ZooKeys 375: 75–99, doi: 10.3897/zookeys.375.6116
Acanoides gen. n., a new spider genus from China with a note on the taxonomic status of Acanthoneta Eskov & Marusik, 1992 (Araneae, Linyphiidae, Micronetinae)
Ning Sun 1,†, Yuri M. Marusik 2,3,‡, Lihong Tu 1,§
1 College of Life Sciences, Capital Normal University, Xisanhuanbeilu Str. 105, Haidian Dist., Beijing, 100048, P. R. China
2 Institute for Biological Problems of the North, FEB Russian Academy of Sciences, Portovaya Str. 18, Magadan, 685000, Russia
3 Zoological Museum, University of Turku, FI-20014 Turku, Finland

Corresponding author: Lihong Tu (tulh@cnu.edu.cn)

Academic editor: D. Dimitrov

received 17 August 2013 | accepted 16 December 2013 | Published 30 January 2014


(C) 2014 Ning Sun. 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.


For reference, use of the paginated PDF or printed version of this article is recommended.

Citation: Sun N, Marusik YM, Tu L (2014) Acanoides gen. n., a new spider genus from China with a note on the taxonomic status of Acanthoneta Eskov & Marusik, 1992 (Araneae, Linyphiidae, Micronetinae). ZooKeys 375: 75–99. doi: 10.3897/zookeys.375.6116

Abstract

A new “micronetine” genus Acanoides gen. n. is erected to accommodate two species from China: Acanoides beijingensis sp. n. as the type species and Acanoides hengshanensis (Chen & Yin, 2000), comb. n., with the females described for the first time. The genitalic characters and somatic features of the new genus were studied by means of light microscopy and scanning electron microscopy (SEM). The monophyly of the new genus was tested by a phylogenetic analysis based on molecular data. Descriptions of the new genus, the new species and the new combination are presented; SEM images and microscopy pictures of somatic and genitalic characters are provided in detail. To distinguish from other genera with similar genitalic characters, we compare the new genus with the species of Acanthoneta Eskov & Marusik, 1992, Epibellowia Tanasevitch, 1996 and Wubanoides Eskov, 1986. Four putative synapomorphies for Acanoides gen. n. are suggested to support its monophyly that could be tested in the future. Furthermore, redescriptions of the epigynal morphology of Acanthoneta aggressa Chamberlin & Ivie, 1943 (Nearctic) and on the male of A. dokutchaevi Eskov & Marusik, 1993 (Far East Asia, firstly recorded from China) are provided. Based on comparison with Poeciloneta, from which Acanthoneta stat. n. was separated by Saaristo and Tanasevitch (1996), a revised diagnosis is proposed to support the generic status.

Keywords

Taxonomy, new species, new genus, genitalic morphology, movable epigynum

Introduction

Micronetinae Hull, 1920 is a fairly large subfamily of Linyphiidae Blackwall, 1859, including 1199 species placed in 90 genera (Tanasevitch 2014). It was redelimited by Saaristo and Tanasevitch (1996), who established eight new genera for 89 species, and raised three subgenera to generic status. Subsequently, a series of revisions were published (e.g. Saaristo and Tanasevitch 2002a, 2002b; Saaristo and Marusik 2004; Saaristo et al. 2006; Tu et al. 2006; Tu and Li 2006) that resulted in many new genera and a new subfamily Ipainae Saaristo, 2007. Results of these series of review works have not been tested in a phylogenetic context; neither Micronetinae nor Ipainae monophyly, as well as that of the genera included in the two subfamilies have been tested so far.

Poeciloneta hengshanensis (Chen & Yin, 2000) from China, originally placed in Lepthyphantes Menge, 1866, has its male palp very similar to that of Poeciloneta (Acanthoneta) aggressa (Chamberlin & Ivie, 1943). Acanthoneta Eskov & Marusik, 1992 is one of the three genera raised from subgeneric status by Saaristo and Tanasevitch (1996) with the type species Acanthoneta aggressa. Tu et al. (2006) transferred Poeciloneta hengshanensis to Acanthoneta based on the similarity of the male palpal morphology. However, raising Acanthoneta to a generic status “was not accompanied by a diagnosis or justification”, and hence not accepted in The World Spider Catalog (Platnick 2014). All members of Acanthoneta are currently placed in Poeciloneta Kulczyński, 1894.

Females of Poeciloneta hengshanensis (previously unknown) were found in new material from China. However, its epigynal conformation is neither congruent with that of Poeciloneta aggressa, nor with any other species of Poeciloneta. Based on the presence of an extensible basal part, the movable epigynum accords with the diagnosis of the subfamily Ipainae Saaristo, 2007 (for example Ipa Saaristo, 2007 and Solenysa Simon, 1894). Additionally, we found another new species with genitalic morphology very similar to that of Poeciloneta hengshanensis: the male palpal morphology similar to Acanthoneta and a movable epigynum in accordance with ipaine type.

A new genus Acanoides gen. n. is erected here for these two species. To test the placement of the new genus within Linyphiidae, a phylogenetic analysis based on newly sequenced molecular data of the two species and that of other linyphiids downloaded from NCBI was conducted. In the present study, the two species and the new genus are described. Characters of copulatory organs and somatic features of both species are illustrated by means of SEM and light microscopy. To distinguish the new genus from other “micronetine” genera with similar male palpal morphology and ipaine genera with a similar movable epigynum, the new genus is compared with the genera Acanthoneta (Micronetinae), Wubanoides Eskov, 1986 and Epibellowia Tanasevitch, 1996 (Ipainae). Due to limited material available for examination, comparisons are largely based on descriptions and illustrations provided by Tanasevitch (1996), Saaristo and Tanasevitch (2000) and Saaristo (2007); images of the epigynum of Acanthoneta aggressa and the male of Acanthoneta dokutchaevi Eskov and Marusik, 1994 are presented here. Four putative synapomorphies are suggested for Acanoides gen. n. that could be tested in future study. In addition, diagnoses for Acanthoneta stat. n. are provided based on comparison with illustrations of genitalic characters provided by Saaristo and Tanasevitch (2000), to support its generic status proposed by Saaristo and Tanasevitch (1996). The composition and monophyly of both Acanoides gen. n. and Acanthoneta stat. n. could be tested in future study.

