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. 2025 Apr 25;1236:85–101. doi: 10.3897/zookeys.1236.139274

Two new species of larval Erythraeidae (Parasitengona) ectoparasites of leafhoppers from Southwestern China

Yan Jiang 1,2, Tian-Ci Yi 1, Si-Yuan Xu 1,✉, Dao-Chao Jin 1,✉
PMCID: PMC12048815  PMID: 40322614

Abstract

In this study, three species were examined. Among them, two new species, Caeculisomataianensissp. nov. and Iguatoniabarboproximasp. nov. from Southwestern China, were described and illustrated based on larvae. The two new species can be distinguished from the known species by the following characteristics: the anterior sensilla are nude, and the gnathosoma has two pairs of nude hypostomalae in C.taianensissp. nov.; the anterior sensilla and posterolateral scutalae are located in the posterior half of the scutum, and the posterior hypostomalae with barbs on the proximal half in I.barboproximasp. nov. In addition, a new host and distribution range of Abrolophusquadrapexicis Xu & Jin, 2022 is reported.

Key words: Abrolophinae, Callidosomatinae, Chongqing, insect, new host, range extension, taxonomy, Yunnan

Introduction

Southwestern China comprises three provinces (Sichuan, Guizhou, and Yunnan), one municipality (Chongqing), and one autonomous region (Xizang (Tibet)). This region is characterized by diverse topography with significant variations in elevation and numerous separated basins (Shi et al. 2018), including the Yunnan-Guizhou Plateau, the Qinghai-Tibetan Plateau, the Hengduan Mountains, and the Sichuan Basin. Simultaneously, the region exhibits diverse climatic conditions, including a subtropical monsoon climate, plateau mountain climate, mountain climate, tropical monsoon climate, dry-hot valley climate and a temperate monsoon climate. The variety of climates provides a wealth of environmental conditions, which play a pivotal role in fostering species diversity in the ecosystem. Therefore, Southwestern China is a key area for fauna-flora biodiversity research in China (Han et al. 2008; Shi et al. 2018).

To date, 13 species in eight genera (Balaustium von Heyden, 1826, Caeculisoma Berlese, 1888, Charletonia Oudemans, 1910, Erythraeus Latreille, 1806, Grandjeanella Southcott, 1961, Leptus Latreille, 1796, Marantelophus Haitlinger, 2011, and Neoabrolophus Khot, 1965) of five subfamilies (Abrolophinae Witte, 1995, Balaustiinae Grandjean, 1947, Callidosomatinae Southcott, 1957, Erythraeinae Robineau-Desovidy, 1828, and Leptinae Billberg, 1820) of Erythraeidae have been documented in Southwestern China (Xu et al. 2017, 2019a, b, 2020, 2022a, 2022b, 2023, 2025). Of them, three species were described based on post-larval forms, one species was recorded based on both larval and post-larval stages, and the remaining species were known only by larvae (Table 1).

Table 1.

Hosts and distribution of the known species of Erythraeidae from Southwestern China.

Species Host Distribution
Balaustiumallomedicagoense Xu & Jin, 2025 [P] Unknown Yunnan
Balaustiumneomedicagoense Xu & Jin, 2025 [P] Unknown Xizang
Caeculisomapenlineatus Xu & Jin, 2019 [L] Mileewamargheritae (female) (Hemiptera, Cicadellidae, Mileewinae), Neuterthronhamuliferum (male) (Hemiptera, Delphacidae), unidentified Alebrini (Hemiptera, Cicadellidae, Typhlocybinae), unidentified Delphacinae (Hemiptera, Delphacidae), unidentified Issidae (Hemiptera, Auchenorrhyncha) Chongqing, Guizhou
Caeculisomasemispinus Xu & Jin, 2019 [L] Shaddai sp. (male) (Hemiptera, Cicadellidae), unidentified Zyginellini (female) (Cicadellidae) Chongqing
Charletoniarectangia Xu & Jin, 2022 [L] Unidentified Acrididae (Orthoptera), unidentified Tettigoniidae (Orthoptera), unknown Chrysomelidae (Coleoptera), unidentified mantis (Mantodea), unidentified moth (Lepidoptera), unidentified stick insect (Phasmatodea) Yunnan
Erythraeus (Zaracarus) mossesus Xu & Jin, 2023 [P, L] Unknown Guizhou
Grandjeanelladianensis Xu & Jin, 2022 [L] Unknown. Yunnan
Leptus (Leptus) bomiensis Xu & Jin, 2022 [L] Unidentified moth (Lepidoptera), unidentified Elateridae (Coleoptera), unidentified Pentatomidae (Hemiptera) Xizang
Leptus (Leptus) striatus Xu & Jin, 2022 [L] Unidentified Opiliones Yunnan
Leptus (Leptus) trisolenidionus Xu & Jin, 2022 [L] Unidentified Cicadellinae (Hemiptera, Cicadellidae) Guizhou
Marantelophusdubifurcatus Xu, Yi & Jin, 2017 [L] Cacopsylla sp. (Hemiptera, Psyllidae), unidentified Psocoptera. Guizhou
Marantelophusneodubifurcatus Xu & Jin, 2023 [L] Unknown Guizhou
Neoabrolophusguizhouensis Xu & Jin, 2025 [P] Unknown Guizhou
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In this checklist, host and distribution data were obtained from Xu et al. 2017, 2019a, b, 2020, 2022a, b, 2023, 2025. [P]: Post-larval form. [L]: Larva.

