Abstract

Hainanthereua albilineata gen. nov. et sp. nov. is described and illustrated based on specimens from Hainan Province, China. Morphological examination showed that these specimens belong to Thereuoneminae and share certain similarities with Thereuopodina Verhoeff, 1905. Phylogenetic reconstruction based on five genes (nuclear 18S and 28S rRNA, mitochondrial 12S and 16S rRNA, and cytochrome c oxidase subunit I) indicated that the specimens form a distinct and well-supported clade that is sister group to Thereuonema Verhoeff, 1904, so a new genus is accordingly established. Combining the evolutionary history and biogeographic framework of Thereuoneminae, this study revealed the evolutionary significance of Hainanthereua gen. nov. in the context of the Peninsular Indian Plate as a biotic ferry implicated in the origin of East and Southeast Asian lineages. Morphological similarities between Hainanthereua and Thereuopodina are symplesiomorphies of a clade that includes these two genera, Thereuonema and Thereuopoda Verhoeff, 1904.

Key words

Chilopoda, morphology, new species, phylogeny, taxonomic key, Thereuoneminae

1. Introduction

Scutigeromorpha is an ancient and morphologically highly specialized monophyletic group within the Chilopoda. It is taxonomically recognized as comprising three families (Edgecombe and Giribet 2009; Giribet and Edgecombe 2006, 2013). Among these, the Scutigeridae exhibits the highest diversity, and is widely distributed across warm temperate, tropical and subtropical regions of the Americas, southern Europe and North/East Africa, and from Asia to Australia (Edgecombe 2011). Displaying a typical transoceanic disjunct distribution pattern, it serves as an ideal model for studying historical biogeography. Manivannan et al. (2024) confirmed that the scutigerid subfamily Thereuoneminae originated in Gondwana by integrating molecular phylogenetics, fossil-calibrated time trees and biogeographic models. Subsequent dispersal events have shaped the group’s current distribution pattern centered on the Indo-Australian region, greatly advancing understanding of its macroevolutionary history.

With the application of methodologies such as scanning electron microscopy, molecular phylogenetics, and biogeographic modeling, the Thereuoneminae is currently recognized to include approximately 17 genera (Butler et al. 2010; Edgecombe 2011; Bonato et al. 2016; Porta and Giribet 2024). This comprises 14 valid genera and three monotypic genera of uncertain taxonomic status: Diplacrophor Chamberlin, 1920 (Solomon Islands), Phanothereua Chamberlin, 1958 (Solomon Islands), and Thereuella Chamberlin, 1955 (Peru).

In this study, an integrative taxonomic approach was employed to analyze a series of specimens collected from Hainan, China. Morphological examination indicates these specimens belong to the Thereuoneminae and share several similarities with Thereuopodina Verhoeff, 1905. Phylogenetic reconstruction based on five gene fragments (18S rRNA, 28S rRNA, 12S rRNA, 16S rRNA, and COI) reveals that these specimens form a distinct, well-supported clade, which is phylogenetically distinct from Thereuopodina and the allied genera Thereuopoda Verhoeff, 1904 and Thereuonema Verhoeff, 1904. This pattern is consistent with the “Out-of-India” biogeographic model (Manivannan et al. 2024). Combining morphological and molecular evidence, we herein establish a new genus to accommodate these specimens.

This paper provides a comprehensive morphological description and illustrations of the new genus and its type species. We present a diagnostic key to all known valid genera characterized by the absence of a pair of spine-bristles at the distal end of tarsus I (the traditionally delimited Thereuoneminae). Based on molecular clock estimates and existing biogeographic frameworks (Manivannan et al. 2024), we further discuss the divergence time of this new lineage and its significance in regional evolution. New evidence is provided for improving the phylogenetic framework of the Thereuoneminae and testing the colonization history of Gondwanan groups in Asia.

2. Material and methods

2.1. Specimens, morphology, terminologies

Specimens were hand-collected and fixed in 75% ethanol. All specimens including the types of the newly described species are deposited in the National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences (CMMI). Specimen identification numbers are in the format ‘CMMI-YYYYMMDDXXX’, where CMMI is an abbreviation of the depository; YYYYMMDD is the date on which the specimen was collected; and XXX is a number given to this specimen according to the sequence of specimens collected on that date. Morphological terminology follows Edgecombe and Giribet (2006), Bonato et al. (2010) and Porta and Giribet (2024). Proportions of the female gonopods follow measurements described by Würmli (1973: fig. 1). Terminology for peristomatic structures (epipharynx and hypopharynx) and antennal trichomes and sensilla is as used by Koch and Edgecombe (2006) and Sombke et al. (2011), respectively. Abbreviations and nomenclature used in this study are listed: T (singular) and TT (plural), stomatotergite(s); PP, Bayesian posterior probabilities; BP, Maximum likelihood bootstrap.

Specimens’ preservation and observation follow Li et al. (2026). They were cleared in 70% lactic acid for observation of the venom glands. Morphological examination and imaging used a Leica DMC 6200 camera attached to a Leica M205 FA microscope and Keyence VHX7000N ultra-depth-of-field microscope. The habitus was photographed with a NIKKOR 105 mm macro lens attached to a Nikon D850. Electron micrographs were taken with a Hitachi S-3400N scanning electron microscope.

