Godoy, Lucas Lorena ¹ ; Prado, Thais Juliane ² and Navia, Denise ³

¹✉ Departamento Entomologia e Acarologia, Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo,13418–900 Piracicaba, SP, Brazil.
²CBGP, INRAE, CIRAD, IRD, Institut Agro Montpellier, University of Montpellier, France.
³CBGP, INRAE, CIRAD, IRD, Institut Agro Montpellier, University of Montpellier, France.

2026 - Volume: 66 Issue: 1 pages: 14-27

ZooBank LSID: A23745C8-D8DB-485D-988B-19AF0902B67B

Original research

Keywords

Abstract

Introduction

Winterschmidtiidae is a mite family in the cohort Astigmatina, currently comprising over 140 species in 24 genera (Schatz et al. 2011) and occupying a diverse range of habitats. The family is divided into four subfamilies, each with distinct ecological associations. Members of Saproglyphinae are commonly found on decaying materials, fungi, and plant leaves (OConnor 2009; Liu & Zhang 2016), while Oulenziinae inhabit plant foliage, nests of vertebrates and stored food products (Fan et al. 2015). The other two subfamilies are associated with insect: Winterschmidtiinae are linked to various wood-boring insects (OConnor 1994; 2009), while Ensliniellinae are exclusively associated with Hymenoptera (Klompen & OConnor 1995; OConnor 2009). Most species in these subfamilies are primarily detritivorous, although some are phytophagous. Winterschmidtiinae are mostly fungivorous, while some Ensliniellinae feed primarily as adults on the haemolymph of developing wasp larvae without causing apparent damage (Baker & Wharton 1952; OConnor 2009).

A recent publication about the association of mites and solitary wasps (Hymenoptera: Vespidae: Eumeninae) in Brazil reported six genera of Ensliniellinae, namely Calvolia Oudemans (only C. summersi Mostafa), Macroharpa Mostafa, Monobiacarus Baker & Cunliffe, Vespacarus Baker, Zethacarus Mostafa and Zethovidia Mostafa (Pereira et al. 2018). While the ecological role of this relationship remains unclear, the presence of acarinaria (specialized structures on the wasps' bodies that house mites) suggests a long-term evolutionary interaction. These structures are thought to play a crucial role in co-cladogenesis, the parallel evolution of Ensliniellinae mites and their wasp hosts (OConnor and Klompen, 1999), reinforcing the hypothesis of a specialized, long-term symbiotic relationship.

The Ensliniellinae genus Sphexicozela, established by Mahunka (1970), is associated with Polistes wasps. The genus Polistes Latreille consists of over 200 species with a cosmopolitan distribution, most commonly found in tropical regions (Kevan & Nunes-Silva, 2021). These wasps are important predators, feeding primarily on soft-bodied insects (Akte 1982; Cervo 2006). Their nests are made of plant fiber, typically umbrella-shaped with open cells where larvae develop under the care of workers or the foundress (Reeve 1991; Cervo 2006). Some Polistes species are obligate social parasites, invading the nests of other Polistes species instead of building their own (Carpenter 1997; Cervo 2006). Ecologically, they play a role in bioregulation and pollination, making them ecosystem service providers in the context of agriculture (Gould & Jeanne, 1984; Oliveira, et al., 2017; Borchardt et al., 2024).

The genus Sphexicozela is currently monospecific. The only known species, Sphexicozela connivens Mahunka, 1970, was found in nests of an uncertain Polistes species collected from Budapest, Hungary. This mite has also been reported in Ukraine (Rusina & Orlova, 2011; Rusina et al., 2013), in association with Polistes dominulus (Christ) and Polistes nimpha (Christ). An undescribed species of Sphexicozela was also discovered in Japan in association with Polistes snelleni de Saussure, with Yamasaki et al. (2010) providing images of the deutonymph. Additionally, another unidentified species of Sphexicozela was reported by OConnor and Klompen (1999) in association with Polistes perplexus Cresson in Mississippi, USA, although no further details were given.

So far there have been no studies of mites associated with social wasps in Brazil. In 2023, during ecological studies conducted in Muzambinho, Brazil, Astigmatina mites were detected in regular inspection of colonies of Polistes simillimus Zikán. These mites were identified as members of the Winterschmidtiidae family, Sphexicozela genus. In order to identify them, type material of the only known species S. connivens were examined. Specimens collected in Brazil do not fit with S. connivens and were identified as a new species for science.

