum (Figs 2, 4). Rostrum rounded. Lamellar lines thin, distinct, directed backwards at ventral ends. Prodorsal leg niches and lateral ridges of prodorsum well-developed. Rostral setae (32– 36) setiform, barbed. Lamellar (8–10) and interlamellar (4) setae setiform, thin, smooth. Bothridial setae (57–65) clavate, with long, smooth stalks and short, rounded and barbed distally heads. Exobothridial setae and their alveoli absent. Sejugal porose areas (12–20 × 6–10) elongate oval, transversely orientated, posterolateral to interlamellar setae. Dorsophragmata long, elongated.
Notogaster (Figs 2, 4–6). Dorsosejugal suture complete, well visible. With 10 pairs of microsetae (c, 2; others 1) and four pairs of porose areas; Aa transversely elongate (length 32–41), slightly triangular, A1, A2, A3 (all 12–20) rounded. Microsetae la located close to the pteromorphal hinges, posterior to Aa. Median pore present in females and males, represented by one part, located between A2. All lyrifissures distinct, im located between lm and lp, ip between p1 and p1, ih anterior to p3, ips lateral to p3. Opisthonotal gland openings located lateral to Al.
Gnathosoma (Figs 7–9). Subcapitulum longer than wide (90–98 × 77–86). Subcapitular setae setiform, smooth, a (18–20) longer and thicker than m (14–16) and h (14–16). Adoral setae (10) setiform, smooth. Palps (65–69) with typical setation 0-2-1-3-9(+1 solenidion). Postpaipai setae (4) spiniform, smooth. Chelicerae (102–110) with two setiform, barbed setae, cha (32–36) longer than chb (20–24). Trägårdh's organ of chelicerae elongate triangular.
Epimeral region (Figs 3, 4). Anterior tectum of epimere I dentate. Pedotecta I and II rounded in ventral view. Discidia triangular. Epimeral setal formula: 1-0-3-1. Epimeral setae (3c, 12–16; 1b, 3a, 3b, 4a, 8–10) setiform, thin, smooth. Circumpedal carinae long, thin, directed to pedotecta I.
Anogenital region (Figs 3–6). Six pairs of genital, one pair of aggenital, two pairs of anal and three pairs of adanal setae similar in length (6–8), setiform, thin, smooth. Anterior edges of genital plates with one seta. Aggenital setae located between genital and anal apertures, equal distanced from them. Adanal lyrifissures located close and parallel to anal plates. Adanal setae ad1 and ad2 postanal, ad3 paraanal and lateral to adanal lyrifissures. Distance ad1–ad2 slightly shorter than ad2– ad3. Unpaired postanal porose area elongate oval (12–16 × 6–10). Ovipositor elongated (151 × 32), blades (69) shorter than length of distal section (beyond middle fold; 82). Each of the three blades with four smooth setae, ψ1 ≈ τ1, (53–57) setiform, longer than thorn-like ψ2 ≈ τa ≈ τb ≈ τc (24–28). Six coronal setae (4) spiniform, smooth.
Legs (Figs 10–13). Median claw distinctly thicker than laterals, all serrate on dorsal side. Dorsoparaxial porose areas on all femora and on trochanters III, IV well visible. Formulas of leg setation and solenidia: I (1-4-3-4-20) [1-2-2], II (1-4-3-4-15) [1-1-2], III (1-2-1-3-15) [1-1-0], IV (1-2-2-3-12) [0-1-0]; homologies of setae and solenidia indicated in Table 1. Solenidion of tibiae IV located in middle of the segments. Famulus of tarsi I inserted posterolateral to solenidion ω1.
Material examined
Holotype (male) and three paratypes (two females and one male): China, Macau Peninsula, Ilha Verde Hill, 22.2117 °N, 113.5375 °E, 5 m a.s.l., locality #12, leaf litter and soil, 13.VIII.2017 (collected by C.-M. Leong). Five paratypes (two females and three males): China, Macau Peninsula, Ilha Verde Hill, 22.2117 °N, 113.5375 °E, 41 m a.s.l., locality #18, leaf litter and soil, 27.VII.2017 (collected by C.-M. Leong).
