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Research Article
Five new species of Exalloniscus Stebbing, 1911 (Crustacea, Isopoda, Oniscidea) from China
expand article infoChao Jiang‡, Chonghui Yao§, Luqi Huang‡, Weichun Li§
‡ China Academy of Chinese Medical Sciences, Beijing, China
§ Jiangxi Agricultural University, Nanchang, China

Corresponding author: Weichun Li ( weichunlee@126.com )

Academic editor: Luiz F. Andrade
© 2024 Chao Jiang, Chonghui Yao, Luqi Huang, Weichun Li.
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: Jiang C, Yao C, Huang L, Li W (2024) Five new species of Exalloniscus Stebbing, 1911 (Crustacea, Isopoda, Oniscidea) from China. Zoosystematics and Evolution 100(2): 373-383. https://doi.org/10.3897/zse.100.115813
ZooBank: urn:lsid:zoobank.org:pub:1B0AF998-E43A-4F82-9F97-F6F8C99133B4
Open Access

Abstract

Exalloniscus Stebbing, 1911 is investigated from China, and eleven species of the genus are now recorded from China. Five of them are described as new: E. duospinatus Li & Jiang, sp. nov., E. curvispinatus Li & Jiang, sp. nov., E. triangulus Li & Jiang, sp. nov., E. tridentatus Li & Jiang, sp. nov. and E. taitii Li & Jiang, sp. nov. A map of China showing the recorded localities of Exalloniscus members is provided.

Key Words

China, distribution, morphology, new species, taxonomy

Introduction

The genus Exalloniscus was established by Stebbing (1911) to allocate Alloniscus coecus Dollfus, 1898 from Indonesia. To date, the genus includes twenty-eight species with Oriental and Palaearctic distribution, occurring in South Asia (India, Sri Lanka, Nepal), Southeast Asia (Cambodia, Laos, Indonesia, Malaysia, Myanmar, Philippines, Singapore, Thailand, Vietnam), and East Asia (China, Korea, Japan) (Manicastri and Argano 1986; Taiti and Ferrara 1988; Manicastri and Taiti 1991; Kwon 1993, 1995; Kwon and Taiti 1993; Nunomura 2000; Nunomura and Xie 2000; Taiti and Gruber 2008; Taiti and Cardoso 2020). The genus has been redefined (Taiti and Ferrara 1988), and its diagnosis has also been amended (Taiti and Cardoso 2020). Prior to this study, six Exalloniscus species were known from China (Taiti and Ferrara 1986; Kwon and Taiti 1993; Nunomura and Xie 2000; Chen 2003; Taiti and Gruber 2008; Taiti and Cardoso 2020). The present study aimed to describe five new species of the genus collected in China.

Materials and methods

Specimens were collected by hand using tweezers and preserved in 75% ethanol. The appendages were stained with acid fuchsin and mounted on micro-preparations in a neutral balsam mounting medium. Habitus were recorded with a Zeiss AxioCam Icc 5 digital camera attached to a Zeiss Stereo Discovery V12 microscope. Illustrations of the appendages were prepared using an Optec DV E3 630 digital camera attached to an Optec BK6000 microscope. GNU Image Manipulation Program (Montesanto 2015) was used for line drawings. The terminology for morphological structures followed Taiti and Cardoso (2020). The specimens are deposited at the Insect Museum, Jiangxi Agricultural University, Nanchang, China (JXAUM) and National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences, Beijing, China (CMMI).

Taxonomic account

Exalloniscus duospinatus Li & Jiang, sp. nov.

https://zoobank.org/3F019AA6-3E86-4AD0-828F-20E115548925
Figs 1A, 2

Type material

Holotype. China: male, Sichuan Province, Qionglai, Datong Town (30°30'N, 103°18'E), alt. 770 m, 16 April 2021, Chao Jiang leg., habitus no. QL2301, prep. slide no. L23098 (JXAUM).

