Deeleman-ReinholdChrista L.AddinkWouterMillerJeremy A.The genera Chrysillaand Phintelloidesrevisited with the description of a new species (Araneae, Salticidae) using digital specimen DOIs and nanopublicationsBiodivers Data J030920241210.3897/BDJ.12.e129438 ED448511-2CFA-5EB5-9F2B-97C0F1B469D1 Phintelloides versicolor https://www.checklistbank.org/dataset/288943/taxon/76Z2N (C. L. Koch, 1846) World Spider Catalog: urn:lsid:nmbe.ch:spidersp:035557 Plexippusversicolor C. L. Koch, 1846 - Koch 1846: vol. 13: 103, fig. 1165 (m) Bintan [Bintang] Island, Indonesia. Attusversicolor (C. L. Koch, 1846) - Walckenaer 1847: 426. Maeviapicta C. L. Koch, 1848 - Koch 1848: vol 14: 72, fig. 1328 (f; juv m according to Thorell 1891) Bintan [Bintang] Island, Indonesia. Chrysillaversicolor (C. L. Koch, 1846) - Thorell 1891: 117 (mf; synonymy with Meviapicta) Indonesia (Bintang, Sumatra), Malaysia (Pinang), Singapore; Workman and Workman 1894: 10, pl. 10 (mf) Indonesia (Pinang, Sumatra, Bintang), Singapore; Simon 1901: 544; Żabka 1985: 211, figs 83-96 (mf) Vietnam. Telamonialeucaspis Simon, 1903 - Simon 1903b: 307 (m) Sumatra. syn. nov.; Prószyński 1978: 336, fig. 11 (m). Phintellaleucaspis (Simon, 1903) - Bohdanowicz and Prószyński 1987: 112, figs 214-215 (m). Phintellaversicolor (C. L. Koch, 1846) - Prószyński 1987: 152, 161 (in part). Phintelloidesversicolor (C. L. Koch, 1846) - Kanesharatnam and Benjamin 2019: 22. Materials

Type status: Other material. Occurrence: recordedBy: W. Corley; individualCount: 2; sex: male; lifeStage: adult; otherCatalogNumbers: https://doi.org/10.3535/C69-M7K-VWC; occurrenceID: 451E13DE-7042-5932-A295-646E928CEC1F; Taxon: scientificName: Phintelloidesversicolor; Location: country: Malaysia; stateProvince: Selangor; locality: Banting; verbatimElevation: 100 m; verbatimCoordinates: 2°48’04”N 101°30’46”E; decimalLatitude: 2.8011111111111; decimalLongitude: 10.512777777778; Event: eventDate: 1983-01-28; fieldNotes: CM 21848, DSC 6302-6327; Record Level: institutionID: https://ror.org/027m9bs27; institutionCode: MMUE; basisOfRecord: PreservedSpecimen

Type status: Other material. Occurrence: catalogNumber: MMUE G7572.6413; recordedBy: F. & J. A. Murphy; individualCount: 1; sex: female; lifeStage: adult; otherCatalogNumbers: https://doi.org/10.3535/3NW-1BX-8BK; occurrenceID: A0A5233F-918E-5755-AB82-D3EE60952018; Taxon: scientificName: Phintelloidesversicolor; Location: country: Singapore; locality: Lim Chu Kang; verbatimCoordinates: 1°26’N 103° 43’E; decimalLatitude: 1.4333333333333; decimalLongitude: 103.71666666667; Event: eventDate: 1991-01-28/29; fieldNotes: CM19264; Record Level: institutionID: https://ror.org/027m9bs27; institutionCode: MMUE; basisOfRecord: PreservedSpecimen