To observe the hypopharynx and epipharynx, mouthparts were removed with tweezers and minute needles. The detached mouthparts (mandibles, first and second maxillae, and forcipules) were cleaned in an ultrasonic bath and fixed with 2.5% glutaraldehyde. The samples were then dehydrated through a graded ethanol series (50%, 70%, 80%, 90%, 95% for 20 min each, followed by three changes of 100% ethanol), then dehydrated ethanol was replaced with anhydrous tert-butanol, and the samples were freeze-dried in tert-butanol. The specimens were mounted on stainless steel stubs using double-sided conductive tape, and examined and electron micrographs taken under a Hitachi S-3400N scanning electron microscope.

SKETCHBOOK 6.0.6 was used on an iPad 9 tablet to prepare line drawings from photographs. Grammarly was used to polish English in the manuscript, subsequently edited by the native-speaking coauthor.

2.2. Phylogenetic methods

Five markers (two nuclear ribosomal RNA genes: 18S and 28S rRNA; two mitochondrial ribosomal RNA genes: 12S and 16S rRNA; and the mitochondrial protein-encoding gene cytochrome c oxidase subunit I: COI) used in previous studies of scutigeromorph phylogeny (Edgecombe and Giribet 2006, 2009; Butler et al. 2010; Giribet and Edgecombe 2013; Manivannan et al. 2024; Porta and Giribet 2024; Ji et al. 2025; Li et al. 2026) were used to estimate the phylogenetic position of the new genus. Primers and PCR amplification procedures for specimens of the new genus and two additional species of Thereuopoda from China collected by us (T. chinensis Verhoeff, 1905 and T. kaijiangensis Ji et al., 2025) follow Giribet and Edgecombe (2013). Sample information, localities, and GenBank accession numbers are shown in Table S1.

Representatives of three centipede orders outside Scutigeromorpha were selected as outgroup taxa to root the tree. The data were analyzed using both Maximum Likelihood (ML) and Bayesian Inference (BI). ML analysis was performed on the IQ-TREE web tool on the Galaxy platform (Nguyen et al. 2015; Trifinopoulos et al. 2016; Minh 2020) and BI performed on the PhyloSuite V1.2.2 platform (Zhang et al. 2020). Sequences for each dataset were aligned using the FFT-NS-2 algorithm in MAFFT (Katoh and Standley 2013, 2014; Katoh et al. 2005, 2019). The resulting alignments were processed with Gblocks (Castresana 2000) under settings allowing gaps within the final blocks, applying less stringent constraints on flanking positions. The trimmed gene alignments were concatenated using the Concatenate datasets tool, with external Ns converted to “?”. The final concatenated alignment comprised five genes, further refined using Gblocks.

The concatenated alignment was analyzed with the IQ-TREE web tool on the Galaxy platform. The best-fit nucleotide substitution model, selected by ModelFinder (Kalyaanamoorthy et al. 2017) under the Bayesian Information Criterion (BIC), was SYM+I+R4. Branch support under the Maximum Likelihood (ML) framework was calculated using the Ultrafast Jackknife method with 200,000 replicates (Hoang et al. 2018) and the Shimodaira–Hasegawa approximate likelihood ratio test (SH-aLRT) with 200,000 replicates (Guindon et al. 2010). Bayesian inference (BI) employed MRBAYES 3.2.6 (Ronquist et al. 2012) in the PhyloSuite v1.2.2 platform (Zhang et al. 2020), with 10,000,000 bootstrap replicates. SYM+I+G4 was chosen as the preferred model for BI, sampling every 1,000 generations and using 25% of the trees as burn-in. A split frequency of less than 0.01 was used to determine stationarity, and the consensus tree was constructed from the remaining trees. The resulting gene phylogenies were visualized in iTOL v5 (Letunic and Bork 2021).

3. Results

3.1. Phylogenetic analyses

Maximum Likelihood (ML) and Bayesian Inference (BI) analyses based on concatenated sequences for the five genes recovered trees within which Scutigeromorpha contained 80 and 60 strongly supported nodes (Maximum Likelihood bootstrap proportion (BP) > 90 or Bayesian posterior probabilities (PP) > 95), respectively, with minor topological differences between them (Fig. 1).

Figure 1.

Maximum likelihood tree (left) and Bayesian consensus tree (right) based on 12S, 16S, 18S, 28S and COI sequences. BP and PP values are shown next to each node, the support values with BP < 90 and PP < 95 were not given.

Relationships between the three monophyletic scutigeromorph families are as in previous molecular phylogenies (Edgecombe and Giribet 2006, 2009; Butler et al. 2010; Giribet and Edgecombe 2013; Benavides et al. 2023; Li et al. 2026), with Scutigerinidae and Scutigeridae being sister groups to the exclusion of Pselliodidae. The ML and BI trees both divided the Scutigeridae into two clades, consistent with the subfamilial circumscriptions newly defined by Li et al. (2026). In both trees, Scutigeridae was divided into: (1) a clade comprising the genera Tachythereua Verhoeff, 1905, Dendrothereua Verhoeff, 1944, and Scutigera Lamarck, 1801; (2) a clade containing Sinothereua Li, Edgecombe & Jiang, 2026, Edgethereua Porta and Giribet, 2024, Lassophora Verhoeff, 1905, Ballonema Verhoeff, 1904 and the traditionally recognized subfamily Thereuoneminae.