In this publication we provide a redescription of S. connivens based on the re-examination of type specimens, along with the description of a new species associated with P. simillimus from Brazil.

Material and methods

The holotype, female, and four paratype deutonymphs of S. connivens were kindly provided by Dr. Eszter Lázányi of the Hungarian Natural History Museum, Budapest, for examination. These specimens were observed, photographed and measured using a differential interference contrast (DIC) microscope (Leica DMLB) provided with an AmScope 0.63x camera and connected to a computer containing the AmScope capturing software at the Laboratory of Acarology, Center for Biology and Management of Populations (UMR CBGP), Montferrier-sur-Lez, France.

Mite specimens of Sphexicozela collected from Brazil were studied at the Laboratory of Acarology,'Luiz de Queiroz' College of Agriculture (ESALQ-USP), Piracicaba, São Paulo, Brazil. The specimens were slide-mounted in Hoyer's medium and examined using differential interference and phase-contrast microscopy (Zeiss Axio Imager.A2).

Illustrations were made using Adobe Illustrator 2020, tracing images taken with a Zeiss AxioCamHRc camera attached to a Zeiss Axio Imager 2 microscope. Leg length was measured from the base of the trochanter to the tip of the tarsus without including the pre-tarsus. All measurements are given in micrometers (µm). The terminology of the idiosomal chaetotaxy follows Griffiths et al. (1990), with corrections proposed by Norton (1998); leg chaetotaxy follows Grandjean (1939) and Griffiths (1970), and gnathosomal chaetotaxy follows Bochkov & Mironov (2011) and Klimov & OConnor (2008). The terminology of gland pores and lyrifissures is based on Norton (1998).

Systematics

Sphexicozela connivens Mahunka

The examination of the holotype female and paratype deutonymphs confirmed the details provided in the original description of the species. Complementary information is subsequently provided.

Holotype adult female

(Figures 1–2)

Gnathosoma — (Figure 1A). Subcapitular seta (subc) setiform (13); palpal supracoxal seta (elcp) absent; dorsal palptibial setae (dTi) (6) and lateral palptibial setae (l′) (7) as well as dorsal palptarsal seta (dTa) (3) setiform; terminal palptarsal solenidion ω very short and stout (1). [The positioning of the chelicerae did not allow for a proper visualization].

Idiosoma — Oval; 430 long and 292 at its widest width (next to coxa III); cuticle mammillate, except on the prodorsal area, smooth, constituting the indistinctly sclerotized trapezoidal prodorsal shield. Supracoxal sclerite and ocelli not discernable. Seta sce and ps₁ whip-like, longest (88 and 198, respectively); vi setiform, of intermediate length (35); other dorsally viewed idiosomal setae (sci, scx, c₁ , c₂ , c_p , d₁ , d₂ , e₁ , e₂ , h₁ and h₂ ) thorn–shaped, minute (4–5). [Setae ve indistinct and setae called ve in the original description corresponds to scx and much shorter than depicted originally]. Sejugal furrow apparent. Opisthonotal gland opening (gla) barely distinguishable.

Coxal apodemes I fused at midline; coxal apodemes II directed posteromedially; sejugal apodeme not distinguishable; apodeme III almost transverse; apodemes IV short, directed anteromedially and joined with posterior apodeme of coxa III. Genital opening inverted V–shaped, between coxae III and IV; epigynal sclerite anterior to genital opening not reaching beyond inner tips of apodemes III.

Ventral setae 1a, 3a and 4b subequal (about 23), longer than c3 (13); g (53) and 4a (40) significantly longer than the other ventral setae [other ventral setae possibly illustrated longer than actual length in original description, may have been broken]. Seta 1a inserted in the central region of coxal plate I; seta 3a laterad genital opening, in the central region of coxal plate III, 4b medially and about in transverse line with 3a; seta g at level of second pair of genital papillae seta 4a about in level with posterior margin of genital sclerite. Seta c₃ near lateral edge between coxae II and III. Anal opening far posteriad genital opening but not reaching the rear end of the idiosoma, half length of genital opening and surrounded by three pairs of pseudanal setae: h₃ the longest (198), ps₂ the shortest (5) and ps₃ of intermediate length (10). Setae f₂ [not depicted in original description] similar to ps₂ in length.