Type status: Other material. Occurrence: catalogNumber: RMNH.ARA.18260; recordedBy: P. R. & C. L. Deeleman; individualCount: 1; sex: female; lifeStage: adult; otherCatalogNumbers: https://doi.org/10.3535/M42-Z4P-DRD; occurrenceID: DE44414C-2104-59CD-9533-5324F9DABE02; Taxon: scientificName: Phintelloidesversicolor; Location: country: Thailand; stateProvince: Kanchanaburi Province; locality: Erawan waterfalls N. P.; verbatimCoordinates: 14°22’N 99°08’E; decimalLatitude: 14.366666666667; decimalLongitude: 99.133333333333; Event: eventDate: 1987-11; Record Level: institutionID: https://ror.org/0566bfb96; institutionCode: RMNH; basisOfRecord: PreservedSpecimen

Type status: Other material. Occurrence: catalogNumber: RMNH.ARA.18261; recordedBy: P. R. & C. L. Deeleman; individualCount: 1; sex: male; lifeStage: adult; otherCatalogNumbers: https://doi.org/10.3535/5MR-J6N-26M; occurrenceID: 41A8211A-6F67-5ED3-9CA4-3F4C6ECADB9D; Taxon: scientificName: Phintelloidesversicolor; Location: country: Thailand; stateProvince: Prachuap Khiri Kan Province; locality: Sam Roi Yot National Park; verbatimCoordinates: 12°14’N 99°56’E; decimalLatitude: 12.233333333333; decimalLongitude: 99.933333333333; Event: eventDate: 1988-12-31; habitat: forest on limestone; Record Level: institutionID: https://ror.org/0566bfb96; institutionCode: RMNH; basisOfRecord: PreservedSpecimen

Type status: Other material. Occurrence: catalogNumber: RMNH.ARA.18262; recordedBy: P. Schwendinger; individualCount: 1; sex: male; lifeStage: adult; otherCatalogNumbers: https://doi.org/10.3535/Q6C-91C-BS5; occurrenceID: 5EEB6839-2EC9-54A3-96FA-70DE59BC8142; Taxon: scientificName: Phintelloidesversicolor; Location: country: Thailand; locality: Chiang Mai; verbatimElevation: 300 m; verbatimCoordinates: 18°47’N 98°57’E; decimalLatitude: 18.783333333333; decimalLongitude: 98.95; Event: eventDate: 1987-07-01; Record Level: institutionID: https://ror.org/0566bfb96; institutionCode: RMNH; basisOfRecord: PreservedSpecimen

Description

Both male and female with flattened white hair on clypeus, in males just a small moustache below AME (Fig. 17c), in females with frontal strip of thick white flattened setae over whole carapace width (Fig. 17d); anterior eye region with patch covered with white setae, thorax with wide broad submarginal band with dark edge (Fig. 17a), in live specimens black with 2 white central patches and several small ones (Koh et al. 2022: 437). In alcohol tiny greenish iridescent pits on head in male and female. In males, legs I dark, with a light ring on tibia, metatarsus and tarsus, other legs pale; in females, legs and palps pale (all these features are also mentioned in the original description of leucaspis by Simon 1903b, here synonymized with versicolor). Abdomen dorsally with elongate black and white scales, side all white, venter in both sexes partly covered with white appressed flattened setae. Male palp pigmented on trochanter and base of femur, rest white; female palps all white. Epigyne of female (from Thailand) with slender, almost straight ducts (Fig. 18c, e).

Measurements. Total length: males Banting 6.30 and 4.40, males (Sam Roi Yot N. P.) 4.70 in mm, , Chiang Mai 5.00. Male Sam Roi Yot: total length 4.70, carapace 2.30 long, 1.80 wide 1.30 high, abdomen 2.30 long, 1.20 wide; palp 0.80 – 0.35 – 0.30 - 0.60, width cymbium 0.23.