The species described herein was consistently and robustly nested within the Thereuoneminae clade, with strong support for a sister-group relationship with Thereuonema in both the ML and BI trees. Based on the combined results, we support recognition of the Hainan species as a distinct new genus, which we hereby describe as Hainanthereua gen. nov. Expanding the scope of Thereuonema to encompass H. albilineata sp. nov. would require a rediagnosis that would not include the distinctive elongate spiculae that have long been used to diagnose Thereuonema (Würmli 1975a; Dyachkov 2026). Thereuonema and Hainanthereua group with Thereuopoda and Thereuopodina in both ML and BI trees, but the two methods differ in whether Thereuopoda and Thereuopodina are each other’s sister group (ML) or if Thereuopoda is more closely related to Thereuonema + Hainanthereua (BI).

3.2. Taxonomy

Order Scutigeromorpha Pocock, 1895

Family Scutigeridae Leach, 1814

Subfamily Thereuoneminae Verhoeff, 1905

3.2.1. Hainanthereua gen. nov.

https://zoobank.org/4E84E1DE-65A3-49FB-82EC-1C2530F0E96F

Chinese name: 琼蚰属

Type species.

Hainanthereua albilineata sp. nov. by monotypy and present designation.

Diagnosis.

Antenna ca. 1.5 times as long as the body. First flagellum of antenna with 55–101 annulations (including node). Anterior projection of cephalic transverse sutures short, triangular, posterior part subparallel. Stomatotergites with a prominent white median longitudinal stripe; bristles (Stachelborsten) associated with elongate, conical, unpaired spines as long as bristles on TT4–7, and short paired spines on all tergites; the ratio of paired spines to unpaired spines associated with Stachelborsten on T6 is approximately 1:1 to 1:2, bristles (Stachelborsten) on borders with short paired spines along posterior margin. Elongate triangular spiculae, relatively sparse. Stomata elongate. Prefemur of legs 3–9 lacking a saw-like row of spines (sometimes legs 1–9). Legs 1–14 with pairs of tarsal papillae on consecutive tarsomeres, short-long alternation of tarsal papillae on legs 1–9, and uniform size of tarsal papillae on legs 10–14 (paired long papillae). Proarthron and metarthron of female gonopods subparallel-sided.

Etymology.

The name Hainanthereua (feminine) is a compound derived from ‘Hainan-’ (Hainan Province, China) and the common scutigerid suffix -thereua, meaning “the scutigerid genus of Hainan Province, China”.

Remarks.

Hainanthereua gen. nov. resembles Thereuonema in the cephalic sutures and the spines on the stomatotergites (all associated with a Stachelborste). Hainanthereua differs from this genus in color, form of spiculae, and stronger Stachelborsten. The dark pigmented band on the stomatotergites of Thereuonema is rather diffuse, while there is a distinct white longitudinal stripe in the middle of the stomatotergites of Hainanthereua. The spiculae of Hainanthereua are short triangular or more elongate, relatively sparse. In marked contrast, the spiculae of Thereuonema are dense, long, needle-like and variably parallel-sided (Verhoeff 1904, 1905; Würmli 1975a; Edgecombe and Giribet 2006: fig. 2f; Edgecombe 2011; Dyachkov 2026).

Hainanthereua gen. nov. resembles Thereuopoda, Thereuopodina and Pilbarascutigera Edgecombe and Barrow, 2007 in the shape of its spiculae (Edgecombe 2007), but those of Hainanthereua are relatively longer than in the other three genera. Thereuopoda possesses a unique kinked cephalic suture, while that of Hainanthereua, Thereuopodina and Pilbarascutigera is of the M-shaped type common in Scutigeridae (Edgecombe 2011). Compared to Pilbarascutigera with stoma-saddles weakly vaulted, the stoma-saddle of Hainanthereua is moderately vaulted, while that of Thereuopoda is strongly vaulted and is approached by that of Thereuopodina (Edgecombe and Barrow 2007). In addition, the stomatotergites bear spines associated with Tastborsten in Thereuopoda and Pilbarascutigera (anterior tergites bearing Stachelborsten and Tastborsten but lacking spines), while Hainanthereua and Thereuopodina bear Stachelborsten. Hainanthereua differs from Thereuopodina in having the margins of the proarthron and mesarthron of the female gonopods diverging in Thereuopodina v. subparallel-sided in Hainanthereua; and the prefemur of legs 2–4 of Thereuopodina lacking a saw-like row of spines whereas Hainanthereua lacks them on legs 3–9 (sometimes 1–9).

Hainanthereua gen. nov. resembles Allothereua and Parascutigera in the spines on the stomatotergites being associated with Stachelborsten and the margin of the proarthron and mesarthron of female gonopods being subparallel-sided. The spiculae of Allothereua and Parascutigera are, however, variably setiform (Edgecombe and Giribet 2006), rather than elongate triangular as in Hainanthereua, and they are considerably denser in the former two genera. The Stachelborsten on TT5–7 of Parascutigera are associated with paired spines, unpaired spines being wholly lacking (Verhoeff 1904, 1905; Edgecombe and Giribet 2006; Edgecombe 2011), but both types of bristles/spines are present on the stomatotergites of Allothereua and Hainanthereua.