Spermatheca (Figure 1B). Copulatory opening barely visible, about 5 in diameter, located terminally on posterior edge of opisthosoma between setae h₃ ; spermathecal duct very long (about 70), winding, particularly thick (thicker than setae h₃ ); basis of spermatheca flat indistinct; sclerites of oviduct strongly sclerotized and shaped as an inverted U, constricted near the basis, 1–2 apart; spermathecal sac indistinguishable.

Legs — (Figure 2). Leg setae smooth; most setae setiform, except as subsequently indicated. Leg I (164) (Figure 2A): trochanter bearing only pR (21); femur, vF (43) whip-like; genu, σ′ (7) and σ″ (11) (both tapering apically), cG (20) and mG (11); tibia, φ (88) whip-like, gT (21) and hT (22); tarsus, ω1 (19), ω2 (11) and ε (4), tapering apically, ω3 (31), d (46) whip-like, wa (28), la (12), ra (25), e (4) and f (9) Leg II (155) (Figure 2B): trochanter bearing pR (23); femur, vF whip-like (48); genu σ (15) slender, tapering apically, setae cG (17) and mG (14); tibia, φ (87) whip-like, gT (22) and hT (13); tarsus, ω (28) tapering apically, d (49) whip-like, wa (26), la (10), ra (17), e (4) and f (9). Leg III (169) (Figure 2C): trochanter bearing sR (22); femur nude; genu, nG (21); tibia, φ (74) whip-like and kT (17); tarsus, d (43) whip-like, w (25), r (16), e (6) and f (6). Leg IV (198) (Figure 2D): trochanter nude; femur, wF (28) setiform; genu nude; tibia, φ (20); tarsus, r (17) and w (22), d (70) whip-like. In all legs, p, q (both 3–4) and s stout (4–5), p and q slightly truncate.

Paratype deutonymph (four specimens examined)

(Figure 3)

Idiosoma — Ellipsoidal, convex, posteriorly constricted, with posteriorly and laterally exposed soft cuticle. 355–400 long and 205–224 at its widest width (next to coxa III); Dorsal shields strongly reticulated and punctated, with a band of smooth, unsclerotized cuticle between the podonotal and opistonotal shields; sejugal furrow distinct. Podonotal shield bearing two pairs of minute setae (probably sci and sce) (3–4) and vi (7–8). Gnathosoma with filiform palpal solenidia ω (15–21) ventrally and seta dm (5–6) apically. Opistonotal shield bearing ten pairs (c₁ , c₂ , c_p , d₁ , d₂ , e₁ , e₂ , f₂ , h₁ and h₂ [originally depicted as h1]) of minute setae (3–4), in addition to one posterior pair (h₃ ) distinctly longer (39–45) [originally depicted as h₂ ], whip-like. Ocelli barely visible, not protruding from the surface. Ventrally one pair of setae is present on coxal plates I and III (barely visible). Minute and indiscernible setae on coxal plate IV. Genital pair of setae (g) are present, very short, adjacent to the genital opening. [The poor visualization may be due to aging of the slide and over clarification of the mite's body structures].

Legs — (Figure 3). Leg setae smooth; most setae setiform, except as subsequently indicated. Leg I (129–134) (Figure 3A): trochanter bearing one short ventral seta (pR) (5–6); femur, vF (32–37) whip-like; genu, setae cG and mG subequal (8–12), σ (9–11); tibia, gT and hT spine-like (11–12), φ whip-like (50–57); tarsus, ω1 and ω3 subequal (20–25), ω2 represented by vestigial alveolus, d and f whip-like (46–50), ra and e subequal (27–30), la and wa spine-like (12–15). Leg II (127–134) (Figure 3B): trochanter bearing one short ventral seta (pR) (5–6); femur, vF (42–49) whip-like; genu, mG (12–14), cG (9–10), σ (11–13); tibia, gT and hT spine-like (11–14), φ whip-like (53-56); tarsus, ω rod-like (26–30), ra and e (22–27), f and d whip-like (50–56) la and wa spine-like (10–12 and 17–19). Leg III (93–101) (Figure 3C): trochanter bearing one short ventral seta (sR) (5–6); femur nude; genu, σ(5–6); tibia, kT thick spine-like(15–18), φ whip-like (36–42); tarsus, f, d and e whip-like (89–98, 100–107 and 160–176), p, q, w, r, s thick, subequal, spine-like (15–18). Leg IV (75–82) (Figure 3D): trochanter nude; femur bearing one short ventral seta (wF) (5–6); genu nude; tibiotarsus, w and d whip-like (197–210 and 211–221), kT (4–5), s short, spine-like (5 –7), p and q thick, subequal, spine-like (13–15), φ (4–5).