Materials and methods

Specimens were examined and measured using a Leica M205A stereomicroscope. Male palps and epigyna were examined after they were dissected from the body. Left structures (e.g. palps, legs, etc.) were depicted. Embolic divisions were excised by breaking the membranous column which connects the suprategulum and radix. Male palps and epigyna were cleared in methyl salicylate. Digital images were taken with a Leica DFC 500 camera, as composites of multiple focus images assembled using the software package LEICA APPLICATION SUITE. Scanning electron microscopy (SEM) images were taken using a S-3400N scanning electron microscope at the China Agricultural University. For SEM examination the specimens were prepared following Álvarez-Padilla and Hormiga (2008). SEM images of the embolic division taken from the right palp were mirrored to match those taken from the left palp. All measurements were taken with a micrometer and are expressed in millimeters. The leg measurements are given in the following sequence: total (femur, patella+tibia, metatarsus, tarsus). All specimens examined here are deposited in the College of Life Sciences, Capital Normal University, China (CNU) and in the College of Life Sciences, Hunan Normal University, China (HNU), except for the female material of Acanthoneta aggressa, the epigynal pictures of which were provided by Don Buckle (Saskatoon, Canada). Distribution data for these species within China are presented at the provincial level. Terminology for the epigynal characters follows Tu and Hormiga (2010) and male palpal and somatic characters follows that of Saaristo and Tanasevitch (1996) and Hormiga (2000). Anatomical abbreviations used in the text and figures:

Somatic morphology

AER anterior eye row

ALE anterior lateral eye(s)

AME anterior median eye(s)

AMEd diameter of AME

PER posterior eye row

PLE posterior lateral eye(s)

Male palp

AX apex of embolus

DM distal membrane of terminal apophysis

DSA distal suprategular apophysis

EM embolic membrane

EP embolus proper

FiG Fickert’s gland

LC lamella characteristica

P paracymbium

PCA proximal cymbial apophysis

PH pit hook

R radix

SE serrated area on embolus

SPT suprategulum

TA terminal apophysis

TH thumb of embolus

Epigynum

CO copulatory opening

CG copulatory groove

DP dorsal plate

EA extensible area of epigynal basal part

EB epigynal basal part

FG fertilization groove

MP median plate

S spermathecae

SC scape

ST stretcher

VP ventral plate

Phylogenetic analysis

Based on the dataset of Arnedo et al. (2009) which includes 34 linyphiid taxa (Erigone dentipalpis was not included as it has only one of the five genes available), newly sequenced data of the two Acanoides and data of another 65 linyphiid taxa downloaded from NCBI were added. A total of 111 taxa were sampled in our matrix, ten outgroup taxa of other araneoid families as in that of Arnedo et al. (2009) and 101 ingroup taxa, which cover the representatives of all the seven subfamilies currently proposed; one Solenysa, as a representative of ipaine, and Acanthoneta were included to test the placement of Acanoides.

Five genes: cytochrome c oxidase subunit I (CO1) and 16S rRNA (16S) and three nuclear genes 18S rRNA (18S), 28S rRNA (28S) and Histone H3 (H3) were sequenced for Acanoides beijingensis sp. n. and Acanoides hengshanensis. Molecular procedures for sequencing follow that of Arnedo et al. (2009), with the same molecular markers to maximize the overlap of dataset. Taxa sampled and sequence accession numbers are presented in Appendix - Table S1. Data were automatic multiple aligned using the computer program Clustal X version 1.81 (Thompson et al. 1997). Gaps were treated as missing data. Maximum Likelihood analysis was performed using RAxML v7.2.7 as implemented on the Cipres Gateway (Miller et al. 2010). Bootstrap support analysis was performed with the commands: raxmlHPC-HYBRID-7.3.1 -T 6 -s infile -n result -x 12345 -p 876 -f a -N 1000 -m GTRCAT -q part.

Results

All five genes were sequenced for Acanoides beijingensis sp. n. and Acanoides hengshanensis (Appendix - Table S1). The monophyly of Linyphiidae and its sister relationship with Pimoidae were not recovered in the result of phylogenetic analysis as two outgroup taxa: cyatholipid Alaranea and theridiosomatid Theridiosoma are embedded within Linyphiidae (Appendix - Fig. S1). Besides some weakly supported deeper branches, four robustly supported clades are recognized: Stemonyphantes clade (clade S), “micronetines-erigonines” clade (clade ME), “linyphiines”-1 clade (clade L1) and “linyphiines”-2 (clade L2). For the seven subfamilies currently proposed, only Stemonyphantinae Wunderlich, 1986 (the Stemonyphantes clade) and Mynogleninae Lehtinen, 1967 are monophyletic, while the mynoglenines clade and the Dubiaranea clade fall into clades L1 and L2 respectively that make Linyphiinae Blackwall, 1859 become a paraphyletic group; taxa of Micronetinae form a paraphyletic group, nested with taxa of Ipainae and Erigoninae within clade ME. The two Acanoides species form a robustly supported monophyly, distantly related to Acanthoneta and Solenysa.

Discussion

The result of the phylogenetic analysis based on molecular data suggests that the new species from Beijing is the sister taxon of Poeciloneta hengshanensis which had ever been transferred to Acanthoneta by Tu et al. (2006). The lineage comprised by the two species is distantly related to Acanthoneta sp. (Appendix - Fig. S1). Accordingly, we erected here Acanoides gen. n. to accommodate the two species: Acanoides beijingensis sp. n. and Acanoides hengshanensis comb. n. The three known Acanthoneta species have very distinct male palpal morphology, only differ from that of the type species in small details (Eskov and Marusik 1992, 1993). Regardless the Acanthoneta taxon is congeneric with, or is the type species Acanthoneta aggresus, the new genus differs from all the three known species of Acanthoneta as well as all other “micronetines” in the females having a movable epigynum (Figs 4G, 5G) and the males having a longer and sharper embolus proper (Figs 2D, 3D) which generally is pointed in “micronetines” (Fig. 6F); Fickert’s gland located in the membranous area outside the radix (Figs 2D, 3D), rather than embedded within the radix as usually the case in “micronetines” (Fig. 6G). This suggests that the two species are not congeneric with Acanthoneta.