Like Seychellonema Butler et al. 2010, Hainanthereua gen. nov. expresses a short-long alternation of tarsal papillae on successive tarsomeres. In Seychellonema, this feature is present on legs 1–7, while legs 8–14 bear paired short papillae of uniform size (Butler et al. 2010). However, in Hainanthereua, the alternation is observed on legs 1–9, and uniform size of tarsal papillae on legs 10–14 (paired long papillae). Würmli (1975b) interpreted such alternation in the holotype of Lassophora madagascariensis Verhoeff, 1905 as unreliable for generic diagnosis because the specimen is a praematurus, and mature specimens have uniform papillae. This morphology is, however, retained in mature specimens of Seychellonema and Hainanthereua. Accordingly, we include it in the generic diagnosis.

3.2.2. Identification key

What follows is a key to all known valid scutigerid genera characterized by the absence of a pair of spine-bristles at distal end of all tarsus I, with emphasis on tergal prominences.

1	Dorsal spine-bristle on prefemur of second maxillae; ventral spine-bristle lacking	2
1’	Dorsal and ventral spine-bristles on prefemur of second maxillae	3
2	Margins of proarthron and mesarthron of female gonopods diverging posteriorly	Edgethereua Porta and Giribet, 2024
2’	Margins of proarthron and mesarthron of female gonopods weakly converging	Thereuoquima Bücherl, 1949
3	Tergites bearing bristles and spines, lacking spiculae or spinulae	Tachythereua Verhoeff, 1905
3’	Tergites bearing bristles, spines and spiculae or spinulae	4
4	All tergites with spinulae	Seychellonema Butler, Edgecombe, Ball and Giribet, 2010
4’	Tergites with simple spiculae	5
5	Spiculae as long as bristles, needle-like	Thereuonema Verhoeff, 1904
5’	Spiculae shorter than bristles	6
6	Tergites bearing few bristles/spines (spines lacking on stoma-saddles of TT6–7)	Prionopodella Verhoeff, 1925
6’	Tergites bearing bristles associated with paired spines	7
6’’	Tergites bearing bristles associated with unpaired spines	9
7	Metarthron of female gonopods massive, hook-shaped	Pesvarus Würmli, 1974
7’	Metarthron of female gonopods slender, tapering distally	8
8	Short paired spines (no unpaired spines); margin of proarthron and mesarthron of female gonopods diverging posteriorly	Prothereua Verhoeff, 1925
8’	Variably long paired spines; margin of proarthron and mesarthron of female gonopods subparallel-sided	Parascutigera Verhoeff, 1904
9	Anterior projection of cephalic sutures with divergent posterior part such that suture is kinked	Thereuopoda Verhoeff, 1904
9’	Anterior projection of cephalic sutures subparallel	10
10	Tergites bearing isolated bristles (Tastborsten), few unpaired spines (spines lacking on stoma-saddles of TT6–7)	Podothereua Verhoeff, 1905
10’	Tergites bearing bristles (Tastborsten), unpaired spines (spines present on stoma-saddles of TT6–7)	11
11	Tergites bearing bristles (Tastborsten and Stachelborsten)	Pilbarascutigera Edgecombe and Barrow, 2007
11’	Tergites bearing bristles (Stachelborsten)	12
12	Margins of proarthron and mesarthron of female gonopods diverging posteriorly	Thereuopodina Verhoeff, 1905
12’	Margins of proarthron and mesarthron of female gonopods subparallel-sided	13
13	Spiculae dense (at margins of most polygonal cuticular scales), variably setiform; uniform size of tarsal papillae on legs 1–14	Allothereua Verhoeff, 1905
13’	Spiculae relatively sparse, elongate triangular; short-long alternation of tarsal papillae on legs 1–9	Hainanthereua gen. nov.

3.2.3. Hainanthereua albilineata gen. nov. et, sp. nov.

https://zoobank.org/F4A46240-140E-43D6-AE9C-70594B6AFF8F

Chinese name: 白线蚰蜒

Material examined.

Holotype: CHINA • ♀; Mingfenggu Valley, Jianfengling, Ledong Li Autonomous County, Hainan Province, China (Fig. 2), 18.7426°N 108.8400°E, 990 m a.s.l., 22 June 2021, leg. Chao Jiang & Tianyun Chen (labelled as CMMI-20210622177). Paratypes: CHINA • 2♀♀ 5♂♂, same as holotype, 22–23 June 2021, leg. Chao Jiang & Tianyun Chen (CMMI-20210622104, CMMI-20210622174, CMMI-20210622176, CMMI-20210622179, CMMI-20210623126, CCMI-20210623127, CMMI-20210623128); CHINA • 2♀♀ 3♂♂, Mt. Diaoluoshan, Benhao Town, Lingshui Li Autonomous County, Hainan Province, China, 18.7272°N 109.8724°E, 900 m a.s.l., 24–25 June 2021, leg. Chao Jiang & Tianyun Chen (CMMI-20210624170, CMMI-20210624171, CMMI-20210625112, CMMI-20210625173, CMMI-20210625175).