Specimens examined

Holotype female: Coll. Mus. Nat. Hung. Holotype Sphexicozela connivens g. n. sp. n. det. Mahunka S.; HNHM Astig-219; Budapest, 1969 VII 18, Ladunga. Paratype deutonymphs: Coll. Mus. Nat. Hung. Paratype Sphexicozela connivens g. n. sp. n. det. Mahunka S.; HNHM Astig-221 / Astig-222 / Astig-226 / Astig-228; Budapest, 1969 VII 18, Ladunga.

Sphexicozela neotropica Godoy & Prado sp. nov.

ZOOBANK: 3D03E4BE-4CC9-4737-B987-BFAC01564810

(Figures 4-8)

Diagnosis

Adult Female — Idiosomal surface mammillate, except for prodorsal shield area; prodorsal shield not distinctly sclerotized, smooth; with a crescent-shaped transverse posterior structure (apparently internal anteriorly). Idiosomal setae sce and h₃ whip-like, 148–160 and 148–176, respectively; vi setiform, of intermediate length (21–28); ve represented by alveoli; other dorsally viewed idiosomal setae (sci, c₁ , c₂ , c_p , d₁ , d₂ , e₁ , e₂ , h₁ and h₂ ) thorn-shaped, minute (4–5). Setae viewed ventrally c₃ , 1a, 3a, 4b, 4a and g (23–30); setae f₂ , ps₂ and ps₃ about as long (5–6) as most dorsally viewed idiosomal setae. Ocelli indiscernible. Basis of spermatheca flat; primary spermaduct very long (64–74) and winding towards base of spermatheca; sclerite of oviducts as an inverted U.

Deutonymph — Opisthonotal region mostly covered with reticulate dorsal shields, with a band of smooth, unsclerotized cuticle between the podonotal and opisthonotal shields. Podonotal shield triangular, leaving unsclerotized cuticle exposed laterally. Posterior dorsal setae h₃ (15–17), longest, appearing ventrally. Ocelli present. Tarsi of legs I and II each bearing two ventral setae (la and wa), not notably thicker than other leg setae. Tarsus III with two whip-like setae (f and d). Tarsus IV with one thick spine-like setae (p).

Male, larva and protonymph. Unknown

Description

Adult female (n = holotype and six paratypes)

(Figures 4–6)

Gnathosoma — (Figure 4A). Chelicera robust; fixed and movable digits with three and four large teeth besides respective apical tooth, respectively; seta cha (2–3) small, stout, conical and chb absent. Subcapitulum (Figure 4C): subcapitular seta (subc) setiform (8–10); palpal supracoxal seta (elcp) absent; dorsal palptibial setae (dTi) (7–9) and lateral palptibial setae (l′) (6–8) as well as dorsal palptarsal seta (dTa) (7–9) setiform; terminal palptarsal solenidion ω very short and stout (2–3).

Idiosoma — (Figure 5). Oval; 456–537 long and 237–321 at its widest width (level of coxa III); cuticle mammillate, except on the prodorsal shield. Prodorsal shield (Figure 5A): 96–99 long and 56–63 wide at ve alveoli level, with a small median constriction slightly above ve, not distinctly sclerotized, smooth, nearly trapezoidal, with a crescent-shaped transverse posterior structure (apparently internal anteriorly). Supracoxal sclerite lightly sclerotized (absence of mammillae); Grandjean's organ not visible. Ocelli indistinct. Setae ve represented by alveoli, situated at about median level of lateral margins of prodorsal shield; idiosomal dorsal setae sce whip-like, longest (148–160); vi setiform, of intermediate length (21–28); scx setiform, very short (5–6); other dorsally viewed idiosomal setae (sci, c₁ , c₂ , c_p , d₁ , d₂ , e₁ , e₂ , f₂ , h₁ and h₂ ) thorn–shaped, minute (4–5). Opisthonotal gland opening (gla) anteromediad e₂ , not distinctly colored.