Species of the genus Abrolophus Berlese, 1891 from China are distributed in the Macao Special Administrative Region, Hainan Province, Zhejiang Province, Shandong Province, Guangxi Zhuang Autonomous Region, Guangdong Province, and Hunan Province (Zheng 2002a; Haitlinger 2006; Xu et al. 2021, 2022a). However, there has been no report of Abrolophus in Southwestern China.

A total of 27 species of Caeculisoma have been reported worldwide, of which 14 were based on the post-larval stage, 12 were based on the larval stage, and only one species was based on both larval and post-larval instars (Mąkol and Wohltmann 2012; Xu et al. 2019a, 2019b, 2020; Saboori et al. 2023; Kohansal et al. 2024; Noei et al. 2024). Among the 27 known Caeculisoma species, four were recorded in China based on the larval stage, with two (C.penlineatus Xu & Jin, 2019, C.semispinus Xu & Jin, 2019) collected from Southwestern China (Zheng 2002b; Xu et al. 2019a, 2019b, 2020).

Hitherto, only two species of the genus Iguatonia have been reported based on their larval stage worldwide (Haitlinger 2004; Xu et al. 2020; Noei et al. 2024): I.barbillae Haitlinger, 2004 from Brazil and I.xinfengi Xu & Jin, 2020 from Hainan Province (Island), China.

In this study, two new species, C.taianensis sp. nov. and I.barboproxima sp. nov., collected from Yunnan Province and Chongqing Municipality, respectively, are described and illustrated based on larvae. Additionally, new data for A.quadrapexicis Xu & Jin, 2022 is provided.

Material and methods

Erythraeid larvae were collected along with their insect hosts using 200 mesh insect nets, and subsequently preserved in small vials containing absolute ethanol. The larvae on the hosts were detached using a fine brush under a stereomicroscope (Nikon SMZ745) in the lab. Then, all larval specimens were cleared in Oudemans’ fluid for about 12 h at 25 °C and slide-mounted in Hoyer’s medium (Walter and Krantz 2009). Figures were drawn with the aid of a drawing tube attached to a Nikon Eclipse Ni-E compound microscope. Genus identification is based on the key to world genera of larval Callidosomatinae of Noei et al. (2024). Terminology and abbreviations are adapted from Noei et al. (2024), Wohltmann et al. (2007) and Xu et al. (2020). Measurements are expressed in micrometers (μm). The standard deviations (SD) are provided with two decimal places. All specimens were deposited at the Institute of Entomology, Guizhou University, Guiyang, China (GUGC).

Results

Abrolophinae Witte, 1995

Abrolophus Berlese, 1891

. Abrolophus quadrapexicis

Xu & Jin, 2022

A26E19A0-AF53-5C5F-961E-59122D327E44

Material examined.

China • one larva (2000–1600-GZ-yj); Guizhou Province, Fanjingshan National Nature Reserve; 27°59'27″N, 108°33'30″E; 673 m; 30 Jun. 2023; Si-Yuan Xu leg.; on an unidentified Psyllidae (Hemiptera).

Distribution.

Guizhou Province (new distribution), Shandong Province, Zhejiang Province.

Note.

This species was collected from plants without a host record in Zhejiang and Shandong (Xu et al. 2022b).

Callidosomatinae Southcott, 1957

Caeculisoma Berlese, 1888

. Caeculisoma taianensis sp. nov.

F0A460C9-EA62-55AC-BE81-E3CE2C1A1F06

https://zoobank.org/F5C1A34A-2366-423A-B816-969D2202B854

Figs 1 , 2 , 3 , 4

Figure 1.

Figure 1.

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Caeculisomataianensis sp. nov., larva A dorsal view of idiosoma B ventral view of idiosoma C dorsal view of coxa I, showing supracoxal seta.

Figure 2.

Figure 2.

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Caeculisomataianensis sp. nov., larva A scutum B ventral view of gnathosoma C dorsal view of the cheliceral bases.

Figure 3.

Figure 3.

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Caeculisomataianensis sp. nov., larva photograph. Scutum, showing the shape of scutum, ASE and PSE.

Figure 4.

Figure 4.

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Caeculisomataianensis sp. nov., larva. Leg I A trochanter—genu B tibia—tarsus; leg II C trochanter—genu D tibia—tarsus; leg III E trochanter—genu F tibia—tarsus.

