Research Article |
Corresponding author: Wayne P. Maddison ( wmaddisn@mail.ubc.ca ) Academic editor: Jeremy Miller
© 2022 Wayne P. Maddison, Gustavo R. S. Ruiz, Paul Y. C. Ng, Ettukandathil Haridas Vishnudas, Ambalaparambil V. Sudhikumar.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Maddison WP, Ruiz GRS, Ng PYC, Vishnudas EH, Sudhikumar AV (2022) Kelawakaju gen. nov., a new Asian lineage of marpissine jumping spiders (Araneae, Salticidae, Marpissina). ZooKeys 1130: 79-102. https://doi.org/10.3897/zookeys.1130.87730
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The genus Kelawakaju Maddison & Ruiz, gen. nov., is described for a lineage of bark-dwelling Asian marpissine jumping spiders that represent a dispersal to Eurasia separate from that of the Marpissa-Mendoza lineage, according to the phylogeny recovered from analysis of four gene regions. All species of Kelawakaju are new to science except Kelawakaju frenata (Simon, 1901), comb. nov., which is transferred from Ocrisiona Simon, 1901. Kelawakaju frenata is known from Hong Kong, Guangdong, Guangxi, and likely Taiwan. The five new species are Kelawakaju mulu Maddison & Ruiz, sp. nov. (type species of Kelawakaju, from Sarawak, Malaysia, ♂♀), K. intexta Maddison & Ruiz, sp. nov. (from Sarawak, ♂), K. leucomelas Maddison & Ng, sp. nov. (Singapore and Johor Bahru, ♂♀), K. sahyadri Vishnudas, Maddison, & Sudhikumar, sp. nov. (India, ♂♀), and K. singapura Maddison & Ng, sp. nov. (Singapore, ♂♀).
Classification, Dendryphantini, molecular phylogeny, new genus, new species, Salticinae, Salticoida, taxonomy
Jumping spiders of the tribe Dendryphantini diversified into more than 700 known species largely in the Americas (
Spider specimens examined for this study are stored in the University of British Columbia Spencer Entomological Collection, Canada (
Preserved specimens were examined under both dissecting microscopes and a compound microscope with reflected light. Drawings were made with a drawing tube on a Nikon ME600L compound microscope. Most photographs of living specimens were made with either a Pentax Optio 33WR digital camera with a small lens glued to it for macro capability or an Olympus OM-D E-M10 II camera with 60 mm macro lens.
All measurements are given in millimeters. Descriptions of color pattern are based on the alcohol-preserved specimen. Carapace length was measured from the base of the anterior median eyes not including the lenses to the rear margin of the carapace medially; abdomen length to the end of the anal tubercle. The following abbreviations are used: PLE, posterior lateral eyes; RTA, retrolateral tibial apophysis.
Molecular data was gathered for four gene regions by traditional Sanger PCR methods and combined with previously published data to compose a dataset of 36 taxa (Table
Maximum-likelihood phylogenetic analyses were performed with IQ-TREE version 1.6.7.1 (
Alignments and trees are deposited in the Dryad data repository (http://dx.doi.org/10.5061/dryad.mw6m9060r).
The reconstructed phylogeny (Fig.
The phylogenetic results emphasize the difficulties faced in recognizing salticid relationships from general appearances. When one author (WPM) first collected members of the K. mulu group, he recorded them as baviines, and assumed that their resemblance to the marpissine Balmaceda Peckham & Peckham, 1894 was convergence for trunk-dwelling. It was only with the molecular data that their identity as marpissines became clear. When other authors (EHV, AVS) first collected K. sahyadri, they also thought it likely to be a baviine.