Setae c₃ (25–28) visible ventrally (Figure 5B), near lateral edge between trochanters I and II. Coxal apodemes I fused at midline; coxal apodemes II arched, directed posteromedially; coxal plate II large, posterior margin short; sejugal apodeme absent. Genital opening inverted V-shaped, situated between coxae III and IV, apodeme III almost transverse; apodemes IV short, directed anteromedially and fused with post–apodeme of coxa III. All podosomatic ventral setae (Figure 4B) subequal (27–35), except for 3a (19–23). Ventral seta 1a inserted in the central region of coxal plate I; setae 3a and 4b approximately aligned transversely along the midline of coxal plate III; seta g at level of posterior pair of genital papillae; seta 4a about in level with posterior margin of genital sclerite. Anal opening positioned far posterior to the genital opening but well anteriad rear end of the idiosoma; about 0.4 as long as nearby seta h₃ and half as long as genital opening and surrounded by three pairs of longitudinally aligned pseudanal setae: h₃ the longest (148–176), ps₂ and ps₃ of similar lengths (4–6).

Spermatheca (Figure 4B): Copulatory opening 5–7 in diameter, located terminally on posterior edge of opisthosoma between lyrifissures ih; spermathecal duct very long (64–74), winding, thin, flaring at the ends near the copulatory opening and the base of the spermatheca; basis of spermatheca indistinct; sclerites of oviduct strongly sclerotized and shaped as an inverted U, slightly constricted near the basis, 1–2 apart; spermathecal sac with a lobed margin, weakly sclerotized.

Legs — (Figure 6). All leg setae smooth, most setae setiform, except as subsequently indicated. Leg I (172–184) (Figure 6A): trochanter bearing only pR (12–15); femur, vF (43–45); genu, σ′ (5–6) and σ″ (8–10) (both tapering apically), cG (17–19) and mG (13–15); tibia, φ (96–98) whip-like, gT (17–18) and hT (10–12); tarsus, ω1 (18–21), ω2 (9–10) and ε (4–5), tapering apically, ω3 (20–25), d (39–43) whip-like, wa (32–34), la (7–8), ra (13–15), e (4–5) and f (10–11). Leg II (170–179) (Figure 6B): trochanter bearing pR (13–16); femur, vF (38–40); genu σ (11–14) slender, tapering apically, setae cG (14–16) and mG (12–16); tibia, φ (93–99) whip-like, gT (16–20) and hT (10–12); tarsus, ω (33–35) tapering apically, d (53–57) whip-like, wa (26–34), la (8–11), ra (19–22) and f (7–9) setiform, e (3–5) minute. Leg III (175–189) (Figure 6C): trochanter bearing sR (14–16); femur nude; genu, nG (36–40); tibia, φ (50–53) whip-like and kT (30–32); tarsus, w (34–37), r (24–25), d (48–55), e (4–5) and f (4–5). Leg IV (181–197) (Figure 6D): trochanter nude; femur, wF (23–26) setiform; genu nude; tibia, φ (28–35); tarsus, r (24–26) and w (24–26) and d (60–71) whip-like. In all legs, p, q (both 3–4) and s (4–6) stout, p and q slightly truncate.

Deutonymph (n = 10)

(Figures 7–8)

Idiosoma — (Figure 7). Dorsal shields (Figures 7A) strongly reticulate, distinctly separate from each other by a band of smooth unsclerotized cuticle which also extends laterad podonotal shields. Ocelli present, highlighted by the presence of darker pigment-spots. Podonotal shield bearing two pairs of barely visible minute setae (sci and sce) (3–4) and vi (6–8). Gnathosoma with filiform palpal solenidia ω (25–29) ventrally and seta dm (5–7) apically. Opistonotal shield bearing eight barely visible pairs (c₁ , c₂ , c_p , d₁ , d₂ , e₁ , e₂ , h₁ and h₂ ) of minute setae (2–3) viewed dorsally, in addition to the often ventrally viewed f₂ (2–3) and h₃ distinctly longer (15–17), spine-like (basically straight). No ventral setae (Figures 7B) observed in coxal plates, though the presence of debris inside the available deutonymphs hampered the precise determination of the characteristics of structures present on the ventral side of the specimens.