Paratypes. China, Sichuan Province: One female, same data as holotype, no. 20210416044; two males, one female, Dayi County, near Xiling Snow Mountain Tunnel (30°37'N, 103°19'E), alt. 860 m, 16 April 2021, Chao Jiang leg., no. 20210416031−202104160033; one female, Dayi County, Heishuihe Nature Reserve, Dafeishui (30°38'N, 103°10'E), alt. 1290 m, 16 April 2021, Chao Jiang leg., no. 20210416026; one male, Chengdu, Jincheng Park (30°34'N, 104°02'E), alt. 450 m, 19 April 2021, Chao Jiang leg., no. 20210419001 (CMMI).

Diagnosis

Male pleopod 1 endopod has two spinelike lobes at apical part of outer margin.

Description

Maximum length: male 3.6 mm and female 4.2 mm.

Body oval, flattened and pale brown. Cephalon with lateral lobes slightly protruding laterally, apex rounded. Eyes with three ommatidia. Pereonites 1–2 with postero-lateral corners nearly right-angled, pereonites 3–7 with postero-lateral corners directed backwards. Pleonites 3–5 with epimera falciform, protruding backwards. Telson triangular, twice as wide as long, lateral margin slightly concave, ending with rounded apex. Uropod exopod as long as endopod (Fig. 1A).

Figure 1. 

Habitus of Exalloniscus species. A. E. duospinatus sp. nov., holotype; B (holotype); C (paratype). E. curvispinatus sp. nov.; D. E. thailandensis, male; E. E. triangulus sp. nov., holotype; F. E. tridentatus sp. nov., holotype; G. E. cortii, male; H. E. taitii sp. nov., holotype. Scale bar: 1 mm.

Antenna with fifth article of peduncle slightly longer than flagellum; ratio of flagellum approximately 3:2:2 (Fig. 2A).

Figure 2. 

Exalloniscus duospinatus sp. nov., holotype. A. Antenna; B. Pereopod 1; C. Pereopod 7; D. Pleopod 1; E. Pleopod 2; F. Pleopod 3 exopod; G. Pleopod 4 exopod; H. Pleopod 5 exopod. Scale bar: 0.1 mm.

Pereopod 1 with long strong setae on sternal margin of merus and carpus, carpus with transversal antennal grooming brush (Fig. 2B). Pereopod 7 with several strong setae on sternal margin; basis with distinct water conducting system; ischium gently concave on rostral surface of base, and straight on sternal margin (Fig. 2C).

Male pleopod 1 exopod almost quadrangular, outer margin convex; endopod with apical part bearing two spinelike lobes on outer margin and four tiny spines on inner margin (Fig. 2D). Pleopod 2 exopod triangular with several setae on outer margin; endopod flagelliform, longer than exopod (Fig. 2E). Pleopods 3−5 exopods triangular with several setae on outer margin (Fig. 2F−H).

Etymology

Latin: prefix duo- = double plus spinatus = spinous. The new species name refers to the male pleopod 1 endopod with two spinelike lobes at the apical part of outer margin.

Remarks

This new species is similar to E. silvestrii Kwon & Taiti, 1993, but it can be distinguished by pereopod 7 ischium slightly concave at the base of the rostral surface, and pleopod 1 endopod bearing two spinelike lobes at the apical part of the outer margin (Fig. 2C, D). In E. silvestrii, the base of pereopod 7 ischium with a large flat rounded lobe on the rostral surface, and the apical part of pleopod 1 endopod with three triangular lobes on the outer margin (Kwon and Taiti 1993: figs 110, 112).

Exalloniscus curvispinatus Li & Jiang, sp. nov.

https://zoobank.org/1CFF7EBE-9FBA-46A3-857C-25511D27B65E
Figs 1B, C, 3

Type material

Holotype. China: male, Yunnan Province, Mengla County, Xiaomo Road (21°14'N, 101°42'E), alt. 650 m, 18 August 2023, Chao Jiang leg., habitus no. ML23002, prep. slide no. L23104 (JXAUM).