Figure 2.

Habitat of Hainanthereua albilineata sp. nov. (from type locality: Mingfenggu Valley, Jianfengling, Ledong Li Autonomous County, Hainan Province, China). A Panoramic view of type locality; B woodland on the mountain of type locality shown in A; C interior habitat of the woodland shown in B, the species was collected in relatively dry leaf litter or under stones; D individual of the species found under a stone; E live habitus of the species. A–D provided by Mr. Quanyu Ji, E provided by Mr. Jiazhou Lu.

Diagnosis.

As for genus.

Description.

Body length (from anterior edge of head capsule to tip of gonopods): 18–23 mm in adult. — Colour: Head capsule brown (Fig. 3A, B); antennal base ferruginous grading distally to pale yellow; stomatotergites with a prominent white median longitudinal stripe, black and brown on both sides, margins dark brown (Figs 2D, 2E, 3C, 3D, 8A–F); stoma saddle red (Fig. 2D, E); legs yellow with conspicuous black bands on prefemur, femur, and tibia (Fig. 2D, E, 3A); sternites pale yellow; female gonopods yellow (Fig. 8G) (based on living specimens). — Head capsule: Anterior projection of the cephalic transverse sutures short, triangular, posterior part parallel; transverse suture long; longitudinal median depression extends from the base of the antennae to between the two eyes (Figs 3A, 3B, 10A, 10C). Antenna ca. 1.5 times as long as body length. First flagellum of antenna with 55–101 annulations (including the first node). Annulations much wider than long (Figs 3A, 3B, 10A, 10C), with three or four whorls of flattened trichomes, a few beak-like sensilla, and a few setae (trichoid sensilla) associated with paired spines connected to the base form a single whorl encircling the distal end of the annulation. Tömösváry’s organ small (Fig. 3B), in typical position between eye and base of antenna (Fig. 2D, E). — Epipharynx: Arrangement typical for Scutigeridae (Koch and Edgecombe 2006: fig. 1) (Fig. 4). Lateral bar of labral trapezoid with narrow longitudinal groove along whole length of bar (Fig. 4A). Labral bristles differentiated into narrow outer band of short, pectinate bristles and wider inner band of longer simple bristles (Fig. 4A, C). Two clusters of sensilla along mid-line of labral trapezoid: more distal unpaired, transverse group of a few dozen bottle-shaped sensilla at termination of median ridge (Fig. 4B); the three proximal sensilla clusters of the labral part are composed of nipple-and bottle-shaped sensilla, median sensilla completely distal to lateral clusters (Fig. 4D). Chevron-shaped spine row of triangular and distally-curved denticles at the border (Fig. 4A, E, F) between labral and clypeal part of epipharynx; immediately proximal to the spine row is a rhomboid branched-spine field having transverse axis approximately 1.5 times the length of longitudinal axis, with a dense median field of pectinate spines (Fig. 4E). Broadly ovate cluster of bottle-shaped sensilla on medial part of clypeal triangle, a short distance behind dense field of branching spines (Fig. 4G). Lateral clusters of nipple-shaped sensilla positioned in a depression within the non-sclerotized area distal to the oblique bar and proximal to the edge extending from the submarginal armature to the transverse bar of the labral trapezoid. — Hypopharynx: Elongate, deeply projecting into preoral chamber (Fig. 5). Bars of lateral sclerotised fork with lateral bulges. Median excavation on proximal part of frontal and distal part of hypopharynx surface bordered laterally by pectinate bristles (Fig. 5A, D). Proximal cluster of sensilla (Fig. 5B, C) composed of a few sparse elements distributed in rows between mouth and a median elevation proximal to the median excavation. Area between the median excavation and the converging flattened bars of the proximal fork is sub-triangular with serrated spines at the proximal end of the carina (Fig. 5B, D). Inner margin of the tongue tip has strong irregular folds (Fig. 5E–G), on which nipple-shaped sensilla are distributed (Fig. 5E–G). — Mandible: Incisor region composed of three teeth, all with surface smooth and three cusps. Approximately 16 pectinate lamellae in gnathal lobe. Molar plate surrounded by a Haarpolster of approximately the same length. (Fig. 6A) — Second maxilla: Dorsal and ventral spine bristles on prefemur, four spine bristles on femur; two dorsal spine bristles on tibia (Fig. 6B). Ventral side of all podomeres and dorsal side of femur with rows of bristles. All spine bristles with similar