Specimens and GenBank accession numbers of four gene regions analyzed. Accession numbers with * indicate already published (
Species | Specimen ID | Locality | 28S | Actin | 16SND1 | CO1 |
---|---|---|---|---|---|---|
Non-marpissoid outgroups | ||||||
Evarcha proszynskii Marusik & Logunov, 1998 | d096/S232 | Canada: British Columbia | DQ665765* | EU522704* | DQ665723* | AY297379* |
Bavia cf. intermedia (Karsch, 1880) | d079 | Malaysia: Sabah | EU815490* | KM032958* | KM032925* | EU815603* |
Myrmarachne sp. | d162 | Malaysia: Pahang | EU815507* | JX145837* | EU815565* | EU815616* |
Simaetha sp. | d027 | Australia: Queensland | EU815477* | JX145839* | EU815546* | EU815592* |
Ballini | ||||||
Afromarengo sp. | MRB262 | Gabon: Ngounié: Waka National Park | JX145758* | JX145842* | JX145905* | JX145682* |
Mantisatta longicauda Cutler & Wanless, 1973 | S209 | Philippines: Luzon | AY297270* | AY296689*/AY297333* | AY297399* | |
Peplometus sp. | d199 | Ghana: N. of Cape Coast, Kakum Forest | EU815515* | JX145843* | EU815572* | EU815621* |
Dendryphantini: Synagelina | ||||||
Admestina sp. | GR057 | U.S.A.: Mississippi | OP605970 | OP700690 | OP700674 | |
Attidops youngi (Peckham & Peckham, 1888) | S97 | U.S.A.: Missouri | AF327933* | AF327961*/AF328020* | AF327990* | |
Peckhamia sp. | GR137 | Dominican Republic: Barahona | OP605980 | OP700699 | OP700683 | |
Synageles sp. | GR056 | U.S.A.: Mississippi | OP605985 | OP700705 | OP700689 | |
Dendryphantini: Dendryphantina | ||||||
Dendryphantes hastatus (Clerck, 1757) | d043 | Poland: Siedlce | EF201646* | KY200848* | KM032927* | KM033228* |
Ghelna canadensis (Banks, 1897) | d005 | U.S.A.: North Carolina | EF201651* | EU522708* | OP700675 | |
d391 | U.S.A.: North Carolina | KT462689 | ||||
Hentzia grenada (Peckham & Peckham, 1894) | GR064 | USA: Florida | OP605971 | OP700691 | OP700676 | |
Phanias albeolus (Chamberlin & Ivie, 1941) | GR049 | Canada: British Columbia | OP605981 | OP700700 | OP700684 | |
Phidippus otiosus (Hentz, 1846) | GR073 | USA: Florida | OP605982 | OP700701 | OP700685 | |
Rhene sp. | MRB081 | China: Guangxi | OP605984 | OP700704 | OP700688 | |
Sassacus papenhoei Peckham & Peckham, 1895 | S295 | U.S.A.: Arizona | AF327953* | AF327982/AF328041* | AF328012* | |
Zygoballus rufipes Peckham & Peckham, 1885 | S142 | U.S.A. and Panama | AF327944* | AF327972*/AF328031* | AF328002* | |
Dendryphantini: Itatina | ||||||
Itata sp. A | S181 | Ecuador: Manabi | AF327932* | AF327960*/AF328019* | AF327989* | |
Itata sp. B | GR107 | Ecuador: Napo | OP605972 | OP700692 | ||
Itata sp. C | ECU11-4724 | Ecuador: Orellana:Yasuní | OP605973 | OP700677 | ||
Dendryphantini: Marpissina | ||||||
Kelawakaju mulu sp. nov. | SWK12-2610 | Malaysia: Sarawak: Mulu | OP605974 | OP700678 | OP606004 | |
Kelawakaju frenata (Simon, 1901) | d224 | China: Guangxi | JX145769* | JX145911* | JX145688* | |
GR048 | China: Guangxi | OP700693 | ||||
Maevia inclemens (Walckenaer, 1837) | d465 | USA: Tennessee | OP700694 | |||
GR126 | USA: North Carolina | OP605975 | ||||
Maevia intermedia Barnes, 1955 | S87 | USA: Alabama | AY297269* | AY296688*/AY297332* | AY297398* | |
Marpissa lineata (C. L. Koch, 1846) | GR055 | USA: Mississippi | OP605977 | OP700696 | OP700680 | |
Marpissa nivoyi (Lucas, 1846) | GR145 | Spain: Sitges | OP605978 | OP700697 | OP700681 | |
Marpissa pikei (Peckham & Peckham, 1888) | S294/S299 | USA: Arizona | AF327936* | AF327964*/AF328032* | AF327993* | |
