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Abstract

Araneus is a large genus containing many species with various features, and phylogenetic analysis has revealed it to be a polyphyletic group. It has been proposed that nine new genera should be established independently from Araneus. One example is a group that includes Araneus mitificus (Simon 1886) and Araneus praesignis (L. Koch 1872). In this study, we examined specimens from several species that are morphologically similar to them, and confirmed their close relationship by phylogenetic analysis using the data of five genes. All targeted species were united in a monophyletic clade, comprising two groups in detail. The epigynes and male palps of the species in these two groups were clearly distinguishable; therefore, we concluded that they should be recognized as two independent genera. Consequently, we describe two new genera, Aoaraneus n. gen. and Bijoaraneus n. gen. Additionally, previously identified Japanese Araneus mitificus specimens were reexamined and concluded to be misidentified. It is described as a new species under the name Bijoaraneus komachi n. sp. in this study.
Introduction
The genus Araneus Clerck 1757 is the largest spider ge-
nus, currently consisting of 575 species (World Spider Cat-
alog 2021). When Clerck established the genus, various spi-
der species, even non-web lycosid, and salticid spiders, were
placed in this genus. Although various new families and
genera have been described, and many species have moved
from Araneus to other genera, the genus presently contains
many species with various characteristics. Its polyphyly was
revealed by recent molecular phylogenetic analyses (Kallal
et al. 2020, Schar et al. 2020). Schar et al. (2020) inferred
the phylogeny of the family Araneidae using ve genes and
suggested that at least nine new genera should be established
independently of Araneus. One example is a group that in-
cludes Araneus miticus (Simon 1886) and A. praesignis (L.
Koch 1872), namely NGEN10 in Scharff et al. (2020). In
this study, we examined specimens from several species that
are morphologically similar to them, and confirmed their
close relationships by molecular phylogenetic analysis using
the same ve genes used by Schar et al. (2020). Next, we
describe two new genera in the present paper.
Simultaneously, the Japanese A. miticus was taxonomi-
cally revised. It was described based on the specimens from
Cambodia and widely known in South, Southeast, and East
Asia, particularly from India to Japan. However, Japanese
specimens are remarkably larger than Thai specimens, and
the shape of male palps and epigynes dier from each other.
This situation led us to revise their taxonomic positions, par-
ticularly in Japanese specimens.
Materials and methods
We examined specimens identified as Araneus amabilis
Tanikawa 2001, A. legonensis (Grassho & Edmunds 1979),
A. pentagrammicus (Karsch 1879), A. postilena (Thorell
1878), and Japanese so-called A. mitificus (abbreviated as
JA), along with A. mitificus from Southeast Asia, because
their general appearance and web style are similar.
The specimens were preserved in 75% ethanol solution at
18–27°C. Morphological characteristics were examined un-
der an M3Z stereoscopic microscope (Wild Heerbrugg AG,
Heerbrugg, Switzerland), and photographs were obtained us-
ing an EOS Kiss X7 with an EF-S 60 mm f2.8 macro photo
lens and an MT-24EX macro twin ash (Canon Inc., Tokyo,
Two new genera of Araneidae (Arachnida: Araneae)
Akio Tanikawa1*, Takeshi Yamasaki2 & Booppa Petcharad3
1 Laboratory of Biodiversity Science, School of Agriculture and Life Sciences,
The University of Tokyo, 1-1-1, Yayoi, Bunkyo-ku, Tokyo, 113-8657 Japan
E-mail: dp7a-tnkw@j.asahi-net.or.jp
2 Institute of Nature and Environmental Sciences, University of Hyogo / Museum of Nature and Human Activities, Hyogo, Yayoigaoka 6,
Sanda-shi, Hyogo, 669-1546 Japan.
Email: yamasaki@hitohaku.jp
3 Faculty of Science and Technology, Thammasat University, Rangsit, Pathum Thani, 12121 Thailand
E-mail: argiope2@sta.tu.ac.th, *Corresponding author
Abstract Araneus is a large genus containing many species with various features, and phylogenetic anal-
ysis has revealed it to be a polyphyletic group. It has been proposed that nine new genera should be estab-
lished independently from Araneus. One example is a group that includes Araneus miticus (Simon 1886)
and Araneus praesignis (L. Koch 1872). In this study, we examined specimens from several species that are
morphologically similar to them, and conrmed their close relationship by phylogenetic analysis using the
data of ve genes. All targeted species were united in a monophyletic clade, comprising two groups in detail.
The epigynes and male palps of the species in these two groups were clearly distinguishable; therefore, we
concluded that they should be recognized as two independent genera. Consequently, we describe two new
genera, Aoaraneus n. gen. and Bijoaraneus n. gen. Additionally, previously identied Japanese Araneus miti-
cus specimens were reexamined and concluded to be misidentied. It is described as a new species under the
name Bijoaraneus komachi n. sp. in this study.
Key words Araneus, Araneini, Aoaraneus, Bijoaraneus, komachi, new genus, new species
Acta Arachnologica, 70 (2): 87–101, December 27, 2021
A. Tanikawa, T. Yamasaki & B. Petcharad
Japan), or attached to the microscope. All measurements are
provided in millimeters. The type specimens designated in
this study were deposited in the National Science Museum
of Nature and Science, Tsukuba, Japan. Specimens collected
in Thailand were deposited in the Natural History Muse-
um, National Science Museum, Thailand, and the Princess
Maha Chakri Sirindhorn Natural History Museum, Prince of
Songkla University, Thailand.
The phylogenetic relationships between specimens were
inferred by sequencing data of the same ve genes used in
Schar et al. (2020), namely, mitochondrial cytochrome ox-
idase subunit I (COI) and 16S-rRNA (16S), nuclear 18S-rR-
NA (18S), 28S-rRNA (28S), and Histone 3 (H3). We added
the sequence data obtained in the present study to the data
set of Schar et al. (2020) and performed a molecular phy-
logenetic analysis. The specimens used for molecular study
are presented in the appendix.