Paratypes. China, Yunnan Province: Two males, one female, same data as the holotype, no. 20230818301; one male, Menglian County, Nayunzhen (22°19'N, 99°35'E), 23 August 2023, Han Qiu leg., habitus no. NYZ23001, prep. slide no. L23105 (JXAUM); one male, Jinghong, Jinuo Village, Bakaxiaozhai (21°57'N, 101°12'E), 15 August 2023, Chao Jiang leg., no. 20230815301; three males, one female, Menglian County, Nayun Town, 23 August 2023, Han Qiu leg., nos. 20230302301 and 20230302202 (CMMI).

Diagnosis

Male pleopod 1 exopod almost pentagonal, and endopod recurved outwards at apical part.

Description

Maximum length: male 5.0 mm and female 4.2 mm.

Body oval, flattened and white. Cephalon with lateral lobes slightly protruding laterally, nearly triangular with blunted apex. Eyes absent. Pereonites with postero-lateral corner progressively more acute, directed backwards. Pleonites 3−5 with epimera falciform, protruding backwards. Telson triangular, twice as wide as long, lateral margin gently concave, ending with rounded apex. Uropod exopod as long as endopod (Fig. 1B).

Antenna with fifth article of peduncle slightly longer than flagellum; ratio of flagellum approximately 4:3:3 (Fig. 3A).

Figure 3. 

Exalloniscus curvispinatus sp. nov., holotype. A. Antenna; B. Pereopod 1; C. Pereopod 7; D. Pleopod 1; E. Pleopod 2; F. Pleopod 3 exopod; G. Pleopod 4 exopod; H. Pleopod 5 exopod. Scale bar: 0.2 mm.

Pereopod 1 with long strong setae on sternal margin of merus and carpus, carpus with basal part as wide as apical part, bearing transversal antennal grooming brush (Fig. 3B). Pereopod 7 with several strong setae on sternal margin; basis with distinct water conducting system; ischium straight on sternal margin (Fig. 3C).

Male pleopod 1 exopod almost pentagonal; endopod with triangular lobe on outer margin, apical part thin and long, recurved outwards (Fig. 3D). Pleopod 2 exopod oval, apical part narrowed toward round apex; endopod flagelliform, longer than exopod (Fig. 2E). Pleopods 3−5 exopods oval with several setae on outer margin (Fig. 3F−H).

Etymology

Latin: prefix curv- = curved plus spinatus = spinous. The new species name refers to the apical apex of male pleopod 1 endopod with a spinelike projection recurved outwards.

Remarks

This species varied in body shapes (Fig. 1B versus 1C). Specimens collected in Mengla (Fig. 1B) are distinctly thinner than the specimens found in Menglian (Fig. 1C).

The new species is similar to E. thailandensis in having a recurved pleopod 1 endopod (Fig. 3 versus Fig. 4). However, it can be distinguished by pereopod 1 with the basal part as wide as the apical part, pleopod 1 exopod almost pentagonal, and endopod without transversal sclerotized projections at the apical part (Fig. 3B, D). In E. thailandensis, the basal part of pereopod 1 is distinctly narrower than the apical part, pleopod 1 exopod is almost rounded, and endopod armed with transversal sclerotized projections at the apical part (Fig. 4B, D). Moreover, the body pigments of the new species (white, Fig. 1B, C) differ from E. thailandensis (brown, Fig. 1D) based on two male specimens from China, Yunnan Province, Mengla County, Guanlei Town, Baojiaoniu Cave. However, the colourless individuals of E. thailandensis have been described previously (see Taiti and Ferrara 1988). Thus, the body shape and colour should not be considered as exact characters to identify species of the genus.

Figure 4. 

Exalloniscus thailandensis (two males, China, Yunnan Province, Mengla, County, Guanlei Town, Baojiaoniu Cave, 17 August 2023, Chao Jiang leg., prep. slide no. L23106). A. Antenna; B. Pereopod 1; C. Pereopod 7; D. Pleopod 1; E. Pleopod 2; F. Pleopod 3 exopod; G. Pleopod 4 exopod; H. Pleopod 5 exopod. Scale bar: 0.2 mm.