Marpissa aff. pikei (Peckham & Peckham, 1888) | GR141 | Dominican Republic: Pedernales | OP605976 | OP700695 | OP700679 | |
Metacyrba pictipes Banks, 1903 | GR140 | Dominican Republic: Pedernales | OP605979 | OP700698 | OP700682 | |
Metacyrba taeniola (Hentz, 1846) | S298 | USA: Arizona | AY297271* | AY296690*/AY297334* | ||
Platycryptus californicus (Pkm & Pkm, 1888) | d316 | Canada: British Columbia | KM033194* | KM032960* | KM033229* | |
d158 | Canada: British Columbia | OP700686 | ||||
Platycryptus undatus (De Geer, 1778) | S72 | U.S.A.: Florida | AF327935* | AF327963*/AF328022* | AF327992* | |
d462 | Canada: Ontario: St. Williams | OP700702 | ||||
Psecas cf. viridipurpureus (Simon, 1901) | S227 | Ecuador: Sucumbios | AY297273* | AY297336* | AY297400* | |
Psecas sp. | GR124 | Ecuador: Napo | OP605983 | OP700703 | OP700687 |
K. mulu Maddison & Ruiz, sp. nov.
K. mulu species group:
Kelawakaju mulu Maddison & Ruiz, sp. nov.
Kelawakaju intexta Maddison & Ruiz, sp. nov.
K. singapura species group:
Kelawakaju singapura Maddison & Ng, sp. nov.
K. frenata species group:
Kelawakaju frenata (Simon, 1901)
Kelawakaju leucomelas Maddison & Ng, sp. nov.
Kelawakaju sahyadri Vishnudas, Maddison, & Sudhikumar, sp. nov.
The name means tree spider in the Berawan language from the area of Long Terawan, Sarawak (kelawak = spider; kaju or kajuh = tree; Syria Lejau Malang, pers. comm.), where the first specimens of K. mulu were found. To be treated grammatically as feminine.
Elongate and flat-bodied salticids, unusual among marpissines for the elongated or enlarged male chelicerae. Retrolateral tibial apophysis of palp long, blade-like, more or less straight and parallel to axis of palp. Embolus relatively short among marpissines, arising more or less terminally on the bulb (9–12 o’clock in ventral view of left palp). Markings cryptic on tree trunks, either mottled or with low-contrast longitudinal bands.
Carapace
flat, narrower (K. mulu group, Figs
We recognize three species groups in the genus.
The mulu species group includes K. mulu, K. intexta, and a third as-yet-undescribed species from Singapore. They are smaller-bodied than other Kelawakaju, with mottled markings, and narrow chelicerae that project forward in the male. The embolus is narrow and forms a smooth curve bending toward the retrolateral. The lower part of the thorax has three vertical stripes of pale scales on each side. Epigynal openings are delicate and the edges difficult to discern (Fig.
Holotype
: male (SWK12-2610) in
From the name of the type locality (a noun in apposition).
Dark with only a dusting of golden scales, unlike the similar but more thoroughly scale-covered K. intexta (Figs
Chelicerae of Kelawakaju species 2–4 oblique view with carapace 5–9 ventral view 2 K. mulu male holotype 3 K. intexta male holotype 4 K. frenata male from Guangxi 5 K. mulu male holotype 6 K. intexta male holotype 7 K. singapura male holotype 8 K. frenata male from Guangxi 9 K. frenata female from Guangxi. Scale bars: 1.0 mm.
Male (based on holotype). Carapace length 2.85; abdomen length 3.05. Carapace dark brown, with white scales around cephalic region, between AME and sparse on thoracic region. Clypeus very narrow. Chelicera dark brown, elongate and projected, with a line of white scales on the prolateral face. Retromarginal tooth with two cusps, the more lateral long and curved (Fig.