surface details, spiniform scales, short proximally, becoming more elongate distally to confer a ridged or fluted surface to distal third of spine bristle. — Forcipular segment: Four spine bristles on anterior margin of coxa approximately equal in length to coxa. Spine comb of tarsus covering the entire inner side of the segment and composed of flattened setae mostly inserted by pairs in the bases and terminating in a rounded bulb. Ventrolateral side of trochanteroprefemur and femur with rows of spine bristles. Venom gland extending to distal part of tibia (Fig. 6C). — Stomatotergites: Stomatotergites with rugose surface (Fig. 7), posterior borders evenly rounded (Figs 2E, 3A, 3C, 3D, 7C, 7G, 8A–F, 10A, 10B). Stoma saddles moderately inflated; stomata elongate; with stomata 0.5–0.7 times the length of the stoma saddles (Fig. 7C, D). Stomatotergites bearing bristles (Stachelborsten) associated with elongate, conical, unpaired spines, as long as the bristles, or short paired spines (Figs 7A–E, 10F); spiculae short triangular or more elongate, emerging between the scutes, each separated by several polygonal scales that lack spiculae, relatively sparse (Figs 7, 10F); spinulae occur exclusively on T8 (Fig. 7G, H). Unpaired spines usually initially occur on T4 (on TT4–7), sparse, and always associated with a Stachelborste; 27–51 bristles associated with a spine on T6 (Figs 8D, 10D), uniformly distributed. Paired spines short; association with Stachelborste on TT1–8, 30–55 bristles associated with spines on T6 (Figs 8D, 10D). In adults, the ratio of paired spines to unpaired spines associated with Stachelborsten on T6 is approximately 1:1 to 1:2. Stomatotergite lateral margins and posterior margin with marginal bristles (Stachelborsten) with short, conical unpaired spines (Figs 7F, 10G). — Legs: Tarsus I and II tarsomeres as follow (range in tarsus I/tarsus II): leg 1, 11–16/25–31; leg 2, 10–11/27–30; leg 3, 10–11/25–30; leg 4, 7–9/24–30; leg 5, 7–9/24–30; leg 6, 7–9/24–27; leg 7, 7–8/19–29; leg 8, 6–7/22–27; leg 9, 6–8/24–28; leg 10, 6–7/28–29; leg 11, 6–8/29–30; leg 12, 6–8/26–32: leg 13, 8/31–34; leg 14, 7–9/29–32. Prefemoral spine-bristles in a 2/1 pattern on legs 1–14; femoral spine-bristles 1/2 on legs 1–14; tibial spine-bristles 1/1 on legs 1–11 and 1/2 on legs 12–14. Prefemur of legs 3–9 lacking a saw-like row of spines (sometimes legs 1–9). Pair of spine-bristles lacking at distal end of tarsus I (Fig. 10I). Pairs of tarsal papillae on legs 1–14, expression of short-long alternation of tarsal papillae on successive tarsomeres on legs 1–9 (sometimes legs 1–10) (Figs 6D, 9A–D, 10J); legs 10–14 (or legs 11–14) with paired long tarsal papillae (Figs 6E, 9E–H, 10K). Setal cluster associated with tarsal papillae sparse, arranged in longitudinal bands of 2–3 setae. Resilient sole hairs originate near posteromedial edge of tarsal papillae, extending to approximately one-third length of the succeeding tarsomere. — Sternites: Sternites with median swelling. Longitudinal median furrow visible, extending two-thirds length of the sternite. All sternites are scattered with setae, no hairs, spines or spiculae. — Gonopods: Female gonopods with maximum length 1.5–2.2 times maximum width; longitudinal median suture in proarthron complete (Figs 8G, 9I, 10E). Lateral margins of proarthron nearly parallel (lateral margins of the proarthron of the juvenile gonopods diverge posteriorly). Subtriangular depression on proarthron deep, lacking setae. Proarthron 1–1.5 times length of mesarthron (Figs 8G, 9I, 10E). Coarsest setae on proarthron slightly thicker than those on mesarthron apart from cluster at distomedial corner of mesarthron, this cluster usually composed of 6–10 setae (Fig. 9J, K). Sinus between mesarthron usually broadly parabolic, its apex typically weakly pointed (Fig. 9J). Width of mesarthron 0.4–0.8 times maximum width of sinus. Proarthron + mesarthron 1.7–2.2 times length of metarthron. Outer margin of metarthron arched (Fig. 9L). Ventral surface of metarthron scattered with small, pointed conical sensilla; outer margin of the metarthron bears 15–16 sensilla coeloconica, arranged roughly in a single row (Figs 9L, 9M, 10H). A few setae on dorsal side of metarthron (Figs 9L, 10E). Subanal plate drop-shaped; smooth, seta-free band in the middle of the subanal plate; setae present on outer surface of subanal plate (Fig. 8G, H). Both pairs of styliform male gonopods densely covered with setae, terminal part with short, spiniform setae between longer setae (Fig. 8H).