Our results suggest an unknown Lepthyphantes species as a sister group to the Acanoides clade. Lepthyphantes Menge, 1866, which includes almost 500 species, is not a natural group (Saaristo and Tanasevitch 1996). All Lepthyphantes species, except five, have been transferred or are waiting to be transferred to other genera (e.g. Saaristo and Tanasevitch 1996, 2002a, b; Saaristo and Marusik 2004; Tu et al. 2006). The position of Lepthyphantes sp. on the tree indicates it is neither Acanthoneta, nor Lepthyphantes. Nevertheless, without morphological data, we fail to determine whether Lepthyphantes sp. is as a sister group of, or a number of Acanoides gen. n., so the close relative of Acanoides remains unresolved.

The genitalic characters of Acanoides make its subfamily placement problematic due to the epigynal character in accordance with Ipainae type, but the male palpal morphology of the “micronetine” type. Redelimitation of Mironetinae (Saaristo and Tanasevitch 1996) and the series of revisions of “micronetine” genera (e.g. Saaristo and Tanasevitch 2002a, 2002b; Saaristo and Marusik 2004; Saaristo et al. 2006; Tu et al. 2006; Tu and Li 2006) resulted in many new genera and even a new subfamily Ipainae (Saaristo 2007). However, none of them has been tested in a phylogenetic framework. Results of the first phylogenetic analysis for linyphiids based on molecular data indicate that neither Micronetinae nor Ipainae is a monophyletic group (Arnedo et al. 2009). Such a result was recovered in the present study too: “micronetine” taxa formed a paraphyletic group, and movable epigynum independently evolved in Acanoides and Solenysa (Appendix - Fig. S1). The extensible solenoid serving as a synapomorphy for Solenysa (Tu & Hormiga, 2011), the ventrally folded extensible epigynal basal part, together with long and sharp embolus proper, slender and unbranched lamella characteristica, and outside radix located Fickert’s gland are four putative synapomorphies for Acanoides gen. n.

With greatly increased ingroup sampling, the result of the present study produce a similar topology with that of Arnedo et al. (2009): four strongly supported clades S, L1, L2 and ME that correspond to theStemonyphantes clade, clades C and D, and the “micronetines-erigonines” clade in the latter (Appendix - Fig. S1). Most newly added taxa fell into the clade ME that enriched the topology. However, the problems left by the previous study (Arnedo et al. 2009), such as the monophyly of Linyphiidae, placements of the weakly supported deeper branches, and taxa of different subfamilies placed together rendering most of the traditionally recognized subfamilies non monophyletic, persist. Six of the seven subfamilies currently proposed are not monophyletic groups. The higher level relationships within linyphiids reflected by phylogenetic result are still far away from the classic subfamily system (see Millidge 1984, 1993; Saaristo and Tanasevitch 1996; Saaristo 2007). Nevertheless, revising the whole higher level linyphiid systematics is beyond the scope of the present study. In the text bellow we keep using Micronetinae and Ipainae following the current taxonomic system.

Although with ingroup sampling about two times increased, the sampling size of the current dataset seems not to be enough to resolve the placements of Acanoides and Acanthoneta, as well as Poeciloneta, from which Acanthoneta were separated (Eskov and Marusik 1992), their close relatives, and the relationships among them. To better understand the higher level phylogenetic relationships of linyphiid spiders, more information, such as morphology and behavior, and a comprehensive sampling design are necessary. Here, we provide four putative synapomorphies for the new genus Acanoides that could be tested in future phylogenetic studies.

Taxonomy
Linyphiidae Blackwall, 1859
Type species.

Acanoides beijingensis sp. n.

Composition.

Two species, Acanoides beijingensis sp. n. and Acanoides hengshanensis (Chen & Yin, 2000) comb. n.

Diagnosis.

The males of Acanoides gen. n. can be distinct from Acanthoneta by the sharp embolus proper, the slender lamella characteristica unbranched, and by the Fickert’s gland located in the membranous area outside the radix (Figs 2D, 3D). The females can be distinguished by the ventrally folded extensible epigynal basal part (Figs 2F, 3F).

Description.

Male total length 2.34–2.73; female total length 2.10–2.42. Carapace yellowish-brown. Male carapace unmodified. AMEs smallest, others subequal; from the dorsal view AER recurved, PER straight, eyes separated by AMEd, ALE and PLE juxtaposed. Chelicerae medium-sized, with strong stridulatory ridges, female fang groove with three promarginal and three retromarginal teeth in Acanoides beijingensis sp. n., and two promarginal and two retromarginal teeth in Acanoides hengshanensis. Chaetotaxy: Ti I–IV: 2-2-2-2; Mt I–IV: 1-1-1-1; Mt I of males with two rows of ventral bristles, one prolateral, one retrolateral (Fig. 1C, 1D); Tm I about 0.25, Tm IV absent. Both species have a haplotracheate system.

Figure 1.

Acanoides beijingensis sp. n. (A–C) and Acanoides hengshanensis (D–F). A male, dorsal B female, dorsal C male, lateral, rectangle indicates ventrolateral rows of bristles on Mt I D male, lateral, rectangle indicates ventrolateral rows of bristles on Mt I E male, dorsal F female, dorsal. [Scale bars: mm].