Diagnosis (larva).

ASE nude and posterior to the level of ML, closer to ML than PL; PSE with barbs on the distal one-third; gnathosoma with two pairs of nude hypostomalae; ISD 56–63; Ti I 188–207; Ti III 264–287.

Description.

Dorsum. Idiosoma lateral cuticle of holotype used for drawing broken slide preparation, almost oval, with 30 (fD = 28–30 in paratypes) barbed setae, a pair of setae located between scutum and eyes (Fig. 1A). Scutum outline pentagonal with rounded angles, length somewhat longer than width, anterior margin slightly concave, anterolateral and posterolateral margins slightly sinuous, posterior margin with small concavity between bases of PSE (Figs 1A, 2A, 3). Three pairs of normal setae (AL, ML and PL), and two pairs of sensilla (ASE and PSE) placed on scutum. AL, ML and PL completely barbed, ASE nude, PSE with fine barbs in distal about one-third. ASE placed between ML and PL, and closer to ML than PL, PSE near posterior margin of scutum. PSE much longer than ASE, ML slightly longer than AL and PL, AL slightly longer than PL, one paratype (c) AL equal to PL (Table 2).

Table 2.

Metric and some meristic data of Caeculisomataianensis sp. nov. (larvae, a–f = paratypes; a: 1088–0275-YN-yl, b: 1082–0271-YN-yl, c: 1083–0271-YN-yl, d: 1084–0271-YN-yl, e: 1085–0272-YN-yl, f: 1086–0273-YN-yl).

Character Holotype a b c d e f SD range
fD 30 28 30 30 28 30 28 0.99 28–30
fV 14 14 12 14 12 14 14 0.90 12–14
NDV 44 42 42 44 40 44 42 1.40 40–44
IL 999 689 1121 901 1069 1061 1543 240.10 689–1543
IW 843 415 840 608 720 678 1028 181.47 415–1028
DS 30–66 33–57 33–59 32–62 36–64 33–60 33–55 1.64–3.58 30–66
DS min. 30 33 33 32 36 33 33 1.64 30–36
DS max. 66 57 59 62 64 60 55 3.58 55–66
PDS 34–66 36–57 37–59 34–62 37–64 36–60 33–55 1.48–3.58 33–66
PDS min. 34 36 37 34 37 36 33 1.48 33–37
PDS max. 66 57 59 62 64 60 55 3.58 55–66
Oc 21 22 22 23 21 20 19 1.25 19–23
1a 35 30 32 27 21 25 24 4.53 21–35
2a 44 47 43 38 39 44 40 3.00 38–47
3a 38 36 38 32 34 35 33 2.17 32–38
1b 74 67 74 66 72 64 71 3.73 64–74
2b1 47 49 43 44 47 48 40 2.97 40–49
2b2 36 34 29 32 33 34 31 2.12 29–36
3b1 48 44 46 44 46 46 40 2.36 40–48
3b2 35 37 36 36 34 34 32 1.55 32–37
L 114 109 109 110 111 114 103 3.46 103–114
W 108 103 103 100 103 102 97 3.10 97–108
AW 57 60 57 53 55 61 56 2.56 53–61
MW 76 71 71 71 74 72 69 2.14 69–76
PW 85 78 79 77 79 80 76 2.70 76–85
MA 41 38 40 43 40 43 38 1.92 38–43
AA 13 12 12 13 12 13 11 0.70 11–13
SB 15 14 15 14 15 16 14 0.70 14–16
ISD 63 60 60 57 59 58 56 2.14 56–63
AP 56 52 54 55 54 53 54 1.20 52–56
AL 46 45 37 44 44 45 40 3.02 37–46
ML 50 50 49 48 46 51 47 1.67 46–51
PL 41 41 37 37 37 36 35 2.19 35–41
ASE 39 34 33 39 31 36 33 2.88 31–39
PSE 71 66 60 70 60 67 57 5.04 57–71
as 15 14 16 15 17 18 16 1.25 14–18
bs 34 27 30 36 35 33 34 2.91 27–36
cs 26 23 20 28 25 22 25 2.47 20–28
PaScFed 74 78 76 81 83 85 73 4.24 73–85
PaScGed 36 33 31 34 37 36 34 1.92 31–37
GL 136 141 132 135 136 140 138 2.85 132–141
Ta I (H) 22 19 22 20 18 18 15 2.29 15–22
Ta I (L) 152 153 160 159 152 160 157 3.44 152–160
Ti I 199 191 200 192 201 207 188 6.22 188–207
Ge I 162 155 166 151 151 162 152 5.76 151–166
TFe I 93 88 93 88 92 96 89 2.81 88–96
BFe I 105 96 100 94 97 101 95 3.61 94–105
Tr I 54 47 48 48 47 44 52 3.11 44–54
Cx I 56 51 57 58 63 64 62 4.27 51–64
Ta II (H) 18 20 19 21 22 17 17 1.81 17–22
Ta II (L) 157 153 158 154 157 159 161 2.56 153–161
Ti II 215 198 214 204 211 216 194 8.17 194–216
Ge II 161 150 161 154 157 163 148 5.39 148–163
TFe II 95 86 85 85 91 90 84 3.78 84–95
BFe II 95 93 96 96 96 102 91 3.16 91–102
Tr II 58 54 50 53 48 51 54 3.02 48–58
Cx II 61 66 69 67 68 74 75 4.44 61–75
Ta III (H) 15 19 14 13 14 18 14 2.12 13–19
Ta III (L) 170 167 170 165 171 169 176 3.19 165–176
Ti III 287 266 277 264 271 277 273 7.17 264–287
Ge III 161 153 164 153 156 166 156 4.87 153–166
TFe III 136 124 132 133 128 131 126 3.89 124–136
BFe III 124 119 121 120 119 128 118 3.28 118–128
Tr III 66 57 57 53 53 54 51 4.61 51–66
Cx III 66 66 67 71 72 78 66 4.20 66–78
Leg I 821 781 824 790 803 834 795 18.28 781–834
Leg II 842 800 833 813 828 855 807 18.34 800–855
Leg III 1010 952 988 959 970 1003 966 20.68 952–1010
IP 2673 2533 2645 2562 2601 2692 2568 56.05 2533–2692
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Venter. All ventral setae, including coxalae, barbed and with pointed ends (Fig. 1B). Three pairs of intercoxal setae (1a, 2a and 3a), 2a longer than 1a and 3a, 3a slightly longer than 1a (Table 2), 14 setae behind coxae III (fV = 12–14 in paratypes). Five pairs of coxalae (1b, 2b1, 2b2, 3b1 and 3b2), 1b much longer than the other coxalae, 2b1 subequal 3b1, 2b2 and 3b2 subequal, 2b1 and 3b1 longer than 2b2 and 3b2, respectively (Table 2). Dorsum of coxa I with a peg-like supracoxal seta (elc I) (Fig. 1C).