Genomic DNA was extracted from leg muscle using a
DNeasy Blood & Tissue kit (Qiagen, Inc., Hilden, Germany)
or a FavorPrep Tissue Genomic DNA Extraction Mini Kit
(Favorgen Biotech Corp, Ping-Tung, Taiwan). The COI par-
tial sequence was amplified using the primer combination
LCO1498: 5’-GGT CAA CAA ATC ATA AAG ATA TTG
G-3’ and HCO2198: 5’-TAA ACT TCA GGG TGA CCA
AAA AAT CA-3’ (Folmer et al. 1994). 16S rRNA was am-
plied using the primer combination 16S-A: 5’-CGC CTG
TTT ATC AAA AAC AT-3’ and 16S-B: 5’-CTC CGG TTT
GAA CTC AGA TCA-3’ (Palumbi et al. 1991). The reac-
tants were initially denatured for 2 min at 94 °C, followed
by 40 cycles of 15 s at 94°C, 20 s at 47°C, and 30 s at 72°C
for both COI and 16S. 18S was amplied using the primer
combination 1F: 5’-TAC CTG GTT GAT CCT GCC AGT
AG-3’ or 3F: GTT CGA TTC CGG AGA GGG A-3’ with
9R: 5’-GAT CCT TCC GCA GGT TCA CCT AC-3’ (Giribet
et al. 1996). The reactants were initially denatured for 2 min
at 94°C, followed by 40 cycles of 30 s at 94°C, 20 s at 47°C,
and 2 min at 72°C. 28S was amplied using the primer com-
bination 28S-B: TCG GAA GGA ACC AGC TAC TA-3’
with 28S-O: 5’-GAA ACT GCT CAA AGG TAA ACG G-3’
(Whiting et al. 1997). The reactants were initially denatured
for 2 min at 94°C, followed by 40 cycles of 30 s at 94°C,
20 s at 50°C, and 45 s at 72°C. H3 was amplied using the
primer combination H3aF: 5’-ATG GCT CGT ACC AAG
CAG ACV GC-3’ with H3aR: 5’-ATA TCC TTR GGC ATR
ATR GTG AC-3’ (Colgan et al. 1998). The reactants were
initially denatured for 2 min at 94°C, followed by 40 cycles
of 30 s at 94°C, 20 s at 47°C, and 15 s at 72°C. PCR prod-
ucts were sequenced by the Takara Bio CDS Center (Shiga,
Japan) or Fasmac Co., Ltd. (Kanagawa, Japan) using an
ABI 3130xl or 3730xl DNA analyzer (Life Technologies
Japan Ltd., Tokyo, Japan), using the primers HCO2198,
16S-A, 4F, 28S-B, and H3aF for COI, 16S, 18S, 28S, and
H3, respectively. The chromatogram obtained was checked
manually using MEGA ver.7 (Kumar et al. 2016). The se-
quence data used by Schar et al. (2020) were downloaded
from the DDBJ/EMBL/GenBank database. Sequence align-
ments were performed using the online version of MAFFT
(Katoh & Standley 2013). We used the L-INS-I option for
coding genes and the Q-INS-I option for RNA gene align-
ments. Alignments of protein-coding genes were translated
into amino acids and checked for stop codons. Nucleotide
sequence data obtained in this study are available in the
DDBJ/EMBL/GenBank databases.
The Perl script KAKUSAN 4 (Tanabe 2011) and
TREEFINDER (Jobb et al. 2004) were used to determine
the appropriate model of DNA evolution by BIC. MrBayes
ver.3.1.2 (Ronquist & Huelsenbeck 2003) was employed for
Bayesian analyses on a combined dataset. Four concurrent
Markov chain Monte Carlo were run for 40 million genera-
tions, saving a tree every 1000 generations. Topologies prior
to ln stabilization (“burn-in”) were discarded, and posterior
clade probabilities were calculated from the remaining trees.
The following abbreviations are used for some collectors:
AT, Akio Tanikawa; BP, Booppa Petcharad; YS, Yuya Suzu-
ki.
Results
JA was morphologically dierent from the holotype of A.
miticus, and is describe here as a new species.
We obtained 122–618 bp, 426–438 bp, 664–700 bp,
550–700 bp, and 287–309 bp of COI, 16S, 18S, 28S, and
H3 partial sequences, respectively, from the specimens for
molecular analysis. The best-t models of sequence evolu-
tion determined by Kakusan 4 for Bayesian analyses were
as follows: HKY85+G for 16S, JC69 homologous for the
third position of H3, GTR+G for other partitions, and gene
proportional and codon proportional, rather than other pat-
terns of mixed models. The phylogeny obtained was almost
congruent with Scharff et al. (2020). The target species of
this study and their sister groups are shown in Fig. 1. All
of the targeted species are united in a monophyletic clade,
comprising two groups in detail, namely, A. pentagrammic-
us + A. amabilis: clade A, and A. miticus + A. legonensis +
JA + A. postilena + A. praesignis: clade B (Fig. 1). Clade B
is consistent with NGEN10 in Schar et al. (2020).
As mentioned in Schar et al. (2020), although the type
species of Araneus, A. angulatus Clerck 1757 is not includ-
ed in the analysis, the clade including A. diadematus may be
the core Araneus, and clade A, B in this study diers from
the core Araneus. We concluded that it should be treated
as an independent genus from Araneus, in agreement with
Schar et al. (2020). Furthermore, the species in clade A and
B are clearly distinguishable by the morphological features
mentioned in the diagnosis given below, so we concluded
that these two groups should be treated as independent gen-
era.
Consequently, we describe two new genera, one consist-
ing of A. pentagrammicus and A. amabilis, and the other
consisting of A. mitificus, JA, A. legonensis, A. postilena,
and A. praesignis. All the species in these two new genera
differ morphologically from A. angulatus the generotype.
Male tibia I and II are similar to each other, but it is modi-
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New genera of Araneidae
ed, essentially sinuating or with a characteristic prong, in
the core Araneus. The embolus of the male palp lacks a cap
but has an embolus cap in the core Araneus.
The clade consisting of genus Eriovixia and Araneus
rotundulus is inferred to be a sister group of these two gen-
era (Fig. 1). Araneus rotundulus should be treated as an
independent genus or included in Eriovixia, however, we
could not examine any specimen of the species, and we will
therefore investigate this in the future. Two new genera are
distinguishable from Eriovixia as follows: the abdomen is
a vertically long oval, median ocular area is wider anterior-
ly, male palpal tibia has two macrosetae, male coxa I has a
dorsal tubercle; while in Eriovixia, the abdomen is inverted
triangle or wide oval, male palpal tibia lacks macroseta,
male coxa I lacks a dorsal tubercle, and median ocular area
is wider posteriorly.