Female (based on paratype SWK 12-2639). Carapace length 2.7; abdomen length 3.55. Color as in male, except when mentioned. Chelicera light brown. Retromarginal tooth with two cusps, the distal one almost twice the size of the other, both acute. Legs II–IV with narrow stripes of white scales. Length of femur I 1.90, II 1.50, III 1.45, IV 1.85; patella + tibia I 2.60, II 1.90, III 1.75, IV 2.60; metatarsus + tarsus I 1.40, II 1.30, III 1.60, IV 1.90. Leg spines as in male, except for femur III, as in II. Abdomen as in male, except for stripes, medially interrupted; ventrally white, with two longitudinal dark brown stripes extending from booklungs to spinnerets. Epigyne with a pair of small copulatory openings distant from the posterior border, which has a medial excavation; internally, copulatory ducts fuse with glandular portions, spiral backwards and enter the large spermathecae, from which fertilization ducts emerge.
Both specimens were collected on tree trunks on a forested slope.
Holotype
: male (SWK12-3752) in
Latin, interwoven, referring to the textile-like pattern of coloured scales on the body.
Body covered with a dense and intricate pattern of pale scales, white on the abdomen and slightly golden on the carapace (Figs
Male (based on holotype). Carapace length 2.45; abdomen length 3.45. Carapace dark brown, with white scales on cephalic region, sparse on thoracic region and with line of white scales along borders of carapace. Clypeus very narrow. Chelicera dark brown, slightly projected, with mastidion. Retromarginal tooth with two cusps, the more lateral larger (Fig.
Female unknown.
The paratype from Bako was found along a trail in a forest.
The singapura species group includes only K. singapura, distinctive for the robust male chelicerae, short and stout embolus, and the long palp tibia (longer than the tibial apophysis). It is larger-bodied, like the frenata group, but has a longer ocular quadrangle, and the abdominal markings are inverted compared to the frenata group: dark laterally, paler medially, similar to K. intexta of the mulu group. There is no clear indication to which of the other two groups K. singapura is more closely related, and hence we keep it separate.
Holotype
: male (JK.21.08.02.0001) in LKCNHM from Singapore: Labrador Nature Reserve, 1.2653°N, 103.8019°E, J.K.H. Koh & P.Y.C. Ng, 2 August 2021. Paratypes: One female (JK.21.05.14.0001) in LKCNHM from Singapore: Labrador Nature Reserve, 1.2664°N, 103.8014°E, J.K.H. Koh & P.Y.C. Ng, 14 May 2021. One male (90.10.21.0002) in LKCNHM from Singapore: Simpang, 1.44°N, 103.85°E, J.K.H. Koh, 21 October 1990. One female (AS19.0023) in
From name of the type locality, Singapura in the Malay language, a noun in apposition.
Distinctive for the abdomen’s central pale longitudinal band with wavy edge (Figs
Male (based on holotype). Carapace length 3.1, width 2.3; abdomen length 3.9. Carapace (Figs
Female (based on specimen AS19.0023). Carapace length 3.2, width 2.2; abdomen length 3.9. Carapace: As in male, but not quite so wide, and with two fine vertical lines of pale scales on lower posterior thorax (Figs
Kelawakaju singapura 35 holotype male JK.21.08.02.001 embolus, ventral 36 paratype male JK.90.10.21.0002 palp, ventral 37 same, retrolateral 38 paratype female AS19.0023 epigyne, ventral 39 same, vulva, dorsal 40–42 holotype male 43 paratype female JK.21.05.14.0001 44, 45 paratype female AS19.0023. Scale bars: 0.1 mm.
The holotype and females from the type locality were found under bark of both small and large trees in open areas at the edge of coastal forest. Female AS19.0023 was found under bark of large tree in roadside clearing.
The frenata species group includes the relatively large-bodied K. frenata, K. leucomelas, and K. sahyadri. They differ from other Kelawakaju in having pale longitudinal bands on the sides of the body, a proportionately shorter ocular quadrangle, and longer first legs in the male. The male chelicerae diverge but do not project as forward as in the mulu group. A narrow band of white scales descends along the front face of the male chelicerae (Figs
Ocrisiona frenata Simon, 1901.
