# Pronematinae (Trombidiformes: Iolinidae) from Brazil, with original and complementary descriptions of taxa as well as with a reappraisal of the subfamily and of Parapronematus

Raf M J de Vis, José Dantas Araujo Lacerda, Gilberto J de Moraes, Edward

A Ueckermann

► To cite this version:

Raf M J de Vis, José Dantas Araujo Lacerda, Gilberto J de Moraes, Edward A Ueckermann. Pronematinae (Trombidiformes: Iolinidae) from Brazil, with original and complementary descriptions of taxa as well as with a reappraisal of the subfamily and of Parapronematus. Acarologia, 2025, 65 (2), pp.331-372. 10.24349/7w23-xsl9. hal-05059133

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# Pronematinae (Trombidiformes: Iolinidae) from Brazil, with original and complementary descriptions of taxa as well as with a reappraisal of the subfamily and of *Parapronematus*

Raf M. J. De Vis IDª, José Dantas Araujo Lacerda IDᵇ, Gilberto J. de Moraes IDᵇ, Edward A. Ueckermann IDᶜ

## Original research

### ABSTRACT

Pronematinae was originally proposed as a subfamily of Tydeidae Kramer, 1877 and later transferred to Iolinidae Pritchard, 1956. Although apparently omnivorous, some species of this group have been reported as predators of mites. New collection records of Pronematinae from Brazil are presented. The original descriptions of *Peridasilcoferla paraensis* De Vis & Ueckermann **n. gen. n. sp.** and *Parapronematus connarus* De Vis & Ueckermann **n. sp.** are presented. Complementary descriptions of *Parapronematus geminus* Meyer & Rodrigues, 1966 and *Naudea richinda* Meyer & Rodrigues, 1966 are also presented, based on the examination of type specimens. A reappraisal of the subfamily and of *Parapronematus* Baker, 1965, is made based on the specimens collected and on literature information. Updated keys to the world genera of Pronematinae and to the world species of *Parapronematus* as well as suggestions of new generic characteristics to be considered in descriptions of new Pronematinae taxa are presented.

**Keywords** taxonomy; Tydeoidea; mites; opisthosomal setae; aggenital setae

**Zoobank** http://zoobank.org/E317A260-87BA-4FD9-A6D3-C57DABBD396A

In memory of the senior author who passed away in December 2024 before the completion of this article.

### Introduction

An effort has been dedicated in Brazil to the knowledge of the mite species found on plants and in the soil. One of the main objectives is to determine the species potentially useful as biological control agents. Studies have focused mainly on taxa belonging to Phytoseiidae, given their recognized importance as biological control agents (Carrillo *et al.* 2015; Lofego *et al.* 2024).

Little effort has been dedicated to the study of mites of the family Iolinidae Pritchard, 1956 in Brazil (Da Silva *et al.* 2017). Although apparently omnivorous, some species of this family have been reported as predators (De Vis *et al.* 2006a; Hernandes *et al.* 2015; Marcossi *et al.* 2024). The present work refers to species of Pronematinae André, 1980, originally described in Tydeidae Kramer, 1877 and transferred to Iolinidae by André and Fain (2000). Ueckermann

Received 03 November 2024
Accepted 31 March 2025
Published 14 April 2025

Corresponding author
Edward A. Ueckermann ID:
edalbert@lantic.net

Academic editor
Akashi Hernandes, Fabio

https://doi.org/10.24349/7w23-xsl9

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

<figure><img src="image_2.png" /></figure>

De Vis R. M. J. *et al.*

Licensed under
Creative Commons CC-BY 4.0

OPEN ACCESS

<footnote>ªResearch Station for Vegetable Production (PSKW), 2860 Sint-Katelijne-Waver, Belgium.</footnote>

<footnote>ᵇDepartamento de Entomologia e Acarologia, Escola Superior de Agricultura "Luiz de Queiroz", Universidade de São Paulo (ESALQ-USP), 13418-900 Piracicaba, São Paulo, Brazil.</footnote>

<footnote>ᶜUnit for Environmental Sciences and Management, Potchefstroom Campus, North-West University, Private Bag X6001, Potchefstroom, 2520, South Africa.</footnote>
et al. (2024) and De Vis et al. (2024a, b) provided new information about Pronematinae, but no recent publication provides a key for the separation of the nearly 20 genera presently placed in this subfamily. Before that is done, it is necessary to confirm the validity of the synonymy between *Pausia* Kuznetsov & Livshits, 1972 and *Naudea* Meyer & Rodrigues, 1966, proposed by Baker and Delfinado (1976). André (1980) did not accept this synonymy.

*Parapronematus* Baker, 1965, the type genus of Pronematinae, was originally described in Tydeidae and transferred to Iolinidae by André and Fain (2000). A recent unpublished evaluation showed that a species of this genus reported by De Vis et al. (2006b) as *Parapronematus* sp. in southern Brazil and deposited at Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo (ESALQ-USP) was recently collected in northern Brazil and it has still not been reported in the literature. Baker (1965) created *Parapronematus* based on a single species, *P. acaciae* Baker, 1965. Six additional species have been described, not always completely complying with all the reported generic characteristics: *P. geminus* Meyer and Rodrigues, 1966, *P. citri* Salviejo, 1969, *P. formosanus* Tseng, 1985, *P. ferox* Gupta, 1991, *P. camelia* Gupta, 1992 and *P. murshidabadensis* Gupta, 1992 (https://species.wikimedia.org/wiki/Pronematinae, accessed on 11/10/2024, André 2021). The genus has distinctive features facilitating its separation from other genera of the same subfamily: setae *la* absent, three pairs of *ag* setae, dorsal setae of femora III and IV forked and the following leg chaetotaxy (from tarsus to trochanter, solenidia in brackets): leg I (8 (ω) – 2 (φ + κ) – 2 – 3 – 1), leg II (6 (ω) – 2 – 2 – 3 – 0), leg III (5 – 2 – 2 – 2 – 1) and leg IV (5 – 2 – 1 – 1 – 0).

The objectives of this work were: a) to present new records of Pronematinae species from Brazil, describing some new taxa and complementarily describing others; b) to conduct a reappraisal of Pronematinae and *Parapronematus*, presenting keys for the separation of the respective genera and species; c) to suggest new generic characteristics for Pronematinae.

## Material and methods

Samples of leaves, inflorescences and fruits of plants of economic interest as well as of surrounding vegetation were collected by J.D.A. Lacerda in July 2023 in rural communities and urban areas of Parauapebas (about 150-165 m altitude), southeastern municipality of Pará State, in the Amazon region of Brazil. These were transported to the Agricultural and Forestry Entomology Laboratory of Universidade Federal Rural do Amazonas, where mites were collected and preserved in 70% ethanol until mounted in Hoyer's medium (Krantz and Walter 2009) at ESALQ-USP. Those belonging to Iolinidae were sent to PSKW where taxonomic studies were conducted. Additionally, slides with *Parapronematus* specimens reported by De Vis *et al.* (2006a, b) and slides of other Brazilian localities deposited at the Mite Reference Collection of ESALQ-USP were also included in the study.

## Morphological examination and principal component analysis

Specimens were measured using a Leica DM750 microscope with a Leica Flexacam c3 camera and Leica LAS-X software. Line drawings were made with a Leica DM750 equipped with a phototube and Wacom Cintiq 16 pen display and drawn using Adobe Illustrator 2024.

Nomenclature of the body regions, setae and organotaxy is that of Kaźmierski (1989). Leg setae nomenclature is that of André (1981b) and Khaustov *et al.* (2020). The tables of Kaźmierski (1989) were used for the interpretation of the setal nomenclature of different authors. In the text, measurements are given in micrometres, following the name of each structure. A principal component analysis (PCA) based on setal lengths (Ueckermann and De Vis *et al.* 2024) of specimens of *Parapronematus connarus* n. sp. was done with PAST (version 4.03).
## Key to developmental stages and sexes of Pronematinae

As in any other mite group, the characterization of the specimens under analysis initially requires the identification of their developmental stage. In this study, that was done basically with the help of the key provided by André (1980), with some additional information considered relevant. That is important because most descriptions in the literature refer to adult females. Larvae are recognized by having only three pairs of legs, as in most mite groups. Other stages as well as adult sexes were separated according to the following characteristics:

A. With a dehiscence line (breakage line for ecdysis) anteriad the prodorsal setae, curved backwards towards *la* (or *ro* setae when *la* are absent; procurved) ...................... nymph

• Leg IV with tarsus bearing five setae or more and other segments nude; reduced number of ventral idiosomatic setae (lacking *mtβ*, *3c*, *3d* and all *ag* setae) ...................... protonymph

• Leg IV with all segments, except trochanter, bearing setae; venter of idiosoma lacking only *mtβ*, *3d*, *ag3* and either *ag4* (in species with four pairs of *ag* setae) or *ag2* (in species with three pairs of *ag* setae) ......................................................... deutonymph

• Leg IV with all segments, often including trochanter, bearing setae; no idiosomatic or leg setae absent (as in adult) ......................................................... tritonymph

B. Without dehiscence line ......................................................... adult

• Genital opening shaped as an inverted T (exceptionally completely opened, then as a large circle), with 2-4 pairs of *ag* setae ......................................................... female

• Genital opening not shaped as an inverted T; with two rounded dorsal structures laterad *e1*; with stout solenidion extending beyond the tip on tarsus I (in *Pronematus*, *Neopronematus*); with a dorso-distal adaxial spur on femur IV: aedeagus T-shaped (usually indistinguishable), with the same number of *ag* setae as in female or less (only one in *Homeopronematus anconai*), at least in *Parapronematus connarus n. sp.*, with two pairs of eugenital setae; smaller than female ......................................................................................... male

## Estimation of length of structures of previously described species in the review section

As we could not have access to the type specimens of *P. acaciae*, *P. citri* and *P. ferox*, the length of different structures of these species was estimated based on conversion factors calculated from the given values of the length and width of the idiosoma of those species and the length of those structures in the drawings. The calculated values are given in bold italics in Table 1. Additionally, for *P. acaciae*, the estimated length of the tarsus of Figure 84 of the original description was used as basis for the estimation of the length of the setae of tarsus I, depicted in Figure 85.
# Results

## New collection records of Pronematinae from Brazil

Iolinidae Pritchard, 1956

Pronematinae André, 1980

Pronematus Canestrini, 1886

Pronematus brasiliensis Ueckermann & De Vis, 2024

**Specimens examined** — all collected in Parauapebas; 5°59'11.68" S, 49°52'34.73" W, on *Vitis labrusca* L. (MZLQ 4415: 1♂); 6° 4'35.49" S, 49°52'51.54" W, on *Senna occidentalis* (L.) Link (MZLQ 4416: one ♂; 6° 4'13.53" S, 49°54'1.40" W, on *Adenanthera pavonina* L. (MZLQ 4417: one protonymph).

**Remarks** — The following arthropods were found in association with *P. brasiliensis*: on leaves of *A. pavonina*, unidentified *Euseius* Wainstein species (Phytoseiidae) and insects of the order Psocoptera; on *V. labrusca*, the pronematids *Dasilcoferla nadirae* (Da Silva et al.(2017) and *Peridasilcoferla paraensis n. gen. n. sp.*, and unidentified *Amblyseius* Berlese and *Iphiseiodes* De Leon species (Phytoseiidae); on *S. occidentalis*, *Eutetranychus banksi* (McGregor) (Tetranychidae).

## Peridasilcoferla De Vis & Ueckermann n. gen.

Zoobank: A2AB2608-895A-4D9D-9FBB-6BF0B4AAE4F8

## Diagnosis

Adult females with four pairs of setae on aspidosoma, 11 pairs on the opisthosoma, one on trochanters I to III, six on tarsus II and five on each of tarsi III and IV. The leg chaetotaxy is the same as in *Neopronematus* Panou, Emmanuel & Kaźmierski, 2000 and *Dasilcoferla* De Vis & Ueckermann, 2024a, the former differing in having nine and the latter ten pairs of opisthosomal setae.

## Description

Adult female. Aspidosoma procurved with four pairs of setae (bo, ro, la and ex); opisthosoma with 11 pairs of setae (c1, c2, d1, e1, f1, f2, h1, h2, ps1, ps2 and ps3) and four pairs of lyrifissures (ia, im, ip and ih). Eye spots absent. Four pairs of aggenital setae. One pair of genital papillae. Leg I without apotele; each of other legs with ciliate empodium and two claws, without empodial hooks. Seta homologous to seta s (see Ueckermann and De Vis et al, 2024) on tarsus I absent. Femur IV undivided. Leg chaetotaxy (tarsus to trochanter): leg I (8 + ω – 3 + φ + κ – 3 – 3 – 1); leg II (6 + ω – 2 – 3 – 3 – 1); leg III (5 – 2 – 2 – 2 – 1); leg IV (5 – 2 – 1 – 2 – 0). Epimeral formula: 3-1-4-2.

## Etymology

The ancient Greek prefix “peri” means “near”; hence, the *Peridasilcoferla* refers to the similarity between the new genus and *Dasilcoferla*.

## Remark

The new genus differs from *Dasilcoferla* (also described from Brazil) by having setae *ps1* (absent in *Dasilcoferla*). Some other characteristics distinguish the new genus from *Dasilcoferla* as well as from *Neopronematus*, but these are not included in the description of the genus (adopting the model of generic characterization proposed by André, 1980). In *Dasilcoferla*, the eupathidia of tarsus I with fine instead of course barbs in *Peridasilcoferla n. gen.* and *Neopronematus* spp. In all *Neopronematus*, except *N. aegeae* Panou, Emmanuel & Kaźmierski,
2000 (Kuznetsov 1972; Panou *et al.* 2000; Sadeghi *et al.* 2012; Darbemamieh *et al.* 2015), and *Dasilcoferla* (De Vis *et al.* 2024a), the eupathidia are thoroughly barbed whereas in *Peridasilcoferla n. gen.* the distal third is smooth. Conversely, the body and leg setae have heavier barbs in *Dasilcoferla* and in *Neopronematus* (except *N. aegeae*), compared to those in *Peridasilcoferla n. gen.* The placement of the *ag* setae in the new genus is similar to that in *Dasilcoferla* (well separate from each other), while in *Neopronematus* species *ag3* setae are inserted close to each other, at the anterior end of the genital opening and near *ag2*. Additionally, *ag3* is simple in the former two genera but forked in *Neopronematus*, except in *N. aegeae*. With the description of other *Neopronematus* species, *N. aegeae* seems to differ in many respects from those species, suggesting that their generic classification should be re-evaluated.

### Type species

*Peridasilcoferla paraensis* n. gen. n. sp.

### Peridasilcoferla paraensis De Vis & Ueckermann n. gen. n. sp.

Zoobank: A004CFA0-A311-4917-B8E5-DC9E3137F14A

(Figures 1–4)

### Adult female (n = 3)

**Idiosoma** — (Figures 1, 4). Ovaloid, tapering posteriorly. Distance *bo-h1* 134 (127–140), width at level of *c1* 118 (106–131). Completely striate, and striae with tubercles.

**Dorsum** — (Figures 1, 4a-b). Aspidosoma with striation diverging towards the *das* (sejugal) furrow (Kaźmierski 1989); dorsal part of opisthosoma with longitudinal striation between *c1* and *d1*; diverging towards *das*, to form a short but wide spindle-shaped striation pattern between *ro* and *c1*; also diverging posteriorly to form a spindle-shaped pattern between *d1* and *e1*; striation transverse between *f1* and *h1* and inversed V-shaped posteriorly, up to *ps1*, and about transverse between *ps1* and the posterior margin.

Most idiosomal setae aciculate (Figures 4a, b, g, h), apparently serrate, their surface at least with a rough aspect (overlapping with striae makes it unclear); trichobothrial setae (*bo*) on aspidosoma longer than *ex*, which in turn is longer than subequal *la* and *ro*; *la* somewhat thinner than others of subequal thickness (Figures 1, 4a); *bo* finely barbed, except for a smooth proximal section (Figure 4f). Dorsal setae *f2* (Figure 4h) and *h1* longest followed by *c1*, *e1* and *ps2*; other setae shorter, with *ps1* very short. Setae *f1* (Figure 4g), *h1* and *ps1* almost in a straight row and decreasing in length and much shorter than respectively *f2*, *h2* and *ps2*, also in a straight row and decreasing in length. Setae *c1* and *d1* not reaching bases of respectively *d1* and *e1* but *e1* about as long as distance between its base and the base of *f1*. Setal lengths and distances between setae: *bo* 27 (24–28), *bo-bo* 26 (26–27), *ro* 12 (12–13), *ro-ro* 21 (20–21), *la* 15 (14–16), *la-la* 27 (27–28), *ex* 19 (19–20), *c1* 15 (14–15, *c2* 15 (15–16), *c1-c1* 33 (31–35), *d1* 15 (15–16), *d1-d1* 25 (22–27), *e1* 16 (15–17), *e1-e1* 34 (33–35), *f1* 12 (11–13), *f1-f1* 10 (9–10), *f2* 24 (22–26), *h1* 10 (9–11), *h2* 22 (21–23), *ps1* 4 (3–4), *ps2* 14 (13–14).