Taxonomy
Family Araneidae
Aoaraneus new genus
(Japanese name: Ao-onigumo-zoku).
Type species. Miranda pentagrammica Karsch 1879 from
Japan.
Diagnosis. Aoaraneus n. gen. differs from the related
Bijoaraneus n. gen. by having long, wrinkled and flexible
scape with bent tip (Figs. 3E, 4E), vs. short and inexible in
Bijoaraneus n. gen. (Figs. 5D, 6C, 7C, 8 B). The male palp
of Aoaraneus n. gen. has long terminal apophysis (Figs. 2A–
B, 3G, 4G), but that of Bijoaraneus either lacking terminal
apophysis (Figs. 2C–E, 5H, 6G, 7G) or it is small and incon-
spicuous (Figs. 2F, 8F). The male palp of Aoaraneus n. gen.
has a subterminal apophysis (Figs. 2A–B, 3G, 4G), lacking
in Bijoaraneus n. gen. (Figs. 2C–F, 5H, 6G, 7G, 8F). The fe-
mur II of Aoaraneus ventrally lacking strong spine (Fig. 3I),
while that of Bijoaraneus n. gen. has a row of strong spines
(Fig. 5C). The coxa I of Aoaraneus has a tubercle anteriorly,
while that of Bijoaraneus either lacking a tubercle, or it is
slightly swollen.
Description. Carapace longer than wide, median ocular
area wider anteriorly than posteriorly. Male palpal femur
basally with tubercle, tibia with 2 macrosetae; bulb with
terminal and subterminal apophyses (Figs. 2A–B), embolus
without cap. Male endite with lateral tooth (Fig. 3I). Male
coxa I with ventral hook (Fig. 3I), dorsal tubercle, and ante-
rior tubercle (Fig. 3I), femur II with groove. Abdomen oval,
longer than wide. Epigynal scape long, wrinkled, flexible,
and tip is bent (Figs. 3C–F, 4C–E); with basal plate.
Web with a free sector, silk yellow, signal line leads to
retreat made by living leaf and silk.
Etymology. Generic name is coined from the Japanese
word “Ao,” and Araneus. The Japanese word “Ao” means
blue or green. Gender is masculine.
Species included. Aoaraneus pentagrammicus (Karsch
1879) n. comb. and Aoaraneus amabilis (Tanikawa 2001) n.
comb.
Aoaraneus pentagrammicus (Karsch 1879) new combination
(Japanese name: Ao-onigumo).
(Figs. 2A, 3A–I)
Miranda pentagrammica Karsch 1879, p. 72, pl. 1, g. 6 (♂), syntypes
Fig. 1. Phylogeny inferred by Bayesian inference. Only the part of targeted species in this study and their sister group are shown, see Schar
et al. (2020) for other parts. Score at each node is posterior probabilities. Clade A is described as Aoaraneus n. gen. and clade B is described as
Bijoaraneus n. gen., JA is described as Bijoaraneus komachi n. sp. in this study. The scale bar shows substitution per site. Asterisk indicates the
data from Schar et al. (2020).
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A. Tanikawa, T. Yamasaki & B. Petcharad
from Japan, preserved in Museum für Naturkunde, Berlin (ZMB),
not examined.
Aranea pentagrammica: Bösenberg & Strand 1906, p. 219, pl. 4,  g.
35, pl. 11,  g. 211 (♀♂).
Catheistela pentagrammica: Yaginuma & Archer 1959, p. 36.
Araneus pentagrammicus: Yaginuma 1986, p. 100, fig. 53.4 (♀♂);
Tanikawa 2007, p. 84,  gs. 248–250, 695–696 (♀♂).
Further literature, see World Spider Catalog (2021).
Specimens examined. JAPAN. GUNMA Pref. 1♀,
Kawauchi-cho, Kiryu-shi, 22-VI-1983, AT leg. SAITAMA
Pref. 1♀, Han-no, Han-no-shi, 12-VII-1998, AT leg. CHI-
BA Pref. 1♀, Orikisawa, Kimitsu-shi, 28-VI-2008, AT leg.
Fig. 2. Male palp. A, Aoaraneus pentagrammicus n. comb. B, Aoaraneus amabilis n. comb. C, Bijoaraneus miti cus n. comb. D, Bijoaraneus
komachi n. sp. E, Bijoaraneus legonensis n. comb. F, Bijoaraneus postilena n.comb. Scales = 0.2 mm.
Abbreviations: C, conductor; E, embolus; MA, median apophysis; TA, terminal apophysis; ST, subterminal apophysis; STII, subterminal
apophysis II.
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New genera of Araneidae
Fig. 3. Aoaraneus pentagrammicus n. comb. A, female habitus; B, male habitus; C–F, epigyne, ventral (C), posterior (D), lateral (E), and dorsal
view (F); G–H, male palp, prolateral (G) and ventral (H) view; I, male leg I and II, ventral view. Scales = 5 mm (A–B); 0.2 mm (C–F); 1 mm (I).
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Fig. 4. Aoaraneus amabilis n. comb. A, female habitus; B, male habitus; C–F, epigyne, ventral (C), posterior (D), lateral (E), and dorsal (F)
view; G–H, male palp, prolateral (G), and ventral view (H). Scales = 5 mm (A–B); 0.2 mm (others).
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Fig. 5. Bijoaraneus miti cus n. comb. A, female habitus; B, female in a hide; C, male leg I and II, ventral view; D–G, epigyne, ventral (D), pos-
terior (E), lateral (F), and dorsal (G) view; H–I, male palp, prolateral (H) and ventral (I) view. Scales = 5 mm (A–B); 1 mm (C); 0.2 mm (others).
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Fig. 6. Bijoaraneus komachi n. sp. A, female habitus; B, male habitus; C–F, epigyne, ventral (C), posterior (D), lateral (E), and dorsal (F) view;
G–H, male palp, prolateral (G) and ventral (H) view. Scales = 5 mm (A–B); 0.2 mm (others).
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Fig. 7. Bijoaraneus legonensis n. comb. A, free sector web; B, female habitus; C–F, epigyne, ventral (C), posterior (D), lateral (E), and dorsal (F)
view; G–H, male palp, prolateral (G), and ventral (H) view. Scales = 5 mm (B); 0.2 mm (others).