Type deposition
The holotype (ethanol with drop of glycerol) is deposited in NTU. Two paratypes (ethanol with drop of glycerol) are deposited in SMNH. Six paratypes (ethanol with drop of glycerol) are deposited in TSUMZ.
Etymology
The specific name ilhaverdeensis refers to the locality of the new species, Ilha Verde, Macao SAR, China.
Differential diagnosis
Dimidiogalumna ilhaverdeensis Ermilov sp. nov. differs from the other species of the genus by the dentate anterior margin of ventral plate (versus smooth). The new species is morphologically most similar Dimidiogalumna comoroensis Mahunka, 1994 from the Comoro Islands in having sejugal suture and transversely elongate (slightly triangular) notogastral porose areas Aa, but differs by the presence of four pairs of notogastral porose areas including A1 (versus three pairs, A1 absent) and one strium on genital plates (versus several stria), and the absence of ridges on anal plates (versus one longitudinal ridge present on each anal plate).
Remarks
The generic diagnosis of Dimidiogalumna was presented by Ermilov and Klimov (2017). One of the morphological characters of the genus is the presence of 10 pairs of notogastral alveoli. However, Dimidiogalumna ilhaverdeensis Ermilov sp. nov. has 10 pairs of microsetae instead alveoli. The presence of alveoli and microsetae in representatives of the same genus is generic trait of some superspecies taxa in the family Galumnidae (for example, Allogalumna Grandjean, 1936, Flagellozetes (Cosmogalumna) Aoki, 1988, Pergalumna Grandjean, 1936), therefore, this character state should be included in diagnosis of Dimidiogalumna.
TABLE 1.
Leg setation and solenidia of adult Dimiodiogalumna ilhaverdeensis Ermilov sp. nov. and Allogalumna bipartita (Aoki & Hu, 1993).
Allogalumna bipartita (Aoki & Hu, 1993) comb. nov.
(Figs 22–33)
Supplementary description
Measurements. Body length: 531–713 (five specimens, three females and two males); notogaster width: 431–564 (five specimens). Females larger than males: 531–564 × 431–448 versus 680–713 × 531–564.
Integument. Body color brown. Body surface microfoveolate (visible in dissected specimens under high magnification, × 1000).
Prodorsum (Figs 22, 24). Rostrum rounded. Sublamellar lines thin, distinct, directed backwards at ventral ends. Prodorsal leg niches and lateral ridges of prodorsum well-developed. Rostral (32– 41), lamellar (41–49) and interlamellar (176–205) setae setiform, slightly barbed. Bothridial setae (90–94) lanceolate, with long, smooth stalks and short, pointed and smooth distally heads. Exobothridial setae and their alveoli absent. Sejugal porose areas (16–24 × 6–10) elongate oval, transversely orientated, posterior to interlamellar setae. Dorsophragmata short, slightly elongated.
Notogaster (Figs 22, 24–26). Dorsosejugal suture interrupted medially. With 10 pairs of vestigial microsetae (1 or shorter) and five pairs of rounded porose areas; Aal (16) smaller than Aam, A1, A2, A3 (all 20–24). Microsetae 1a located close to the pteromorphal hinges, posterior to Aal and Aam. Median pore absent in females and males. All lyrifissures distinct, im located between lm and A1, ip between p1 and p1 and closer to p1, ih and ips anterior to p3, Opisthonotal gland openings located lateral to A1.
Gnathosoma (Figs 27–29). Subcapitulum longer than wide (139–143 × 114–123). Subcapitular setae setiform, barbed, a (28–32) and h (28–32) longer than m (16–20). Adoral setae (16) setiform, barbed. Palps (90–94) with typical setation 0-2-1-3-9(+1 solenidion). Postpalpal setae (6) spiniform, smooth. Chelicerae (160–164) with two setiform, barbed setae, cha (49–53) longer than chb (32–36). Trägårdh's organ of chelicerae elongate triangular.
Epimeral region (Figs 23, 24). Anterior tectum of epimere I smooth. Pedotecta I and II rounded in ventral view. Discidia triangular. Epimeral setal formula: 1-0-2-2. Epimeral setae (3b, 20–24; 1b, 3a, 4a, 4b, 16–20) setiform, thin, smooth. Circumpedal carinae short, thin, not reaching the acetabula IV.