Legs — (Figures 8). Leg setae smooth; most setae setiform, except as subsequently indicated. Leg I (140–152) (Figures 8A): trochanter bearing one short ventral seta (pR) (5–6); femur, vF (52–63) whip-like; genu, setae cG and mG subequal (6–10), σ (8–12); tibia, gT thick conical (17–20), hT (9–11), φ whip-like (70–76); tarsus, ω1 thick conical (20–25), ω3 (30–35), ω2 absent, f (10–13), d (15–17), ra and e subequal (31–36), wa (31–35),la (6–8). Leg II (129–136) (Figures 8B): trochanter bearing one short ventral seta (pR) (5–6); femur, vF (49–55) whip-like; genu, cG and mG subequal (7–10), σ (10–12); tibia, gT slightly thick and conical (16–20), hT (9–11), φ whip-like(54–59); tarsus, ω thick (25–28), f (27–34), d (25–28), ra and e subequal (16–18), wa (25–28), la (6–8). Both tarsi I and II exhibit a depression extending from the broad tip of the tarsus, gradually tapering down to the level of seta la. Leg III (83–95) (Figures 8C): trochanter bearing one short ventral seta (sR) (5–6); femur nude; genu, nG (5–6); tibia, kT (8–10), φ whip-like (40–46); tarsus, f and d whip-like (31–45 and 110–123), e (6–7), p, q, w, r, s thick, subequal, spine-like (9–12). Leg IV (68–76) (Figures 8D): trochanter nude; femur bearing one short ventral seta (wF) (4–6); genu nude; tibiotarsus, w and d whip-like (137–151 and 231–257), s and kT subequal (4–5), p thick spine-like (13–15), φ (1–2).

Egg (n= 4)

Elliptical, smooth, longer (173–180) than wider (110–125).

Differential diagnosis

Adult females of S. neotropica sp. nov. differ from S. connivens by the following characteristics: a) ventral setae g about 2.5 times shorter than that in S. connivens; b) seta h₃ about 2.0 times as long as length of anal opening; c) seta ps₃ and ps₂ both short and subequal in length; d) thinner primary spermaduct. Deutonymphs of S. neotropica sp. nov. differ from S. connivens by having: a) dorsal shields more extensive, not leaving a band of unsclerotized lateral and posterior cuticle in a dorsal view of slide mounted specimens; b) idiosoma not posteriorly constricted; c) posterior idiosomal membrane smoothly rounded, not triolobed as in S. connivens; d) seta h₂ (basically straight) about 2.5 times as shorter than S. connivens (whip-like); e) space between setae h₂ 3.5 times as long as the setae, whereas about as long as the setae in S. connivens; f) tarsi I and II without notably thicker (spine-like) ventral setae; g) pair of whip-like setae on tarsi I about 3.0 time as shorter as in S. connivens; h) tarsus III with only two whip-like dorsal setae; i) tibiotarsus IV with only one thick spine-like ventral setae; j) second basalmost free segment of leg IV bearing two short ventral setae.

Type specimens

All type specimens (ten deutonymphs and seven adult females) were collected by A. R. de Souza in Muzambinho, state of Minas Gerais, Brazil (Latitude 21°22′ 33S, Longitude 46°31′ 33W), on adults and pupae of Polistes simillimus Zikán, 1951 (Hymenoptera: Vespidae, Polistinae) and in its nests. Deutonymphs and one adult female were collected in March 2023, and other adult females in November 2023. The holotype adult female and 6 paratype females and 10 paratype deutonymphs were deposited in the mite collection of Escola Superior de Agricultura ''Luiz de Queiroz″, Universidade de São Paulo, Piracicaba, São Paulo state, Brazil (specimens' numbers: 5660–5676).

Etymology

The species name neotropica refers to its geographical origin, as it was found in the Neotropics (Brazil), representing the first known record of the genus Sphexicozela outside the Nearctic and Palearctic regions.

Discussion

The description of S. neotropica sp. nov. expands the known diversity of Sphexicozela, previously represented by a single described species from eastern Europe. This discovery not only represents the first record of the genus in South America but also highlights a previously undocumented association between Ensliniellinae mites and the social wasp P. simillimus. These findings underscore the biogeographic and host-association plasticity within the Ensliniellinae, suggesting a broader distribution and ecological range than previously assumed.

Morphologically, S. neotropica is distinguishable from S. connivens by several consistent traits in both adult females and deutonymphs, particularly in the chaetotaxy and morphology of setae, leg proportions, and the configuration of the spermatheca. These differences are significant enough to warrant specific status and are reinforced by the distinct host and geographic locality. Both species differ from an undescribed Sphexicozela species reported by Yamasaki et al. (2010) from Japan, at least by the presence of a third whip-like seta on tarsus IV of the Japanese deutonymph, as visible in the published image; however, a thorough examination of Japanese specimens would be necessary to confirm the differences.