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Fig. 8. Bijoaraneus postilena n. comb. A, female habitus; B–E, epigyne, ventral (B), posterior (C), lateral (D), and dorsal (E) view; F–H, male
palp, prolateral (F), ventral (G), and retrolateral (H) view. Scales = 5 mm (A); 0.2 mm (others).
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TOKYO, 1♂, Midori-cho, Nishi-tokyo-shi, 9-VII-1995,
AT leg. 1♀, Motohachiouji-machi, Hachiouji-shi, 21-V-
1989, AT leg. KANAGAWA Pref. 1♀, Noba-cho, Konan-ku,
Yokohama-shi, 9-V-1981, AT leg. 1♀1♂, Hitorizawa-cho,
Isogo-ku, Yokohama-shi, 29-IV-1987, AT leg. 1♀, Numama,
Zushi-shi, 19-VI-1974, K. Kumada leg. 1♀, Iriya-higashi,
Zama-shi, 12-V-2000, AT leg. 1♀, Obara, Midori-ku, Sag-
amihara-shi, 18-V-2021, AT leg. 1♀1♂, Negoya, Midori-ku,
Sagamara-shi, 10-V-1987, AT leg. 1♀, Hasugesan, Aika-
wa-machi, Aiko-gun, 12-V-2021, AT leg. 1F, Iriya-higashi,
Zama-shi, 12-V-2000, AT leg. 1♀, Nanasawa, Atsugi-shi,
6-VI-1993, AT leg. 1♂, Zenba, Isehara-shi, 28-IV-2021, AT
leg. SHIZUOKA Pref. 1♀, Minamiizu-cho, Kamo-gun, 25-
VII-1992, AT leg. 1♀, Iwamoto, Fuji-shi 19-V-2014, AT leg.
1♀, Kitamatsuno, Fuji-shi, 19-V-2014. AT leg. 1♀, Yainaba,
Fujieda-shi, 16-V-2013, AT leg. 1♀, Hagino, Kakegawa-shi,
26-VI-2015, AT leg. 1♀1♂, Ryokukeidai, Tenryu-ku, Ham-
amatsu-shi, 10-IV-2013, YS leg. AICHI Pref. 1♀, Iwasa-
ki-cho, Toyohashi-shi, 25-VIII-1993, AT leg. MIE Pref.
1♀, Kogawaguchi, Kiwa-cho, 30-VIII-1990, AT leg. 2♀,
WAKAYAMA Pref. Kumanogawa-cho, Shingu-shi, 3-VIII-
1991, AT leg. SAGA Pref. 1♀, Kinryu, Kinryu-machi, Sa-
ga-shi, 31-VII-2005, AT leg. KAGOSHIMA Pref. 1♀, Miya-
noura, Yakushima Is., 15-VII-1990, AT leg. 1♀, Kusukawa,
Yakushima Is., 17-VII-1990, AT leg.
Diagnosis. Aoaraneus pentagrammicus resembles A. am-
abilis. However, they are distinguishable as follows: basal
plate of the epigyne is small and invisible in ventral view in
A. pentagrammicus (Fig. 3C), while large and visible in the
ventral view in A. amabilis (Fig. 4C). Subterminal apophy-
sis is leaf-shaped in A. pentagrammicus (Figs. 2A, 3G), vs.
boot-shaped in A. amabilis (Figs. 2B, 4G).
Description. Based on a female and a male from Japan.
Coloration and markings when living as in Figs. 3A, B: car-
apace greenish orange; abdomen dorsally whitish green with
dark blue and black markings, ventrally green; legs greenish
orange with black annulation.
Measurements. Female / male. Body 9.70 / 6.70 long.
Carapace 4.06 / 3.40 long; 3.25 / 2.80 wide. Length of legs
[tarsus, metatarsus, tibia, patella, femur = total]: I, 1.63,
4.06, 3.19, 1.81, 3.94 = 14.63 / 1.32, 3.68, 3.04, 1.32, 3.64
= 13.00; II, 1.44, 3.50, 2.69, 1.75, 3.69 = 13.07 / 1.20, 3.12,
2.32, 1.20, 3.40 = 11.24; III, 1.00, 1.75, 1.38, 1.19, 2.50
= 7.82 / 0.80, 1.56, 1.24, 0.84, 2.24 = 6.68; IV, 1.06, 3.19,
2.50, 1.63, 3.69 = 12.07 / 1.00, 2.76, 2.08, 1.12, 3.12 =
10.08. Abdomen 5.63 / 3.84 long; 5.19 / 3.28 wide.
Body and legs. Male / female. Carapace length / width
ratio 1.25 / 1.21. Median ocular area length / width ratio 0.95
/ 0.86; anterior width / posterior width ratio 1.29 / 1.40. La-
bium length / width ratio 0.90 / 0.84. Sternum length / width
ratio 1.26 / 1.40. Length of leg I / length of carapace ratio 3.60
/ 3.82. Abdomen 1.08 / 1.17 times longer than wide.
Copulatory organs. Epigyne as in Figs. 3C–F; heart
shaped and with hole at the center in posterior view; scape
longer than median plate, wrinkled, flexible, with bent
tip. Male palp as in Figs. 2A, 3G–H, terminal apophysis
reaching conductor; with 2 subterminal apophyses, median
apophysis with apical and basal projections.
Range. Japan (Honshu, Shikoku, Kyushu), Korea, China,
Taiwan.
Aoaraneus amabilis (Tanikawa 2001) new combination
(Japanese name: Chura-onigumo)
(Figs. 2B, 4A–H)
Araneus amabilis Tanikawa 2001, p. 82, gs. 69–72, 75–76, holotype
♀ from Iriomotejima Is., Japan, preserved in National Science Mu-
seum of Nature and Science, Tsukuba (NSMT), examined.
Further literature, see World Spider Catalog (2021).
Specimens examined. JAPAN. KAGOSHIMA Pref.
1♀, Yakushima Is., 14-VII-2006, AT leg. 1♀, Nakanoshi-
ma Is., Tokara Isls., 5-VII-2006, AT leg. 1♀, Setouchi-cho,
Amami-oshima Is., 21-II-2005, AT leg. 1♂, Nishiakina,
Tokunoshima Is., 12-IX-2005, AT leg. OKINAWA Pref. 1♀,
Nishime, Kumejima Is., 15-VI-2004, AT leg. 1♀1♂, Urau-
chi, Iriomotejima Is., Okinawa Pref., 1-I-2000, AT leg.