Exalloniscus triangulus Li & Jiang, sp. nov.

https://zoobank.org/1E234552-69F1-4128-BACA-9FC8609F649F
Figs 1E, 5

Type material

Holotype. China: male, Hubei Province, Jingshan County, Kongshandong Scenic Area (30°58'N, 113°02'E), alt. 100 m, 9 April 2021, Zhidong Wang and Tianyun Chen leg., habitus no. KSD2302, prep. slide no. L23097 (JXAUM).

Diagnosis

Male pleopod 1 exopod distinctly convex on outer margin, and endopod with a triangular lobe and a seta recurved inwards at apical part.

Description

Length 2.9 mm.

Body oval, flattened and white mixed with pale brown. Cephalon with lateral lobes slightly protruding laterally, apex rounded. Eyes with three ommatidia. Pereonites 1 with postero-lateral corner nearly right-angled, pereonites 2−7 with postero-lateral corners directed backwards. Pleonites 3−5 with epimera falciform, protruding backwards. Telson triangular, twice as wide as long, lateral margin gently concave, ending with rounded apex. Uropod exopod almost as long as endopod (Fig. 1E).

Antenna with fifth article of peduncle longer than flagellum; ratio of flagellum approximately 3:2:2 (Fig. 5A).

Pereopod 1 with long strong setae on sternal margin of merus and carpus, carpus with transversal antennal grooming brush (Fig. 5B). Pereopod 7 with several strong setae on sternal margin; basis with distinct water conducting system; ischium slightly convex on sternal margin (Fig. 5C).

Figure 5. 

Exalloniscus triangulus sp. nov., holotype. A. Antenna; B. Pereopod 1; C. Pereopod 7; D. Pleopod 1; E. Pleopod 2; F. Pleopod 3 exopod; G. Pleopod 4 exopod; H. Pleopod 5 exopod. Scale bar: 0.1 mm.

Male pleopod 1 exopod basal half broad, distal half narrowed towards blunted apex, distinctly convex on outer margin; endopod ending with triangular lobe and seta recurved inwards (Fig. 5D). Pleopod 2 exopod triangular with several setae on outer margin; endopod flagelliform, longer than exopod (Fig. 5E). Pleopods 3 and 4 exopods almost oval, pleopod 5 triangular with several setae on outer margin (Fig. 5F−H).

Etymology

Latin: triangulus = triangular. The new species name refers to the male pleopod 1 endopod ending with a triangular lobe.

Remarks

This new species is similar to E. malaccensis Taiti & Ferrara, 1988 in having a triangular lobe at the apical part of male pleopod 1 endopod. But it can be distinguished by pleopod 1 exopod distinctly convex on the outer margin, and the apical part of endopod with a triangular lobe and a seta recurved inwards (Fig. 5D). In E. malaccensis, pleopod 1 exopod is distinctly concave on the outer margin, and the apical lobe of endopod is thinner and recurved outwards (Taiti and Ferrara 1988: fig. 23D).

Exalloniscus tridentatus Li & Jiang, sp. nov.

https://zoobank.org/C7A742F7-5BCF-4A53-AA09-099FB4E8DDB3
Figs 1F, 6

Type material

Holotype. China: male, Shaanxi Province, Zhashui County, Dongshan Forestry Park (33°42'N, 109°01'E), alt. 1020 m, 12 May 2021, Chao Jiang leg., habitus no. ZS2301, prep. slide no. L23095 (JXAUM).

Paratypes. China: two males, two females, same data as the holotype, nos. 20210512007−20210512009 (CMMI).

Diagnosis

Male pleopod 1 exopod straight on outer margin, and endopod bearing three well-developed dentations at apical part.

Description

Maximum length: male 2.5 mm and female 2.8 mm.

Body oval, slightly convex. Colour white mixed with pale brown. Cephalon with lateral lobes slightly protruding laterally, apex rounded. Eyes with three ommatidia. Pereonites with postero-lateral corners directed backwards. Pleonites 3−5 with epimera falciform, protruding backwards. Telson triangular, twice as wide as long, lateral margin slightly concave, apex rounded. Uropod exopod approximately twice as long as endopod (Fig. 1F).

Antenna with fifth article of peduncle slightly longer than flagellum; ratio of flagellum approximately 3:2:2 (Fig. 6A).

Figure 6. 