From a taxonomic perspective, the clear morphological differentiation between the two species reinforces the diagnostic utility of deutonymphal traits in Ensliniellinae systematics. This developmental stage, although frequently under reported, can provide critical insights into both interspecific variation and ecological adaptations.

Future research should explore the possibility of additional cryptic species of Sphexicozela existing among other Polistes hosts in South America and beyond, as Brazil is home to a diverse array of 43 Polistes wasp species (Brito et al., 2024). Additionally, molecular phylogenetic studies would be valuable to test hypotheses of coevolution and biogeographic divergence within the genus. Broader surveys may reveal the diversity of Sphexicozela mites associated with social wasps around the world, including Neotropics, and whether S. neotropica is host-specific to P. simillimus or part of a larger, yet undocumented, complex of species associated with other Neotropical Polistes.

Acknowledgements

Special thanks to Dr. André Rodrigues de Souza and Sírcio Alison dos Santos (FFCLRP-USP) for collecting and providing the specimens of the new species here described. To José Dantas Araújo Lacerda, for slide-mounting the specimens. To Prof. G.J. de Moraes (ESALQ-USP), for his major contribution in the conduction of this work, including the preparation of the manuscript. To Dr. Eszter Lázányi and Dr. Viktória Szőke of the Hungarian Natural History Museum, for kindly lending the holotype and paratypes of Sphexicozela connivens. To Dr. Pavel Klimov and Dr. Barry OConnor, for kindly assisting with the species chaetotaxy. To the São Paulo Research Foundation (FAPESP) for the scholarship granted to L.L.G. (2024/21481-1), which provided financial support for this work.