Male palp (Figs 2A–E, 3A–E, 4A–B, 5A–B). Cymbium with proximal apophysis. Paracymbium medium to large-sized, with one tooth on lateral margin. Distal suprategular apophysis not modified as pit hook, or absent. Embolic division: radix long and narrow, Fickert’s gland located in the membranous area connecting radix and embolus; embolus wide and strongly sclerotized with serrated area, embolus proper sharp with a thumb and an apex at each side; lamella characteristica unbranched, long and narrow with sharp sclerotized apex, almost parallel to radix; terminal apophysis with distal membrane.

Figure 2.

Acanoides beijingensis sp. n. A male palp, prolateral B male palp, prolateral, with embolic division removed C male palp, retrolateral D embolic division, ventral E embolic division, dorsal F epigynum, ventral G epigynum, dorsal H epigynum, lateral. CG copulatory groove; CO copulatory opening; DP dorsal plate; EA extensible area of epigynal basal part; EM embolic membrane; EP embolus proper; FG fertilization groove; FiG Fickert’s gland; LC lamella characteristica; MP median plate; P paracymbium; PCA proximal cymbial apophysis; R radix; S spermathecae; TA terminal apophysis; TH thumb of embolus; VP ventral plate. [Scale bars: mm].

Figure 3.

Acanoides hengshanensis. A male palp, prolateral B male palp, ventral C male palp, retrolateral, arrow indicates pointed tooth on posterolateral margin D embolic division, ventral E embolic division, dorsal F epigynum, ventral G epigynum, dorsal. CG copulatory groove; CO copulatory opening; DP dorsal plate; EA extensible area of epigynal basal part; EM embolic membrane; EP embolus proper; FG fertilization groove; FiG Fickert’s gland; LC lamella characteristica; P paracymbium; PCA proximal cymbial apophysis; R radix; S spermatheca; TA terminal apophysis; TH thumb of embolus; VP ventral plate. [Scale bars: mm].

Figure 4.

Acanoides beijingensis sp. n. A palp (embolic division removed), prolateral B palp, retrolateral, arrow indicates half rounded lateral tooth on paracymbium C embolic division, ventral D embolic division, dorsal E detail of D F detail of C G epigynum, ventral H epigynum, dorsal. AX apex of embolus; CG copulatory groove; CO copulatory opening; DM distal membrane of terminal apophysis; DSA distal suprategular apophysis; EA extensible area of epigynal basal part; EM embolic membrane; EP embolus proper; FG fertilization groove; LC lamella characteristica; MP median plate; P paracymbium; PCA proximal cymbial apophysis; R radix; S spermatheca; SE serrated area on embolus; SPT suprategulum; TA terminal apophysis; TH thumb of embolus; VP ventral plate. [Scale bars: mm].

Figure 5.

Acanoides hengshanensis. A palp (embolic division removed), prolateral B palp, retrolateral, arrow indicates pointed tooth on posterolateral margin C embolic division, ventral D embolic division, dorsal E detail of D F detail of C G epigynum, ventral H epigynum, dorsal. AX apex of embolus; CG copulatory groove; CO copulatory opening; DM distal membrane of terminal apophysis; EA extensible area of epigynal basal part; EM embolic membrane; EP embolus proper; FG fertilization groove; LC lamella characteristica; P paracymbium; PCA proximal cymbial apophysis; R radix; S spermatheca; SPT suprategulum; TA terminal apophysis; TH thumb of embolus; VP ventral plate. [Scale bars: mm].

Epigynum (Figs 2F–H, 3F–G, 4G–H, 5G–H). Protruding, with deeply wrinkled basal part, extensible and ventrally folded in constricted state. Epigynum well sclerotized, epigynal cavity present (in Acanoides beijingensis sp. n.) or absent (in Acanoides hengshanensis), both scape and stretcher absent.

Etymology.

The genus name, Acanoides, is a combination of the first four letters of “Acanthoneta” and the last five letters of “Wubanoides”. “-oides” itself in Latin means “similar to”, masculine in gender.

Phylogenetics.

Due to limitations of the current dataset the monophyly of Acanoides could not be tested explicitly in our phylogenetic analyses, however it is supported by the following four putative synapomorphies: sharp embolus proper, slender and unbranched lamella characteristica, outside radix located Fickert’s gland and ventrally folded extensible epigynal basal part.

Distribution.

China (Beijing, Hunan, Hebei) (Fig. 7).

Remarks.

The males of Acanoides gen. n. have the palp of a “micronetine” type: presence of the Fickert’s gland, the boat-shaped radix, the trunk-like embolus with a pointed proper and a thumb, as well as the well developed terminal apophysis and lamella characteristica (Saaristo and Tanasevitch 1996). However, these sclerites in Acanoides (Fig. 2D) have some features different from the normal “micronetine” type (Fig. 6F, Saaristo and Tanasevitch 1996): Fickert’s gland is not embedded within the radix, but located in the membranous area connecting the radix and the embolus; and the embolus has a wide, strongly sclerotized body, with a longer and sharper embolus proper, whereas in most “micronetines” the embolus is usually trunk-like with a pointed embolus proper. The male palp of both Acanoides and Acanthoneta, have a long and slender lamella characteristica parallel to the long radix, but with an additional long and thin branch in Acanthoneta (Fig. 6F), unbranched in Acanoides (Figs 2D, 3D). The epigynum of Acanthoneta is in a normal “micronetine” type, with a sigmoid scape surrounded by an epigynal cavity (Fig. 6H), but with an extensible basal part in Acanoides.

Figure 6.

Acanthoneta dokutchaevi (A–G) and Acanthoneta aggressa (H–J). A male, dorsal B male, lateral C male palp, prolateral D male palp, ventral E male palp, retrolateral F embolic division, ventral G embolic division, dorsal H epigynum, ventral I epigynum, posterior J epigynum, lateral (H–J photos provided by Don Buckle). EB epigynal basal part; EM embolic membrane; EP embolus proper; FiG Fickert’s gland; LC lamella characteristica; P paracymbium; PCA proximal cymbial apophysis; PH pit hook; R radix; SC scape; ST stretcher; TA terminal apophysis; TH thumb of embolus. [Scale bars: mm].

Figure 7.