**Lyrifissure ia** located between setae *c1* and *d1*, slightly laterad hypothetical line connecting the bases of these setae; **lyrifissure im** immediately anteriad *e1*, in hypothetical line between *d1* and *e1*; **lyrifissure ip** immediately anteriad *f2*, slightly laterad hypothetical line between *e1* and *f2*.

**Venter** — (Figures 1, 4 c-d). Striation transverse immediately behind gnathosoma, longitudinal between coxae I and II and between coxae III and IV; striation spindle-shaped between setae *pt* and *mta* and anteriad *ag1*, the former much larger than the latter; striae converging from *ag1* to level of genital opening.

With four pairs of *ag* setae; distances between *ag1-ag1* 11 (11–12), *ag2-ag2* 21 (17–25), *ag3-ag3* 18 (17–19) and *ag4-ag4* 30 (30–31). Setae *ag1* positioned within the spindle-shaped striation pattern. One pair of genital papillae. Seta *ps3* very short, anterolaterad anal opening,
<figure><img src="image_3.png" /><figcaption>**Figure 1** *Peridasilcoferla paraensis* De Vis & Ueckermann **n. gen. n. sp.**, dorsum (left) and venter (right) of holotype female.</figcaption></figure>

without small excrescence in between them (which is present in other genera). Lyrifissure *ih* laterad genital opening.

Setal lengths and distances: *1b* 13 (11–14), *1c* 12 (11–14), *pt* 8 (8–9), *pt-pt* 14 (13–14), *2a* 17 (16–17), *mta* 9 (9–10), *mta-mta* 26 (22–31), *3b* 14 (13–14), *3c* 14 (14–16), *3d* 15 (14–16), *mtß* 8 ± (8–9), *mtß-mtß* 19 (16–24), *4b* 14 (13–15), *ag1* 7 (7–8), *ag2* 7 (7–8), *ag3* 5 (4–5), *ag4* 10 (9–11), *ps3* 4 (4–5).

**Gnathosoma** — (Figure 2). Chelicera (abaxial view) with parallel diagonal striae converging posteriorly; stylet 17 (16–18). Subcapitulum with striae in section anteriad *sc2* almost transverse; posteriad *sc2* diverging posteriorly, except behind palp, where they are transverse; with two pairs of oral setae, *or1* 5 and *or2* 5 and two pairs of scapular setae, *sc1* 8, *sc2* 11. Palp tarsus 7 (7–8) long, with lobed longitudinal striae, bearing distally a lanceolate and smooth eupathidium *pζ* 4 (4–5); setae *l'* and *l''* curved towards eupathidium, seta *v* stout, longer than other setae and finely serrated (barely distinguishable); setae *l'*, *l''* and *d* smooth, *l'* a bit shorter
<figure><img src="image_4.png" /><figcaption>**Figure 2** *Peridasilcoferla paraensis* De Vis & Ueckermann **n. gen. n. sp.** Female. a – Dorsal view of gnathosoma; b – Ventral view of gnathosoma; c – Palptarsus.</figcaption></figure>

than the other two setae, solenidion and *ba* not visible; *l'*, *l''* and *d* inserted at about same level, slightly more proximal than *v*; chaetotaxy from distal to proximal segment 5-1-2. Femorogenu 21 long and 8 in diameter.

**Legs** — (Figure 3). Leg lengths (from base of trochanter to base of claws): I 96 (95–98), II 81 (79–85), III 78 (77–80), IV 88 (83–93), tarsus I length 20 (20–21), tibia I length 19 (18–20). Leg I without pretarsus; each of other legs with ciliate empodium and two claws, without empodial hooks. All leg setae aciculate, finely serrate. Ventral setae of all tibiae stouter than other leg setae, serration hardly visible. Leg I with solenidion ωI not reaching tip of tarsus, eupathidia ft"ζ, tc'ζ, tc"ζ (Figure 4e), p'ζ and p"ζ barbed, except distal third; subequal prorals about 0.6 as long as subequal tectals, and a pair of small forked unguinal setae proximal to prorals on ventral side, seta homologous to *s* absent (see Ueckermann and De Vis *et al.* 2024). Tibia I with bifurcated famulus κ, about half as long as solenidion φI. Femur IV undivided. Setal lengths: **leg I**: ωI 4 (4–5), ft'ζ 11 (11–12), ft"ζ 18 (18–19), tc'ζ 19 (19–20), tc"ζ 20 (19–22), p'ζ 12 (11-12), p"ζ 11 (10–11), *u'* 3 (2–3), *u''* 2 (2–3), φI 3 (2–3), κ" 1 (1–2); **leg II**: ωII 2 (1–2), tibia *v'* 23 (22–23), femur *v''* 18 (16–19); **leg III**: tibia *v'* 21 (20–22), tibia *d* 17 (16–18); **leg IV**: tibia *v'* 21 (19–22), tibia *d* 21 (20–22), femur *ev'* 18 (17–18), femur *d* 17 (16–18).

**Protonymph (n = 1)**

Idiosomal chaetotaxy as in female but *mtβ* and all aggenital setae absent, two setae on coxa III (3d absent) and 4b absent, hence epimeral formula 3-1-3-0. Leg chaetotaxy like female but with no setae on trochanters I and II (instead of one in each), and segments of leg IV nude except for tarsus IV (as in adult female). Hence, protonymph with the following chaetotaxy: I (8+ω-3+k+φ-3-3-0), II (6+ω-2-3-3-0), III (5-2-2-2-1), IV (5-0-0-0-0).

**Male and other immature stages**

Not found.

**Differential diagnosis**

This is the only species in this genus, but differences with related species of other genera are subsequently mentioned. The placement of the *ag* setae is similar to that of the closely related *D. nadirae*, but it differs by having 11 pairs of opisthosomal setae, seta *l''* on palp not sigmoid, setae *p* subequal to each other, *tc* also subequal to each other, setae *tc* about twice as long as setae *p*; eupathidia not completely barbed, spindle-form striae near *ag1* small, only a bit broader
<figure><img src="image_5.png" /><figcaption>**Figure 3** *Peridasilcoferla paraensis* De Vis & Ueckermann **n. gen. n. sp.**, legs of holotype female. a – leg I; b – Leg II; c – Leg III; d – Leg IV.</figcaption></figure>

than distance *ag1-ag1*. It can be distinguished from the closely related *Neopronematus* species
by the presence of 11 pairs of opisthosomals, non-forked *ag3* and the position of *ag* setae.

Type locality and habitat

Collected by J. D. A. Lacerda on July 13, 2023, on leaves of an unidentified *Bauhinia* L.
(Fabaceae) species at Fazenda Vitória (5°58'6.79" S, 49°47'39.71" W, and 166 m altitude),
Palmares II, Parauapebas, Pará State, Brazil.

Type specimens and depository

Holotype female (MZLQ 4382), two paratype females (MZLQ 4383 and MZLQ 4384) and one paratype protonymph (MZLQ 4385) deposited at the Mite Reference Collection of
<figure><img src="image_6.png" /><figcaption>**Figure 4** *Peridasilcoferla paraensis* De Vis & Ueckermann **n. gen. n. sp**: a – Aspidosoma; b – Posterior region of dorsum; c – Anterior region of venter with transversal spindle-shaped striae (within circle); d – Posterior region of venter with transversal spindle-shaped striae (within ellipse) and genital area (bar on each image = 20 µm); e–h setae: e – *te*''ζ, f – *bo*, g – *f1* and h – *f2*.</figcaption></figure>
Departamento de Entomologia e Acarologia, Escola Superior de Agricultura “Luiz de Queiroz” (ESALQ-USP), Universidade de São Paulo (USP), Piracicaba, São Paulo state, Brazil, ESALQ (Zhang 2018).

## Etymology

The species name refers to Pará, the Brazilian state where the type specimens were collected.

## Parapronematus Baker, 1965

### Parapronematus connarus De Vis & Ueckermann n. sp.

Zoobank: 8DEAC753-7D23-4907-B906-A6D7D596FCC6

(Figures 5–11)

**Measurements** — Measurements of different structures of each life stage shown in Table 1.

### Adult female (n = 20)

**Idiosoma** — (Figures 5, 8). Broad, pyriform, tapering posteriorly. Completely covered with fine tuberculate striae, less discernible than in other pronematid genera. Aspidosoma with three pairs of setae (bo, ro and ex; la absent) and opisthosoma with nine pairs of setae (c1, c2, d1, e1, f1, f2, h1, h2 and ps2; ps1 and ps3 absent); lyrifissures ia, im, ip and ih present, but mostly not discernible in adult female, most distinguishable in adult males and immatures.

<figure><img src="image_7.png" /><figcaption>**Figure 5** *Parapronematus connarus* De Vis & Ueckermann n. sp. Female: dorsum (left) and venter (right).</figcaption></figure>
<figure><img src="image_8.png" /><figcaption>**Figure 6** *Parapronematus connarus* De Vis & Ueckermann **n. sp.** Female: a – Dorsal view of gnathosoma; b – Ventral view of gnathosoma; c – Palptarsus.</figcaption></figure>

**Dorsum** — (Figures 5, 8a-b). Aspidosoma with diverging striation towards *das furrow* (Kaźmierski 1989); dorsal part of opisthosoma with longitudinal striation between *c1* and *d1*; diverging towards *das* to form a short but wide spindle-shaped striation pattern between *ro* and *c1*; also diverging posteriorly to form a spindle-shaped pattern between *d1* and *e1*; striation transverse between *f1* and *h1* and inverted V-shaped posteriorly.

Idiosomal setae aciculate and distinctly serrate (Figures 5, 8a-b, 9c-d); trichobothrial setae *bo* (Figures 5, 9b) on aspidosoma longer than *ex*, which is in turn longer than *ro*; *ex* and *ro* stouter than *bo* (Figures 1, 4b); *bo* finely barbed, barbs longer in distal half (Figure 9b). Dorsal setae *f2* (Figure 9d) and *h2* longest, followed by *f1* (Figure 9c) and *h1*, and then *c2* and *ps2*; *c1* shortest. Setae *c1*, *d1* and *e1* not reaching bases of respectively *d1*, *e1* and *f1*, but *f1* passing base of *h1*.

Lyrifissures not always distinguishable, *ia* located between setae *c2* and *d1*; *lyrifissure im* immediately anteriad *e1*; *lyrifissure ip* immediately anteriad *f2*.

**Venter** — (Figures 5, 8c-d). Striae transverse between coxae I; concave outwards in the region of *pt* setae; about longitudinal medially between levels of *mta* and *4b*; spindle-shaped between setae *pt* and *mta* and anteriad *ag1*; converging posteriorly to level of genital opening, around which it is radiating and behind of which it is about transverse.

With three *ag* setae. One pair of genital papillae, only occasionally distinct. *Lyrifissure ih* between genital opening and seta *ps2*.

**Gnathosoma** — (Figure 6). Chelicera with parallel diagonal striae in dorsal view. Striae in section anteriad *sc2* of subcapitulum almost transverse, posteriad *sc2* almost longitudinal medially, diverging posteriorly elsewhere; with two pairs of oral setae (*or1* 4 and *or2* 4) and two subcapitular setae (*sc1* 10, stout and barbed and *sc2* 13, thin and smooth). Palp tarsus long and narrow, with lobed longitudinal striae, bearing distally a slightly forked eupathidium *pζ*; setae *l'* and *l''* curved towards eupathidium; seta *d* distally forked; seta *v* stout and barely longer than eupathidium; all setae smooth, solenidion *ω* and *ba* not distinguishable; *l'* inserted more proximal than *d*, which is more proximal than *l''*, which is about at the same level as *v*; chaetotaxy (5-1-2). Femorogenu 21 long and 12 in larger diameter.

**Legs** — (Figure 7). All leg setae aciculate, finely serrate with dorsal setae, in general, stouter and longer than ventral setae of the same leg segment (although less clearly on leg III). Ventral setae on tibiae gradually varying from stout and long on tibia I to fine and short on tibia IV. Leg I with solenidion *ω* not reaching tip of tarsus, eupathidia *f1*"ζ, *tc*"ζ, *tc"ζ*, *p*"ζ and *p"ζ* barbed for their entire length, except for smooth tip 2 µm, tectals subequal, prorals shorter and unequal, with *p"ζ* the shortest, and a pair of small forked unguinal setae proximal to prorals on ventral side, seta homologous to *s* absent (see Ueckermann and De Vis *et al.* 2024). Tibia I with
<figure><img src="image_9.png" /><figcaption>**Figure 7** *Parapronematus connarus* De Vis & Ueckermann **n. sp.** legs: a–d female: a – Leg I; b – Leg II; c – Leg III with variation on the number of setae on tarsus III (five or six setae); d – Leg IV; e – Male, leg IV.</figcaption></figure>

bifurcated famulus κ but this not always visible, and tip can also appear as a knot (bifurcation not visible), about half as long as solenidion φ. Tarsus III with six setae, but two specimens with only five (specimens 113 and 112 in Table 1; probably shortest, most distal *ft'* absent and no setal base visible); these two specimens lack the other leg III, so double checking is not possible. Femur IV undivided. Dorsal setae *d* on both femora III and IV forked.

Male (n = 7) (Figures 7, 10)

**Idiosoma** — Most characters as in female, except: dorsal setae shorter, serrate; with one pair of rounded structures of 11 µm in diameter posterolaterad *el*, genital opening with two pairs of eugenital setae. Femora III and IV each with forked seta relatively shorter than in female. Femur IV with a small and pointed dorso-distal adaxial spur inserted on a thumb-like process (Figure 7).
<figure><img src="image_10.png" /><figcaption>**Figure 8** *Parapronematus connarus* De Vis & Ueckermann **n. sp.**: a – Aspidosoma; b – Posterior region of dorsum of idiosoma; c – Anterior region of venter of idiosoma with transverse spindle-shaped striae (within ellipse); d – Posterior region of venter with transverse spindle-shaped striae (within ellipse) and genital area; e–h *Parapronematus geminus* Meyer & Rodrigues: e – Aspidosoma; f – Posterior region of dorsum of idiosoma; g – Anterior region of venter of idiosoma, transverse spindle-shaped striae not visible; h – posterior region of venter, transversal spindle-shaped striae not visible and genital area. Scalebar for a–d = 20 µm.</figcaption></figure>
<figure><img src="image_11.png" /><figcaption>**Figure 9** Different setae of *Parapronematus connarus* De Vis & Ueckermann **n. sp.**: a – *tc"ζ*, b – *bo*, c – *f1*, d – *f2*; and *Parapronematus geminus* Meyer & Rodrigues, 1966: e – *tc"ζ*, f – *bo*, g – *f1*, h – *f2*.</figcaption></figure>

**Tritonymph (n = 5) (Figure 11)**

Dehiscence line anteriad *bo*, *ro* and *ex*, projecting toward *ro*; idiosomal and leg chaetotaxy as in adults. Lyrifissures clearly visible. Epimeral formula as in female. With longitudinal invagination connecting genital and anal openings. With distal excrescence, inserted anteriad anal opening and bending over it. Seta *ft'* on tarsus III present. Dorsal setae on femur III not forked or with one very short branch (1 µm); femur IV with dorsal setae not forked.