Diagnosis. Aoaraneus amabilis resembles A. pentagram-
micus see the diagnosis of the latter species for the discrimi-
nating points.
Description. See Tanikawa (2001).
Range. Japan (Nansei Islands).
Bijoaraneus new genus
(Japanese name: Bijo-onigumo-zoku)
Type species. Epeira mitica Simon 1886, from Cambo-
dia.
Diagnosis. See diagnosis in Aoaraneus n. gen.
Description. median ocular area wider anteriorly than pos-
teriorly. Male palpal femur basally with tubercle, tibia with 2
macrosetae. Male endite with lateral tooth. Male coxa I with
ventral hook (Fig. 5C), and dorsal tubercle, femur II with
groove, femur II with a row of ventral spines (Fig. 5C). Abdo-
men oval, longer than wide. Epigynal scape short, well sclero-
tized, inexible (Figs. 5D, 6C, 7C, 8B); without dierentiation
between median plate and lateral lamella. Male palp without
terminal apophysis, or small and not reaching conductor.
Web with a free sector (Fig. 7A), silk yellow, with signal
line lead to retreat made by living leaf and silk (Fig. 5B).
Etymology. Generic name is coined from the Japanese
word “Bijo” and Araneus. The Japanese word “Bijo” means
beautiful lady. Gender is masculine.
Species included. Bijoaraneus miticus (Simon 1886) n.
comb., Bijoaraneus komachi n. sp., Bijoaraneus legonensis
(Grassho & Edmunds 1979) n. comb., Bijoaraneus postile-
na (Thorell 1878) n. comb., and Bijoaraneus praesignis (L.
Koch 1872) n. comb.
97
Acta Arachnologica, 70 (2), December 2021 Arachnological Society of Japan
A. Tanikawa, T. Yamasaki & B. Petcharad
Bijoaraneus miticus (Simon 1886) new combination
(Thai name: Mangmoum-Yai-Klom-Tong-Lai-Huajai)
(Figs. 2C, 5A–I)
Epeira mitica Simon 1886, p. 150, holotype ♀ from Cambodia, pre-
served in Muséum Nationa d’Histoire Naturelle, Paris, examined.
Araneus miticus: Simon 1909, p. 109.
Further literature, see World Spider Catalog (2021).
Specimens examined. Holotype with label ‘Pavie 28-
85’, ‘Cambodia’. TAIWAN. 1♂, Tunghai University, Xitun
District, Taichung City, 27-III-2012, Tatsumi Suguro leg.
THAILAND. 1♀, Ban Pangdang, Tambon Chiang Dao,
Amphoe Chiang Dao, Chiang Mai Prov., 3-VII-2014, AT
& BP leg. 1♀1♂, Tambon Khlong Sam, Amphoe Khlong
Luang, Pathum Thani Prov., 13-XI-2018, AT & BP leg. 1♀,
Tambon Ronpiboon, Amphoe Ronpiboon, Nakhon Si Tham-
marat Prov., 7-X-2013, AT & BP leg. 1♀, Tambon Bo Dan,
Amphoe Sathing Phra, Songkhla Prov., 22-II-2016, AT & BP
leg. 1♀, Tambon Sakom, Amphoe Chana, Songkhla Prov.,
15-XI-2013, BP leg. 1♀, Tambon Sakom, Amphoe Chana,
Songkhla Prov., 14-II-2015, BP leg. 1♀, Tambon Khu Tao,
Amphoe Hat Yai, Songkhla Prov., 28-II-2016, AT & BP leg.
1♀, Ban Thung Chang, Tambon Phatong, Amphoe Hat Yai,
Songkhla Prov., 10-III-2018, AT & BP leg. SINGAPORE.
1♂, Pasir Ris Park, 6-VI-2016, Nicky Bay & Joseph K. H.
Koh leg. (Collection permit no. NP/RP12-070-3).
Diagnosis. Bijoaraneus mitificus resembles B. komachi
n. sp. and B. legonensis (Figs. 5A, 6A–B, 7B). Bijoaraneus
mitificus females are distinguishable by the heart-shaped
epigyne in ventral view (cf. Fig. 5D and Figs. 6C, 7C). Bi-
joaraneus miticus males can be separated from B. komachi
n. sp. by the apically bent embolus (cf. Fig. 2C and 2D) and
can be separated from B. legonensis by the shape of median
apophysis; in the prolateral view, it is an inverted triangle in
B. legonensis, but not in B. miticus (cf. Fig. 2C and 2E).
Description. Based on a female and a male specimen
from Thailand. Coloration and markings when living (Fig.
5A): carapace greenish orange; abdomen dorsally whitish
green with black markings, ventrally green; legs green.
Measurements. Female / male. Body 5.94 / 4.19 long.
Carapace 2.80 / 1.88 long; 2.24 / 1.60 wide. Length of legs
[tarsus, metatarsus, tibia, patella, femur = total]: I, 1.00, 2.48,
2.12, 1.24, 2.64 = 9.48 / 0.80, 1.88, 1.64, 0.84, 2.04 = 7.20;
II, 0.84, 2.00, 1.68, 1.12, 2.32 = 7.96 / 0.64, 1.56, 1.12, 0.72,
1.80 = 5.84; III, 0.60, 1.00, 0.84, 0.76, 1.52 = 4.72 / 0.48,
0.72, 0.60, 0.48, 1.16 = 3.44; IV, 0.68, 1.84, 1.56, 1.12, 2.70
= 7.90 / 0.52, 1.28, 1.04, 0.64, 1.60 = 5.08. Abdomen 3.52 /
2.15 long; 3.00 / 1.95 wide.
Body and legs. Male / female. Carapace 1.25 / 1.17 times
longer than wide. Median ocular area length / width ratio
0.84 / 0.94, anterior width / posterior width ratio 2.38 / 1.45.
Labium length / width ratio 0.68 / 0.62. Sternum 1.25 / 1.24
times longer than wide. Length of leg I / carapace length ra-
tio 3.39 / 3.83. Abdomen 1.17 / 1.10 times longer than wide.
Copulatory organs. Epigyne as in Figs. 5D–G, well
sclerotized, heart shaped in ventral view, inverted triangle in
posterior view. Male palp as in Figs. 2C, 5H–I, without ter-
minal apophysis, embolus apically bent.