References

1. Akte R.D. 1982. Social wasps. In: Hermann H. (Ed). The Social Insects. Vol. 4. New York: Academic Press, pp. 1-105. https://doi.org/10.1016/B978-0-12-342204-0.50008-5
2. Baker E.W., Wharton G.W. 1952. An Introduction to Acarology. New York: The Macmillan Company, pp. 465.
3. Bochkov A.V., Mironov S.V. 2011. Phylogeny and systematics of mammal-associated psoroptidian mites (Acariformes: Astigmata: Psoroptidia) derived from external morphology. Invertebr. Syst., 25(1): 22-59. https://doi.org/10.1071/IS10023
4. Borchardt K.E., Holthaus D., Soto Méndez P.A., Toth A.L. 2024. Debunking wasp pollination: Wasps are comparable to bees in terms of plant interactions, body pollen and single-visit pollen deposition. Ecol. Entomol., 49(4): 569-584. https://doi.org/10.1111/een.13329
5. Brito S., Oliveira M.L., Carpenter J.M., Somavilla A. 2024. The genus Polistes Latreille, 1802 (Hymenoptera: Vespidae) of Brazil: morphological diagnoses, identification key, and geographical distribution. Zootaxa, 5496(4): 509-545. https://doi.org/10.11646/zootaxa.5496.4.3
6. Carpenter J.M. 1997. Morphological and molecular evidence for the evolutionary history of Polistes social parasites. Ann. Zool. Fenn., 34: 101-110.
7. Cervo R. 2006. Polistes wasps and their social parasites: an overview. Ann. Zool. Fenn., 43(5/6): 531-549.
8. Fan Q.-H., George S., Li D.-M., Zhang Z.-Q. 2015. Establishment of Oulenziella gen. nov. for Oulenzia bakeri (Hughes, 1962) (Acari: Winterschmidtiidae). Zootaxa, 3949: 191-202. https://doi.org/10.11646/zootaxa.3949.2.2
9. Grandjean F. 1939. La chaetotaxie des pattes chez les Acaridiae. Bull. Soc. Zool. Fr., 64: 50-60.
10. Gould W.P., Jeanne R.L. 1984. Polistes wasps (Hymenoptera: Vespidae) as control agents for lepidopterous cabbage pests. Environ. Entomol., 13(1): 150-156. https://doi.org/10.1093/ee/13.1.150
11. Griffiths D.A. 1970. A further systematic study of the genus Acarus L., 1758 (Acaridae, Acarina) with a key to species. London: British Museum (Natural History).
12. Griffiths D.A., Atyeo W.T., Norton R.A., Lynch C.A. 1990. The idiosomal chaetotaxy of astigmatid mites. J. Zool., 220(1): 1-32. https://doi.org/10.1111/j.1469-7998.1990.tb04291.x
13. Kevan P., Nunes-Silva P. 2021. Pollination and agriculture. In: Encyclopedia of Social Insects. Cham: Springer International Publishing, pp. 736-745. https://doi.org/10.1007/978-3-030-28102-1_176
14. Klimov P.B., OConnor B.M. 2008. Morphology, evolution, and host associations of bee-associated mites of the family Chaetodactylidae (Acari: Astigmata). Ann Arbor: Museum of Zoology, University of Michigan.
15. Klompen J.S.H., OConnor B.M. 1995. Systematic relationships and the evolution of some life history aspects in the mite genus Ensliniella Vitzthum, 1925 (Acari: Winterschmidtiidae). J. Nat. Hist., 29: 111-135. https://doi.org/10.1080/00222939500770061
16. Liu J.-F., Zhang Z.-Q. 2016. Effects of short-term exposure to low temperature on survival, development and reproduction of banana-associated Oulenziella bakeri (Acari: Winterschmidtiidae). Syst. Appl. Acarol., 21: 1078-1086. https://doi.org/10.11158/saa.21.8.8
17. Mahunka S. 1970. Sphexicozela connivens gen. n., sp. n. (Acari, Acaridoidea); a new mite from wasp nest. Parasitol. Hung., 3: 77-86.
18. Norton R.A. 1998. Morphological evidence for the evolutionary origin of Astigmata (Acari: Acariformes). Exp. Appl. Acarol., 22: 559-594. https://doi.org/10.1023/A:1006135509248
19. OConnor B.M. 1994. Life-history modifications in astigmatid mites. In: Houck M.A. (Ed). Mites. Boston: Springer, pp. 136-159. https://doi.org/10.1007/978-1-4615-2389-5_6
20. OConnor B.M. 2009. Cohort Astigmatina. In: Krantz G.W., Walter D.E. (Eds). A Manual of Acarology. 3rd ed. Lubbock: Texas Tech University Press, pp. 565-657.
21. OConnor B.M., Klompen J.S.H. 1999. Phylogenetic perspective on mite-insect associations: the evolution of acarinaria. In: Needham G.R. (Ed). Acarology IX. Vol. 2. Columbus (OH): The Ohio State University Press, pp. 63-71.
22. Oliveira M.M., Gomes F.B., Somavilla A., Krug C. 2017. Polistes canadensis (Linnaeus, 1758) (Vespidae: Polistinae) in the Western Amazon: a potential biological control agent. Sociobiology, 64(4): 477-483. https://doi.org/10.13102/sociobiology.v64i4.1936
23. Pereira M.C., Hermes M.G., Bernardi L.F. 2018. An overview of the mite fauna (Acari) associated with eumenine wasps (Hymenoptera: Vespidae) found in Brazilian collections. J. Nat. Hist., 52(47-48): 3017-3038. https://doi.org/10.1080/00222933.2019.1568602
24. Reeve H.K. 1991. Polistes. In: Ross K.G., Matthews R.W. (Eds). The Social Biology of Wasps. Ithaca: Cornell University Press, pp. 99-148. https://doi.org/10.7591/9781501718670-007
25. Rusina L.Y., Orlova E.S. 2011. The relationship between phenotypic variability in future foundresses of Polistes nimpha (Christ) (Hymenoptera: Vespidae, Polistinae) and infestation of their larvae by the mite Sphexicozela connivens Mahunka (Acari, Astigmata, Winterschmidtiidae). Entomol. Rev., 91(6): 685-691. https://doi.org/10.1134/S0013873811060017
26. Rusina L.Y., Firman L.A., Orlova E.S. 2013. Relation between the male reproductive strategies of Polistes dominula (Christ) (Hymenoptera, Vespidae) and invasion with the mite Sphexicozela connivens (Acari, Astigmata, Winterschmidtiidae) in the Black Sea Biosphere Reserve. Prirod. Alm. Biol. Nauki, 18: 135-146.
27. Schatz H., Behan-Pelletier V.M., OConnor B.M., Norton R.A. 2011. Suborder Oribatida van der Hammen, 1968. Zootaxa, 3148: 141-148. https://doi.org/10.11646/zootaxa.3148.1.26
28. Yamasaki K., Takahashi J., Okabe K., Makino S., Tsuchida K. 2010. The first record of Sphexicozela sp. (Acari: Winterschmidtiidae) phoretic on the paper wasp, Polistes snelleni (Hymenoptera: Vespidae). Jpn. J. Syst. Entomol., 16(2): 311-312.

instructions