**Deutonymph (n = 1)**

Dehiscence line as in tritonymph. Idiosomal chaetotaxy as in female but with only one *ag* seta instead of three. Epimeral formula as in female. With longitudinal invagination connecting genital and anal openings. With distal excrescence as in tritonymph. Leg chaetotaxy as in female, but with trochanter I nude instead of with one seta; with one dorsal seta on tarsus III instead of two (*ft*' lacking); and setae on femora III and IV not forked. Hence, leg chaetotaxy as follows: leg I (8 + ω – 2 + φ + κ – 2 – 3 – 0); leg II (6 + ω – 2 – 2 – 3 – 0); leg III (5 – 2 – 2 – 2 – 1); leg IV (5 – 2 – 1 – 1 – 0).
<figure><img src="image_12.png" /><figcaption>**Figure 10** *Parapronematus connarus* De Vis & Ueckermann **n. sp.**: dorsum (left) and venter (right) of male.</figcaption></figure>

**Protonymph (n = 2)**

Dehiscence line as in tritonymph. Idiosomal chaetotaxy as in female, but without *ag* setae, with only two setae on coxa III and without setae *mtβ* and *4b*, hence epimeral formula 3-1-3-0. With longitudinal invagination connecting genital and anal opening. With distal excrescence as in tritonymph. Leg chaetotaxy as in female but trochanter I nude instead of with one seta, with only five setae on tarsus III instead of six, and leg IV segments nude except tarsus, as in female. Hence, protonymph with the following chaetotaxy: leg I (8 + ω – 2 + φ + κ – 2 – 3 – 0), leg II (6 + ω – 2 – 2 – 3 – 0); leg III (5 – 2 – 2 – 2 – 1); leg IV (5 – 0 – 0 – 0 – 0).

**Larva (n = 2)**

Idiosomal chaetotaxy as female but all *ag* setae absent and coxa III with only one seta (*3c* and *3d* absent); epimeral formula 3-1-2. Leg chaetotaxy as in female, but with no unguinal setae (*u*), trochanter III nude instead of with one seta. Thus, chaetotaxy: leg I (6 + ω – 2 + φ + κ – 2 – 3 – 0), leg II (6 + ω – 2 – 2 – 3 – 0), leg III (5 – 2 – 2 – 2 – 0). Tarsus I with small pad like empodium, without claws (*om*), setae *p* and *ft* not eupathidial, setae *tcζ* eupathidial.
<figure><img src="image_13.png" /><figcaption>**Figure 11** *Parapronematus connarus* De Vis & Ueckermann **n. sp.** Tritonymph: dorsum (left) with dehiscence line anterior to aspidosomal setae; venter (right), arrow indicates distal excrescence bending over the anal opening.</figcaption></figure>

**Differential diagnosis**

This species differs from *P. geminus* by having three setae on aspidosoma instead of four; from *P. formosanus* by having three setae on each of femora I and II instead of two; from *P. citri* by having dorsal setae *ro*, *ex*, *c1*, *c2*, *d* and *e2* longer than 13 µm; from *P. acaciae* by having setae *f1* 30 ± 0,9 (25–41) 19, *h1* 33 ± 0,7 (28–43), *h2* 56 ± 0,6 (51–63) shorter and *p''ζ* 42 ± 0,6 (38–47), instead of *f1*≈49, *h1*≈49, *h2*≈68 and *p''ζ*≈34; and from *P. camelia*, *P. ferox* and *P. murshidabadensis* by having *tc''* /tarsus I > 3 instead of < 2 (see Table 3).

**Remarks**

*Parapronematus connarus* De Vis & Ueckermann **n. sp.** shows great variability in relation to the number of setae on tarsus III and some setal lengths. Two specimens (112 and 113) have five setae (*fi*' absent) on tarsus III instead of six. Together with specimen 121, these have longer setae *ro*, *c2*, *f1*, *f2*, *h1*, *3d*, *ag2* and *ag3* than the other specimens of the same new species (Table 1), with specimen 122 (MZLQ 4386) showing other aberrations. Still, other characteristics of these specimens were the same as in other specimens. The bold setae of specimens 112 (MZLQ
4387), 113 (MZLQ 4388) and 121 (MZLQ 4404) do not differ from the holotype. Therefore, we do not consider these three specimens to belong to a different species. Baker (1965) and André (1980) were not clear on that, mentioning the presence of five or six setae on tarsus III in species of this genus. Additionally, although the differences in setal lengths might support the thesis that they belong to a different species, the two specimens with five setae on tarsus III were found together with three specimens with six setae on a single host plant. Seta *ft'* appears during ontogeny in the tritonymph.

The seta on trochanter I and the second dorsal seta on tarsus III also appear in the tritonymph of *Homeopronematus*, *Quasihomeopronematus* (André 1980; De Vis and Ueckermann 2024b), *Pseudopronematulus* (De Vis and Ueckermann 2024a) and *Proctotydaeus* (André 1980). This seems to be a general pattern.

## Type specimens and depository

Parauapebas, Pará, J.D. Lacerda, on *Connarus* sp., July 2023, holotype female (MZLQ 4386); on *Clusia rosea*, July 2023, 5 paratype females (MZLQ 4387–4391); on *Eucalyptus* spp., July 2023, paratype female (MZLQ 4392), 2 paratype deutonymphs (MZLQ 4393 and MZLQ 4394); on *Andropogon gayanus*, July 2023, paratype female (MZLQ 4395); on *Arrabidaea chica*, July 2023, paratype female (MZLQ 4396); on *Musa paradisiaca*, July 2023, paratype male (MZLQ 4397); on *Sida rhombifolia*, July 2023, paratype tritonymph (MZLQ 4398); on *Stachytarpheta* sp., July 2023, paratype female (MZLQ 4399); on *Connarus* sp., July 2023, paratype male (MZLQ 4400); on *Syzygium malaccense*, July 2023, 2 paratype males (MZLQ 4401 and MZLQ 4402). Piracicaba, São Paulo State, R.M.J De Vis, on *Hevea brasiliensis*, July, 2002, paratype male (MZLQ 4403); on *H. brasiliensis*, April-May, 2003, 4 paratype females (MZLQ 4404, MZLQ 4405, MZLQ 4406, MZLQ 4407, MZLQ 4408), 2 paratype larvae (MZLQ 4409 and MZLQ 4410), 2 paratype protonymphs (MZLQ 4409), 3 paratype tritonymphs (MZLQ 4404, MZLQ 4411 and MZLQ 4412). Pontes e Lacerda, Mato Grosso State, N.J. Ferla, on *H. brasiliensis*, date unknown, paratype female (MZLQ 4413); *H. brasiliensis*, September-October 1999, 2 paratype females (MZLQ 5658 and MZLQ 5659), paratype male (MZLQ 5659). Itambé, Pernambuco, M.G.C. Gondim Jr., on *Acrocomia intumescens*, July 1998, paratype female (MZLQ 4414), paratype tritonymph (MZLQ 4473). All types deposited at the Mite Reference Collection, ESALQ-USP, Piracicaba, São Paulo State, Brazil.

## Etymology

The species is named after *Connarus*, the genus of the host plant of the holotype.

## PCA analysis

The characteristics that define the major differences between the specimens are setae *f1*, *f2*, *h1*, *3d*, *ag3*, *tc'*ζ, *tc''ζ* and *p'*ζ (Figure 12A, Table 1). Specimens 112, 113 and 121 stand out from the other specimens (Figure 12B).

## Reappraisal of *Parapronematus*

Most of the descriptions of species of this group are rather incomplete. Several of them do not have measurements and none indicates the variability of measurements. Most do not provide scale bars in illustrations. Characters of doubtful value have been used to distinguish species while characters of recognizable value are not always considered. Although complementary descriptions should be done, in this work we could only redescribe *P. geminus*, as in most cases the museums in which they are deposited do not allow their shipment.

None of the descriptions mentions lyrifissures. However, in our specimens, lyrifissures could be seen in most males and all immatures, despite not always visible in adult females.

Similarly, striae are very fine and indistinct in part of the body of several species. We suppose that they are present but not always visible. In our specimens, digital augmentation shows striae in specimens where striae are not distinguishable under phase contrast microscopic
<figure><img src="image_14.png" /><figcaption>**Figure 12** Principal component analysis (PCA) of measurements of *Parapronematus connarus* De Vis & Ueckermann **n. sp.** females. A. Loading of component one for different characters. B. Scatter plot of 20 individual females (including 112: MZLQ 4387, 113: MZLQ 4388, 121: MZLQ 4404).</figcaption></figure>

examinations at 1000 X. It seems that this characteristic is influenced by mounting conditions. We therefore do not consider this a reliable characteristic for distinguishing species based on publications of distinct authors (as used for the characterization of *P. citri*, e.g.).

As the species are not sclerotized, distances between setae are influenced by the state of the mite when mounted, mounting conditions, as the amount of medium used and the 'pressure' on the specimens, the presence of folds, etc. These aspects will lead to inaccurate measurements of distance between setae. Thus, we also consider this characteristic inadequate to separate species.

Likewise, length of the dorsal setae on femora III and IV in relation to the width of the segment seems a characteristic of questionable value. We believe that the width or height of
<table><caption>**Table 1** Length and distances between dorsal and ventral setae of *Parapronematus connarus* De Vis & Ueckermann **n. sp.** stages (mean, min-max). Aberrant values of specimens 112 (MZLQ 4387), 113 (MZLQ 4388) and 121 (MZLQ 4404) are given in bold. Specimen 122 (MZLQ 4386) is the holotype.</caption><thead><tr><th rowspan="2"></th><th colspan="4">Female</th><th rowspan="2">Male</th><th rowspan="2">Tritonymph</th><th rowspan="2">Deutonymph</th><th rowspan="2">Protonymph</th><th rowspan="2">Larva</th></tr><tr><th>All</th><th>Spc. 112</th><th>Spc. 113</th><th>Spc. 121</th><th>Spc. 122</th></tr></thead><tbody><tr><td>bo-h1</td><td>190 (170-231)</td><td>191</td><td>198</td><td>182</td><td>188</td><td>134 (126-146)</td><td>156 (146-172)</td><td>112</td><td>108 (97-120)</td><td>89 (88-89)</td></tr><tr><td>width (at level c1)</td><td>184 (155-212)</td><td>198</td><td>194</td><td>168</td><td>196</td><td>124 (109-138)</td><td>151 (147-158)</td><td>133</td><td>110 (103-116)</td><td>102 (101-103)</td></tr><tr><td>bo</td><td>44 (39-50)</td><td>41</td><td>39</td><td>50</td><td>40</td><td>33 (32-35)</td><td>42 (40-44)</td><td>39</td><td>36 (35-37)</td><td>33 (33-34)</td></tr><tr><td>bo-bo</td><td>52 (46-62)</td><td>58</td><td>62</td><td>51</td><td>56</td><td>37 (33-45)</td><td>46 (40-50)</td><td>44</td><td>37 (37-38)</td><td>29 (29-30)</td></tr><tr><td>ro</td><td>18 (16-22)</td><td><strong>22</strong></td><td><strong>22</strong></td><td><strong>20</strong></td><td>16</td><td>10 (8-12)</td><td>11 (9-13)</td><td>117</td><td>9 (8-9)</td><td>9 (8-10)</td></tr><tr><td>ro-ro</td><td>27 (23-36)</td><td>34</td><td>36</td><td>28</td><td>28</td><td>19 (16-21)</td><td>24 (21-26)</td><td>23</td><td>21 (19-22)</td><td>16 (15-16)</td></tr><tr><td>ex</td><td>23 (20-27)</td><td>27</td><td>25</td><td>26</td><td>23</td><td>13 (10-15)</td><td>15 (13-17)</td><td>14</td><td>11 (10-11)</td><td>12 (11-12)</td></tr><tr><td>c1</td><td>21 (18-29)</td><td>21</td><td>21</td><td><strong>29</strong></td><td>19</td><td>12 (9-15)</td><td>15 (13-17)</td><td>17</td><td>13 (13-14)</td><td>14 (13-14)</td></tr><tr><td>c2</td><td>22 (18-26)</td><td><strong>26</strong></td><td><strong>24</strong></td><td><strong>26</strong></td><td>18</td><td>9 (8-11)</td><td>14 (13-16)</td><td>13</td><td>11 (11-12)</td><td>14 (13-16)</td></tr><tr><td>c1-c1</td><td>52 (39-66)</td><td>48</td><td>49</td><td>48</td><td>54</td><td>33 (31-35)</td><td>41 (36-46)</td><td>35</td><td>34 (34-35)</td><td>35 (34-37)</td></tr><tr><td>d1</td><td>21 (19-26)</td><td>23</td><td>22</td><td><strong>26</strong></td><td>20</td><td>13 (11-16)</td><td>15 (13-17)</td><td>15</td><td>12 (12-13)</td><td>12 (11-13)</td></tr><tr><td>d1-d1</td><td>51 (34-60)</td><td>55</td><td>59</td><td>53</td><td>60</td><td>32 (31-33)</td><td>40 (38-43)</td><td>34</td><td>31 (31-32)</td><td>32 (31-32)</td></tr><tr><td>e1</td><td>21 (19-26)</td><td>22</td><td>20</td><td><strong>26</strong></td><td>19</td><td>13 (10-15)</td><td>15 (13-19)</td><td>15</td><td>11 (10-11)</td><td>10 (10-11)</td></tr><tr><td>e1-e1</td><td>42 (36-63)</td><td>43</td><td>49</td><td>42</td><td>43</td><td>25 (22-27)</td><td>30 (28-34)</td><td>26</td><td>22 (21-22)</td><td>24 (24-25)</td></tr><tr><td>f1</td><td>30 (25-41)</td><td><strong>39</strong></td><td><strong>41</strong></td><td><strong>35</strong></td><td>30</td><td>19 (17-21)</td><td>19 (16-22)</td><td>17</td><td>13 (13-14)</td><td>11 (11-12)</td></tr><tr><td>f1-f1</td><td>36 (29-45)</td><td>44</td><td>45</td><td>37</td><td>34</td><td>24 (24-26)</td><td>26 (23-33)</td><td>20</td><td>17 (16-17)</td><td>12 (12-13)</td></tr><tr><td>f2</td><td>73 (67-81)</td><td><strong>81</strong></td><td><strong>77</strong></td><td><strong>79</strong></td><td>70</td><td>47 (45-52)</td><td>52 (49-55)</td><td>45</td><td>34 (33-344)</td><td>26 (24-28)</td></tr><tr><td>h1</td><td>33 (28-43)</td><td><strong>43</strong></td><td><strong>39</strong></td><td><strong>38</strong></td><td>32</td><td>22 (21-24)</td><td>22 (19-26)</td><td>19</td><td>14 (14-15)</td><td>11 (10-11)</td></tr><tr><td>h2</td><td>56 (51-63)</td><td>58</td><td>56</td><td><strong>63</strong></td><td>51</td><td>34 (32-37)</td><td>36 (35-37)</td><td>19</td><td>21 (20-22)</td><td>15 (14-15)</td></tr><tr><td>ps2</td><td>24 (21-29)</td><td><strong>29</strong></td><td><strong>28</strong></td><td>25</td><td>21</td><td>16 (14-18)</td><td>17 (15-21)</td><td>19</td><td>9 (8-9)</td><td>4 (3-4)</td></tr><tr><td>lb</td><td>18 (17-20)</td><td>19</td><td>17</td><td>19</td><td>18</td><td>13 (12-15)</td><td>13 (12-13)</td><td>12</td><td>9 (9-10)</td><td>9 (9-10)</td></tr><tr><td>lc</td><td>8 (6-10)</td><td>9</td><td>7</td><td>8</td><td>6</td><td>6 (5-7)</td><td>6 (5-9)</td><td>5</td><td>4 (4-5)</td><td>5 (4-5)</td></tr><tr><td>pt</td><td>11 (10-13)</td><td>11</td><td>11</td><td>13</td><td>11</td><td>9 (8-10)</td><td>8 (7-9)</td><td>6</td><td>6 (5-6)</td><td>5 (5-6)</td></tr><tr><td>pt-pt</td><td>16 (11-21)</td><td>16</td><td>21</td><td>15</td><td>17</td><td>13 (11-16)</td><td>16 (15-17)</td><td>14</td><td>14 (14-15)</td><td>15 (14-15)</td></tr><tr><td>2a</td><td>21 (17-23)</td><td>22</td><td>21</td><td>23</td><td>21</td><td>15 (12-16)</td><td>14 (12-16)</td><td>12</td><td>11 (10-12)</td><td>10 (10-11)</td></tr><tr><td>mta</td><td>9 (6-11)</td><td>9</td><td>11</td><td>10</td><td>9</td><td>7 (6-8)</td><td>6 (5-8)</td><td>5</td><td>4 (4-5)</td><td>4 (4-5)</td></tr><tr><td>mta-mta</td><td>31 (22-42)</td><td>33</td><td>37</td><td>30</td><td>22</td><td>22 (18-25)</td><td>26 (219-31)</td><td>25</td><td>23 (22-24)</td><td>24 (23-25)</td></tr><tr><td>3b</td><td>20 (17-22)</td><td>22</td><td>19</td><td>20</td><td>19</td><td>13 (11-14)</td><td>13 (11-15)</td><td>12</td><td>9 (9-10)</td><td>10 (9-10)</td></tr><tr><td>3c</td><td>32 (28-36)</td><td>33</td><td>30</td><td>35</td><td>30</td><td>18 (15-20)</td><td>22 (20-24)</td><td>22</td><td>15 (15-16)</td><td>Absent</td></tr><tr><td>3d</td><td>26 (21-32)</td><td><strong>32</strong></td><td><strong>31</strong></td><td><strong>32</strong></td><td>25</td><td>16 (14-17)</td><td>18 (13-21)</td><td>14</td><td>Absent</td><td>Absent</td></tr><tr><td>mtβ</td><td>7 (5-9)</td><td>8</td><td>9</td><td>7</td><td>6</td><td>5 (4-7)</td><td>5 (4-5)</td><td>4</td><td>Absent</td><td>Absent</td></tr><tr><td>mtβ-mtβ</td><td>21 (17-26)</td><td>20</td><td>-</td><td>19</td><td>22</td><td>15 (13-18)</td><td>16 (15-20)</td><td>19</td><td>Absent</td><td>Absent</td></tr><tr><td>4b</td><td>15 (13-18)</td><td>17</td><td>-</td><td>17</td><td>13</td><td>11 (9-14)</td><td>10 (8-12)</td><td>8</td><td>Absent</td><td>Absent</td></tr><tr><td>ag1</td><td>7 (6-9)</td><td>8</td><td>9</td><td>7</td><td>6</td><td>5 (5-,6)</td><td>4 (4-6)</td><td>4</td><td>Absent</td><td>Absent</td></tr><tr><td>ag1-ag1</td><td>17 (13-24)</td><td>18</td><td>24</td><td>15</td><td>15</td><td>9 (9-10)</td><td>11 (8-14)</td><td>11</td><td>Absent</td><td>Absent</td></tr><tr><td>ag2</td><td>8 (7-12)</td><td><strong>12</strong></td><td><strong>11</strong></td><td>8</td><td>7</td><td>5 (4-6)</td><td>5 (3-7)</td><td>absent</td><td>Absent</td><td>Absent</td></tr><tr><td>ag2-ag2</td><td>29 (14-47)</td><td>22</td><td>24</td><td>29</td><td>30</td><td>20 (18-22)</td><td>20 (18-22)</td><td>absent</td><td>Absent</td><td>Absent</td></tr><tr><td>ag3</td><td>15 (12-22)</td><td><strong>22</strong></td><td><strong>21</strong></td><td>14</td><td>20</td><td>8 (8-10)</td><td>7 (2-10)</td><td>absent</td><td>Absent</td><td>Absent</td></tr><tr><td>ag3-ag3</td><td>37 (22-65)</td><td>23</td><td>22</td><td>34</td><td>45</td><td>28 (24-32)</td><td>21 (18-24)</td><td>absent</td><td>Absent</td><td>Absent</td></tr></tbody></table>