Diagnosis

The abdomen in males is easily recognizable by the dark central band flanked by a pair of lateral white bands (yellow in life; Fig. 17a, Koh et al. 2022: 437), in reverse to that in most Chrysilla and Phintella species and similar to Phintelloidesscandens; this is a reliable character also valid in material preserved in alcohol. This feature apparently is expressed in the latin name: reversal of pale and dark. The shape of the white central area on the thorax is variable in shape and width (compare Fig. 17a from Thailand with Koh et al. 2022: 437 from Singapore). Just like in representatives of Chrysilla and Phintelloides, the embolus is filiform and relatively short, straight and then slightly curved and bent near the base at an angle of 90° with the retrolateral distal edge of the tegulum (Fig. 19a). For a difference in tegulum see diagnosis of munita. Females differ from males by the different carapace, having a pair of black semi-rings on a light background on the posterior part of the thorax; they differ in abdomen pattern which is dorsally pale with irregular cinnamon-brown blotches and a central white band (Fig. 17b; Koh et al. 2022: 437). Epigyne (Fig. 18a, c, e): the copulatory duct is uniform in diameter, parallel, at the anterior end the ring-like copulatory opening in a 90° inward bend, the outer edge is prolonged as a fold or rim; the left and righthand folds are directed mesally, relatively short, the tips crossing. See P.munita for differences with that species.

Distribution

Sumatra, Bintang Island, Singapore, Malaysia, Thailand, Brunei, Vietnam.

Taxon discussion

The previous generic assignment to Phintella of this species is doubtful, as the embolus does not conform to the definition of that genus (see for example Żabka 1985: fig. 403), nor of Phintelloides (see for example Kanesharatnam and Benjamin 2019: fig. 18A). But despite the generic ambiguity, Phintelloidesversicolor is difficult to distinguish from its close relative Phintelloidesminuta, particularly based on the male copulatory organ. Phintelloidesminuta was removed from synonymy with P.versicolor without argumentation (Prószyński 2016; https://salticidae.pl/salticidae.php?adres=specimen.php?id=12129), and in lieu of justification this has not been adopted by World Spider Catalog (2024). However, we agree with the validity of both species and shall try to provide the missing arguments.

This species complex is a taxonomic snake in the grass. In the various papers listed, the identity of this species is full of contradictions. Prószyński (2017), in his pragmatic classification, used this species as the representative of the genus Phintella; this is misleading. The World Spider Catalog (2024) cites 62 taxonomic treatments of P.versicolor. The species including its synonyms has been placed in 11 different genera over nearly 175 years. Recently, it was assigned to Phintelloides (Kanesharatnam and Benjamin 2019, p. 22). The genus assignment of versicolor through 130 years has commuted between Chrysilla or Phintella by authors with authority (Simon, Zabka, Song, Prószyński) which suggests that the species fits in neither of them satisfactorily.

Koch’s description of the male from a small islant between Singapore and Sumatra is mostly an enumeration of colours of the various body parts: black, white, and yellow, and the central abdominal band rusty red, which also fits our specimens. He mentioned that the female is unknown, but described one two years later as Maeviapicta from the same locality. Then, starting in the 1970s, records attributed to this poorly known tropical species stated to appear from Japanese localities on the latitude of southern Europe (Yaginuma 1977, Prószyński 1973). Prószyński (1973) was the first to provide detailed drawings of the male’s genital organs, in a paper on salticid type specimens from Japan present in the Berlin Museum. He stated that he found the male type specimen of Koch’s “Plexippus” versicolor from Bintang Island as well as male and female specimens labeled Hasariusversicolor Koch from Japan (the latter name combination does not appear elsewhere in the taxononmic literature; World Spider Catalog 2024). In the description, Prószyński (1973) focused on the colouration and the abdomen pattern and apparently decided for some reason that the specimens from Sumatra and Japan are conspecific. No female was available from Bintang. Genitals, male and female, he drew from Japanese specimens only (Prószyński 1973, figs 1-7).