Collecting localities of Acanoides species and Acanthoneta aggressa, Acanoides beijingensis sp. n. (Beijing, Hebei); Acanoides hengshanensis (Hunan, Beijing); Acanthoneta dokutchaevi (Jilin).

The result of phylogenetic analysis based on molecular data indicates that Ipainae is not a monophyletic group as the movable epigynum independently evolved in Acanoides and Solenysa (Appendix - Fig. S1). This is also supported by the tracheal characters: haplotracheate type in Acanoides, but intermediate type in Solenysa, with the median pair extending into the prosoma (Tu and Hormiga 2011). We infer that the extensible basal part of the epigynum may have also evolved convergently with that in other ipaines. In Acanoides it differs by being ventrally folded, while it forms a solenoid in Solenysa (Tu & Hormiga, 2011), and folds inwards in other ipaines, e.g. Ipa (Saaristo 2007: fig. 29), Wubanoides and Epibellowia (Tanasevitch 1996: figs 7–9). Furthermore, the male palp of typical ipaines has filiform embolus proper (Saaristo 2007: fig. 7; Tanasevitch 1996: figs 1, 4) much longer than that of Acanoides (Fig. 2D).

Type-locality.

China, Beijing: Mt. Yangtaishan, 39°20.15'N, 115°34.52'E, alt. ca 320m, 15 Oct. 2007, L. Tu leg.

Type-specimens.

Holotype, ♂ (CNU), China, Beijing, Mt. Yangtaishan, 39°20.15'N, 115°34.52'E, alt. ca 320 m, 15 Oct. 2007, L. Tu leg. Paratypes, 2 ♂♂ and 3 ♀♀ (CNU), same data as holotype.

Additional material examined.

1 ♂ and 2 ♀♀ (CNU), China, Hebei Province, Mt. Wulingshan, 40°33.61'N, 117°29.69'E, alt. ca 1100 m, 12 Aug. 2009, L. Tu leg.

Diagnosis.

The male of Acanoides beijingensis sp. n. can be distinguished from Acanoides hengshanensis by the spine-shaped lamella characteristica (Figs 2D, 4C), ribbon-like in the latter (Figs 3D, 5C); by the hook-shaped terminal apophysis (Fig. 4C), straight in the latter (Fig. 5D); and by the presence of a distal suprategular apophysis (Fig. 4A), absent in the latter. The female is distinct by having the epigynum two times longer than wide (Fig. 2F), shorter than wide in Acanoides hengshanensis (Fig. 3F); and by the presence of a remnant epigynal cavity (Fig. 2G), totally absent in Acanoides hengshanensis (Fig. 3G).

Description.

Male holotype (Fig. 1A, C): Total length 2.69. Carapace 1.22 long, 1.01 wide. Abdomen 1.39 long, 0.88 wide. Lengths of legs: I 3.88 (1.05 + 1.18 + 0.99 + 0.66); II 3.02 (1.03 + 0.73 + 0.69 + 0.57); III 2.66 (0.87 + 0.88 + 0.51 + 0.40); IV 3.78 (1.12 + 1.09 + 0.93 + 0.64). Female (Fig. 1B): Total length 2.12. Carapace 0.93 long, 0.78 wide. Abdomen 1.25 long, 0.83 wide. Lengths of legs: I 6.10 (1.68 + 2.04 + 1.43 + 0.95); II 5.43 (1.56 + 1.74 + 1.24 + 0.89); III 4.39 (1.24 + 1.13 + 1.10 + 0.75); IV 5.88 (1.79 + 1.78 + 1.46 + 0.83). Tm I: 0.20. For other somatic features see description of the genus.

Male palp (Figs 2A–C, 4A–B). Cymbium with proximal apophysis. Paracymbium narrow, half rounded lateral tooth strongly sclerotized. Distal suprategular apophysis blunt, not modified as pit hook. Embolic division: radix long and narrow; Fickert’s gland located in the membranous area connecting radix and embolus; embolus main body short and wide, strongly sclerotized, with serrated area on ventral surface; embolus proper sharp with pointed thumb and tail-like apex at each side; unbranched lamella characteristica long and slender, with sharp and strongly sclerotized apex; terminal apophysis hook-shaped with distal membrane.

Epigynum (Figs 2F–H, 4G–H). Two times longer than wide, wrinkled basal part extensible and ventrally folded in constricted state. Median plate and epigynal cavity present, without scape and stretcher. Copulatory openings opened dorsally.

Etymology.

The species name refers to the type locality.

Variation.

Males (n = 3). Total length 2.61–2.73. Carapace: 1.13–1.27 long, 0.95–1.05 wide. Abdomen 1.34–1.45 long, 0.71–0.99 wide.

Females (n = 3). Total length 2.10–2.23. Carapace: 0.90–0.96 long, 0.74–0.78 wide. Abdomen: 1.10–1.38 long, 0.79–0.88 wide.

Distribution.

China (Beijing, Hebei) (Fig. 7).

Remarks.

Although Acanoides beijingensis sp. n. looks quite different from Acanoides hengshanensis in the shape of the male paracymbium and in terms of female epigynal morphology, the strongly sclerotized embolus main body and the sharp embolus proper, the location of Fickert’s gland, the presence of a ventrally folded extensive area of the epigynal basal part and the absence of a scape and stretcher, shared by the two species suggest they are closely related. A close relationship between the two species is additionally supported by the phylogenetic analysis (Appendix - Fig. S1).

Acanoides hengshanensis (Chen & Yin, 2000), comb. n.

http://species-id.net/wiki/Acanoides_hengshanensis

Figs 1D–F, 3, 5
Lepthyphantes hengshanensis Chen & Yin, 2000: 87, figs 12–16 (♂)
Acanthoneta hengshanensis: Tu et al. 2006: 412, figs 24–27 (♂).
Type-specimen.

Holotype of Lepthyphantes hengshanensis Chen & Yin, 2000, ♂ (HNU), China, Hunan Province, Mt. Hengshan, 27°18'N, 112°42'E, 1–7 Aug. 1995, C. Yin leg. (examined).