femora are also influenced by the mounting conditions. So, any parameter comparing the length of the dorsal setae with the width of the segment may be inaccurate and should preferably not be used to separate species. This type of problem seems of lesser importance in relating the length of eupathidia and length of tarsus I. Hence, in this publication the ratio between the lengths of *tc"ζ* and of tarsus I is taken into account in the separation of species.

The length of individual setae, even on the same species, can differ significantly, so relative length can also vary, for practical reasons. The tips of the setae can break off and that may be overlooked. We therefore recommend measuring setae of Pronematinae at 1000 X for higher confidence on measurements.

André (1979: 204) considered the description of a new genus based on a single specimen to be unadvisable, except when the specimen exhibits very special characteristics. We propose to extend this suggestion to the description of new species. For Phytoseiidae, Tixier (2012) concluded that measurement of at least ten specimens is necessary to express variability reliably. We, therefore, suggest describing new taxa only when several specimens are available, unless the specimen exhibits special characteristics assumed to be of lower variability.

The position of the solenidion on tarsus I is used as a distinctive character in the *Paraprone-
<table><caption>**Table 1** Continued.</caption><thead><tr><th rowspan="2"></th><th colspan="4">Female</th><th colspan="2">Male</th><th rowspan="2">Deutonymph</th><th rowspan="2">Protonymph</th><th rowspan="2">Larva</th></tr><tr><th>All</th><th>Spc. 112</th><th>Spc. 113</th><th>Spc. 121</th><th>Spc. 122</th><th>Tritonymph</th></tr></thead><tbody><tr><td>eu1</td><td>Absent</td><td>absent</td><td>absent</td><td>absent</td><td>absent</td><td>3 (3-4)</td><td>Absent</td><td>Absent</td><td>Absent</td></tr><tr><td>eu2</td><td>Absent</td><td>absent</td><td>absent</td><td>absent</td><td>absent</td><td>4 (3-5)</td><td>Absent</td><td>Absent</td><td>Absent</td></tr><tr><td>palptarsus</td><td>23 (20-26)</td><td>24</td><td>24</td><td>26</td><td>21</td><td>16 (14-18)</td><td>20 (19-22)</td><td>13</td><td>13 (12-13)</td><td>12 (12-13)</td></tr><tr><td>palp p'ζ</td><td>6 (4-8)</td><td>8</td><td>7</td><td>6</td><td>7</td><td>4 (4-5)</td><td>5 (5-6)</td><td>3</td><td>3 (3-4)</td><td>3 (3-4)</td></tr><tr><td>stylet</td><td>13 (11-15)</td><td>11</td><td>14</td><td>14</td><td>14</td><td>10 (10-11)</td><td>12 (10-12)</td><td>10</td><td>10 (10-11)</td><td>9 (8-9)</td></tr><tr><td>leg I</td><td>142 (129-154)</td><td>140</td><td>139</td><td>147</td><td>141</td><td>110 (100-123)</td><td>118 (106-132)</td><td>84</td><td>68 (67-70)</td><td>67 (66-68)</td></tr><tr><td>leg II</td><td>121 (112-134)</td><td>116</td><td>114</td><td>128</td><td>122</td><td>99 (94-112)</td><td>108 (101-118)</td><td>83</td><td>66 (65-66)</td><td>64 (64-65)</td></tr><tr><td>leg III</td><td>119 (109-147)</td><td>125</td><td>113</td><td>147</td><td>124</td><td>89 (82-101)</td><td>109 (100-125)</td><td>77</td><td>70 (68-71)</td><td>69 (67-70)</td></tr><tr><td>leg IV</td><td>123 (111-135)</td><td>127</td><td>111</td><td>131</td><td>127</td><td>95 (86-107)</td><td>108 (91-129)</td><td>76</td><td>65 (63-67)</td><td>Absent</td></tr><tr><td>tarsus I length</td><td>24 (23-26)</td><td>24</td><td>23</td><td>24</td><td>23</td><td>21 (18-24)</td><td>22 (20-24)</td><td>18</td><td>16 (15-16)</td><td>17 (16-17)</td></tr><tr><td>tibia I length</td><td>22 (20-25)</td><td>22</td><td>20</td><td>24</td><td>22</td><td>18 (17-20)</td><td>19 (18-21)</td><td>14</td><td>12 (12-13)</td><td>12 (11-12)</td></tr><tr><td>ωI</td><td>5 (4-6)</td><td>4</td><td>4</td><td>6</td><td>4</td><td>4 (3-5)</td><td>3 (3-4)</td><td>2</td><td>2 (1-2)</td><td>2 (2-3)</td></tr><tr><td>ft'ζ</td><td>12 (9-16)</td><td>9</td><td>9</td><td>16</td><td>-</td><td>9 (7-11)</td><td>10 (8-11)</td><td>8</td><td>6 (6-7)</td><td>6 (6-7)</td></tr><tr><td>ft"ζ</td><td>69 (63-73)</td><td>69</td><td>69</td><td>-</td><td>63</td><td>539 (49-61)</td><td>44 (38-52)</td><td>29</td><td>11 (10-11)</td><td>9 (9-10)</td></tr><tr><td>tc'ζ</td><td>72 (67-79)</td><td>67,5</td><td>67</td><td>-</td><td>-</td><td>56 (47-63)</td><td>51 (47-59)</td><td>41</td><td>31 (31-32)</td><td>22 (21-23)</td></tr><tr><td>tc"ζ</td><td>70 (65-76)</td><td>66,8</td><td>65</td><td>-</td><td>-</td><td>55 (51-60)</td><td>49 (44-55)</td><td>39</td><td>32 (32-33)</td><td>30 (29-31)</td></tr><tr><td>p'ζ</td><td>51 (44-58)</td><td>46,7</td><td>47</td><td>-</td><td>48</td><td>39 (36-45)</td><td>34 (31-40)</td><td>26</td><td>19 (19-20)</td><td>7 (6-7)</td></tr><tr><td>p"ζ</td><td>42 (38-47)</td><td>39,1</td><td>40</td><td>-</td><td>38</td><td>31 (26-345)</td><td>25 (23-32)</td><td>20</td><td>16 (16-17)</td><td>6 (6-7)</td></tr><tr><td>u'</td><td>4 (2-5)</td><td>3</td><td>3</td><td>4</td><td>2</td><td>3 (2-5)</td><td>3 (2-3)</td><td>4</td><td>3 (2-3)</td><td>Absent</td></tr><tr><td>u"</td><td>4 (2-5)</td><td>3</td><td>3</td><td>3</td><td>2</td><td>3 (2-4)</td><td>2 (2-3)</td><td>3</td><td>3 (2-3)</td><td>Absent</td></tr><tr><td>φI</td><td>3 (2-4)</td><td>3</td><td>3</td><td>3</td><td>2</td><td>2 (2-3)</td><td>3 (2-4)</td><td>-</td><td>-</td><td>2 (1-2)</td></tr><tr><td>κ"</td><td>2 (2-3)</td><td>3</td><td>2</td><td>3</td><td>-</td><td>2 (1-2)</td><td>2 (1-2)</td><td>2</td><td>-</td><td>2 (2-3)</td></tr><tr><td>ωII</td><td>3 (2-4)</td><td>3</td><td>2</td><td>3</td><td>2</td><td>2 (2-3)</td><td>1 (1-2)</td><td>13</td><td>2 (1-2)</td><td>1 (1-2)</td></tr><tr><td>tibia II v'</td><td>32 (30-34)</td><td>31</td><td>30</td><td>34</td><td>31</td><td>24 (22-25)</td><td>24 (22-27)</td><td>22</td><td>18 (17-18)</td><td>16 (15-17)</td></tr><tr><td>tibia II d</td><td>19 (17-22)</td><td>18</td><td>19</td><td>22</td><td>18</td><td>12 (11-13)</td><td>13 (12-15)</td><td>10</td><td>8 (7-8)</td><td>7 (6-7)</td></tr><tr><td>femur II v"</td><td>36 (31-40)</td><td>37</td><td>39</td><td>40</td><td>33</td><td>25 (23-28)</td><td>26 (24-27)</td><td>23</td><td>17 (16-18)</td><td>17 (16-17)</td></tr><tr><td>femur II d</td><td>25 (23-29)</td><td>24</td><td>25</td><td>27</td><td>23</td><td>14 (12-17)</td><td>20 (18-23)</td><td>15</td><td>13 (12-13)</td><td>13 (12-13)</td></tr><tr><td>tarsus III ft'</td><td>10 (8-11)</td><td>-</td><td>-</td><td>10</td><td>9</td><td>8 (7-8)</td><td>7 (5-9)</td><td>absent</td><td>Absent</td><td>Absent</td></tr><tr><td>tarsus III ft"</td><td>19 (17-21)</td><td>19</td><td>17</td><td>20</td><td>17</td><td>14 (14-15)</td><td>14 (13-15)</td><td>11</td><td>9 (9-10)</td><td>10 (9-10)</td></tr><tr><td>tibia III v'</td><td>20 (18-23)</td><td>23</td><td>20</td><td>22</td><td>19</td><td>15 (13-16)</td><td>15 (15-16)</td><td>10</td><td>9 (8-9)</td><td>8 (8-9)</td></tr><tr><td>tibia III d</td><td>16 (13-17)</td><td>16</td><td>16</td><td>17</td><td>13</td><td>11 (9-13)</td><td>12 (11-13)</td><td>9</td><td>8 (8-9)</td><td>8 (8-9)</td></tr><tr><td>femur III ev'</td><td>31 (28-34)</td><td>34</td><td>32</td><td>32</td><td>28</td><td>21 (21-23)</td><td>22 (21-24)</td><td>18</td><td>15 (15-16)</td><td>13 (12-13)</td></tr><tr><td>femur III d</td><td>36 (32-40)</td><td>36</td><td>35</td><td>40</td><td>34</td><td>17 (13-20)</td><td>30 (27-32)</td><td>27</td><td>22 (22-23)</td><td>20 (20-21)</td></tr><tr><td>long fork</td><td>16 (10-21)</td><td>16</td><td>14</td><td>21</td><td>16</td><td>7 (3-10)</td><td>11 (8-15)</td><td>absent</td><td>Absent</td><td>Absent</td></tr><tr><td>short fork</td><td>12 (8-15)</td><td>9</td><td>11</td><td>15</td><td>10</td><td>5 (4-8)</td><td>2 (1-3)</td><td>absent</td><td>Absent</td><td>Absent</td></tr><tr><td>tibia IV v'</td><td>16 (14-22)</td><td>16</td><td>-</td><td>22</td><td>17</td><td>12 (9-13)</td><td>12 (11-14)</td><td>9</td><td>Absent</td><td>Absent</td></tr><tr><td>tibia IV d</td><td>17 (16-20)</td><td>18</td><td>19</td><td>17</td><td>16</td><td>12 (11-14)</td><td>13 (11-15)</td><td>10</td><td>Absent</td><td>Absent</td></tr><tr><td>femur IV d</td><td>39 (34-44)</td><td>38</td><td>34</td><td>44</td><td>36</td><td>15 (13-18)</td><td>33 (26-36)</td><td>24</td><td>Absent</td><td>Absent</td></tr><tr><td>long fork</td><td>19 (14-25)</td><td>17</td><td>15</td><td>25</td><td>15</td><td>8 (6-9)</td><td>7 (10-15)</td><td>absent</td><td>Absent</td><td>Absent</td></tr><tr><td>short fork</td><td>14 (10-18)</td><td>13</td><td>12</td><td>18</td><td>13</td><td>5 (4-6)</td><td>3 (6-9)</td><td>absent</td><td>Absent</td><td>Absent</td></tr></tbody></table>

matus key of Gupta (2002). However, this is not always clearly presented in illustrations of different species.

The number of setae on tarsi III and IV varies between species, so it could be considered a reliable feature to separate species. However, in *P. connarus* n. sp., we found two specimens with five setae on tarsi III and IV instead of six in the majority of the specimens. This characteristic is not always clearly stated in the descriptions, and we see discrepancies between the text of descriptions and the respective illustrations (e.g. *P. acaciae* Baker, 1965) and in all descriptions but that of *P. formosanus*, leg figures are unclear or incomplete.

It is concluded that a revision of the whole genus is necessary, starting with a redescription of its type species and the creation of eventual new genera according to the generic concept of André (1980). Types of *P. acaciae* can only be studied in the museum where they are deposited. Thus, in this publication we selected characteristics that allow the separation of the species, although many uncertainties remain concerning other characteristics. A provisional re-characterization of *Parapronematus* is subsequently provided, based on what is available in the literature, the examination of the type specimens of the new species here described (*P. connarus* n. sp.) and the complementary description of *P. geminus*, provided in this publication.
Additionally, a discussion about the other species placed in the genus is presented, concerning their main characteristics and the limitation of the information available in the literature.

**Diagnosis of *Parapronematus***

Adult females with three pairs of setae on the aspidosoma (la lacking, except in one species), nine pairs on the dorsum of the opisthosoma (ps1 and ps3 lacking), three pairs of ag, three setae on tibia I and two on genu I instead of respectively four and three in all other pronematid genera. Two setae on genu II, trochanter II nude, and one seta on femur IV. Only two other genera have three ag setae, but they have different leg I setation. Aberrations of some species concerning the type species: one species with four setae (la present) on aspidosoma, three of the seven species with five setae on tarsus III instead of six (the species described below has five or six setae on tarsus III), one species with two setae on each of femora I and II instead of three.