Twelve years later, in the magnificent work by Żabka (1985) on Salticidae from Vietnam appear excellent drawings of a male palp by Prószyński of the alleged holotype of versicolor (Żabka 1985, figs 91, 92) side by side with Zabka’s drawings of versicolor from the same specimen, but apparently opposite pedipalp; Żabka 1985, figs 88-90) along with a specimen from Vietnam (Żabka 1985, figs 83-86). Prószyński’s 1973 identification of versicolor from Japan was followed by Yaginuma (1977) and since then a number of authors cited, re-described and illustrated males and females of versicolor from various material from Japan and China. It has to be admitted that the morphology of palps from the Malay and Japanese specimens is very similar, and warrants further comparative study. However, as is the case also in certain other chrysilloid genera, it is the females that express their identity more clearly than do males by differences in structure in the epigyne. The drawings of the epigyne from Vietnam (Żabka 1985, figs 93-95) differ consistently from those from the Japanese specimens, and better agree with that from specimens we collected in Malaysia and Thailand, representing versicolor. In Bohdanowicz and Prószyński (1987), the latter author presented illustrations of the palp of Phintellaleucaspis (Simon) from Sumatra (figs 214, 215), which looks identical to drawings of tropical Southeast Asian versicolor specimens and apparently leucaspis is a new synonym of versicolor. Although Japanese specimens according to drawings of palpal structure can hardly be distinguished from that of specimens from Bintang Island, Sumatra, Malaysia, Thailand and Vietnam, the epigynes drawn from Japan, China and Hong Kong (e.g., Fig. 18b, d, f; Zhu and Zhang 2011, fig. 362a, b; Prószyński 1973, figs 6, 7) are incompatible with female specimens from the Malay Region, which have not been figured in detail previously (Fig. 18a, c, e). The population represented in Japan and China cannot be maintained in versicolor; the oldest name available is munitus Boesenberg & Strand 1906, which name we propose to remove from synonymy.

Notes

The World Spider Catalog (2024) erroneously lists Maeviapicta as Maeviapicta C. L. Koch 1846: 72; it should be C. L. Koch 1848: 72 (Brignoli 1985).

Phintelloidesversicolor (C. L. Koch, 1846) and Phintelloidesmunita (Bösenberg & Strand, 1906), photographs of preserved male and female specimens

2805352C-A369-5750-9F5A-47569CFA3B0210.3897/BDJ.12.e129438.figure17a

Phintelloidesversicolor (C. L. Koch, 1846), male habitus, dorsal view, RMNH.ARA.18261

https://binary.pensoft.net/fig/1060990
7136C1F2-08CF-56C9-B814-69C1FE9C2B3410.3897/BDJ.12.e129438.figure17b

Phintelloidesversicolor (C. L. Koch, 1846), female habitus, dorsal view, CM 19264

https://binary.pensoft.net/fig/1060991
8E527DD5-B919-5672-8B98-B2E2A9CDCEDB10.3897/BDJ.12.e129438.figure17c

Phintelloidesversicolor (C. L. Koch, 1846), male face, RMNH.ARA.18261

https://binary.pensoft.net/fig/1060992
5BA7F213-CFE8-5BF4-A21B-FB3EFF0F96D110.3897/BDJ.12.e129438.figure17d

Phintelloidesversicolor (C. L. Koch, 1846), female face, CM 19264

https://binary.pensoft.net/fig/1060993
36B8D8CE-B2DE-5B34-BF92-F359151CD5E610.3897/BDJ.12.e129438.figure17e

Phintelloidesmunita (Bösenberg & Strand, 1906), female habitus, dorsal view, CM 15605

https://binary.pensoft.net/fig/1060994
B0DE3A4C-DCCA-5C11-B4A0-0409E0929C6810.3897/BDJ.12.e129438.figure17f

Phintelloidesmunita (Bösenberg & Strand, 1906), female face, CM 15605

https://binary.pensoft.net/fig/1060995
8E527DD5-B919-5672-8B98-B2E2A9CDCEDB10.3897/BDJ.12.e129438.figure17c

Phintelloidesversicolor (C. L. Koch, 1846), male face, RMNH.ARA.18261

https://binary.pensoft.net/fig/1060992
5BA7F213-CFE8-5BF4-A21B-FB3EFF0F96D110.3897/BDJ.12.e129438.figure17d

Phintelloidesversicolor (C. L. Koch, 1846), female face, CM 19264

https://binary.pensoft.net/fig/1060993
2805352C-A369-5750-9F5A-47569CFA3B0210.3897/BDJ.12.e129438.figure17a