Gnathosoma (Fig. 2B, C). Dorsal view of the cheliceral base punctated. One pair of galealae (cs) and two pairs of hypostomalae (as and bs) nude; bs longer than cs and much longer than as (Table 2). Hypostomal lip with fimbriation. Palpfemur and palpgenu each with one barbed, pointed dorsal seta (PaScFed and PaScGed). Palptibia with three barbed setae, one on ventral surface, odontus bifid. Palptarsus with seven setae, five nude, one solenidion (ω) and one eupathidium (ζ). fPp = 0-B-B-3B2-5Nωζ. Palpal supracoxal seta (elcp) peg-like.

Legs (Figs 1B, 3). With seven segments (femora divided). IP = 2533–2692 (Holotype and six paratypes). Claws hook-like and posterior claw with few ciliations, and empodium claw-like. Normal setae on legs barbed and pointed. Leg setal formula: leg I: Cx—1n; Tr—1n; Bfe—4n; Tfe—5n; Ge—1σ, 1κ, 12n; Ti—2φ, 1κ, 1Cp, 18n; Ta—1ω, 1ε, 2ζ, 1Cp, 27n. leg II: Cx—2n; Tr—1n; Bfe—4n; Tfe—5n; Ge—1κ, 12n; Ti—2φ, 19n; Ta—1ω, 1ζ, 28n. leg III: Cx—2n; Tr—1n; Bfe—2n; Tfe—5n; Ge—12n; Ti—1φ, 19n; Ta—1ζ, 28n. The morphometric data of the legs is listed in Table 2.

Etymology.

The new species’ name is derived from Taian Town, where the holotype and paratype were collected.

Material examined.

Holotype. China • a larva (1087–0275-YN-yl); Yunnan Province, Yulong County, Taian Town; 26°37'8"N, 100°02'9"E; 2502 m; 8 Aug. 2021; Yan Jiang leg.; from Atkinsoniella sp. (Hemiptera, Cicadellidae). Paratypes China • one larva (1088–0275-YN-yl), same data as the holotype. China • three larvae (1082–0271-YN-yl, 1083–0271-YN-yl, 1084–0271-YN-yl); Yunnan Province, Yulong County, Taian Town; 26°37'8"N, 100°02'9"E; 2502 m; 8 Aug. 2021; Yan Jiang leg.; from unidentified Atkinsoniella (Hemiptera: Cicadellidae). China • one larva (1085–0272-YN-yl); Yunnan Province, Yulong County, Taian Town; 26°37'8"N, 100°02'9"E; 2502 m; 8 Aug. 2021; Yan Jiang leg.; from an unknown nymph of Cicadellidae. China • one larva (1086–0273-YN-yl); Yunnan Province, Yulong County, Taian Town; 26°37'8"N, 100°02'9"E; 2502 m; 8 Aug. 2021; Yan Jiang leg.; from an unidentified nymph of Cicadellidae.