**Complementary description**

Adult female. Aspidosoma procurved, with 3-4 pairs of setae (bo, ro and ex; la absent in all but one species); opisthosoma with nine pairs of setae (c1, c2, d1, e1, f1, f2, h1, h2 and ps2; ps1 and ps3 absent) and four pairs of lyrifissures (ia, im, ip and ih). Eye spots absent. Three pairs of ag setae. One pair of genital papillae. Leg I without pretarsus; each of other legs with ciliate empodium and two claws, without empodial hooks. Seta homologous to s on tarsus I absent. Femur IV undivided. Leg chaetotaxy (tarsus to trochanter; aberrations from type species in bold): leg I (8 + ω - 2 + φ + κ - 2 - 2 or 3 - 1); leg II (6 + ω - 2 - 2 - 2 or 3 - 0); leg III (5 or 6 - 2 - 2 - 2 - 1); leg IV (5 - 2 - 1 - 1 - 0). Epimeral formula 3-1-4-2.

**Remarks**

According to the concept of André (1980), new genera should be defined for: i) species having setae *la*; ii) species having two setae on each of femora I and II; and iii) species having five setae on tarsus III. However, to describe a new genus, a full description of the type species is needed. A redescription of *P. geminus* (*seta la* present) is presented underneath based on two paratypes; but more specimens are needed to determine variability. Type specimens of other species could not be studied, and the respective descriptions are sometimes ambiguous, with contradictions between the text and the illustrations, including the type species of *Parapronematus*. So, we prefer not to create new genera, given the impossibility of making a complete description of the type species, adapting the generic descriptions of Baker (1965) and André (1980). The latter author mentioned 5-6 setae on tarsus IV in his diagnosis of the genus. None of the descriptions of *Parapronematus* species mentions tarsus IV with six setae. Hence, in the complementary description of this genus in this publication, the variation of six setae on tarsus IV is not mentioned, despite being detected in some of the specimens of the new species described in this work, as this could be just a case of aberration.

We suppose that the following setae are absent in *Parapronematus* (not mentioned in original descriptions): tibia I, *d*; genu I, *v*'; tarsus II *tc"*; genu II, *v*'; trochanter II, *v*'; tarsus III, *ft* '(shortest seta); tarsus IV, *ft* '(shortest seta); and femur IV, *ev*'.

**Type species**

*Parapronematus acaciae* Baker 1965.

**Parapronematus acaciae Baker, 1965**

*Parapronematus acaciae* Baker, 1965: 116.

*Parapronematus acacia*, Gupta 2002: 121.

The original description of this species is extremely basic, with no measurements or details in the illustration. In the complementary description of Gupta (2002), he used part of the original description and all drawings of the original description, with no new measurements.
The type specimens could not be studied because they could not be loaned; hence, in this
publication, the length of setae of the dorsal part of opisthosoma and of tarsus I were estimated
(Table 2) based on the illustrations provided in the original description.

We found the following inconsistencies between the description and the illustrations: tarsus III is depicted with five setae, instead of six in the description of the genus. Also, according to the original description, "The forked seta of femur IV is at least twice as long as the width of the segment", but in the illustration, the length of the seta and the width of the segment are similar. For the moment, we retain the genus description with six setae on tarsus III and the estimated setal lengths as distinguishing characteristics and we consider the relative length of the dorsal setae of femora III and IV to the width of the segment as an unreliable characteristic. However, we are convinced that the species needs a redescription, especially for being the type of the genus.

As distinguishing characters, we retain: *f1* 49, *h1* 49 and *h2* 68, which are clearly longer than in *P. camelia*, *P. formosanus*, *P. murshidabadensis* and *P. connarus n. sp*. Eupathidia long, *tc"ζ/tarsus I≈3.0* but seta *p"ζ* 34 is shorter than in *P. connarus n. sp*. (37.5–47.3 µm), tarsus III with six and tarsus IV with five setae, according to the text of the original description of the genus.

**Parapronematus camelia Gupta, 1992**

*Parapronematus camelia* Gupta, 1992: 126.

*Parapronematus camelliae*, Gupta 2002: 121.

The description of this species is basic and lacks many measurements and detailed figures. We see inconsistencies between the textual description and the illustrations: Figure 58 shows a decrease in length of setae *f2*, *h2*, *f1* and *h1*; seta *ps2* is not depicted, while in the description the lengths of these setae are *D4* (*f1*) = 31, *L2* (*f2*) 56, *D5* (*h1*) 36 and *L3* (*h2*) 23. It is unclear which setal nomenclature is followed. On basis of the figure, we would switch the length of setae mentioned in the text (*h*) to *h1* 23 and *h2* 36. Figure 59 seems to be leg I instead of palp and the limit between tarsus and tibia as well as between femur and trochanter are not depicted, nor are the solenidia. In the text, "*ωI* thick and *φI* very small", but neither of them is illustrated. We suppose that *φI* has been confounded with *κ*. So, we cannot use the form of *φI* as a distinguishing parameter.

As distinguishing characters, we retain: *f2* short (56 µm); seta *c2* stout and lanceolate, other setae thin and pointed; eupathidia intermediate, *tc"*/tarsus *I≈2.2*; tarsus III with six and tarsus IV with five setae, according to the illustration of the original description.

In the original description, the species was named *Pronematus camelia* (the designation *camelia* is a name in apposition). In his monograph, Gupta (2002) changed the name to *P. camelliae*. We believe that that change is not justified.

**Parapronematus citri Salviejo, 1969**

*Parapronematus citri* Salviejo, 1969: 272.

The description of this species is also basic and lacks measurements, although the illustrations are rather detailed, showing some important characteristics, as follows: setae *ro*, *ex*, *c1*, *c2*, *d1* and *e1* very short (less than half as long as *f1*), thick and lanceolate; *f1* about 2/3 as long as *h1*; *f2* slightly over twice as long as *f1*; *h2* about 1.2 times as long as *h1* (Table 1). The number of setae on tarsus IV is mentioned in the text to be "as for the genus" (five setae), whereas the illustration seems to show six setae (not totally clear). We suppose it to be five.

We consider to be distinctive characteristics of this species: setae *ro*, *ex*, *c1*, *c1*, *d1* and *e1* 11–13, shorter than all other setae; eupathidia long (*tc"*/*tarsus *I≈3.0*); tarsi III and IV respectively with six and five setae.
**Parapronematus ferox Gupta, 1991**

*Parapronematus ferox* Gupta, 1991: 231.

*Parapronematus ferox*, Gupta, 2002: 123.

The description of this species is very basic and lacks most measurements and detailed
illustrations. The long *f1* in relation to *f2* in Figure 58 of the original description is noteworthy.
Double checking it would be convenient; if correct, this characteristic would be sufficient to
distinguish this from all other species. The lengths of the dorsal setae of femora III and IV were
not provided, but they seem to be longer than in other species. There is a contradiction between
the descriptive text and the illustration concerning the number of setae on tarsus III; in the text
it is mentioned to be the same as reported for the genus, while the illustration shows both tarsi
III and IV with five setae.

Distinguishing characteristics: *f1* almost twice as long as *f2* (the opposite in all other species); setae *ro, ex, c1, c2, d1* stout and lanceolate; other dorsal setae on opisthosoma, thin and pointed; eupathidia short (*tc"*/*tarsus I*≈1.5); tarsi III and IV each with five setae (based on the illustration of the original description).

**Parapronematus formosanus Tseng, 1985**

*Parapronematus formosanus* Tseng, 1985: 85.

The description of this species is detailed, missing few measurements, although information
on variability was not provided. The illustrations are detailed, though without scale bars. Striae
are shown on dorsum and venter of the idiosoma. The ventral setae are illustrated in detail,
varying from the very broad and leaflike *pt* to the lanceolate *mtβ*.

Calls attention the statement “legs strongly sclerotized, brown in colour”. As far as we
know no other sclerotised species have been reported in Iolinidae. Two inconsistencies were
detected between the descriptive text and the illustrations: in the differential diagnosis with *P.*
*acaciae*, femora III and IV are mentioned to have two pairs of setae each. We suppose that this
should refer to femora I and II, as described in the table with the number of setae on the legs
and the depicted number of setae in the illustrations. Secondly, the text mentions the presence
of striae in a diamond pattern anteriad *ag1*, which seems to be correct, while the pattern is
illustrated in Figure 130 anteriad *mtβ*.

As distinguishing characteristics, we retain: setae *ro, ex, c1, c2, d1* and *e1* stout and lanceolate, other dorsal setae on opisthosoma long, thin and pointed; eupathidia of intermediate length (*tc"*/*tarsus I*≈2.2); two setae on each of femora I and II (instead of the usual three); tarsi III and IV each with five setae; setae *pt, mta* and *mtβ* barbed, from leaflike to lanceolate.

**Parapronematus geminus Meyer & Rodrigues, 1966**

*Parapronematus geminus* Meyer & Rodrigues, 1966: 21.

(Figures 8g-h, 13–15)

**Adult female (n = 2)**

**Measurements** — Measurements of different structures are shown in Table 2.

**Idiosoma** — (Figure 13). Ovaloid, tapering posteriorly. Completely covered with fine striae with tiny tubercles, not clearly visible. Aspidosoma with four pair of setae (*bo, ro, la* and *ex*), of which *la* very short, and opisthosoma with nine pairs of setae (*c1, c2, d1, e1, f1, f2, h1, h2* and *ps2; ps1* and *ps3* absent).

**Dorsum** — (Figures 8e-f, 13). Striae barely visible, in parts of the dorsum. Aspidosoma with striae longitudinal medially, progressively more divergent towards *das* furrow laterally. Opisthosoma with median longitudinal striation, divergent towards *das* furrow, but only patches of striae visible, divergent between levels of *d1* and *e1*, transverse between setae *e1* and *h1* and then divergent towards posterior margin.
<figure><img src="image_15.png" /><figcaption>**Figure 13** *Parapronematus geminus* Meyer & Rodrigues 1966: dorsum (left) and venter (right) of paratype female.</figcaption></figure>

Idiosomal setae acute and clearly serrate (Figures 8e-f, 9g-h, 13); trichobothrial seta (bo) longer than ex, which is in turn longer than ro; bo barbed (Figures 8e, 9b, 13); seta la very short and smooth. Setae f2 (Figures 9h, 13) and h2 longest followed by f1 (Figures 9g, 13) and h2, and then ps2, d1 and c1, with c2 shortest. Setae c1, d1 and e1 not reaching bases of d1, e1 and f1, but f1 longer than distance between its base and the base of h1. Seta ps1 absent.

**Venter** — (Figures 8g-h, 13). Area immediately posteriad gnathosoma with transverse striation; striation longitudinal between setae *pt* and between setae *mta* and *mtβ*; two areas of spindle-shaped striation pattern between *pt* and *mta* and immediately anteriad *ag1*.

With three pairs of *ag* setae anterolaterad anal opening; genital papillae not distinguishable.
Seta *ps3* absent. Lyrifissure *ih* not distinguishable.

**Gnathosoma** — (Figure 14).

With two pairs of scapular setae, *sc1* 8 (8) and *sc2* 10 (12), both serrate and slender; setae *or1* and *or2* not distinguishable; striae in section anteriad *sc1* almost transverse; in section between *sc1* and *sc2* converging anteromedially. Palptarsus elongate, with a distal forked eupathidium *pζ*; setae *l'* and *l''* curved towards eupathidium; seta *d* forked at tip; *l'* the most proximal, followed by seta *d*; setae *l''* and *v* at about same levels; all setae smooth and of similar thickness; solenidia *ω* and *ba* not distinguishable, chaetotaxy (5-1-2). Femorogenu length 20
<figure><img src="image_16.png" /><figcaption>**Figure 14** *Parapronematus geminus* Meyer & Rodrigues 1966. Female: ventral view of gnathosoma with palp of paratype female.</figcaption></figure>

(23), width 11 (11).

**Legs** — (Figure 15). Chaetotaxy: leg I (8 + ω - 2 + φ + k - 2 - 3 - 1); leg II (6 + ω - 2 - 2 - 3 - 0); leg III (5 - 2 - 2 - 2 - 1); leg IV (5 - 2 - 1 - 1 - 0). Epimeral formula: 3 - 1 - 4 - 2. All non-solenidion leg setae acute, serrate and stout. Leg I with solenidion ω not reaching tip of tarsus, eupathidia *ft"ζ*, *tc'ζ*, *tc"ζ* (Figure 9a), *p'ζ* and *p"ζ* barbed for the entire length, tectals subequal, shorter prorals not as long, with *p"ζ* the shortest; with a pair of small forked unguinal setae proximal to prorals on ventral side; seta homologous to *s* between prorals absent (see Ueckermann and De Vis *et al.* 2024). Tibia I with bifurcated famulus κ but this is not always visible, about same length as solenidion φ. Seta *d* on femur III forked. Femur IV undivided and with seta *d* also forked.

**Adult male (n = 2 paratypes, in poor condition)**

**Idiosoma** — Most characters same as in female. Dorsal setae shorter. Distance *bo-h1* 140, width at level *c1* 134, *bo* 30; *bo-bo* 43; *ro* 12; *ro-ro* 21; *la* 4; *la-la* 38; *ex* 16; *c1* 16; *c1-c1* 36; *c2* 10; *d1* 14; *d1-d1* 31; *e1* 12; *e1-e1* 30; *f1* 21; *f1-f1* 24; *f2* 40; *h1* 23; *h2* 37; *ps2* 17, *lb* 12; *lc* 5; *pt* 9; *pt-pt* not possible to measure; *2a* 16; *mta* 6; *mta-mta* 24, remaining setae cannot be measured.

**Legs** — Femora III and IV with forked setae. Femur IV with a dorso-distal, adaxial pointed spur (Figure 15e). Other measurements: leg I 102; tarsus I 19; tibia I 16; tarsus I setae: *ft'* 8; *ft"ζ* 42; *tc'ζ* 53; *tc"ζ* 48 *p'ζ* 37; *p"ζ* 31.

**Differential diagnosis**

This species closely resembles *P. connarus* n. sp., by having similar setal lengths and shapes. It can be distinguished from *P. connarus* n. sp. and other species of the same genus by having seta *la* and five setae on each of tarsi III and IV.
<figure><img src="image_17.png" /><figcaption>**Figure 15** *Parapronematus geminus* Meyer & Rodrigues, 1966: a–d, legs of female; a – Combination of holotype and paratype leg I; b – Holotype leg II; c – Holotype leg III; d – Holotype leg IV; e – Paratype male leg IV.</figcaption></figure>

Type specimens and depository

Holotype female, from *Gossypium* sp., R. Quetxoaio, Mozambique, April 16, 1964, M.C.
Rodrigues; paratype female, from *Mangifera indica*, Morrumbala, Mozambique, March 25,
1964, M.K. P. Meyer; paratype male, from *Gossypium* sp., Kaidundjua, Mozambique, April
17, 1964, M.C. Rodrigues; paratype male, from shrub, Raraga, Mozambique, March 18, 1964,
M.K.P. Meyer.

Holotype female (accession number Acy 65/30) and paratype female (accession number Acy 65/137), paratype male (accession number Acy 65/37), paratype male (accession number Acy 65/142) at Mite Collection, ARC-Plant Health Protection, Pretoria, South Africa, NCA (Zhang 2018).
## *Parapronematus murshidabadensis* Gupta, 1992

*Parapronematus murshidabadensis* Gupta, 1992: 126.

*Parapronematus murshidabadensis*, Gupta, 2002: 123.

The description of this species is basic, missing many measurements and detailed illustrations. In contrast with the description of *P. camelia*, in the description of *P. murshidabadensis* the relative lengths of the setae in the description coincide with what is shown in the illustration: *D4* (*f1*) 44, *L2* (*f2*)78, *D5* (*h1*) = 40, *L3* (*h2*) 56. The used nomenclature does not coincide with that of Baker (1965). Solenidion ω*I* stout, φ*I* extremely small and blunt. We suppose that in this case φ*I* is confused with κ. As an inconsistency within the text of the original description, it is stated in the key to species that the solenidion on tarsus I is placed in the middle of the segment, whereas in Figure 65 of Gupta (1992) it appears in the distal half.

Distinguishing characteristics: seta *f1* nearly twice as long as *e1* (44 and 21). Setae *f1*, *f2*, *h1* and *h2* (44, 78, 40, 56) longer than in other species and *ft"ζ* relative short, less than half length of *p"ζ* (figure 65 of Gupta, 1992). Setae *ro*, *ex*, *c1*, *d1* and *e1* thick and lanceolate; other dorsal setae of opisthosoma long, thin and pointed (*c2* unclear); eupathidia intermediate; *tc"* / *tarsus* I≈1.7, but tarsus I seems long; tarsi III and IV each with five setae.

## Key to the *Parapronematus* species

Based on the previous considerations, a key is subsequently provided to separate the species of this genus. This should be considered provisional, until a thorough re-examination of type specimens or newly collected specimens is done.