Phintelloidesversicolor (C. L. Koch, 1846), male habitus, dorsal view, RMNH.ARA.18261

https://binary.pensoft.net/fig/1060990

Phintelloidesversicolor (C. L. Koch, 1846) and Phintelloidesmunita (Bösenberg & Strand, 1906), photographs and illustrations of female reproductive structures

CC6FE693-57A0-5A6A-93C4-C08DB2DA658D10.3897/BDJ.12.e129438.figure18a

Phintelloidesversicolor (C. L. Koch, 1846), female epigynum, ventral view, CM 19264, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126727
FC9AD106-0A04-59DE-9FC2-D4BEEC087B1910.3897/BDJ.12.e129438.figure18b

Phintelloidesmunita (Bösenberg & Strand, 1906), female epigynum, ventral view, CM 15605, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126728
77377D66-526F-568D-93D3-4BD46ED2725510.3897/BDJ.12.e129438.figure18c

Phintelloidesversicolor (C. L. Koch, 1846), female vulva, dorsal view, RMNH.ARA.18260, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126729
C0066621-88DE-5D2B-942F-35CEEB26861810.3897/BDJ.12.e129438.figure18d

Phintelloidesmunita (Bösenberg & Strand, 1906), female vulva, dorsal view, CM 15605, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126730
203705E3-8180-597B-89CF-26838CDE6E3E10.3897/BDJ.12.e129438.figure18e

Phintelloidesversicolor (C. L. Koch, 1846), female epigynum, dorsal view, illustration, scale bar 0.2 mm

https://binary.pensoft.net/fig/1126731
264E6387-B8E7-54D3-8A29-7A9FDB4535EC10.3897/BDJ.12.e129438.figure18f

Phintelloidesmunita (Bösenberg & Strand, 1906), female epigynum, dorsal view, illustration, scale bar 0.2 mm

https://binary.pensoft.net/fig/1126732
77377D66-526F-568D-93D3-4BD46ED2725510.3897/BDJ.12.e129438.figure18c

Phintelloidesversicolor (C. L. Koch, 1846), female vulva, dorsal view, RMNH.ARA.18260, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126729
203705E3-8180-597B-89CF-26838CDE6E3E10.3897/BDJ.12.e129438.figure18e

Phintelloidesversicolor (C. L. Koch, 1846), female epigynum, dorsal view, illustration, scale bar 0.2 mm

https://binary.pensoft.net/fig/1126731

Phintelloidesversicolor (C. L. Koch, 1846), photographs of male pedipalp

D28F99C9-5948-5782-A319-4E7742E8EF3310.3897/BDJ.12.e129438.figure19a

Phintelloidesversicolor (C. L. Koch, 1846), male pedipalp, ventral view, RMNH.ARA.18262

https://binary.pensoft.net/fig/1060996
88A7A632-84A2-566A-A2CF-49C14ED2D27C10.3897/BDJ.12.e129438.figure19b

Phintelloidesversicolor (C. L. Koch, 1846), male pedipalp, retrolateral view, RMNH.ARA.18262

https://binary.pensoft.net/fig/1060997
D28F99C9-5948-5782-A319-4E7742E8EF3310.3897/BDJ.12.e129438.figure19a

Phintelloidesversicolor (C. L. Koch, 1846), male pedipalp, ventral view, RMNH.ARA.18262

https://binary.pensoft.net/fig/1060996
7136C1F2-08CF-56C9-B814-69C1FE9C2B3410.3897/BDJ.12.e129438.figure17b

Phintelloidesversicolor (C. L. Koch, 1846), female habitus, dorsal view, CM 19264

https://binary.pensoft.net/fig/1060991
CC6FE693-57A0-5A6A-93C4-C08DB2DA658D10.3897/BDJ.12.e129438.figure18a

Phintelloidesversicolor (C. L. Koch, 1846), female epigynum, ventral view, CM 19264, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126727
FC9AD106-0A04-59DE-9FC2-D4BEEC087B1910.3897/BDJ.12.e129438.figure18b