Exalloniscus tridentatus sp. nov., holotype. A. Antenna; B. Pereopod 1; C. Pereopod 7; D. Pleopod 1; E. Pleopod 2; F. Pleopod 3 exopod; G. Pleopod 4 exopod; H. Pleopod 5 exopod. Scale bar: 0.1 mm.

Pereopod 1 with long strong setae on sternal margin of merus and carpus (Fig. 6B). Pereopod 7 with several strong setae on sternal margin; basis without water conducting system; ischium almost straight on sternal margin (Fig. 6C).

Male pleopod 1 exopod oval, straight on outer margin; endopod with apical portion bearing rounded hyaline lobe and three well-developed dentations (Fig. 6D). Pleopod 2 exopod triangular; endopod flagelliform, longer than exopod (Fig. 6E). Pleopods 3−5 nearly triangular with several setae on outer margin (Fig. 6F−H).

Etymology

Latin: tridentatus = trident. The new species name refers to the male pleopod 1 endopod with three dentations at apical portion.

Remarks

This new species is similar to E. cortii Arcangeli, 1927 by body shape and male pleopod 1 endopod bearing a rounded hyaline lobe at the apical portion (Figs 1F, 6D versus Figs 1G, 7D). But it can be distinguished by pereopod 7 basis without water conducting system, pleopod 1 exopod straight on the outer margin, and endopod bearing three well-developed dentations at the apical portion (Fig. 6C, D). In E. cortii, pereopod 7 basis has a well-developed water conducting system, pleopod 1 exopod is sinuous on the outer margin, and endopod with two spinelike projections at the apical apex (Fig. 7C, D).

Figure 7. 

Exalloniscus cortii (one male, China, Hunan Province, Yongzhou, Puliqiao Town, 21 March 2019, Chao Jiang leg., prep. slide no. L23099). A. Antenna; B. Pereopod 1; C. Pereopod 7; D. Pleopod 1; E. Pleopod 2; F. Pleopod 3 exopod; G. Pleopod 4 exopod; H. Pleopod 5 exopod. Scale bar: 0.1 mm.

Exalloniscus taitii Li & Jiang, sp. nov.

https://zoobank.org/D814EDC9-41DF-44F6-A6D6-FD5B4C9D9EF8
Figs 1H, 8

Type material

Holotype. China: male, Yunnan Province, Gengma County, Daxing Village (23°45'N, 99°46'E), alt. 2270 m, 29 May 2021, Chao Jiang leg., habitus no. DXX2302, prep. slide no. L23100 (JXAUM).

Diagnosis

Male pleopod 1 endopod conspicuously concave near middle of outer margin and bearing a tiny spine at apical part of inner margin, pleopods 3 concave on outer margin.

Description

Length 2.0 mm.

Body oval, slightly convex. Colour pale brown. Cephalon with lateral lobes slightly protruding laterally, apex rounded. Eyes with three ommatidia. Pereonites with postero-lateral corners directed backwards. Pleonites 3−5 with epimera falciform, protruding backwards. Telson triangular, twice as wide as long, lateral margin slightly concave, apex rounded. Uropod exopod approximately twice as long as endopod (Fig. 1H).

Antenna with fifth article of peduncle slightly shorter than flagellum; ratio of flagellum approximately 3:3:4 (Fig. 8A).

Figure 8. 

Exalloniscus taitii sp. nov., holotype. A. Antenna; B. Pereopod 1; C. Pereopod 7; D. Pleopod 1; E. Pleopod 2; F. Pleopod 3 exopod; G. Pleopod 4 exopod; H. Pleopod 5 exopod. Scale bar: 0.1 mm.

Pereopod 1 with long strong setae on sternal margin of merus and carpus, carpus without transversal antennal grooming brush (Fig. 8B). Pereopod 7 with several strong setae on sternal margin; basis without water conducting system; ischium almost straight on sternal margin (Fig. 8C).