1. Aspidosomal seta *la* present (despite very short, ≈5–7 µm).................................................. *P. geminus*
   — Aspidosomal seta *la* absent.................................................................................... 2

2 (1). Two setae on each of femora I and II; setae *ro*, *ex*, *c1*, *c2*, *d1* and *e1* (narrowly) lanceolate and serrate; setae *pt* *mtα* and *mtβ* barbed, from leaflike to lanceolate.................................................. *P. formosanus*
   — Three setae on each of femora I and II; dorsal and ventral setae thinner.................................................... 3

3 (2). Dorsal setae *ro*, *ex*, *c1*, *c2*, *d1* and *e2* short (11–13 µm).................................................. *P. citri*
   — Dorsal setae *ro*, *ex*, *c1*, *c2*, *d1* and *e2* longer than 13 µm.................................................... 4

4 (3). Ratio *tc"* / *tarsus* I at least 2.9.................................................................................... 5
   — Ratio *tc"* / *tarsus* I at most 2.2.................................................................................... 6

5 (4). Setae *f1* 30 ± 0,9 (25–41) 19, *h1* 33 ± 0,7 (28–43), *h2* 56 ± 0,6 (51–63), and *p"ζ* 42 ± 0,6 (38–47).................................................................................. *P. connarus* De Vis & Ueckermann **n. sp.**
   — Setae *f1*≈**49**, *h1*≈**49**, *h2*≈**68** longer and *p"ζ*≈**34** shorter.................................................. *P. acaciae*

6 (4). Seta *bo*≈31, *f1*≈31 and *h1*≈23; six setae on tarsus III.................................................. *P. camelia*
   — Seta *bo*, *f1* and *h1* at least 40% longer than above; five setae on tarsus III.................................................... 7

7 (6). Seta *f1* twice as long as *f2*; seta *h1*≈**31**.................................................................................. *P. ferox*
   — Seta *f1* slightly shorter than *f2*; seta *h1*≈40.................................................................................. *P. murshidabadensis*

In addition to the key provided, Table 3 brings an overview of selected characteristics to separate the *Parapronematus* species.

## Reappraisal of Pronematinæ

As presently conceived, Pronematinæ is composed of 19 genera, depending upon a definition of the validity of the synonymy of *Pausia* Kuznetsov & Livshitz, 1972 under *Naudea* Meyer
<table><caption>**Table 2** Comparison of *Parapronematus* species female characteristics as described by the respective author(s). Bold-italic values are estimated based on converting factors calculated from the given values of the length and width of the idiosoma of those species and the given lengths of illustrated structures (see Materials & Methods).</caption><thead><tr><th>Characteristic</th><th>P. acaciae<br>Baker (1965)</th><th>P. camelia<br>Gupta (1992, 2002)</th><th>P. citri<br>Salviejo (1969)</th><th>P. ferox<br>Gupta (1991, 2002)</th><th>P. formosanus<br>Tseng (1985)</th><th>P. geminus*<br>This paper</th><th>P. murshidabadensis<br>Gupta (1992, 2002)</th><th>P. connarus n. sp.<br>This paper</th></tr></thead><tbody><tr><td>body length</td><td>280</td><td>224</td><td>265</td><td>266</td><td>326</td><td></td><td>278</td><td></td></tr><tr><td>bo-h1</td><td></td><td></td><td></td><td></td><td></td><td>192 (180)</td><td></td><td>190 (170-231)</td></tr><tr><td>width at level c1</td><td>97</td><td>145</td><td>152</td><td>156</td><td>179</td><td>172 (165)</td><td>168</td><td>184 (155-212)</td></tr><tr><td>bo</td><td><strong>41</strong></td><td>31</td><td><strong>30</strong></td><td>49</td><td>35</td><td>40 (39)</td><td>47</td><td>44 (39-50)</td></tr><tr><td>bo-bo</td><td><strong>51</strong></td><td></td><td><strong>48</strong></td><td><strong>40</strong></td><td></td><td>56 (52)</td><td></td><td>52 (46-62)</td></tr><tr><td>ro</td><td><strong>20</strong></td><td>25</td><td><strong>12</strong></td><td>24</td><td>18</td><td>19 (19)</td><td>22</td><td>18 (16-22)</td></tr><tr><td>ro-ro</td><td><strong>33</strong></td><td></td><td><strong>22</strong></td><td><strong>21</strong></td><td></td><td>33 (31)</td><td></td><td>27 (23-36)</td></tr><tr><td>la</td><td>Absent</td><td>Absent</td><td>Absent</td><td>Absent</td><td>Absent</td><td>7 (5)</td><td>Absent</td><td>Absent</td></tr><tr><td>la-la</td><td>Absent</td><td>Absent</td><td>Absent</td><td>Absent</td><td>Absent</td><td>55 (55)</td><td>Absent</td><td>Absent</td></tr><tr><td>ex</td><td><strong>27</strong></td><td>22</td><td><strong>13</strong></td><td><strong>20</strong></td><td>24</td><td>23 (26)</td><td>24</td><td>23 (20-27)</td></tr><tr><td>c1</td><td><strong>23</strong></td><td>22</td><td><strong>13</strong></td><td><strong>16</strong></td><td>19</td><td>23 (23)</td><td>22</td><td>21 (18-29)</td></tr><tr><td>c2</td><td><strong>23</strong></td><td>22</td><td><strong>11</strong></td><td></td><td></td><td>63 (48)</td><td>24</td><td>22 (19-26)</td></tr><tr><td>c1-c1</td><td><strong>56</strong></td><td></td><td><strong>47</strong></td><td><strong>47</strong></td><td></td><td>22 (21)</td><td></td><td>52 (39-66)</td></tr><tr><td>d1</td><td><strong>23</strong></td><td></td><td><strong>13</strong></td><td><strong>19</strong></td><td>19</td><td>25 (25)</td><td>21</td><td>21 (19-26)</td></tr><tr><td>d1-d1</td><td><strong>51</strong></td><td></td><td><strong>38</strong></td><td><strong>53</strong></td><td></td><td>62 (51)</td><td></td><td>51 (34-60)</td></tr><tr><td>e1</td><td><strong>26</strong></td><td>22</td><td><strong>13</strong></td><td><strong>23</strong></td><td>16</td><td>23 (22)</td><td>21</td><td>21 (19-26)</td></tr><tr><td>e1-e1</td><td><strong>44</strong></td><td></td><td><strong>32</strong></td><td><strong>62</strong></td><td></td><td>42 (39)</td><td></td><td>42 (36-63)</td></tr><tr><td>f1</td><td><strong>49</strong></td><td>31</td><td><strong>27</strong></td><td><strong>82</strong></td><td>40</td><td>34 (38)</td><td>44</td><td>30 (25-41)</td></tr><tr><td>f1-f1</td><td><strong>44</strong></td><td></td><td><strong>29</strong></td><td><strong>36</strong></td><td></td><td>41 (38)</td><td></td><td>36 (29-45)</td></tr><tr><td>f2</td><td>77</td><td></td><td><strong>59</strong></td><td><strong>45</strong></td><td>70</td><td>69 (68)</td><td>78</td><td>73 (67-81)</td></tr><tr><td>h1</td><td><strong>49</strong></td><td>23</td><td><strong>35</strong></td><td><strong>31</strong></td><td>26</td><td>35 (39)</td><td>40</td><td>33 (28-43)</td></tr><tr><td>h2</td><td><strong>68</strong></td><td></td><td><strong>42</strong></td><td><strong>54</strong></td><td>54</td><td>57 (58)</td><td>56</td><td>56 (51-63)</td></tr><tr><td>ps2</td><td><strong>26</strong></td><td></td><td><strong>24</strong></td><td><strong>24</strong></td><td>26</td><td>27 (29)</td><td></td><td>25 (21-29)</td></tr><tr><td>1b</td><td></td><td></td><td></td><td></td><td></td><td>18 (19)</td><td></td><td>18 (17-20)</td></tr><tr><td>1c</td><td></td><td></td><td></td><td></td><td></td><td>6 (7)</td><td></td><td>8 (6-10)</td></tr><tr><td>pt</td><td></td><td></td><td></td><td></td><td>11</td><td>11 (12)</td><td></td><td>11 (10-13)</td></tr><tr><td>pt-pt</td><td></td><td></td><td></td><td></td><td>16</td><td>16 (14)</td><td></td><td>16 (11-21)</td></tr><tr><td>2a</td><td></td><td></td><td></td><td></td><td>13</td><td>19 (21)</td><td></td><td>21 (17-23)</td></tr><tr><td>mta</td><td></td><td></td><td></td><td></td><td>34</td><td>9 (10)</td><td></td><td>9 (6-11)</td></tr><tr><td>mta-mta</td><td></td><td></td><td></td><td></td><td></td><td>34 (32)</td><td></td><td>31 (22-42)</td></tr><tr><td>3b</td><td></td><td></td><td></td><td></td><td></td><td>20 (23)</td><td></td><td>20 (17-22)</td></tr><tr><td>3c</td><td></td><td></td><td></td><td></td><td></td><td>29 (29)</td><td></td><td>32 (28-36)</td></tr><tr><td>3d</td><td></td><td></td><td></td><td></td><td></td><td>26 (24)</td><td></td><td>26 (21-32)</td></tr><tr><td>mtβ</td><td></td><td></td><td></td><td></td><td>7</td><td>7 (7)</td><td></td><td>7 (5-9)</td></tr><tr><td>mtβ-mtβ</td><td></td><td></td><td></td><td></td><td>22</td><td>27 (19)</td><td></td><td>21 (17-26)</td></tr><tr><td>4b</td><td></td><td></td><td></td><td></td><td></td><td>13 (15)</td><td></td><td>15 (13-18)</td></tr><tr><td>ag1</td><td></td><td></td><td></td><td></td><td>7</td><td>6 (7)</td><td></td><td>7 (6-9)</td></tr><tr><td>ag1-ag1</td><td></td><td></td><td></td><td></td><td></td><td>19 (16)</td><td></td><td>17 (13-24)</td></tr><tr><td>ag2</td><td></td><td></td><td></td><td></td><td>7</td><td>8 (8)</td><td></td><td>8 (7-12)</td></tr><tr><td>ag2-ag2</td><td></td><td></td><td></td><td></td><td></td><td>26 (24)</td><td></td><td>29 (14-47)</td></tr><tr><td>ag3</td><td></td><td></td><td></td><td></td><td></td><td>14 (16)</td><td></td><td>15 (12-22)</td></tr><tr><td>ag3-ag3</td><td></td><td></td><td></td><td></td><td>18</td><td>32 (31)</td><td></td><td>37 (22-65)</td></tr></tbody></table>

& Rodrigues, 1966. The latter two genera were synonymized by Baker and Delfinado (1976), who considered them to differ only in the number of opisthosomal setae (ten pairs in *Pausia* and nine in *Naudea*). André (1980) considered both valid, without referring to Baker and Delfinado (1976) and providing a brief characterization of each.

Thus, the structure of this section starts with a characterization of the sole species included in *Naudea*, followed by a comparative characterization of the genera, subgenera (of *Proctotydaeus* Berlese, 1911) and species of this subfamily, ending with the presentation of a key to separate the genera and subgenera.

Our examination of the holotype female, one paratype female and the paratype male of the single known species of *Naudea* led to the following characterization.

**Naudea richinda Meyer & Rodrigues, 1966**

*Naudea richinda* Meyer & Rodrigues, 1966: 23.

*Pausia richinda*, Baker & Delfinado, 1976: 35.
<table><caption>**Table 2** Continued.</caption><thead><tr><th>Characteristic</th><th>P. acaciae<br>Baker (1965)</th><th>P. camelia<br>Gupta (1992, 2002)</th><th>P. citri<br>Salviejo (1969)</th><th>P. ferox<br>Gupta (1991, 2002)</th><th>P. formosanus<br>Tseng (1985)</th><th>P. geminus*<br>This paper</th><th>P. murshidabadensis<br>Gupta (1992, 2002)</th><th>P. connarus n. sp.<br>This paper</th></tr></thead><tbody><tr><td>palptarsus</td><td></td><td></td><td></td><td></td><td>29</td><td>29 (28)</td><td></td><td>23 (20–26)</td></tr><tr><td>palp p'ζ</td><td></td><td></td><td></td><td></td><td></td><td>4 (5)</td><td></td><td>6 (4–8)</td></tr><tr><td>stylet</td><td></td><td></td><td></td><td></td><td></td><td>10 (11)</td><td></td><td>13 (11–15)</td></tr><tr><td>leg I</td><td></td><td></td><td></td><td></td><td></td><td>120 (141)</td><td></td><td>142 (129–154)</td></tr><tr><td>leg III</td><td></td><td></td><td></td><td></td><td></td><td>118 (120)</td><td></td><td>121 (112–134)</td></tr><tr><td>leg III</td><td></td><td></td><td></td><td></td><td></td><td>121 (126)</td><td></td><td>119 (109–147)</td></tr><tr><td>leg IV</td><td></td><td></td><td></td><td></td><td></td><td>127 (131)</td><td></td><td>123 (111–135)</td></tr><tr><td>tarsus I length</td><td>23</td><td></td><td>18</td><td></td><td>29</td><td>23 (24)</td><td></td><td>24 (23–26)</td></tr><tr><td>tibia I length</td><td>22</td><td></td><td>15</td><td></td><td>26</td><td>19 (21)</td><td></td><td>22 (20–25)</td></tr><tr><td>ωI</td><td>5</td><td>Stout</td><td></td><td></td><td></td><td>7 (6)</td><td></td><td>5 (4–6)</td></tr><tr><td>ft'ζ</td><td>12</td><td></td><td>20</td><td></td><td></td><td>9 (12)</td><td></td><td>12 (9–16)</td></tr><tr><td>ft"ζ</td><td>69</td><td></td><td>54</td><td></td><td></td><td>73 (72)</td><td></td><td>69 (63–73)</td></tr><tr><td>tc'ζ</td><td>71</td><td></td><td>67</td><td></td><td></td><td>75 (absent)</td><td></td><td>72 (67–79)</td></tr><tr><td>tc"ζ</td><td>70</td><td></td><td>59</td><td></td><td></td><td>71 (absent)</td><td></td><td>70 (65–76)</td></tr><tr><td>p'ζ</td><td>43</td><td></td><td>47</td><td></td><td></td><td>49 (51)</td><td></td><td>51 (44–58)</td></tr><tr><td>p"ζ</td><td>34</td><td></td><td>31</td><td></td><td></td><td>40 (39)</td><td></td><td>42 (38–47)</td></tr><tr><td>u'</td><td></td><td></td><td></td><td></td><td></td><td>0 (3)</td><td></td><td>4 (2–5)</td></tr><tr><td>u"</td><td></td><td></td><td></td><td></td><td></td><td>2 (3)</td><td></td><td>4 (2–5)</td></tr><tr><td>φI</td><td></td><td></td><td></td><td></td><td></td><td>2 (2)</td><td></td><td>3 (2–4)</td></tr><tr><td>κ"</td><td></td><td></td><td></td><td></td><td></td><td>2 (2)</td><td></td><td>2 (2–3)</td></tr><tr><td>ωII</td><td></td><td></td><td></td><td></td><td></td><td>3 (3)</td><td></td><td>3 (2–4)</td></tr><tr><td>tibia II v'</td><td></td><td></td><td></td><td></td><td></td><td>24 (31)</td><td></td><td>32 (30–34)</td></tr><tr><td>tibia II d</td><td></td><td></td><td></td><td></td><td></td><td>16 (18)</td><td></td><td>19 (17–22)</td></tr><tr><td>femur II v"</td><td></td><td></td><td></td><td></td><td></td><td>39 (37)</td><td></td><td>36 (31–40)</td></tr><tr><td>femur II d</td><td></td><td></td><td></td><td></td><td></td><td>25 (24)</td><td></td><td>25 (23–29)</td></tr><tr><td>tarsus III ft'</td><td>Present</td><td>Present</td><td>Present</td><td>Absent</td><td>Absent</td><td>Absent</td><td>Absent</td><td>10 (8–11)</td></tr><tr><td>tarsus III ft"</td><td></td><td></td><td></td><td></td><td></td><td>17 (18)</td><td></td><td>19 (17–21)</td></tr><tr><td>tibia III v'</td><td></td><td></td><td></td><td></td><td></td><td>18 (18)</td><td></td><td>20 (18–23)</td></tr><tr><td>tibia III d</td><td></td><td></td><td></td><td></td><td></td><td>12 (15)</td><td></td><td>16 (13–17)</td></tr><tr><td>femur III ev'</td><td></td><td></td><td></td><td></td><td></td><td>29 (31)</td><td></td><td>31 (28–34)</td></tr><tr><td>femur III d</td><td></td><td></td><td></td><td></td><td></td><td>33 (26)</td><td></td><td>36 (32–40)</td></tr><tr><td>long part</td><td></td><td></td><td></td><td></td><td></td><td>17–20</td><td></td><td>16 (10–21)</td></tr><tr><td>short part</td><td></td><td></td><td></td><td></td><td></td><td>13–15</td><td></td><td>12 (8–15)</td></tr><tr><td>tibia IV v'</td><td></td><td></td><td></td><td></td><td></td><td>13 (15)</td><td></td><td>16 (14–22)</td></tr><tr><td>tibia IV d</td><td></td><td></td><td></td><td></td><td></td><td>13 (17)</td><td></td><td>17 (16–20)</td></tr><tr><td>femur IV d</td><td></td><td></td><td></td><td></td><td></td><td>32 (37)</td><td></td><td>39 (34–44)</td></tr><tr><td>long part</td><td></td><td></td><td></td><td></td><td></td><td>17–20</td><td></td><td>19 (14–25)</td></tr><tr><td>short part</td><td></td><td></td><td></td><td></td><td></td><td>13–14</td><td></td><td>14 (10–18)</td></tr></tbody></table>

## Adult female

(Figures 18, 19)

**Measurements** — (Holotype followed by paratype, the latter in brackets).

**Idiosoma** — (Figure 16). Elongate-oval. Distance *bo-h1* 137 (140), width at level *c1* 113 (112). Completely covered with fine striae with tiny tubercles, not clearly visible. Aspidosoma with four pairs of setae (*bo*, *ro*, *la* and *ex*) and opisthosoma with ten pairs of setae (*c1*, *c2*, *d1*, *e1*, *f1*, *f2*, *h1*, *h2*, *ps2* and *ps3*); *ps1* absent.