The type specimen of Ocrisiona frenata Simon, 1901 has not been found, neither in the Oxford Natural History Museum (O. Pickard Cambridge collection;
Differs from other Kelawakaju in the embolus bending suddenly toward the retrolateral, the epigynal atria with sclerotized edge both anteriorly and posteriorly (not just medially or posteriorly), and posterior notch of epigyne narrow and distinct.
Male (based on specimen from Dongxing City). Carapace length 3.1; abdomen length 4.1. Carapace dark brown, with sparse white scales. Clypeus very narrow. Chelicera dark brown, with a line of white scales on the prolateral face. One retromarginal tooth (Fig.
Kelawakaju frenata species group, genitalia 46–54 K. frenata 46 male from Dongxing, palp, ventral 47 same, retrolateral 48 same, dorsal 49 male from Hong Kong, embolus, ventral 50
Kelawakaju frenata 73, 74 male from Tai Tam County Park, Hong Kong (© 2020 Artur Tomaszek) 75 female from Guangdong, Gaotan Town 76 male from Guangxi, Dongxing, dorsal 77 female from Dongxing, dorsal 78 same, ventral. Specimen in 73, 74 not examined microscopically; inferred as K. frenata by appearance and locality.
Female (based on specimen from Dongxing City). Carapace length 3.45; abdomen length 5.55. Color as in male, except when mentioned. Chelicerae dark. One retromarginal tooth. Leg I light brown, with median third of femur, distal of patella and proximal and distal of tibia dark brown; tarsus yellow; II yellow, with same markings as I; III and IV as II, but with patellae entirely yellow and prolateral side of tibia dark brown. Tibia with three pairs of ventral macrosetae. Abdominal pattern as in male; no scutum. Epigyne with a pair of small copulatory openings distant from the posterior border, which has a medial excavation; internally, copulatory ducts fuse with glandular portions, spiral backwards and enter the large spermathecae, from which fertilization ducts emerge.
One male and two females in
Holotype : male (JK.20.11.13.003) in LKCNHM from Singapore: Lorong Pang Sua 1.3833°N, 103.7567°E, 13 xi 2020, J.K.H. Koh & P. Y. C. Ng. Paratypes: Two females (JK.20.11.13.0001 and JK.20.11.13.0002) with same data as holotype. One male (JK.13.12.10.0001) from Singapore: Pulau Tekong, 1.4072°N, 104.0283°E, 10 December 2013, J.K.H. Koh.
Refers to the longitudinal bands of white scales (leuco, Greek for white) on either side of the body contrasting against the black median (melas, Greek for black), formed not as an adjective but more simply as the two colours themselves (and thus without an expectation of agreement with the genus name).
Carapace wider and ocular quadrangle shorter (Fig.
Male (based on holotype). Carapace length 3.6, width 2.6; abdomen length 4.2. Carapace: Relatively flat, broad, depressed around fovea. Integument black to dark brown, clothed loosely with white scales in two broad longitudinal bands along sides, below and behind PME. Below these bands, thorax is black, without marginal white scales. Clypeus very narrow, dark, with some black hairs. Chelicerae diverging, projecting forward slightly, with a bulge anteriolaterally (as in K. frenata, Fig.
Female (based on paratype JK.20.11.13.0001). Carapace length 3.2, width 2.4; abdomen length 4.3. Carapace: As in male, but narrower. Clypeus as in male. Chelicerae with bulge and tubercles, but less prominent than in male. Dark, with black setae. Two promarginal and one unident retromarginal tooth, similar to those of K. frenata (Fig.
Male (JK.19.08.18.0010) in LKCNHM from MALAYSIA: Johor Bahru, Kota Tinggi, Panti Recreational Forest, 1.7872°N, 103.9425°E, 18 August 2019, P.Y.C. Ng.