The holotype and paratypes are deposited in the Institute of Entomology, Guizhou University, Guiyang, China (GUGC).

Remarks.

Based on the description of larvae, the genus Caeculisoma includes 13 species so far; of them, four species are from Australia and four from China, two species are from Brazil, while the remaining three species are found in Iran, the Republic of South Africa and New Zealand, respectively (Mąkol and Wohltmann 2012; Xu et al. 2020; Kohansal et al. 2024; Noei et al. 2024).

Similar species to C.taianensis sp. nov. are currently known as C.darwiniense Southcott, 1961, C.mouldsi Southcott, 1988, C.pouyani Noei & Kohansal, 2024 and C.sparnoni Southcott, 1972 based on the key to species of Caeculisoma in Kohansal et al. (2024).

The new species differs from C.darwiniense by the shape of ASE (nude vs barbed), posterior hypostomalae (bs) (nude vs barbed), galealae (cs) (nude vs barbed), the number of normal setae on fn Ge I–III (12-12-12 vs 11-12-13), the number of normal setae on fn Ti I–III (18-19-19 vs 17-18-20), longer L (103–114 vs 87–88), Ti I (188–207 vs 102), and Ti III (264–287 vs 143); differs from C.mouldsi by the shape of ASE (nude vs barbed), shape of posterior hypostomalae (nude vs barbed), gnathosoma with two pairs hypostomalae (vs one pair hypostomalae), the number of normal setae on fn Ge I–III (12-12-12 vs 12-11-12), the number of normal setae on fn Ti I–III (18-19-19 vs 18-19-18), longer 1b (64–74 vs 37–58), PaScFed (73–85 vs 42), leg II (800–855 vs 750), leg III (952–1010 vs 915), IP (2533–2692 vs 2455); differs from C.pouyani by the shape of ASE (nude vs barbed), galealae (nude vs barbed), hypostomalae (nude vs barbed), palp tarsus with five nude normal setae (vs with five barbed normal setae), longer W (97–108 vs 75–87), 1b (64–74 vs 40–46), Ti I (188–207 vs 85–95), Ti II (194–216 vs 85–92), Ti III (264–287 vs 115–130), IP (2533–2692 vs 1360–1505) and differs from C.sparnoni by shape of ASE (nude vs barbed), cheliceral bases without striations (vs with lengthwise striations), longer L (103–114 vs 83), W (97–108 vs 85), 1b (64–74 vs 28), Ta I (150–160 vs 77), Ti I (188–207 vs 83), Ta III (165–176 vs 79), Ti III (264–287 vs 100), IP (2533–2692 vs 1220).

The differences between the new species and the four present species of Caeculisoma found in China are as follows: C.taianensis sp. nov. differs from C.allopenlineatus by the positions of ASE (ASE closer to ML than PL vs ASE closer to PL than ML), shape of scutum (pentagonal vs oval), ASE (nude vs barbed), the longer Ti I (188–207 vs 164–170), Ti II (194–216 vs 151–161), Ti III (264–287 vs 227–239), IP (2533–2692 vs 2061–2115), the shorter ML (46–51 vs 114–120); differs from C.hunanica by the number of solenidia on Ti II (2 vs 1), the number of normal setae on TFe III (5 vs 4), the number of normal setae on fn Ti I–III (18-19-19 vs 16-16-18), longer leg I (781–834 vs 655), leg II (800–855 vs 621), leg III (952–1010 vs 728), IP (2533–2692 vs 2004); differs from C.penlineatus by ASE base location (closer to ML than PL vs in line with the level of PL), BFe I and II with four barbed setae (vs with three barbed setae and one nude seta), palptarsus with one eupathidium (vs with two eupathidia), longer Ti I (188–207 vs 143–167), Ti II (194–216 vs 150–179), Ti III (264–287 vs 213–239), leg I (781–834 vs 627–709), leg II (800–855 vs 645–724), leg III (952–1010 vs 768–878), IP (2533–2692 vs 2060–2298), data based on Xu et al. (2019b, 2020) and differs from C.semispinus by the shape of ASE (nude vs with barbs on distal halves), palptibia with three barbed setae (vs with one barbed seta and two nude setae), palptarsus with five nude setae (vs with two barbed setae and three nude setae), longer L (103–114 vs 78–84), W (97–108 vs 71–80), ISD (56–63 vs 41–44), Ti I (188–207 vs 149–163), Ti III (264–287 vs 213–227), IP (2533–2692 vs 2041–2100).

Iguatonia Haitlinger, 2004

. Iguatonia barboproxima sp. nov.

BB63528E-EA2F-5BBD-B8AD-E7FFC9B27022

https://zoobank.org/B7AA6D5B-ADD7-4F98-9245-1789A345EFD3

Figs 5 , 6 , 7 , 8

Figure 5.

Figure 5.