**Dorsum** — Aspidosoma with striae longitudinal centrally, diverging toward *das* furrow laterally; dorsal part of opisthosoma with median longitudinal striation, diverging toward *das* furrow, about transverse from the level between *d1* and *im* to the level of *f1* and then diverging toward posterior margin.

Eye spots absent. Idiosomal setae aciculate and smooth; seta *bo* about as long as *ex*, which in turn is longer than *ro* and *la*, the latter two subequal. Setae *f2* and *h2* subequal to *bo* and *ex*, longer than other dorsal idiosomal setae; *ro*, *f2* and *h2* the shortest. Setae *c1*, *d1*, *e1* and *f1* not reaching bases of respectively *d1*, *e1*, *f1* and *h1*.

Setal lengths and distances between setae: *bo* 27 (27), *bo-bo* 33 (33), *ro* 15 (15), *ro-ro* 18 (14), *la* 17 (18), *la-la* 30 (31), *ex* 25 (27), *c1* 21 (17), *c1-c1* 25 (27), *c2* 17 (16), *d1* 17 (17), *d1-d1* 25 (26), *e1* 19 (18), *e1-e1* 45 (44), *f1* 14 (13), *f1-f1* 15 (15), *f2* 25 (21), *h1* 14 (14), *h2* 25

<footnote>* Holotype, paratype between brackets</footnote>
(29), ps2 16 (15) and ps3 6 (7). Lyrifissures *ia* located between setae *c1* and *d1*, *im* immediately anteriad *e1* and *ip* immediately anteriad *f2*.

**Venter** — Striation mostly longitudinal, except for transverse region immediately behind gnathosoma, between *pt* and *mta* and immediately anteriad *ag1*; in the latter two regions, striation forming a spindle-shaped pattern, respectively wider than the distance between setae *mta* and wider than the distance between setae *ag1*.

With four pairs of *ag* setae anterolaterad anogenital area; genital papillae and lyrifissure *ih* not visible.

Setal lengths and distances between setae: *1b* 19 (16), *1c* 13 (11), *pt* 13 (13), *2a* 17 (20), *mta* 11 (13), *3b* 16 (15), *3c* 15 (16), *3d* 17 (16), *mtβ* 11 (11), *4b* 16 (14), *ag1* 9 (10), *ag2* 8 (8), *ag3* 9 (0), *ag4* 12 (0), *pt*–*pt* 19 (18), *mtα*–*mtα* 37 (31), *mtβ*–*mtβ* 29 (24), *ag1*–*ag1* 25 (22), *ag2*–*ag2* 12 (12). Anogenital area in both holotype and paratype ruptured, hence distances *ag3*–*ag3* and *ag4*–*ag4* are not possible to measure.

**Gnathosoma** — (Figure 17). Chelicera, with short stylets 14 (17). Subcapitulum with striae almost transverse in section anteriad *sc2* and diverging posteriorly; with two pairs of smooth scapular setae, *sc1* 11 (11) and *sc2* 13 (13); orals *or1* and *or2* not distinguishable. Palptarsus elongate, with a distal eupathidium *pζ* 6 (8), setae *l', l'', v* and *d* long and smooth, *l'* inserted more proximal than *d* and *l''* (both at about same level); solenidion *ω* and *ba* not distinguishable, chaetotaxy (5-1-2). Femorogenu length 18 (17), width 8 (9).

**Legs** — (Figure 18). Chaetotaxy: leg I (8 + ω - 3 + φ + k - 3 - 3 - 1); leg II (7 + ω - 2 - 3 - 3 - 1); leg III (7 - 2 - 2 - 2 - 1); leg IV (7 - 2 - 1 - 1 + 1 - 0). Epimeral formula: 3 - 1 - 4 - 2. All leg setae aciculate and smooth, some ventral setae stouter. Tarsus I without claws, but with empodium; *ft''ζ*, *tc'ζ* and *tc''ζ* the longest setae. Tibia I with bifurcated famulus κ, 2(2), which is not always visible, about half length of solenidion φI, 2 (3); and seta *v'* very long and stout. Leg II with ωI 5(5), tibia with ventral seta *v'* very long 25(24) and stout, seta *d* 16 (16); femur seta *v''* 23(21) and *d* 20(18). Leg III with setae *v'* and *d* of tibia 25(24) and 19(19), respectively, and *ev'* and *d* of femur 17(17) and 16(17), respectively. Leg IV with femur divided, with setae *ev'* 22(18) and *d* 18(16), tibia with setae *v'* and *d* 23(25) and 19(19), respectively. Legs II to IV each with an empodium and two claws distally.

<table><caption>**Table 3** Distinguishing characteristics of *Parapronematus* species.</caption><thead><tr><th rowspan="2">Characteristic</th><th colspan="7">Parapronematus species</th></tr><tr><th>acaciae</th><th>camelia</th><th>citri</th><th>ferox</th><th>formosanus</th><th>geminus</th><th>murshidabadensis</th><th>connarus n. sp.</th></tr></thead><tbody><tr><td>Presence/absence setae</td><td>la<br/># setae on femora I and II</td><td>Absent<br/>3</td><td>Absent<br/>3</td><td>Absent<br/>3</td><td>Absent<br/>3</td><td>Absent<br/>2</td><td>Present<br/>3</td><td>Absent<br/>3</td><td>Absent<br/>3</td></tr><tr><td># setae on tarsus III</td><td></td><td>6</td><td>6</td><td>6</td><td>5</td><td>5</td><td>5</td><td>6 (5 in few specimens)</td></tr><tr><td>Shape of setae</td><td>ro, ex, c1, c2, d1, el<br/>pt mtα mtβ</td><td>Stout, serrate lanceolate</td><td>Stout, lanceolate</td><td>Stout, lanceolate<br/>Barbed, stout to very thin</td><td>Stout, barbed<br/>Narrow, lanceolate, barbed<br/>Barbed, leaflike to lanceolate</td><td>Narrow, setose<br/>Barbed, stout to very thin</td><td>Stout, lanceolate</td><td>Aciculate, broader at base, barbed<br/>Barbed, stout to very thin</td></tr><tr><td>Length of setae</td><td>ro, ex, c1, c2, d1, el<br/>bo<br/>el<br/>fl<br/>f2<br/>h1<br/>h2<br/>p''ζ<br/>tc''/tarsus I</td><td>23-27<br/>≈41*<br/>≈26<br/>≈49<br/>≈77<br/>≈49<br/>≈68<br/>≈34<br/>≈3.0</td><td>22-25<br/>31<br/>22<br/>31<br/>23<br/>Short (11-13)<br/>≈2.2</td><td>≈30<br/>≈13<br/>≈27<br/>≈59<br/>≈35<br/>≈42<br/>≈31<br/>≈3.0</td><td>≈16-24<br/>≈1.5</td><td>16-24<br/>≈2.2</td><td>19-26<br/>≈3.0</td><td>21-24<br/>≈1.7</td><td>16-25<br/>44 (38-50)<br/>21 (19-26)<br/>30 (25-41)<br/>73 (67-81)<br/>33 (28-43)<br/>56 (51-63)<br/>42 (38-47)<br/>≈2.9</td></tr></tbody></table>

<footnote>*The italic bold values are estimated from the respective illustrations (see Material & Methods)</footnote>
<figure><img src="image_18.png" /><figcaption>**Figure 17** *Naudea richinda* Meyer & Rodrigues, 1966. Female. Ventral view of gnathosoma (left) and palp (right).</figcaption></figure>

Holotype female, paratype female and paratype male (all under the same accession number Acy 65/123), at Mite Collection, ARC-Plant Health Protection, Pretoria South Africa, NCA (Zhang 2018).

## Remarks

The characteristics of the type specimens examined led to the conclusion that the types of the only species of this genus differ from the four described *Pausia* species by the number of setae on tarsus II (seven, instead of six), tarsus III (seven, instead of six), the number of setae on tarsus IV (seven, instead of 5-6) and the number of opisthosomal setae (ten instead of 10-11). Moreover, *Pausia* shows several variations in number of setae on different leg segments and on the opisthosoma among species. *Pausia litchiae* Silva, Da-Costa & Ferla is the only species with 10 pairs of setae on the opisthosoma, five setae on tarsus III (instead of six) and two setae on genu IV (instead of one). *Pausia magdalenae* Baker & Definado has only two setae on tibia I (l' ? absent) and femur II (bv" absent) instead of three in the type species (Baker and Delfinado 1976, Da Silva *et al.* 2017). We recommend studying and/or redescribing, if necessary, *P. litchiae* and *P. magdalenae*, to confirm their aberrant leg chaetotaxy and eventually to transfer them to new genera. Hence, in this publication *Naudea* and *Pausia* are considered different genera. As a result, Pronematinae presently congregates 19 genera, one of which is composed of four subgenera (Kazmierski 1998a), based on Baker (1968b, 1969), Każmierski (1998a) and André (2004).

## Characterization of genera of Pronematinae

Table 4 shows the diagnostic characteristics of the taxa presently included in Pronematinae, mostly based on an evaluation of the literature, but also including our examination of the specimens collected in this study as well as of type specimens of two previously described species that are redescribed in this publication.
<figure><img src="image_19.png" /><figcaption>**Figure 19** *Naudea richinda* Meyer & Rodrigues, 1966. Dorsum (left) and venter (right) of male.</figcaption></figure>

4 (3). Genu II with three setae; genu III with two and tarsus III with 5-6 setae, Brazil, Crimea, Greece, South Africa. *Pausia* Kuznetsov & Livshitz, 1972
— Genua II and III each with only one seta; tarsus III with five setae; Mexico
*Pronecupulatus* Baker, 1944

5 (1). One pair of aggenital setae (known only from males)
— More than one pair of aggenital setae (known from males and females)
6
7

6 (5). Ten pairs of opisthosomal setae (*ps1* absent); Brazil
*Quasihomeopronematus* De Vis & Ueckermann, 2024b (*Q. nordestinus* male)
— Eleven pairs of opisthosomal setae; worldwide except Africa and Oceania
*Homeopronematus* André, 1980 (*Homeopronematus anconai* (Baker, 1943) male)
<table><caption>**Table 4** Differential characteristics of pronematid genera: number of species within a genus (Sp. #), number of setae on aspidosoma (Asp. #), opisthosoma, number of *ag* setae (*ag* #), presence of apotele on tarsus I (1 = present, 0 = absent), number of setae on leg segments tarsus (ta), tibia (ti), genu (ge), femur (fe) and trochanter (tr), total number of setae on leg segments (total #, excl. ω, κ, φ) and division of femur IV (1 = divided, 0 = undivided). For variable genera (*Parapronematus*, *Pausia* and *Proctotydaeus*), characteristics are given per groups of species within each.</caption><thead><tr><th rowspan="2">Genus/ species</th><th colspan="3">Asp.</th><th colspan="2">Opisthosoma</th><th colspan="2">ag</th><th colspan="2">Leg I*</th><th colspan="2">apotele</th><th colspan="2">ta</th><th colspan="2">Ti</th><th colspan="2">ge</th><th colspan="2">tr</th><th colspan="2">ta</th><th colspan="2">ge</th><th colspan="2">fe</th><th colspan="2">tr</th><th colspan="2">ta</th><th colspan="2">ge</th><th colspan="2">ta</th><th colspan="2">ge</th><th colspan="2">fe</th><th colspan="2">re</th><th rowspan="2">Total #</th></tr><tr><th>Sp.</th><th>Asp.</th><th>Opisthosoma</th><th>#</th><th>#</th><th>#</th><th>#</th><th>#</th><th>apotele</th><th>ta</th><th>Ti</th><th>ge</th><th>tr</th><th>ta</th><th>ge</th><th>fe</th><th>tr</th><th>ta</th><th>ge</th><th>tr</th><th>ta</th><th>ge</th><th>fe</th><th>tr</th><th>ta</th><th>ge</th><th>tr</th><th>ta</th><th>ge</th><th>fe</th><th>tr</th><th>ta</th><th>ge</th><th>re</th><th>div.</th></tr></thead><tbody><tr><td>Naudea richinda</td><td>1</td><td>4</td><td>10-ps1</td><td>4</td><td>1</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>7+ω</td><td>3</td><td>3</td><td>1</td><td>7</td><td>2</td><td>7</td><td>1</td><td>2</td><td>1</td><td>60</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>7</td><td>2</td><td>7</td><td>1</td><td>2</td><td>1</td><td>56</td></tr><tr><td>Neonaudea gossypii</td><td>1</td><td>4</td><td>11</td><td>4</td><td>1</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>7+ω</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>2</td><td>1</td><td>56</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>2</td><td>1</td><td>56</td></tr><tr><td>Pausia***</td><td>4</td><td>4</td><td>10 or 11</td><td>4</td><td>1</td><td>8+ω</td><td>or3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>2or3</td><td>1</td><td>5 or 6</td><td>2</td><td>5</td><td>1or2</td><td>2</td><td>1</td><td>56</td><td>2</td><td>11</td><td>3+κ+φ</td><td>3</td><td>3</td><td>1</td><td>7</td><td>2</td><td>7</td><td>1</td><td>2</td><td>1</td><td>56</td></tr><tr><td>taurica, colonus</td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>magdaleneae</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>litchiae</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Pronecupulatus anahuacensis</td><td>1</td><td>4</td><td>11</td><td>?</td><td>1</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>1</td><td>3</td><td>1</td><td>5</td><td>1</td><td>5</td><td>1</td><td>2</td><td>0</td><td>52</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>2</td><td>1</td><td>59</td></tr><tr><td>Duoparus hyeresensis</td><td>1</td><td>4</td><td>11</td><td>2</td><td>0</td><td>9+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>7+ω</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>1</td><td>59</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>6</td><td>1</td><td>2</td><td>1</td><td>55</td></tr><tr><td>Metapronematus leucohippeus</td><td>1</td><td>4</td><td>10-ps1</td><td>3</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>5</td><td>1</td><td>1</td><td>0</td><td>55</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>6</td><td>1</td><td>2</td><td>1</td><td>49-52</td></tr><tr><td>Parapronematus</td><td>8</td><td>3 or 4</td><td>9-ps1-ps3</td><td>3</td><td>0</td><td>8+ω</td><td>2+κ+φ</td><td>2</td><td>1</td><td>6+ω</td><td>2</td><td>2 or3</td><td>0</td><td>5 or 6</td><td>2</td><td>6</td><td>1</td><td>1</td><td>0</td><td>49-52</td><td>3</td><td>3</td><td>3</td><td>3</td><td>1</td><td>5</td><td>1</td><td>5</td><td>1</td><td>1</td><td>1</td><td>52</td></tr><tr><td>acaciae, camelia, citri</td><td>3</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>connarus n. sp.</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>geminus</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>ferox, murshidabadensis</td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>formosanus**</td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Apopronematus bakeri</td><td>1</td><td>4</td><td>11</td><td>3</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>2</td><td>1</td><td>6+ω</td><td>2</td><td>3</td><td>1</td><td>5</td><td>1</td><td>5</td><td>1</td><td>1</td><td>1</td><td>1?</td><td>51</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>7</td><td>2</td><td>7</td><td>1</td><td>2</td><td>1/0</td><td>60</td></tr><tr><td>Proctotydaeus</td><td>22</td><td>4</td><td>10 or 11</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>7+ω</td><td>3</td><td>3</td><td>1</td><td>7</td><td>2</td><td>7</td><td>1</td><td>2</td><td>1/0</td><td>60</td><td>5</td><td>11</td><td>3</td><td>3</td><td>1</td><td>7</td><td>2</td><td>6</td><td>1</td><td>2</td><td>1</td><td>0</td><td>60</td></tr><tr><td>P. (Proctotydaeus)</td><td>5</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>P. (Oriola)</td><td>6</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>P. (Neotydeolus)</td><td>4</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>P. (Proctotydulus)</td><td>7</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Pronematulus</td><td>3</td><td>4</td><td>11</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>7+ω</td><td>2</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>1?</td><td>57</td><td>5</td><td>11</td><td>4</td><td>4</td><td>0</td><td>9+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>59</td></tr><tr><td>Pseudopronematulus</td><td>5</td><td>4</td><td>11</td><td>4</td><td>0</td><td>9+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>7+ω</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>1</td><td>59</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>5</td><td>1</td><td>1</td><td>1</td><td>55</td></tr><tr><td>Ueckermanniella cataracta</td><td>1</td><td>4</td><td>11</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>5</td><td>1</td><td>1</td><td>1</td><td>55</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>0</td><td>58</td></tr><tr><td>Homeopronematus anconai</td><td>1</td><td>4</td><td>11</td><td>1 or 4</td><td>0</td><td>9+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>0</td><td>58</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>0</td><td>58</td></tr><tr><td>Quasihomeopronematus<br>nordestinus</td><td>1</td><td>4</td><td>10-ps1</td><td>4</td><td>0</td><td>9+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>0</td><td>58</td><td>1</td><td>7</td><td>3</td><td>3</td><td>1</td><td>6</td><td>2</td><td>6</td><td>1</td><td>2</td><td>0</td><td>58</td></tr><tr><td>Pronematus</td><td>8</td><td>4</td><td>10-ps2</td><td>4</td><td>0</td><td>9+ω</td><td>3+κ+φ</td><td>3</td><td>0</td><td>6+ω</td><td>3</td><td>3</td><td>0</td><td>5</td><td>2</td><td>5</td><td>1</td><td>2</td><td>0</td><td>54</td><td>2</td><td>4</td><td>11</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>2</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>54</td></tr><tr><td>Lourus</td><td>2</td><td>4</td><td>11</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>2</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>2</td><td>0</td><td>54</td><td>4</td><td>11</td><td>4</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>55</td></tr><tr><td>Neapronematus</td><td>8</td><td>4</td><td>9-ps1, ps2</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>2</td><td>0</td><td>54</td><td>1</td><td>4</td><td>10-ps1</td><td>4</td><td>0</td><td>8+ω</td><td>3+κ+φ</td><td>3</td><td>1</td><td>6+ω</td><td>3</td><td>3</td><td>1</td><td>5</td><td>2</td><td>5</td><td>1</td><td>55</td></tr><tr><td>Dasilcoferla nadirae</td><td>1</td><td>4</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Peridasilcoferla paraensis<br>n. gen. n</td><td>1</td><td>4</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody><tfoot><tr><td>Total 20 genera</td><td colspan="2">71 species</td></tr></tfoot></table>