Additional material examined.

3 ♂♂ and 4 ♀♀, China, Beijing, Mt. Yangtaishan, Dajue Temple, 40°03.06'N, 116°05.97'E, alt. 50 m, 15 Oct. 2007, L. Tu leg.

Diagnosis.

See diagnosis for Acanoides beijingensis sp. n.

Description.

Male (Fig. 1D–E): Total length 2.39. Carapace 1.02 long, 0.78 wide. Abdomen 1.37 long, 0.78 wide. Lengths of legs: I 5.03 (1.37 + 1.56 + 1.32 + 0.78), II 3.33 (0.98 + 0.98 + 0.83 + 0.54), III 3.47 (0.98 + 1.07 + 0.88 + 0.54), IV 4.63 (1.27 + 1.41 + 1.22 + 0.73). Tm I: 0.24. Female (Fig. 1F): Total length 2.42. Carapace 0.96 long, 0.78 wide. Abdomen 1.80 long, 1.25 wide. Lengths of legs: I 4.21 (1.18+ 1.42 + 0.96 + 0.65), II 3.19 (0.98 + 1.06 + 0.66 + 0.49), III 2.81 (0.84 + 0.85 + 0.68 + 0.44), IV 3.70 (1.08 + 1.19 + 0.89 + 0.54). Tm I: 0.23. For other somatic characters see description of the genus.

Male palp (Figs 3A–C; 5A–B). Cymbium with distinct proximal apophysis pointing backwards. Paracymbium wide and U-shaped, with triangular tooth on posterolateral margin. Distal suprategular apophysis absent. Embolic division: radix long and narrow; Fickert’s gland located in the membranous area connecting radix and embolus; embolus main body large and strongly sclerotized with serrated area; embolus proper sharp with large thumb and pointed apex; lamella characteristica long and slender with bifurcated ends, one sharp and sclerotized, one membranous; terminal apophysis straight, with distal membrane.

Epigynum (Figs 3F–G, 5G–H). Short and wide, box-shaped, strongly sclerotized; wrinkled basal part extensible and ventrally folded in constricted state. Neither median plate nor epigynal cavity present. Copulatory openings located on ventral surface, slits of epigynal grooves extending laterally, passing from ventral to dorsal surface, then convergent mesally. No scape, no stretcher.

Variation.

Males (n = 3). Total length 2.34–2.41. Carapace: 1.09–1.12 long, 0.72–0.93 wide. Abdomen 1.14–1.42 long, 0.68–0.83 wide.

Females (n = 4). Total length 2.32–2.42. Carapace: 0.87–1.01 long, 0.75–0.81 wide. Abdomen: 1.63–1.82 long, 0.76–1.22 wide.

Distribution.

China (Beijing, Hunan) (Fig. 7).

Genus Acanthoneta Eskov & Marusik, 1992, stat. n.
Acanthoneta Eskov & Marusik, 1992: 34. Described as a subgenus of Poeciloneta.
Acanthoneta: Saaristo and Tanasevitch 1996: 175. Raised to generic status without any comments or argumentation.
Type species.

Poeciloneta aggressus (Chamberlin & Ivie, 1943).

Composition.

Three species: Acanthoneta aggressa Chamberlin & Ivie, 1943 (Nearctic), Acanthoneta dokutchaevi Eskov & Marusik, 1993 (Far East Asia) and Acanthoneta furcata Emerton, 1913 (Nearctic).

Comments.

Originally Acanthoneta was described as a subgenus of Poeciloneta, including two species: Poeciloneta (Acanthoneta) aggressa and Poeciloneta (Acanthoneta) furcata. One additional species Acanthoneta dokutchaevi was assigned to the subgenus by Eskov and Marusik (1993). Saaristo and Tanasevitch (1996) raised Acanthoneta to genus status without any argumentations and hence the new status was not accepted by Platnick (2014). Here we provide a diagnosis for Acanthoneta and a comparison with Poeciloneta.

Diagnosis.

Males of Acanthoneta differ from Poeciloneta by the long radix almost parallel with the long lamella characteristica (Fig. 6F), in the latter the radix is normal boat-shaped, lamella characteristica large and ribbon-like (Saaristo and Tanasevitch 2000: fig. 11). Females of the two genera differ by the epigynum in Acanthoneta having a sigmoid scape surrounded by an epigynal cavity, the lateral wall of which is posteriorly extended (Fig. 6H), whereas in Poeciloneta the scape is exposed, enlarged and strongly sclerotized (Saaristo and Tanasevitch 2000: fig. 18).

Acanthoneta aggressa (Chamberlin & Ivie, 1943)

http://species-id.net/wiki/Acanthoneta_aggressa

Fig. 6H–J
Lepthyphantes aggressus Chamberlin & Ivie, 1943: 14, figs 19–20.
Poeciloneta aggressa: Crawford 1988: 19.
Acanthoneta aggressa: Saaristo and Tanasevitch 1996: 175.
Poeciloneta aggressa: Paquin and Dupérré 2003: 147, figs 1623–1625.
Material examined.

No material examined, epigynum pictures were provided by Don Buckle (Saskatoon, Canada): 1 ♀, Canada, Alberta, Chinook Lake, under log in spruce or fir woods, 49°40'N, 114°30'W, 25 Jul. 1988, D. J. Buckle leg.

Description.

Epigynum (Fig. 6H–J). Slightly protruding, without extensible area at basal part. Epigynal cavity, with posteriorly extended lateral wall, surrounding sigmoid folded scape; scape long and narrow, with well developed lateral lobes hosting copulatory openings and distal stretcher.

Distribution.

Across North America from Washington State to Québec (Buckle et al. 2001; Paquin and Dupérré 2003).

Acanthoneta dokutchaevi Eskov & Marusik, 1993

http://species-id.net/wiki/Acanthoneta_dokutchaevi

Fig. 6A–G
Poeciloneta (Acanthoneta) aggressa non Chamberlin & Ivie, 1943: Eskov and Marusik 1992: 34–35, figs 11–13 (♂).
Poeciloneta (Acanthoneta) dokutchaevi: Eskov and Marusik 1993: 52, figs 49–51 (♂).
Material examined.