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Iguatoniabarboproxima sp. nov., larva A dorsal view of idiosoma B ventral view of idiosoma C dorsal view of coxa I, showing supracoxal seta.

Figure 6.

Figure 6.

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Iguatoniabarboproxima sp. nov., larva A scutum B ventral view of gnathosoma C dorsal view of the cheliceral bases.

Figure 7.

Figure 7.

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Iguatoniabarboproxima sp. nov., larva photograph. Scutum, showing the shape of scutum, ASE and PSE.

Figure 8.

Figure 8.

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Iguatoniabarboproxima sp. nov., larva. Leg I A trochanter—genu B tibia—tarsus; leg II C trochanter—gen D tibia—tarsus; leg III E trochanter—genu F tibia—tarsus.

Diagnosis (larva).

ASE and PL located in posterior half of scutum; ASE and PSE with fine barbs on distal halves; two pairs hypostomalae barbed; ISD 30–42.

Description.

Idiosoma almost oval, with 32 (fD = 32–34 in paratypes) barbed setae, a pair of setae located between scutum and eyes at level with PSE bases (Fig. 5A). Scutum about trapezoid outline with rounded angles, wider than long, anterior margin concave, lateral margins arcuate obviously, posterior margin convex in median and with small concave between bases of PSE (Figs 6A, 7). Scutum with three pairs of normal setae (AL, ML and PL) and two pairs of sensilla (ASE and PSE). AL, ML and PL completely barbed, AL slightly shorter than PL, and ML longer than both, PW > MW > AW (Table 3). ASE and PSE with setules in distal half, ASE bases posterior to PL bases, PL placed in posterior half of scutum, PSE near posterior border of scutum and longer than ASE (Fig. 6A, 7).

Table 3.

Metric and some meristic data of Iguatoniabarboproxima sp. nov. (larvae, a–c = paratypes; a: 2024–1767-CQ-wx, b: 2025–1767-CQ-wx, c: 2026–1768-CQ-wx).

Character Holotype a b c SD range
fD 32 34 32 32 0.87 32–34
fV 12 12 12 12 0.00 12–12
NDV 44 46 44 44 0.87 44–46
IL 1928 1998 1672 1733 134.32 1672–1998
IW 1213 1320 1366 1276 56.45 1213–1366
DS 46–77 42–70 47–78 48–77 2.28–3.20 42–78
DS min. 46 42 47 48 2.28 42–48
DS max. 77 70 78 77 3.20 70–78
PDS 54–77 53–70 51–78 52–77 1.12–3.20 51–78
PDS min. 54 53 51 52 1.12 51–54
PDS max. 77 70 78 77 3.20 70–78
Oc 20 20 21 23 1.22 20–23
1a 24 28 25 26 1.48 24–28
2a 32 37 34 37 2.12 32–37
3a 33 31 34 34 1.22 31–34
1b 54 63 55 65 4.82 54–65
2b1 42 40 41 44 1.48 40–44
2b2 40 39 40 39 0.50 39–40
3b1 40 39 40 42 1.09 39–42
3b2 34 34 37 36 1.30 34–37
L 103 100 109 106 3.35 100–109
W 124 123 137 130 5.59 123–137
AW 72 70 81 77 4.30 70–81
MW 91 88 98 97 4.15 88–98
PW 99 98 109 104 4.39 98–109
MA 60 59 63 62 1.58 59–63
AA 14 15 16 16 0.83 14–16
SB 14 14 18 17 1.79 14–18
ISD 30 34 42 37 4.38 30–42
AP 47 47 56 60 5.68 47–60
AL 56 53 61 59 3.03 53–61
ML 70 68 76 71 2.95 68–76
PL 59 57 64 62 2.69 57–64
ASE 44 46 46 45 0.83 44–46
PSE 55 59 66 61 3.96 55–66
as 18 16 15 20 1.92 15–20
bs 37 35 37 41 2.18 35–41
cs 32 34 31 35 1.58 31–35
PaScFed 81 76 77 80 2.06 76–81
PaScGed 41 37 46 40 3.24 37–46
GL 136 146 137 141 3.94 136–146
Ta I (H) 18 19 19 19 0.43 18–19
Ta I (L) 159 154 147 157 4.55 147–159
Ti I 167 166 163 166 1.50 163–167
Ge I 135 133 131 135 1.66 131–135
TFe I 83 82 78 78 2.28 78–83
BFe I 89 90 89 91 0.83 89–91
Tr I 50 51 56 51 2.35 50–56
Cx I 73 66 67 63 3.63 63–73
Ta II (H) 19 16 18 18 1.09 16–19
Ta II (L) 150 147 143 152 3.39 143–152
Ti II 178 169 165 174 4.92 165–178
Ge II 132 130 127 132 2.05 127–132
TFe II 74 74 72 79 2.59 72–79
BFe II 91 86 88 89 1.80 86–91
Tr II 60 56 57 57 1.50 56–60
Cx II 81 86 78 79 3.08 78–86
Ta III (H) 17 16 17 19 1.09 16–19
Ta III (L) 165 164 154 163 4.39 154–165
Ti III 252 251 246 254 2.95 246–254
Ge III 150 144 139 147 4.06 139–150
TFe III 115 117 118 111 2.68 111–118
BFe III 117 120 121 114 2.74 114–121
Tr III 67 61 56 54 5.02 54–67
Cx III 80 86 86 77 3.90 77–86
Leg I 756 742 731 741 8.90 731–756
Leg II 766 748 730 762 14.10 730–766
Leg III 946 943 920 920 12.30 920–946
IP 2468 2433 2381 2423 31.01 2381–2468
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Venter. All ventral setae, including coxalae, barbed and with pointed ends (Fig. 5B). Dorsum of coxa I with a peg-like supracoxal seta (elc I) (Fig. 5C). Three pairs of intercoxal setae (1a, 2a and 3a), 1a posterior to level of posterior edge of coxae I, 2a between coxae II, and 3a at a line with anterior edges of coxae III. 2a and 3a subequal and both slightly longer than 1a (Table 3). Five pairs of coxalae (1b, 2b1, 2b2, 3b1 and 3b2), 1b longest, 2b1, 3b1, and 2b2 subequal and all slightly longer than 3b2 (Table 3). 12 setae behind coxae III (fV = 12 in paratypes).