7 (5). Two pairs of aggenital setae, France. *Duoparus* De Vis & Ueckermann, 2024a
— More than two pairs of aggenital setae. 8

8 (7). Three pairs of aggenital setae. 9
— Four pairs of aggenital setae. 11

9 (8). Genua I and II each with three setae; USA. *Metapronematus* André, 1980
— Genua I and II each with two setae. 10

10 (9). Seta *la* reduced or absent; tibia II with two setae + φ + κ; genu III with two setae; Brazil, Democratic Republic of Congo, India, Mozambique, Philippines, Taiwan. *Parapronematus* Baker, 1965
— Seta *la* normal; tibia II with three setae + φ + κ; genu III with one seta; USA. *Apopronematus* André, 1980 (*A. bakeri* André, 1980)

11 (8). Tarsi III and IV each with seven setae; in general insect associates. 12
— Tarsi III and IV each with at most six setae, in general not associated with insect. 15

<footnote>*Do not vary among genera: tibiae II, III and IV, each with two setae, femur I and III with respectively three (except **)) and two setae and trochanters III and IV respectively with one seta and nude, totalling 12 setae on these segments.</footnote>

<footnote>**Parapronematus formosanus has two setae on each femur I and II instead of three.</footnote>

<footnote>***We include the species of *Pausia* in this table because the chaetotaxy of the species belonging to that genus differs from each other.</footnote>
12 (11). *Seta bo* club-shaped; ten pairs of opisthosomal setae (*ps2* absent); femur IV not divided; Brazil .................................... *Proctotydaeus (Neotydeolus)* Flechtmann & Camargo, 1979
— *Seta bo* whip-shaped; 11 pairs of opisthosomal setae (*ps2* present), femur IV divided or not13

13 (12). Paraproctal suckers distinctly developed (especially in females); anus terminally surrounded by sucker-like papillae; tectal setae (*tc*) not longer than tarsus; unguinal setae (*u*) on tarsus I normal; femur IV not divided; Galapagos Islands, Indonesia .................................... *Proctotydaeus (Proctotydaeus)* Berlese, 1911
— Paraproctal suckers weakly developed; setae (*tc*) longer than tarsus; unguinal setae (*u*) on tarsus I shorter than other setae; femur IV divided or not .................................... 14

14 (13). Femur IV not divided; males with some bifurcate setae; Brazil, Canada, Poland, USA .................................... *Proctotydaeus (Oriola)* (Baker, 1968b)
— Femur IV divided; males without bifurcate setae; Crimea, Iran, Mozambique, South Africa, Ukraine, US .................................... *Proctotydaeus (Proctotydulus)* Kaźmierski, 1998a

15 (11). Tarsus II with seven setae .................................... 16
— Tarsus II with six setae .................................... 17

16 (15). Genu II with two setae; Crimea, Taiwan, USA, Uzbekistan .................................... *Pronematulus* Baker, 1965
— Genu II with three setae; Belgium, Brazil, China, Mozambique, South Africa .................................... *Pseudopronematulus* Fan & Li, 1992

17 (15). Tarsus III and femur IV respectively with six and one seta; South Africa .................................... *Ueckermanniella* Beron, 2012
— Tarsus III and femur IV respectively with five and two setae .................................... 18

18 (17). Trochanters I and II each with one seta .................................... 19
— Trochanters I and II nude; worldwide .................................... *Pronematus* Canestrini, 1886

19 (18). Femur II with two setae; opisthosoma with 11 pairs of setae; Crimea, Mozambique .................................... *Lourus* Ueckermann and Grout 2007
— Femur II with three setae; opisthosoma with 9–11 pairs of setae .................................... 20

20 (19). Nine pairs of opisthosomal setae (*ps1* and *ps2* absent); Greece, Hungary, Crimea, Iran .................................... *Neopronematus* (Panou, Emmanouel & Kaźmierski, 2000)
— More than nine pairs of opisthosomal setae .................................... 21

20 (21). Ten pairs of opisthosomal setae (*ps1* absent); Brazil .................................... *Dasilcoferla* De Vis & Ueckermann, 2024a
— Eleven pairs of opisthosomal setae; Brazil .................................... *Peridasilcoferla* De Vis & Ueckermann **n. gen.** (This paper)

For the following genera, keys to the species have been developed (reference in brackets): *Neopronematus* (Darbemamieh *et al.* 2015), *Parapronematus* (this work), *Proctotydaeus* (Kaźmierski 1998a); *Pronematus* (Ueckermann *et al.* 2024) and *Pseudopronematulus* (De Vis *et al.* 2024a).

## Discussion

Presence/absence of one or more setae is a key feature in the taxonomy of Pronematinae. Thus, an important aspect is to decide which dorsal seta/setae is/are absent: *ps1*, *ps2* and/or *h2*?
confirmation, and new genera should be described within the current generic concept if these
characteristics are confirmed. If the presence of two setae on genu IV is confirmed, that would
be the first species within the Pronematinae to show that characteristic (André 1980).

We think that other aspects of the morphology could be introduced in the genus descriptions
and might even have priority over the presence or absence of setae. The difficulty in this
is that early described genera and species have been poorly defined and redescriptions of
those are therefore needed before these characters can be used. An additional problem is
that the types of at least some species are in inadequate condition to be further studied. We
have recently described new species in the following genera of Pronematinae: *Pronematus*
(Ueckermann *et al.* 2024), *Duoparus*, *Pseudopronematulus* and *Dasilcoferla* (De Vis *et al.*
2024a), *Homeopronematus* and *Quasihomeopronematus* (De Vis *et al.* 2024b), *Peridasilcoferla*
**n. gen.** and *Parapronematus* (this paper). While doing so, we were convinced that the following
characteristics seemed of generic value:

a) in the idiosoma:

* Presence or absence of tubercles on the striae: this seems to be fixed within at least two genera: absent in *Pseudopronematulus* (five species examined) and present in *Pronematus* (eight species examined).

* Presence/absence of spindle-shaped pattern of striation anteriad *el*: present in *Parapronematus*, absent in all other genera (compare Figures 1 and 5 of this publication).

* Extent of the spindled pattern of striae anteriad *mta*: in *Peridasilcoferla n. gen.* it extends anteriorly to level of *pt*; in others (*Pronematus*, *Homeopronematus*; *Quasihomeopronematus*, *Dasilcoferla*, *Parapronematus*) it almost reach level of *pt*; in still others (*Duoparus*, *Pseudopronematulus*) it is much smaller. Unfortunately, this characteristic is not pointed out in the description of many species.

* Extent of the spindled pattern of striae anteriad *ag1*: Smaller than distance between setae *ag1* (*Duoparus*), a bit wider than the distance between those setae (*Pseudopronematulus*, *Peridasilcoferla n. gen*, *Parapronematus*) or much broader, about twice the distance between those setae (*Pronematus*, *Homeopronematus*). Also in this case, the characteristic is not properly detailed in the description of many species.

* Presence and form of *la*: Somewhat thinner than *bo* and *ex* (most genera), thin and small (*Parapronematus geminus*) or absent (all other *Parapronematus* spp.). In the generic concept of André (1980), a new genus should be described for species with *la* absent.

* Forked *ag3*: present in *Homeopronematus*, *Quasihomeopronematus* and *Neopronematus* (except *N. aegeae*, Panou *et al.* 2000; in fact, the generic placement of *N. aegeae* should be reevaluated based on that characteristic).

* Positioning of *ag* setae: *Pseudopronematulus*-type, positioned in two eccentric lines (*Pseudopronematulus*, *Duoparus*); or *Pronematus*-type with distances *ag2*-*ag2* and *ag3*-*ag3* short and near base of the longitudinal part of the genital opening (*Neopronematus* (Panou *et al.* 2000), *Pronematus*, *Homeopronematus*, *Quasihomeopronematus*); or *Dasilcoferla*-type with *ag1* and *ag3* close to each other while *ag2* and *ag4* widely separated (*Dasilcoferla* and *Peridasilcoferla n. gen.*).

* The (relative) lengths of setae 3b, 3c and 3d: 3c longer than the other two subequal setae (*Pseudopronematulus*), 3d longer than the other two subequal setae (*Homeopronematus*, *Quasi-homeopronematus*); subequal 3c and 3d and these longer than 3b (*Pronematus*, *Dasilcoferla*); 3d longer than 3c which is longer than 3b (*Duoparus*, *Peridasilcoferla n. gen.*). As differences may not be much, measurement of several specimens seems necessary.

b) in the gnathosoma:

* Status of seta v on palptarsus: smooth (*Pronematus*, *Homeopronematus*, *Quasihomeopronematus*, *Pseudopronematulus*, *Dasilcoferla*), finely barbed (*Peridasilcoferla n. gen.*) or heavily barbed (*Duoparus*). This characteristic is very clear in *Duoparus* but not in other genera under phase microscopy.
c) in the legs:

* Presence of seta homologous to *s* (Ueckermann *et al.* 2024), present in *Pronematus*, *Homepronematus* and *Quasihomeopronematus* (De Vis *et al.* 2024b); vestigial or absent in *Pseudopronematulus* and *Duoparus* (De Vis *et al.* 2024a); absent in *Dasilcoferla*, *Peridasilcoferla n. gen.*, and *Parapronematus* (De Vis *et al.* 2024a and this paper); unknown for other genera.

* Length of ωI. We prefer to use the absolute length as the width of leg is dependent on mounting conditions. In most species, the relative length of the eupathidia: (tc) subequal (most species) or not (Dasilcoferla); (p) subequal (Duoparus, *Pseudopronematulus*), p"ζ shorter (*Pronematus*, *Homepronematus* *Quasihomeopronematus*, *Peridasilcoferla n. gen.*, *Parapronematus*) or longer than p'ζ. (Dasilcoferla, *Neopronematus*).

* The status of the eupathidia on tarsus I: smooth (*Duoparus*, some *Pseudopronematulus*), partially barbed (*Homepronematus*, *Quasihomeopronematus*, *Peridasilcoferla n. gen.*, *Neopronematus*, some *Pseudopronematulus*), or completely barbed (*Dasilcoferla*, *Pronematus* (but not tip), *Parapronematus*, some *Pseudopronematulus*); the type of barbs on the eupathidia also differ.

* Presence of forked dorsal setae: present on femur III only (at least some *Pseudopronematulus*; to be confirmed in others); present on femora III and IV (*Parapronematus*); or absence (all other genera).

* Pending upon confirmation of studies to be conducted, other characteristics seemingly important to consider in generic characterization include: the ornamentation of eupathidia of tarsus I, as well as the distance along the main axis of the two most distal setae of tarsus IV. Still, given the small dimensions of those setae, their usefulness as diagnostic features remains to be determined.

Summarizing our vision of the important characteristics in the definition of taxa of Pronematinae, we retain the listed characteristics as of generic importance and recommend them to be part of genus descriptions and not of species description (although further study of described species should confirm this). For body characteristics: the presence or absence of tubercles on the striae, the presence/absence of spindle form striae anterior to setae *el* on the dorsum; the form of spindle form striae anterior to setae *mta*; the form of spindle form striae anterior to setae *ag1* on the venter; the presence and form of seta *la*; the form of *ag3* (forked or not). For gnathosoma characteristics: the status of seta *v* on palptarsus. For leg characteristics: the relative length of the eupathidia.

## Acknowledgements

The seed of this work was planted during the post-doctoral fellowship nr 01/13725-6 at ESALQ provided by FAPESP (State of São Paulo Research Foundation) to the first author in 2002. The Biodiversity Virtual Institute Program (BIOTA) of FAPESP ([http://www.biotasp.org.br](http://www.biotasp.org.br)); it was in part supported by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES), grant No. 88887.827149/2023-00, awarded to J. Lacerda; Universidade Federal Rural da Amazônia Campus Parauapebas provided logistic support; and Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) issued collection authorization (nr.89189-2) and provided logistical assistance in expeditions.

Arne Jansen and Irene Cardoso were so kind to transport slides of Brazilian mites to the first author and Christine Muggler was our postal hub in Rio de Janeiro.

## Distribution of work

Raf M.J. De Vis collected samples in 2002–2003, made the description of *P. connarus n. sp.*, developed keys and wrote the text. José P. Lacerda collected samples and mounted the mites
in Brazil; Gilberto de Moraes supervised the work of José P. Lacerda and this work as senior acarologist. Edward A. Ueckermann redescribed *P. geminus* and *N. richinda* and supervised the work as senior acarologist and Pronematinae specialist. We want to acknowledge explicitly Henri André for his initiative to build the Tydeoidea database in the wikispecies platform (André 2021). It was an invaluable tool during the preparation of this manuscript and all previous ones.

## ORCID

Raf M. J. De Vis https://orcid.org/0009-0000-6954-5824

José Dantas Araujo Lacerda https://orcid.org/0000-0003-4438-2322

Gilberto J. de Moraes https://orcid.org/0000-0002-5587-1781

Edward A. Ueckermann https://orcid.org/0000-0003-4213-4309

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