Figure 3.

Hainanthereua albilineata sp. nov., holotype. A Habitus, dorsal view, body length 23 mm; B head, dorsal view; C T1, dorsal view; D T2, dorsal view. Abbreviations: lmd, longitudinal median depression; ap, anterior projection of the (cephalic) transverse suture; To, Tömösváry’s organ; t, (cephalic) transverse suture; ce, compound eye.

Figure 4.

Hainanthereua albilineata sp. nov., paratypes. A Internal view of epipharynx, the labels *B–*G indicate the corresponding positions shown in panels B–G, and the lateral orientation is marked; B detail of A, distal part of the epipharynx, distal cluster of sensilla of the medial labral part; C lateral (left) part of the epipharynx, labral bristles, present on both sides; D detail of A, proximal part of the epipharynx, proximal cluster of sensilla of the medial labral part; E detail of A, field of branching spines on clypeal part of epipharynx; F detail of E, showing the chevron-shaped spine row; G sensilla proximal to dense field of branching spines on clypeal triangle, corresponding to the proximal part of the structure shown in panel E; H detail of A, sensilla on lateral (right) part of epipharynx, present on both sides, situated within a depression.

Figure 5.

Hainanthereua albilineata sp. nov., paratype. A Frontal view of hypopharynx, the labels *B, *D, and *E indicate the corresponding positions shown in panels B, D, and E respectively, and the proximal and distal orientations are also marked; B detail of A, proximal part of the hypopharynx, mouth and area between the median excavation and the converging flattened bars of the proximal fork; C detail of B, showing sensilla; D detail of A, flattened bristles of proximal part of excavation; E distal end of hypopharynx, showing flattened bristles and sensilla, the labels *F and *G indicate the corresponding positions shown in panels F and G; F detail of E, showing nipple-shaped sensilla; G detail of E, showing pectinate bristles on distal portion.

Figure 6.

Hainanthereua albilineata sp. nov., paratype. A Mandible, internal view; B second maxilla (left), prolateral view; C part of the forcipular segment, ventral view; D tarsal papillae on tarsomeres of leg 8 tarsus II, posterior view, showing the short-long alternation of tarsal papillae; E tarsal papillae on tarsomeres of leg 10 tarsus II, posterior view, showing long papillae of consistent size. Triangles represent long papillae, and circles represent short papillae.

Figure 7.

Hainanthereua albilineata sp. nov., paratype. A T2, dorsal view; B T5, dorsal view. C T6, dorsal view; D stoma saddles of T6, dorsal view; E bristles (Stachelborsten), spines, spiculae on T6, dorsal view; F paired spines with bristle on margin of T6, dorsal view. G T8, dorsal view; H detail of G, showing spiculae and spinulae on T8, the spiculae clearly emerging between the scutes, dorsal view. Abbreviations: St, Stachelborsten; p.s., paired spines; s, spine; sc, spiculae; sn, spinulae.

Figure 8.

Hainanthereua albilineata sp. nov., A–G holotype, H paratype. A T3, dorsal view; B T4, dorsal view; C T5, dorsal view; D T6, dorsal view; E T7, dorsal view; F T8, (right-)dorsal view; G postpedal segments, including female gonopods and subanal plate, with relevant structures labeled, left-ventral view; H postpedal segments, including male gonopods, left-ventral view.

Figure 9.

Hainanthereua albilineata sp. nov., paratype. A Tarsus II of leg 6, posterior view; B detail of A, tarsal papillae on tarsomeres of leg 6 tarsus II, lateral view; C detail of B, showing long papillae of consistent size, lateral view; D detail of B, showing short papillae of consistent size, lateral view; E tarsus II of leg 12, posterior view; F detail of E, tarsal papillae on tarsomeres of leg 12 tarsus II, lateral view; G, H detail of F, showing long papillae of consistent size, lateral view; I female gonopods and subanal plate, ventral view; J detail of I, showing mesarthron, ventral view; K detail of I, distal end of mesarthron, ventral view; L detail of I, showing metarthron, ventral view; M detail of L, showing sensillum on metarthron, ventral view.

Figure 10.

Hainanthereua albilineata sp. nov., A–E and I holotype, F–H and J–K paratype. A, B Habitus, showing head and TT1–8, dorsal view; C head, dorsal view; D T6, dorsal view; E female gonopods, (right-)ventral view; F detail of T6, dorsal view; G detail of posterior border of T6, right side, dorsal view; H right metarthron of female gonopods, dorsal view; I leg 6 (left) from prefemur to claw, posterior view; J detail of tarsal papillae on tarsus II of leg 6, lateral view; K detail of tarsal papillae on tarsus II of leg 10, lateral view. Triangles (J, K) represent long papillae, circles (J) represent short papillae. Scale bars: 500 μm (A–E, I); 300 μm (F, H); 100 μm (G, J, K).

Distribution.

China (Hainan).

Etymology.

The specific epithet albilineata is a Latin feminine adjective. It is a compound formed from “albi-”, derived from the stem of the Latin adjective albus, meaning “white”, and “-lineata”, the feminine past participle of the Latin verb lineare (to draw a line), meaning “lined” or “striped.” The name means “white-striped,” describing the distinct white longitudinal stripe in the middle of the tergites of this species.

4. Discussion

Scutigeromorphs in the Asia-Australia region are highly diverse, with morphological similarity between species, making identification difficult (Edgecombe and Barrow 2007). Phylogenetic analyses show that Hainanthereua gen. nov. is closely related to Thereuonema, but there are significant morphological differences between the two genera. Although Hainanthereua gen. nov. is morphologically similar to Thereuopodina, they can be distinguished by the shape of margins of the proarthron and mesarthron of female gonopods and the saw-like row of spines of the prefemur of different ranges of legs. In addition, molecular phylogenetic analysis strongly supports Hainanthereua gen. nov. as an independent evolutionary clade. Based on a combination of diagnostic morphological features and molecular evidence, we classify the described specimens as a new genus.