Gnathosoma (Fig. 6B) with a pair of nude galealae (cs), two barbed anterior hypostomalae (as) and two posterior hypostomalae with barbs on proximal half, bs slightly longer than cs, and both longer than as (Table 3). Hypostomal lip fimbriated. Cheliceral bases punctate on the dorsal surface (Fig. 6C). Palpfemur and palpgenu, each with one barbed, pointed dorsal seta. Palptibia with one nude ventral seta, one barbed ventral seta, and one barbed dorsal seta, odontus bifid. Palptarsus with seven setae, three barbed, two nude, one solenidion and one eupathidium. fPp = 0-B-B-2BN2-3B2Nωζ. Palpal supracoxal seta (elcp) peg-like.

Legs (Figs 5B, 8) with seven segments (femora divided). IP = 2381–2468 (Holotype and three paratypes) (Table 3). Anterior and posterior claws hook-like, subequal in length, and anterior claw with few ciliations. Claw-like empodium falciform, longer and slenderer than lateral claws. Normal setae on legs barbed and pointed. Leg setal formula: Leg I: Cx—1n; Tr—1n; Bfe—4n; Tfe—5n; Ge—1σ, 1κ, 12n; Ti—2φ, 1κ, 1Cp, 18n; Ta—1ω, 1ε, 2ζ, 1Cp, 29n. leg II: Cx—2n; Tr—1n; Bfe—4n; Tfe—5n; Ge—1κ, 12n; Ti—2φ, 19n; Ta—1ω, 1ζ, 30n. leg III: Cx—2n; Tr—1n; Bfe—2n; Tfe—5n; Ge—12n; Ti—1φ, 19n; Ta—1ζ, 30n.

Etymology.

The specific epithet of the new species refers to the posterior hypostomalae, which exhibit fine barbs on their proximal half.

Material examined.

Holotype China • a larva (2023–1767-CQ-wx); Chongqing Municipality, Wuxi County, Shuangyang Town; 31°31'29"N, 109°50'12"E; 1151 m; 30 Jun. 2022; Yan Jiang leg.; from an unidentified nymph of Cicadellidae (Hemiptera). Paratypes China • two larvae (2024–1767-CQ-wx, 2025–1767-CQ-wx), the same data as the holotype. China • one larva (2026–1768-CQ-wx); Chongqing Municipality, Wuxi County, Shuangyang Town; 31°29'28"N, 109°49'44"E; 1132 m; 30 Jun. 2022; Xiao-Li Xu leg.; from an unidentified nymph of Cicadellidae (Hemiptera).

The holotype and paratypes are deposited in the Institute of Entomology, Guizhou University, Guiyang, China (GUGC).

Remarks.

To date, two species of Iguatonia have been documented based on larvae, one from Brazil and another one from China (Haitlinger 2004; Noei et al. 2024; Xu et al. 2020).

Iguatoniabarboproxima sp. nov. differs from I.barbillae by the shape of scutum (about trapezoid vs quadrilateral), PL location (in posterior half of scutum vs in anterior half of scutum), ASE location (closer to PL than PSE vs far from PL and near PSE), longer ISD (30–42 vs 10), Ti I (163–167 vs 76–78), and Ti III (246–254 vs 116–126) and differs from I.xinfengi by the shape of scutum (about trapezoid vs sub-rounded), shape of hypostomalae (barbed vs nude), positions of ASE (far from PL and closer to PL than PSE vs almost at the same line with PL), longer ISD (30–42 vs 20–21), Ti I (163–167 vs 114–117), and Ti III (246–254 vs 194–197).