Range. Thailand, Cambodia, Singapore, Taiwan, China.
Bijoaraneus komachi new species
Japanese name: Bijo-onigumo
(Figs. 2D, 6A–H)
Aranea mitica: Bösenberg & Strand 1906, p. 221, pl. 4, g. 20, pl.
11, g. 207 (♀). Misidentication.
Araneus miticus: Saito 1939, p. 19, g. 3; Saito 1959, p. 87, pl. 11,
g. 94a, pl. 13, g. 94b; Yaginuma 1986, p. 100, g. 53.5; Chikuni
1989, p. 66, g. 14; Namkung 2002, p. 246, gs. 19.8a–b; Tanika-
wa 2007, p. 85, figs. 256, 699–701; Tanikawa 2009, p. 457, figs.
275–277; Kim & Lee 2012, p. 24, gs. 12A–C. Misidentication.
Afraranea mitica: Yaginuma & Archer 1959, p. 37. Misidentication.
Types. Holotype. ♀, Oda, Tsukuba-shi, Ibaraki Pref., Ja-
pan, 8-IX-2020, YS leg. (COI barcode = LC637702). Para-
types: same locality as holotype. 1♂, 16-IX-2019, AT & YS
leg. 1♂, 14-IX-2020, AT & YS leg. 1♀, 14-IX-2020, AT &
YS leg.
Other specimens examined. JAPAN. IBARAKI Pref.
1♂, Tsukuba-shi, 29-IX-1975, Toshiyuki Takai leg. CHIBA
Pref. 2♀, Abiko-shi, 7-XII-2013, Yasunori Hagino leg. TO-
KYO. 1♂, Midori-cho, Nishitokyo-shi, 15-X-1995, AT leg.
KANAGAWA Pref. 1♂1♀, Terayama-cho, Midori-ku, Yoko-
hama-shi, 9-IX-1989, AT leg. 3♀, Izumi-ku, Yokohama-shi,
19-IX-1998, AT leg. 4♀, 25-VIII-1980, AT & Miyako Mori
leg. 4♀, 28-VIII-1980, AT & Hisako Iijima leg. 1♀, 13-IX-
1980, Hisako Iijima leg. 4♀, 18-IX-1982, AT & Kazuaki
Sato leg. 1♀, 13-XI-1983, AT leg. 1♀, 14-IX-1985, Sachiko
Tazoe leg. Noba-cho, Yokohama-shi. 1♂, Ikeko, Zushi-shi,
23-IX-1983, Kazuaki Sato leg. 1♀, Iiyama, Atsugi-shi, 21-
IX-1986, AT leg. 1♀, Tsurumaki, Hadano-shi, 15-IX-1989,
AT leg. 1♀, Ikusawa, Oiso-Machi, Naka-gun, 17-IX-2020,
AT leg. 1♀, Kuno, Odawara-shi, 14-X-2012, AT leg. SHI-
ZUOKA Pref. 1♀, Izu-shi, 18-X-1986, AT leg. AICHI Pref.
1♂, Fuso-cho, Toyota-shi, 28-VIII-2014, Kiyoto Ogata leg.
Diagnosis. Bijoaraneus komachi n. sp. resembles B.
miticus. in general appearance, and has been misidentied
as the latter species by many authors as shown in the syn-
onym list, but is distinguishable from the latter by larger
body size, epigyne, and the male palp: range of body size
in B. komaci is: female (N = 26) = 7.70–11.60; male (N =
8) = 5.60–6.50. In B. miticus, female (N = 8) = 5.50–9.20;
male (N = 4) = 4.00–4.30. In B. miticus, the contour on the
anterior side of the epigyne is double-humped, so it appears
heart shaped, but not in B. komachi. The posterior view of
the epigyne of B. miticus appears as an inverted triangle,
while in B. komachi, the outline is constricted and appears as
snowman shape. Male palpal median apophysis of B. koma-
chi is vertically long in the prolateral view (Figs. 2D, 6G),
but horizontally long in B. miticus (Figs. 2C, 5H); embolus
of B. komachi is straight (Figs. 2D, 6G), but apically bent in
98
Acta Arachnologica, 70 (2), December 2021 Arachnological Society of Japan
New genera of Araneidae
B. miticus (Figs. 2C, 5H).
Description. Based on the holotype female and a male
paratype. Coloration and markings when living (Figs. 6A–
B): carapace greenish orange; abdomen dorsally whitish
green with black markings, ventrally green; legs greenish
orange with dark colored annulation; sternum green.
Measurements. Female / male. Body 8.60 / 6.50 long.
Carapace 4.00 / 3.40 long; 3.25 / 2.72 wide. Length of legs
[tarsus, metatarsus, tibia, patella, femur = total]: I, 1.36,
3.76, 3.28, 1.72, 3.92 = 14.04 / 1.32, 3.56, 3.00, 1.36, 3.52
= 12.76; II, 1.16, 3.00, 2.48, 1.56, 3.40 = 11.60 / 1.08, 2.84,
1.88, 1.12, 3.16 = 10.08; III, 0.84, 1.48, 1.20, 1.12, 2.20
= 6.84 / 0.76, 1.32, 1.04, 0.84, 1.92 = 5.88; IV, 0.92, 2.72,
2.24, 1.56, 3.28 = 10.72 / 0.84, 2.36, 1.84, 1.12, 2.76 = 8.92.
Abdomen 5.44 / 3.36 long; 5.00 / 2.76 wide.
Body and legs. Female / male. Carapace length / width
ratio 1.23 / 1.25. Median ocular area length / width ratio 0.96
/ 0.87; anterior width / posterior width ratio 1.35 / 1.53. La-
bium length / width ratio 0.85 / 0.85. Sternum length / width
ratio 1.25 / 1.31. Length of leg I / length of carapace ratio
3.51 / 3.75. Abdomen length / width ratio 1.09 / 1.22.
Copulatory organs. Epigyne as in Figs. 6C–F, snowman
shaped in posterior view (Fig. 6D). Male palp as in Figs. 2D,
6G–H, embolus apically straight, median apophysis with a
projection towards anteriorly (Figs. 2D, 6G–H).
Etymology. The specific epithet was derived from the
Japanese word “Komachi” meaning “a beautiful lady”.
Range. Japan (Hokkaido, Honshu, Shikoku, Kyushu),
Korea.