Anogenital region (Figs 23–26). Six pairs of genital, one pair of aggenital, two pairs of anal and three pairs of adanal setae similar in length (12–16), setiform, thin, smooth. Anterior edges of genital plates with two setae. Aggenital setae located between genital and anal apertures, closer to genital aperture. Adanal lyrifissures located close and diagonally to anal plates. Adanal setae ad1 and ad2 postanal, ad3 paraanal and posterior to adanal lyrifissures. Distance ad1–ad2=ad2–ad3. Unpaired postanal porose area elongate oval (41–55 × 16–20).
Legs (Figs 30–33). Median claw distinctly thicker than laterals, all serrate on dorsal side. Dorsoparaxial porose areas on all femora and on trochanters III, IV well visible. Formulas of leg setation and solenidia: I (1-4-3-4-20) [1-2-2], II (1-4-3-4-15) [1-1-2], III (1-2-1-3-15) [1-1-0], IV (1-2-2-3-12) [0-1-0]; homologies of setae and solenidia indicated in Table 1. Solenidion of tibiae IV located in anterior part of the segments. Famulus of tarsi I inserted anterolateral to solenidion ω1.
Material examined
Two specimens (two females): China, Coloane Island, Ka Ho Reservoir, 22.1341 °N, 113.5766 °E, 56 m a.s.l., locality #5, leaf litter and soil, 27.1.2018 (collected by C.-M. Leong). Three specimens (one female and two males): China, Macau Peninsula, Mong Ha Hill, 22.2085 °N, 113.5476 °E, 32 m a.s.l., locality #7, leaf litter and soil, 18.11.2018 (collected by C.-M. Leong).
Remarks
1. Allogalumna bipartita was described by Aoki and Hu (1993) as the representative of the genus Acrogalumna Grandjean, 1956. The main generic trait of Acrogalumna is the absence of median pore in females and its presence (represented numerous parts) in males. However, this species has no median pore in females and males, and all other morphological character (see Ermilov & Klimov 2017) corresponds to the genus Allogalumna Grandjean, 1936, therefore, this species should be combined: Allogalumna bipartita (Aoki & Hu, 1993) comb. nov.
2. Specimens of Allogalumna bipartita from southern China are morphologically very similar to the original description (Aoki & Hu 1993), and we did not observe clear differences, except for two points: (a) Chinese specimens with vestigial notogastral setae versus setal alveoli in Japanese specimens; (b) Chinese specimens slightly smaller than Japanese specimens (531–713 × 431–564 versus 760–765 × 600–610). We believe these differences represent intraspecific variability.
3. Based on the original description (Aoki & Hu 1993) and supplementary description of Allogalumna bipartita from China, we propose the following diagnostic morphological traits for this species: body size: 531–765 × 431–610; rostral, lamellar and interlamellar setae setiform, barbed; ro shortest, in longest. Bothridial setae lanceolate, smooth. With 10 pairs of notogastral alveoli or microsetae and five pairs of rounded porose areas, Aa divided into two parts. Median pore absent. Circumpedal carinae short, not reaching the acetabula I. Epimeral and anogenital setae short, setiform, thin, smooth. Postanal porose area elongate oval. Solenidion of tibiae IV located in anterior part of the segments.
Acknowledgements
We cordially thank two anonymous reviewers for the valuable comments. We are grateful for the assistance of Chi-Man Leong's family and friends (e.g., Mr. Ken Kwan, Mr. Eric Kwan, Mr. Chi-Keong Chan, Mr. Martinho G. Oxalá, Dr. Feng-long Jia, Mr. Zhen-Yu Yang, Mr. Man-Cheng Choi, Ms. Ying-Cong Lin, Mr. Si-Chong Leong, Mr. Kai-Leung Kwok, Ms. Weng-I Leong, and Mr. Alexander WK Leung) in Macao SAR, China. The field work was supported by the crowdfunding project (DOI: 10.18258/10715) as well as Dr. Shiuh-Feng Shiao (National Taiwan University) and Dr. Benoit Guénard (University of Hong Kong), and the support of Direcção dos Serviços de Educação e Juventude, Gabinete de Apoio ao Ensino Superior, and Fundação Macau.
This study partially was funded by the Russian Foundation for Basic Research (RFBR) according to the research project № 18-04-00097.