Phintelloidesmunita (Bösenberg & Strand, 1906), female epigynum, ventral view, CM 15605, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126728
C0066621-88DE-5D2B-942F-35CEEB26861810.3897/BDJ.12.e129438.figure18d

Phintelloidesmunita (Bösenberg & Strand, 1906), female vulva, dorsal view, CM 15605, scale bar 0.1 mm

https://binary.pensoft.net/fig/1126730
264E6387-B8E7-54D3-8A29-7A9FDB4535EC10.3897/BDJ.12.e129438.figure18f

Phintelloidesmunita (Bösenberg & Strand, 1906), female epigynum, dorsal view, illustration, scale bar 0.2 mm

https://binary.pensoft.net/fig/1126732
KochC. L.1846Die ArachnidenDreizehnter BandJ. L. LotzbeckNürnberg23410.5962/bhl.title.43744WalckenaerC. A.1847Dernier Supplément (365-596)WalckenaerC. A.GervaisP.Histoire naturelles des Insects. AptèresTome quatrièmeRoretParis623KochC. L.1848Die ArachnidenVierzehnter BandJ. L. LotzbeckNürnberg8810.5962/bhl.title.43744ThorellT.1891Spindlar från Nikobarerna och andra delar af södra AsienKongliga Svenska Vetenskaps-Akademiens Handlingar2421149WorkmanT.WorkmanM. E.1894 Malaysian spider Published by the authorBelfast9-2410.5962/bhl.title.101972SimonE.1901Histoire naturelle des araignéesDeuxième éditiontome secondRoretParis381-66810.5962/bhl.title.51973ŻabkaM.1985Systematic and zoogeographic study on the family Salticidae (Araneae) from Viet-NamAnnales Zoologici, Warszawa39197485SimonE.1903Etudes arachnologiques. 34e Mémoire. LIV. Arachnides recueillis à Sumatra par M. J. BouchardAnnales de la Société Entomologique de France72301310PrószyńskiJ.1978Distributional patterns of the Palaearctic Salticidae (Araneae)Symposia of the Zoological Society of London42335343BohdanowiczA.PrószyńskiJ.1987Systematic studies on East Palaearctic Salticidae (Araneae), IV. Salticidae of JapanAnnales Zoologici, Warszawa4143151PrószyńskiJ.1987Atlas rysunków diagnostycznych mniej znanych Salticidae 2Zeszyty Naukowe Wyższej Szkoly Rolniczo-PedagogicznejSiedlcach172KanesharatnamN.BenjaminS. P.2019Multilocus genetic and morphological phylogenetic analysis reveals a radiation of shiny South Asian jumping spiders (Araneae, Salticidae)ZooKeys83918110.3897/zookeys.839.283KohJ. K.H.CourtD. J.AngC. S.P.NgP. Y.C.2022A Photographic Guide to Singapore SpidersNational Parks BoardSingaporePrószyńskiJ.2016Monograph of Salticidae (Araneae) of the World 1995-2015. Part II. Global Species Database of Salticidae (Araneae). Version October 30th, 2016online at: http://www.salticidae.plCatalogWorld Spider2024World Spider Catalog. Version 25.0Natural History Museum Bern, online at http://wsc.nmbe.ch, accessed on 01/03/202410.24436/2PrószyńskiJ.2017Pragmatic classification of the world's Salticidae (Araneae)Ecologica Montenegrina12113310.37828/em.2017.12.1YaginumaT.1977A list of Japanese spiders (revised in 1977)Acta Arachnologica27 (Spec. No.)36740610.2476/asjaa.27.Specialnumber_367PrószyńskiJ.1973Systematic studies on east Palaearctic Salticidae, II. Redescriptions of Japanese Salticidae of the Zoological Museum in BerlinAnnales Zoologici, Warszawa3097128ZhuM. S.ZhangB. S.2011Spider fauna of Henan: Arachnida: AraneaeScience PressBeijing558BrignoliP. M.1985On the correct dates of publication of the arachnid taxa described in some works by C. W. Hahn and C. L. Koch (Arachnida)Bulletin of the British Arachnological Society69414416