Bijoaraneus legonensis (Grassho & Edmunds 1979) new
combination
(Thai name: Mangmoum-Yai-Klom-Tong-Lai-Kangkaw)
(Figs. 2E, 7A–H)
Araneus legonensis Grasshoff & Edmunds 1979, p. 303, figs. 1–11
(♂♀), holotype male from Ghana, preserved in Senckenberg Muse-
um, Frankfurt am Main, not examined.
Specimens examined. THAILAND. 1♀, Tambon Huai
Kaeo, Amphoe Mae On, Chiang Mai Prov., 6-VII-2014, AT
& BP leg. 1♀1♂, Tambon Tha Kha-nun, Amphoe Thong
Pha Phum, Kanchanaburi Prov., 16-XII-2019, AT & BP leg.
Diagnosis. Bijoaraneus legonensis resembles B. mitifi-
cus in general appearance but is distinguishable from the
latter by the shape of epigyne and male palp. the contour
on the anterior side of the epigyne in B. mitificus, is dou-
ble-humped, and appears as heart-shaped (Fig. 5D), unlike
in B. legonensis (Fig. 7C). The posterior view of the epigyne
of B. mitificus appears as an inverted triangle (Fig. 5E),
while in B. legonensis, the outlines of both sides are almost
parallel (Fig. 7D). median apophysis of B. miticus is trans-
versal in prolateral view (Figs. 2C, 5H); however, is an in-
verted triangle in B. legonensis (Figs. 2E, 7G).
Description. Based on a female and a male specimen
from Thailand. Coloration and markings when living (Fig.
7B): carapace greenish orange; abdomen dorsally whitish
green with black markings, ventrally green; legs greenish
orange with weak annulation.
Measurements. Female / male. Body 6.40 / 3.64 long.
Carapace 2.64 / 2.05 long; 2.20 / 1.70 wide. Length of legs
[tarsus, metatarsus, tibia, patella, femur = total]: I, 0.92, 2.60,
2.08, 1.16, 2.60 = 9.36 / 0.80, 2.08, 1.78, 0.83, 2.23 = 7.72;
II, 0.80, 2.04, 1.60, 1.04, 2.24 = 7.72 / 0.68, 1.68, 1.15, 0.75,
1.93 = 6.19; III, 0.60, 1.00, 0.80, 0.76, 1.44 = 4.60 / 0.48,
0.80, 0.60, 0.50, 1.18 = 3.56; IV, 0.64, 1.88, 1.48, 1.04, 2.20
= 7.24 / 0.53, 1.40, 1.08, 0.65, 1.68 = 5.34. Abdomen 4.06 /
1.90 long; 3.56 / 1.38 wide.
Body and legs. Female / male. Carapace length / width
ratio 1.20 / 1.21. Median ocular area length / width ratio 0.94
/ 0.88; anterior width / posterior width ratio 1.31 / 1.33. La-
bium length / width ratio 0.76 / 0.83. Sternum length / width
ratio 1.21 / 1.27. Length of leg I / length of carapace ratio
3.55 / 3.77. Abdomen length / width ratio 1.14 / 1.38.
Genital organs. Epigyne as in Figs. 7C–F, an outline of
both sides almost parallel in ventral and posterior view (Fig.
7C–D). Male palp as in Figs. 2E, 7G–H; median apophysis
inverted triangle in prolatetal view, embolus apically bent.
Range. Thailand, Ghana.
Notes. The specimens from Thailand show a slight dier-
ence from the original description. But the dierence is not
distinct enough to separate Thai specimens from B. legonen-
sis. The type locality of the species is in Ghana, namely far
apart from Thailand, so the dierence might be geographical
variation. Although the species has never been recorded be-
tween Ghana and Thailand, it might be overlooked or mis-
identied in this area. A future faunal survey in this area is
desired.
Bijoaraneus postilena (Thorell 1878) new combination
(Thai name: Mangmoum-Yai-Klom-Tong-Lai-Kled)
(Figs. 2F, 8A–H)
Epeira postilena Thorell 1878, p. 70, syntypes ♂♀ from Amboina, not
examined.
Araneus postilena: Simon 1895, p. 813.
Specimens examined. THAILAND. 1♂, Tambon Na Ch-
aluai, Amphoe Na Chaluai, Ubon Ratchathani Prov., 27-VII-
2013, BP leg. 1♂, Tambon Tha Yaek, Amphoe Mueang, Sa
Kaeo Prov., 16-XI-2018, AT & BP leg. 2, Tambon Bu Fai,
Amphoe Prachantakham, Prachin Buri Prov., 11-XII-2019,
AT & BP leg. 1♂, Tambon Chong, Amphoe Na Yong, Trang
Prov., 22-II-2016, AT & BP leg. 1♀, Ban Khuan Khao
Wang, Tambon Chalung, Amphoe Hat Yai, Songkhla Prov.,
30-VIII-2012, BP leg. 1♀, Ban Ton Nga Chang, Tambon
Thung Tam Sao, Amphoe Hat Yai, Songkhla Prov. 26-II-
2008, BP leg.
Diagnosis. Bijoaraneus postilena resembles B. mitifi-
cus, B. komachi n. sp., and B. legonensis, but can be dis-
tinguished from them as follows: epigynal scape of B.
postilena is inverted triangle in ventral view (Fig. 8B), but
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Acta Arachnologica, 70 (2), December 2021 Arachnological Society of Japan
A. Tanikawa, T. Yamasaki & B. Petcharad
those of the latters are small and knob like shape in ventral
view; male palpal median apophysis of B. postilena is side-
ways-protruding (Figs. 8F, H), but those of the latters are
short and not protruding (Figs. 5H–I, 6G–H, 7G–H).
Description. Coloration and markings when living (Fig.
8A): carapace greenish orange; abdomen dorsally whitish
green with black markings, ventrally green; legs greenish
orange with weak annulation.
Measurements. Female / male. Body 5.06 / 3.60 long.