Approximately ten adult and juvenile specimens were seen on tree bark at the type locality, including the holotype. The male from Kota Tinggi was found on tree bark in a sunny area near the entrance of Panti Recreational Forest.
Holotype : male (AS19.4895 = NCBS IBC-BP847) in NCBS from India: Karnataka: Kodagu: Yavakapadi, Honey Valley area, 12.2224°N, 75.6553°E, 1045 m elev., 27 June 2019, W. Maddison WPM#19-083. Paratypes: Female (AS19.4934 = NCBS IBC-BP848) in NCBS with data as holotype except 12.2214°N, 75.6556°E and 1130 m elev. One male and one female in CATE from India: Kerala: along state highway 21 east of Chalakudy, 10.296°N, 76.685°E, 26 June 2021, Vishnudas & Sudhikumar CATE9826705. One female with same data but 17 July 2021.
From the Sanskrit for ’from the Western Ghats mountains’, where this species lives.
Embolus differs in shape from that of the similar K. leucomelas: embolus tip with retrolateral flange less distinct and smaller (Figs
Male (based on holotype). Carapace length 3.0, width 2.1; abdomen length 3.9. Carapace: Relatively flat; area around fovea slightly depressed. Dark brown, loosely clothed with white scales below and behind PLE forming an indistinct longitudinal band on each side. Clypeus very narrow, dark, with black setae. Chelicerae diverging slightly, lacking the distinct bulge of K. frenata and K. leucomelas, but with hair-bearing tubercles. Narrow stripes of white scales form inverted V as in other frenata group species (Fig.
Female (based on specimen NCBS IBC-BP848). Carapace length 4.0, width 2.9; abdomen length 5.0. Carapace, Clypeus as in male. Chelicerae dark, with black hairs arising from small tubercles. Two promarginal and one larger triangular retromarginal teeth. Legs: First leg darkest, but all legs have dark patches, especially the sides of the femora and near the joints. Length of femur I 2.4, II 1.9, III 1.8, IV 2.1; patella + tibia I 3.3, II 2.5, III 2.3, IV 3.3; metatarsus + tarsus I 2.1, II 1.6, III 2.0, IV 2.3. Leg spination reduced: femur I d0-1-0, p0-3-0 or 2-0, II d1-1-0, p0-0-1, III d1-1-0, p0-0-1, IV d1-1-0, 0; patella I–IV 0; tibia I v2-2-2 (asymmetrical), II v1r-1r-1p, III 0, IV 0; metatarsus I–II v2-2, III v0-0-1p, IV v0-0-1p. Abdomen long, narrow, dark medially and pale laterally. Epigyne with two crescent-shaped openings behind a pale desclerotized area (Fig.
The holotype and female paratype from Kodagu were found under loose bark of large trees standing in a field, beside a small road. The specimens from Kerala were found in crevices in the bark of Swietenia mahogany trees.
For their assistance with field work, with thanks: in Sarawak, Malaysia, Edyta Piascik, Alex Ang, Ch’ien Lee, Syria Lejau Malang, Andyson Raymond Laman, the Sarawak forestry department, and the staff at Mulu and Lambir Hills National Parks; in Singapore, Joseph Koh and Peifen Koh; in Chalakudy, India, Ramesh V.T.; and in Karnataka, India, Kiran Marathe and Abijith A.P.C. We are grateful to Krushnamegh Kunte (NCBS, Bengaluru) for arranging access to facilities and specimens, and to Wendy Wang and Peter Ng (LKCNHM, Singapore) for loan of specimens. Alex Ng kindly offered the specimen of K. frenata from Hong Kong. E. Piascik assisted with DNA sequencing. We thank Chris Ang and Artur Tomaszek for permission to use their photographs, and the developers of iNaturalist for giving us access to the observations of naturalists exploring the landscape. G. Azarkina, J Schubert, and an anonymous reviewer provided helpful comments on an earlier version of the manuscript. Funding to WPM was provided by an NSERC Canada Discovery Grant. Funding to AVS was provided by a DST-SERB grant (Major Research Project EEQ/2021/000453) and an eGrantz from the government of Kerala.