Furthermore, based on the phylogenetic trees constructed in this study, Edgethereua is stably positioned within the subfamily Thereuoneminae. This differs from the findings of Porta and Giribet (2024) and Li et al. (2026), which reported conflicting results between the ML and BI analyses. Lassophora Verhoeff, 1905, from Madagascar and Mozambique, and Ballonema Verhoeff, 1904, from New Guinea, are here allied to the Thereuoneminae, instead of forming a clade or grade excluded from either of the subfamilies Scutigerinae and Thereuoneminae (Butler et al. 2010; Giribet and Edgecombe 2013; Porta and Giribet 2024). This result confirms that the traditional Thereuoneminae and Scutigerinae were reciprocally paraphyletic groups (Edgecombe and Giribet 2006; Edgecombe and Giribet 2009; Butler et al. 2010; Giribet and Edgecombe 2013; Li et al. 2026), indicating that relying only on a single morphological character (presence or absence of tarsus I spine bristles) is insufficient to reflect the true evolutionary history and prompting us to re-examine the early evolution and biogeographic patterns of Thereuoneminae.

The biogeographic reconstruction by Manivannan et al. (2024) indicates that the ancestors of scutigeromorphs were widely distributed on Gondwana, and their major early lineage diversification occurred before the complete isolation of the main continental blocks. Following Li et al. (2026), the affinities of Lassophora and Ballonema to the Thereuoneminae allow the effective divergence time of this subfamily to be traced back to at least the Early Cretaceous, around 145 million years ago (= mya). The transoceanic dispersal events through which Lassophora and Ballonema originated at this time may be more reasonably explained as an early divergence within the widely distributed ancestral group of the Thereuoneminae, rather than strictly corresponding to a specific continental breakup event. This conclusion differs from the view in Giribet and Edgecombe (2013), which preferred to explain the phylogenetic relationships of this clade by vicariance during the Early Jurassic to Early Cretaceous. The ancestral group of Thereuoneminae had already achieved initial diversification on the drifting Indian Plate during the early to middle Cretaceous.

Estimates based on the molecular clock (Manivannan et al. 2024) suggest that the likely divergence time of Hainanthereua gen. nov. is comparable to that of its sister group, Thereuonema, but slightly later than that of Thereuopodina (the Indian diversification of which dates to around 78 mya), likely occurring between 60 and 72 mya. This temporal sequence is highly consistent with the geological history of the Peninsular Indian Plate. The ancestral divergence of Thereuopodina occurred in the late stage of the isolated drift of the Indian Plate. In contrast, the divergence between the new genus and Thereuonema is inferred to have occurred during the critical window of the initial collision between the Indian Plate and the Eurasian Plate (about 60 to 50 mya). It is inferred that the ancestors of Hainanthereua gen. nov. were pioneer groups that dispersed northward around the time of the plate collision. They have evolved independently to this day due to geographical isolation. The morphological similarity between Hainanthereua gen. nov. and Thereuopodina originates from their common ancestor on the Indian Plate during the Cretaceous Period, the similarities between these two genera corresponding to symplesiomorphies of the clade that unites them with Thereuonema and Thereuopoda. This also indicates that the diversification within Asian lineages is more complex than previously thought, which may involve multiple dispersal and vicariance events toward the edge of East Asia.

The discovery of the new genus provides key empirical evidence from East Asia for testing and refining the “Indian Plate as a biological ferry” hypothesis. It also extends the evolutionary history of endemic Asian Thereuoneminae to older geological periods, laying a more solid foundation for understanding the global distribution pattern of this ancient group. Future research should conduct more systematic surveys of scutigeromorph species in Southeast Asia and southern China to trace the complete trajectory of this “northward dispersal” route.

5. Declarations

Authors’ contributions. Qing Li: Conceptualization, Methodology, Software, Data Curation, Writing − Original Draft, Writing − Review & Editing. Gregory D. Edgecombe: Data Curation, Supervision, Writing − Review & Editing. Chao Jiang: Conceptualization, Resources, Data Curation, Methodology, Supervision, Writing − Review & Editing, Project administration, Funding acquisition.

Conflict of interest. The authors declare that there is no conflict of interest.

6. Acknowledgements

We sincerely thank editors Martin Schwentner, Klaus-Dieter Klass, reviewer Andy Sombke and anonymous reviewers for reviewing the manuscript and providing valuable comments. We are grateful to Mr. Jiazhou Lu (Shaanxi Vocational Academy of Art, China) and Mr. Quanyu Ji (Hebei University, China) for providing habitat photos of the new species. We also thank Mr. Feiyu Huang (Northeast Forestry University, China) for help with the molecular work. The research was supported by the CACMS Innovation Fund (nos. CI2024E003, nos. CI2024G00-09) and the Key Project at Central Government Level: the Ability Establishment of Sustainable Use for Valuable Chinese Medicine Resources (nos. 2060302).

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Supplementary material

Supplementary material 1

Table S1

Li Q, Edgecombe GD, Jiang C (2026)

Data type: .docx

Explanation notes: Scutigeromorph and outgroup sampling, MCZ or other accession numbers, country/region of origin and sequenced gene fragments..

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

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