Carapace 2.00 / 1.80 long; 1.70 / 1.45 wide. Length of legs
[tarsus, metatarsus, tibia, patella, femur = total]: I, 0.83, 2.00,
1.80, 0.95, 2.13 = 7.71 / 0.75, 1.78, 1.68, 0.73, 1.93 = 6.87;
II, 0.70, 1.58, 1.33, 0.83, 1.83 = 6.27 / 0.63, 1.38, 1.13, 0.63,
1.63 = 5.40; III, 0.48, 0.80, 0.65, 0.63, 1.23 = 3.79 / 0.48,
0.70, 0.60, 0.45, 1.10 = 3.33; IV, 0.58, 1.45, 1.25, 0.85, 1.83
= 5.96 / 0.48, 1.23, 1.03, 0.63, 1.50 = 4.87. Abdomen 2.92 /
1.68 long; 2.64 / 1.25 wide.
Body and legs. Female / male. Carapace length / width
ratio 1.18 / 1.24. Median ocular area length / width ratio 1.00
/ 0.94; anterior width / posterior width ratio 1.25 / 1.45. La-
bium length / width ratio 0.75 / 0.58. Sternum length / width
ratio 1.27 / 1.22. Length of leg I / length of carapace ratio
3.86 / 3.82. Abdomen length / width ratio 1.11 / 1.34.
Genital organs. Epigyne as in Figs. 8B–E; scape inverted
triangle in ventral view (Fig. 8B). Male palp as in Figs. 2F,
8F–H; with terminal apophysis, median apophysis protrud-
ing sideways.
Range. Thailand, Indonesia.
Bijoaraneus praesignis (L. Koch 1872) new combination
Epeira praesignis L. Koch 1872, p. 110, pl. 9, g. 3, holotype ♀ from
Queensland, Australia, preserved in Zoologisches Museum Ham-
burg, Hamburg, not examined.
Notes. Although no specimen of this species is available
in this study, its phylogenetic position (Fig. 1, Schar et al.
2020) shows that it should be included in this genus.
Acknowledgments
We wish to express our heartfelt thanks to the following people for
their kind help with our study: Joe Dulyapat, Thailand; Yuki G. Baba,
Institute for Agro-Environmental Sciences, NARO; Yuya Suzuki,
Kagoshima University; Tatsumi Suguro, Keio Yochisha Elementary
School; Joseph K. H. Koh and Nicky Bay, Singapore; I-Min Tso,
Tunghai University; Christine Rollard, Muséum National d’Histoire
Naturale; Kiyoto Ogata, Aichi; Yasuhiro Hagino, the Natural History
Museum and Institute, Chiba; Miyako Mori, Hisako Iijima, Kazuaki
Sato, and Sachiko Tazoe, former Noba Senior High Shool; Toshiyuki
Takai, former University of Tsukuba, and Tadashi Miyashita, the Uni-
versity of Tokyo. This research was partly funded by the Department
of Biotechnology, Faculty of Science and Technology, Thammasat
University, Thailand.
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Received July 9, 2021 / Accepted August 2, 2021
Appendix
Species, specimen numbers, locality, and accession numbers of specimens used for molecular work in this study.
Accession numbers
Species and specimen number locality COI 16S H3 18S 28S
Aoaraneus pentagrammicus 1345 Chiba, Japan LC637703 LC637712 LC637721 LC637730 LC637739
Aoaraneus pentagrammicus 71 Kanagawa, Japan LC637704 LC637713 LC637722 LC637731 LC637740
Aoaraneus amabilis 72 Iriomotejima Is., Japan LC637705 LC637714 LC637723 LC637732 LC637741
Bijoaraneus miticus 4799 Pathum Thani, Thailand LC637706 LC637715 LC637724 LC637733 LC637742
Bijoaraneus komachi 1346 Tokyo, Japan LC637707 LC637716 LC637725 LC637734 LC637743
Bijoaraneus komachi 80 Kanagawa, Japan LC637708 LC637717 LC637726 LC637735 LC637744
Bijoaraneus legonensis 4936 Kanchanaburi, Thailand LC637709 LC637718 LC637727 LC637736 LC637745
Bijoaraneus postilena 4801 Sa Kaeo, Thailand LC637710 LC637719 LC637728 LC637737 LC637746
Bijoaraneus postilena 4935 Prachin Buri, Thailand LC637711 LC637720 LC637729 LC637738 LC637747
101
Acta Arachnologica, 70 (2), December 2021 Arachnological Society of Japan
... The family Araneidae Clerck, 1757 is one of the most diverse group of spiders, including 3,090 species worldwide, and currently 81 species belonging to 25 genera have been recorded in Korea (World Spider Catalog, 2022). After a taxonomic review of Araneidae from Korea , eleven species were discovered newly, including three new species (Araniella robusta Lee, Yoo and Kim 2021; Neoscona flavida Kim, Lee and Yoo, 2019; and Neoscona jindoensis Kim, Lee and Ji, 2016) and three species which were recorded as a result of correcting errors in previous literatures: Bijoaraneus komachi Tanikawa, Yamasaki and Petcharad, 2021 [formerly identified as Araneus mitificus (Simon, 1886), corrected by Tanikawa et al. (2021)]; Cyrtarachne akirai Tanikawa, 2013 [formerly Cyrtarachne inaequalis Thorell, 1895, corrected by Tanikawa (2013)]; Larinioides jalimovi (Bakhvalov, 1981) [formerly Larinioides sclopetarius (Clerck, 1757), corrected by Šestáková et al. (2014)]. However, they were not included in the recent checklist of Korean spiders (National Institute of Biological Resources, 2019). ...
... The family Araneidae Clerck, 1757 is one of the most diverse group of spiders, including 3,090 species worldwide, and currently 81 species belonging to 25 genera have been recorded in Korea (World Spider Catalog, 2022). After a taxonomic review of Araneidae from Korea , eleven species were discovered newly, including three new species (Araniella robusta Lee, Yoo and Kim 2021; Neoscona flavida Kim, Lee and Yoo, 2019; and Neoscona jindoensis Kim, Lee and Ji, 2016) and three species which were recorded as a result of correcting errors in previous literatures: Bijoaraneus komachi Tanikawa, Yamasaki and Petcharad, 2021 [formerly identified as Araneus mitificus (Simon, 1886), corrected by Tanikawa et al. (2021)]; Cyrtarachne akirai Tanikawa, 2013 [formerly Cyrtarachne inaequalis Thorell, 1895, corrected by Tanikawa (2013)]; Larinioides jalimovi (Bakhvalov, 1981) [formerly Larinioides sclopetarius (Clerck, 1757), corrected by Šestáková et al. (2014)]. However, they were not included in the recent checklist of Korean spiders (National Institute of Biological Resources, 2019). ...
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