1 ♂, China, Jilin Province, Mt. Changbaishan, Ski. 42°01.54'N, 128°04.25'E, alt. ca 1260 m, 31 July 1971.

Description.

Male (Fig. 6A–B). Chelicera long, with strong stridulatory ridges. Chaetotaxy: Ti I–IV: 2-2-2-2; Mt I–IV: 1-0-0-1; Tm I about 0.80, Tm IV present. For other somatic characters see description by Eskov and Marusik (1993).

Male palp (Fig. 6C–E). Cymbium with proximal apophysis erected. Paracymbium wide, with two pointed teeth on lateral margin. Distal suprategular apophysis modified as pit hook. Embolic division: radix long and narrow; Fickert’s gland located within radix; embolus main body trunk-like with serrated area, pointed embolus proper and well developed thumb; lamella characteristica fork-like branched, long and slender, almost parallel to radix; terminal apophysis with distal membrane and two strongly sclerotized teeth on ventral side.

Female. Unknown.

Remarks.

The male of this species is similar to the type species Acanthoneta aggressa. It differs only by the shape of the paracymbium. For a detailed comparison see Eskov and Marusik (1993).

Distribution.

Far East Asia: Magadan Area (Eskov and Marusik 1993) and China (Fig. 7) (new record).

Acknowledgements

We would like to thank Lyubomir Penev, Dimitar Dimitrov, Nikolaj Scharff, Gustavo Hormiga and Jeremy Miller for their comments on an earlier version of this paper. We would like to thank Don Buckle for providing the epigynum pictures of Acanthoneta aggressa, Fang Wang for helping molecular analysis and Chen Wei for field assistance. English of the final draft was kindly checked by David Penney. The study was supported by Natural Sciences Foundation, China (NSFC-30670244, NSFC-30970314, and NSFC-30911120070) and Russian Foundation for Basic Research grants (No.11-0401716 and 12-04-01548).

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Appendix
Figure S1.

Linyphiid phylogeny resulting from Maximum Likelihood analysis based on molecular data. Numbers at the nodes are bootstrap value. Branches in color indicate the four robustly supported clades within linyphiids: S Stemonyphantes clade (blue) L1 “linyphiines”-1 clade (pale green) L2 “linyphiines-2” clade (dark blue) ME “micronetines-erigonines” clade (red, with “Distal erigonines” clade in green). Taxa in different colors sampled from different groups: grey, outgroup; blue, Stemonyphantinae; pale green, Linyphiinae; dark blue, Mynogleninae; pink, Dubiaraneinae; black, Micronetinae; red, Ipainae, Acanthoneta and Acanoides gen. n.; green, Erigoninae. Red stars indicate the two out-group taxa: cyatholipid Alaranea and theridiosomatid Theridiosoma embedded within Linyphiidae.

Table S1.

GenBank accession numbers. Data of the taxa labeled with “#” are newly sequenced; the taxa labeled with “*” come from Arnedo et al. 2009.