Male pleopod 1 exopod base narrow, distinctly broaden towards apical part, almost straight on outer margin; endopod base broad, narrowed towards point apex, outer margin conspicuously concave near middle, apical part with tiny spine on inner margin (Fig. 8D). Pleopod 2 exopod triangular, outer margin concave; endopod flagelliform, longer than exopod (Fig. 8E). Pleopods 3−5 nearly triangular, pleopods 3 concave on outer margin, pleopods 4−5 almost straight on outer margin (Fig. 8F−H).

Etymology

The new species name honors Dr. Stefano Taiti for his invaluable contribution to the taxonomy of terrestrial isopods.

Remarks

This new species is similar to E. burmaensis Taiti & Cardoso, 2020 by the traits of male pleopod 1. But it can be distinguished by the ratio of flagellum approximately 3:3:4; pereopod 1 carpus without transversal antennal grooming brush, and pereopod 7 basis without water conducting system; pleopod 1 endopod conspicuously concave near the middle of the outer margin, and bearing a tiny spine at the apical part of the inner margin, pleopods 3 concave on the outer margin (Fig. 8A−D, F). In E. burmaensis, flagellum is diminishing in length from the first segment to the third segment; pereopod 1 carpus with a transversal antennal grooming brush, and pereopod 7 basis has a well-developed water conducting system; pleopod 1 endopod is almost straight on the outer margin, and the apical part without spine on the inner margin, pleopods 3 is straight on the outer margin (Taiti and Cardoso 2020: Fig. 17H, 18B−D, F).

Biogeographical considerations

This taxonomic work describes five new species of the genus Exalloniscus from China. To date, a total of eleven species of the genus occur in China (Fig. 9). All the Exalloniscus species from China described here are distributed southward to the 0 °C isotherm of the coldest month (January) of the year, and their habitus almost locates in a humid area with annual precipitation above 800 mm. Among them, only Exalloniscus cortii Arcangeli, 1927 shows a wide distribution. In addition to South China, it is also recorded from sub-humid areas near 40°N of North China (Fig. 9). Based on our available specimens and historical records (Taiti and Ferrara 1988), E. cortii has the northernmost distribution in China reaching Shanhai Pass of Hebei Province, the border between Northeast China and North China (Fig. 8). Thus, all Chinese species of this genus are humidity and warmth dependent except for E. cortii, which has demonstrated to have some level of cold tolerance.

Figure 9. 

Map of China showing the localities where Exalloniscus species are recorded.

Furthermore, the worldwide records indicate the occurrence of Exalloniscus species from South Asia, Southeast Asia to East Asia (Manicastri and Argano 1986; Taiti and Ferrara 1988; Manicastri and Taiti 1991; Kwon 1993, 1995; Kwon and Taiti 1993; Nunomura 2000; Nunomura and Xie 2000; Taiti and Gruber 2008; Taiti and Cardoso 2020). This geographical region is strongly connected to climatic variables, in which the precipitation and minimum temperature are very close to the geographic distribution in China. This suggests that the precipitation and minimum temperature may be the important factors affecting the geographical distribution of Exalloniscus members. In addition, most Exalloniscus species live in association with ants or termites, but the available data are still only fragmentary and insufficient for the biology of many species (Taiti and Ferrara 1988; Taiti and Cardoso 2020). Further research is necessary to precisely identify biotic and abiotic requirements for Exalloniscus species as well as to describe the largely unstudied diversity of the genus in China.

Acknowledgements

We are grateful to Dr. Stefano Taiti (Istituto per lo Studio degli Ecosistemi, Italy), Dr. Christian Schmidt (Senckenberg Naturhistorische Sammlungen Dresden, Germany), and Dr. Noboru Nunomura (Kanazawa University, Japan) for providing important references. We are greatly appreciative of the efforts of Christina Yuan (University of San Francisco, USA) for her help in improving the English. We are also indebted to Han Qiu, Zhidong Wang and Tianyun Chen for collecting specimens. Special thanks are given to Dr. Luiz F. Andrade and two anonymous reviewers for their insightful comments and suggestions on the manuscript. This research was supported by the National Natural Science Foundation of China (nos. 31960100, 82073972) and the Fundamental Research Funds for the Central Public Welfare Research Institutes (no. ZZ13-YQ-089-C1).

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