Discussion

Considering previously published data and the present study, 13 species of Caeculisoma have been documented based on the larval stage and one species was recorded based on both the larval and post-larval instars. Of them, only five species (C.brazilensis Noei & Šundić, 2024; C.carmenae Haitlinger, 2008; C.nestori Haitlinger, 2004; C.hunanica Zheng, 2002; and C.sparnoni Southcott, 1972) are without host records (Southcott 1972; Zheng 2002b; Haitlinger 2004, 2008; Kohansal et al. 2024; Noei et al. 2024). The hosts of Caeculisoma larvae were recorded in Insecta, comprising three orders (Lepidoptera, Orthoptera, Hemiptera), and seven families (Acrididae, Cicadellidae, Cicadidae, Delphacidae, Geometridae, Issidae, Miridae) (Southcott 1961, 1972, 1988; Stroiński et al. 2013, Xu et al. 2019a, b, 2020; Kohansal et al. 2024). Only C.pouyani Noei & Kohansal, 2024 has a host distribution across three families (Acrididae, Cicadellidae, Miridae) in two orders (Hemiptera, Orthoptera); and C.penlineatus Xu & Jin, 2019 has a host distribution across three families (Cicadellidae, Delphacidae and Issidae). And the host of each of the remaining species was only recorded in a single family: three species (C.allopenlineatus Xu & Jin, 2020; C.semispinus Xu & Jin, 2019; C.taianensis sp. nov.) with hosts were recorded in Cicadellidae (Hemiptera), two species (C.cooremani Southcott, 1972; C.darwiniense Southcott, 1961) with hosts were recorded in Acrididae (Orthoptera), C.mouldsi Southcott, 1988 with a host was recorded in Cicadidae (Hemiptera), and C.huxleyi Southcott, 1972 with a host was recorded in Geometridae (Lepidoptera). According to the available data, there is a higher probability of finding larvae of the genus Caeculisoma ectoparasitic on Hemiptera compared to Orthoptera and Lepidoptera.

Iguatoniabarbillae Haitlinger, 2004 from an unidentified Homoptera insect (Hemiptera), I.xinfengi Xu & Jin, 2020 from an unknown Delphacidae (Hemipteran) and I.barboproxima sp. nov. from an unidentified Cicadellidae species (Hemiptera) indicate that the host’s spectrum of Iguatonia species is limited to the order Hemiptera.

Only three families of Hemiptera, including Cicadellidae, Delphacidae and Issidae were documented in China for Caeculisoma hosts. A similar situation occurs in the genus Iguatonia, whose hosts are from two families (Cicadellidae, Delphacidae) of Hemiptera.

According to the present information of these two genera the hosts are limited, which may be related to the locations, times, and methods of collection. Therefore, in future studies of their taxonomy, more attention should be paid to geographical ranges and the use of diverse collection methods at different periods in order to know more about the host.

Supplementary Material

XML Treatment for Abrolophus quadrapexicis
XML Treatment for Caeculisoma taianensis
XML Treatment for Iguatonia barboproxima

Acknowledgments

We sincerely thank the editor and reviewers for their time, effort, and valuable suggestions during the review process. Their insightful comments and constructive suggestions greatly improved the quality and clarity of our manuscript.

Citation

Jiang Y, Yi T-C, Xu S-Y, Jin D-C (2025) Two new species of larval Erythraeidae (Parasitengona) ectoparasites of leafhoppers from Southwestern China. ZooKeys 1236: 85–101. https://doi.org/10.3897/zookeys.1236.139274

Funding Statement

National Natural Science Foundation of China

Contributor Information

Si-Yuan Xu, Email: syxuxsy@163.com.

Dao-Chao Jin, Email: dcjin@gzu.edu.cn.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

This work was supported by the National Natural Science Foundation of China (32300373, 32470478).

Author contributions

Data curation: TCY. Investigation: YJ. Resources: SYX. Supervision: DCJ. Writing - original draft: YJ. Writing - review and editing: SYX, DCJ.

Author ORCIDs

Yan Jiang https://orcid.org/0000-0002-8980-5344

Tian-Ci Yi https://orcid.org/0000-0002-9953-3709

Si-Yuan Xu https://orcid.org/0000-0001-6467-6120

Dao-Chao Jin https://orcid.org/0000-0003-2727-5621

Data availability

All of the data that support the findings of this study are available in the main text.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Abrolophus quadrapexicis
zookeys.1236.139274-treatment1.xml (4.7KB, xml)
XML Treatment for Caeculisoma taianensis
zookeys.1236.139274-treatment2.xml (55.3KB, xml)
XML Treatment for Iguatonia barboproxima
zookeys.1236.139274-treatment3.xml (38.3KB, xml)

Data Availability Statement

All of the data that support the findings of this study are available in the main text.

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