Family Genus Species 16s 18s 28s COI H3
Araneidae Argiope trifasciata FJ525386 FJ525368 FJ525316 FJ525335
Cyatholipidae Alaranea merina* AY230942 AY230890 AY231074 AY231022 AY230982
Mysmenidae Maymena ambita GU456746 GU456765 GU456824 GU456876 GU456921
Nesticidae Nesticus cellulanus EU746444 AF005447 AF124961 EU746435
Pimoidae Pimoa haden GU338640 GU338524 EF128112 EF128155
Pimoidae Pimoa sp.* AY230940 AY230893 AY231072 AY231025 AY230985
Synotaxidae Synotaxus sp. AY230943 AY230894 AY231076 AY231026 AY230986
Theridiidae Steatoda bipunctata* AY230951 AY230926 AY231084 AY231057 AY231014
Theridiidae Theridion varians* AY230976 AY230932 AY231111 AY231063 AY231017
Theridiosomatidae Theridiosoma gemmosum HM030408 HM030417 HM030428 HM030436 HM030443
Linyphiidae Acanoides beijingensis# KJ027589 KJ027587 KJ027580 KJ027582 KJ027583
Acanoides hengshanensis# KJ027585 KJ027588 KJ027584 KJ027586 KJ027581
Acanthoneta sp. GU338479 GU338560 GU338678
Agyneta sp. GU338621 GU338529
Agyneta ramosa* FJ838670 FJ838694 FJ838717 FJ838648 FJ838740
Anguliphantes karpinskii GU338516 GU338566 GU338680
Asperthorax communis GU338482 GU338545 GU338684
Asthenargus sp. GU338493 GU338561
Australolinyphia remota* FJ838671 FJ838695 FJ838718 FJ838649 FJ838741
Bathyphantes floralis GU338604 GU338465 GU338583 GU338659
Bathyphantes gracilis* FJ838672 FJ838696 FJ838719 FJ838650 FJ838742
Bathyphantes gracilis GU338630 GU338464 GU338582 GU338689
Bolyphantes alticeps* AY078660 AY078667 AY078678 AY078691 AY078700
Capsulia sp. GU338470 GU338586
Centromerus trilobus GU338599 GU338468 GU338571 GU338656
Collinsia inerrans GU338601 GU338518 GU338645
Collinsia plumose GU338638 GU338499 GU338543
Denisiphantes sp. GU338619 GU338508 GU338563 GU338669
Dicymbium sinofacetum GU338614 GU338487 EF128119 GU338665
Diplocentria bidentata GU338629 GU338494 GU338542 GU338688
Diplocephalus cristatus GU338637 GU338490 GU338696
Diplostyla concolor* FJ838673 FJ838697 FJ838720 FJ838651 FJ838743
Diplostyla concolor GU338639 GU338467 GU338585 GU338697
Doenitzius peniculus GU338631 GU338469 GU338690
Doenitzius pruvus GU338632 GU338474 GU338691
Drapetisca socialis* FJ838674 FJ838698 FJ838721 FJ838652 FJ838744
Dubiaranea aysenensis FJ838675 FJ838699 FJ838722 FJ838653 FJ838745
Dubiaranea distincta GU338624 GU338459 GU338579 GU338648
Dubiaranea propinquua GU338627 GU338460 GU338580 GU338675
Dubiaranea similis GU338521 GU338581 GU338681
Erigone edentate GU338486 GU338540 GU338686
Erigone prominens GU338498 GU338539 GU338679
Floronia bucculenta* FJ838676 FJ838700 FJ838723 FJ838654 FJ838746
Frontinella communis GU338628 GU338517 GU338573
Frontinella communis* FJ838677 FJ838701 FJ838724 FJ838655 FJ838747
Fusciphantes hibanus GU338512 GU338570 GU338683
Gnathonarium dentatum GU338593 GU338477 EF128120 GU338651
Gnathonarium taczanowskii GU338620 GU338480 GU338547 GU338670
Gonatium japonicum GU338613 GU338492
Gonatium rubellum* FJ838679 FJ838703 FJ838726 FJ838656 FJ838749
Gongylidiellum vivum* FJ838678 FJ838702 FJ838725 FJ838748
Grammonota sp. GU338491 GU338685
Haplinis diloris* FJ838680 FJ838704 FJ838727 FJ838657 FJ838750
Helophora insignis* FJ838681 FJ838705 FJ838728 FJ838658 FJ838751
Himalaphantes azumiensis GU338522 GU338564 GU338677
Hylyphantes graminicola GU338595 GU338478 GU338550 GU338653
Hylyphantes sp. GU338618 GU338481 GU338549 GU338668
Labulla thoracica* AY078662 AY078674 AY078680 AY078694 AY078707
Laetesia sp.* FJ838682 FJ838706 FJ838729 FJ838659 FJ838752
Lepthyphantes minutus* AY078663 AY078673 AY078681 AY078689 AY078705
Lepthyphantes leprosus GU338488 GU338565 GU338682
Lepthyphantes sp. GU338610 GU338509 GU338562 GU338664
Linyphia triangularis* AY078664 AY078668 AY078682 AY078693 AY078702
Linyphia sp. GU338597 GU338461 GU338572 GU338654
Macrargus alpinus GU338505 GU338559
Agyneta nigra GU338608 GU338504 GU338577 GU338662
Agyneta rurestris* FJ838683 FJ838707 FJ838730 FJ838660 FJ838753
Microlinyphia dana* AY078665 AY078677 AY078683 AY078690
Microneta sp. GU338609 GU338472 GU338538 GU338663
Microneta viaria* FJ838684 FJ838708 FJ838731 FJ838661 FJ838754
Microneta viaria GU338598 GU338502 GU338537 GU338655
Moebelia rectangular GU338591 GU338485 GU338557
Mughiphantes nigromaculatus GU338600 GU338510 GU338527 GU338644
Nematogmus sanguinolentus GU338635 GU338489 GU338544 GU338694
Neomaso patagonicus GU338626 GU338473 GU338578 GU338674
Neriene japonica GU338633 GU338462 GU338575 GU338692
Neriene radiata* AY078710 AY078670 AY078684 AY078696 AY078709
Neriene radiate GU338623 GU338463 GU338574 GU338672
Neriene variabilis* AY078711 AY078669 AY078685 AY078699 AY078706
Nesioneta ellipsoidalis GU338519 GU338532 GU338687
Nippononeta kantonis GU338634 GU338471 GU338530 GU338693
Nippononeta sp. GU338602 GU338520 GU338531 GU338657
Notholepthyphantes australis* FJ838685 FJ838709 FJ838732 FJ838662 FJ838755
Novafroneta vulgaris* FJ838686 FJ838710 FJ838733 FJ838663 FJ838756
Oedothorax apicatus* FJ838687 FJ838711 FJ838664 FJ838757
Orsonwelles* malus AY078737 AY078676 AY078795 AY078697 AY078708
Orsonwelles* polites AY078725 AY078671 AY078786 AY078755 AY078701
Ostearius melanopygius* FJ838688 FJ838712 FJ838735 FJ838758
Paikiniana sp. GU338617 GU338495 GU338555 GU338647
Parameioneta bilobata GU338605 GU338503 GU338533 GU338660
Parasisis sp. GU338592 GU338500 GU338534 GU338650
Pityohyphantes costatus* AY078666 AY078675 AY078695
Pocobletus sp.* FJ838689 FJ838713 FJ838736 FJ838665 FJ838759
Porrhomma sp. GU338607 GU338466 GU338584 GU338661
Prosoponoides sinensis GU338606 GU338576 GU338649
Pseudafroneta incerta* FJ838690 FJ838714 FJ838737 FJ838666 FJ838760
Ryojius sp. GU338611 GU338536
Sisicottus montanus GU338625 GU338497 GU338541 GU338673
Solenysa sp. GU338616 GU338506 GU338528 GU338667
Sphecozone bicolor GU338622 GU338496 GU338553 GU338671
Stemonyphantes lineatus* FJ838691 FJ838715 FJ838738 FJ838667 FJ838761
Stemonyphantes sibiricus* FJ838692 FJ838668 FJ838762
Syedra oii GU338615 GU338513 GU338569 GU338666
Tapinopa guttata GU338511 GU338558 GU338676
Tenuiphantes ancatus GU338515 GU338567
Tenuiphantes sp. GU338612 GU338514 GU338568 GU338646
Tenuiphantes tenuis* FJ838693 FJ838716 FJ838739 FJ838669 FJ838763
Ummeliata feminea GU338594 GU338475 GU338551 GU338652
Ummeliata insecticeps GU338476 GU338552
Walckenaeria clavicornis GU338596 GU338483 GU338554
Walckenaeria keikoae GU338636 GU338484 GU338556 GU338695