# Redescriptions of Amblyseius cucurbitae Rather and A. paraaerialis Muma (Parasitiformes: Phytoseiidae) with new synonymies

Ismail Döker, Isabelle Atchia, Jude Gasparro-Brewer, Xiao-Duan Fang, Samuel J Bolton

► To cite this version:

Ismail Döker, Isabelle Atchia, Jude Gasparro-Brewer, Xiao-Duan Fang, Samuel J Bolton. Redescriptions of Amblyseius cucurbitae Rather and A. paraaerialis Muma (Parasitiformes: Phytoseiidae) with new synonymies. Acarologia, 2025, 65 (3), pp.934 - 946. 10.24349/2yok-2sif. hal-05386925

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# Redescriptions of *Amblyseius cucurbitae* Rather and *A. paraaerialis* Muma (Parasitiformes: Phytoseiidae) with new synonymies

Ismail Döker<sup>a</sup>, Isabelle Atchia<sup>b</sup>, Jude Gasparro-Brewer<sup>b</sup>, Xiao-Duan Fang<sup>c</sup>, Samuel J. Bolton<sup>b</sup>

## Original research

### ABSTRACT

The taxonomy of the family Phytoseiidae remains problematic, largely due to insufficient original descriptions and the lack of comprehensive redescriptions. In many cases, new phytoseiid species have been described without the provision of sufficient differential diagnosis in relation to previously known taxa. This has led to repeated descriptions of the same species under different names, particularly in neighboring countries. *Amblyseius cucurbitae*, Rather and *A. paraaerialis*, Muma were originally described from India based on undetailed illustrations, without clear diagnostic characters. As a result, both species have been described multiple times under different names. Based on a comprehensive literature review, we conclude that eight species, *A. amorphalae*, Sadanandan & Ramani, *A. bhadrakshae*, Sadanandan & Ramani, *A. coffeae*, Sadanandan, *A. indirae*, Gupta, *A. kundurukkae*, Anithalatha & Ramani, *A. mohanasundarami*, Anithalatha and *A. velayudhani*, Santhosh & Sadanandan, are junior synonyms of *A. cucurbitae*. Furthermore, *A. hainanensis*, Wu and *A. brachycalyx*, Karmakar, Bowmik & Sherpa are also considered junior synonyms of *A. paraaerialis*. Therefore, we provide detailed redescriptions of *A. cucurbitae* and *A. paraaerialis* based on their type specimens to clarify their taxonomic status and facilitate future identification.

**Keywords** taxonomy; predatory mites; synonymy; Amblyseiini; Indomalayan realm

**Zoobank** http://zoobank.org/88858CD9-23F5-4DCD-9787-0FA819D1E168

### Introduction

Phytoseiid mites, a diverse group within the superorder Parasitiformes, are widely studied due to their importance as biological control agents (McMurtry *et al.* 2013). However, the taxonomy of this family remains problematic, largely due to insufficient original descriptions and the lack of comprehensive redescriptions. Early descriptions often lack critical morphological detail, and new species are frequently described using limited comparisons with the known taxa. These issues have led to the repeated description of the same species under different names, particularly in geographically similar regions.

The genus *Amblyseius* Berlese, established in 1914, represents one of the most species-rich and globally distributed groups within Phytoseiidae, with over 400 nominal species (Demite *et al.* 2014; Ferragut and Navia 2015; Kreiter *et al.* 2021; Khaustov *et al.* 2023). *Amblyseius*

Received 11 June 2025
Accepted 29 August 2025
Published 16 September 2025

Corresponding author
Ismail Döker ID: idoker@cu.edu.tr

Academic editor
Kreiter, Serge

https://doi.org/10.24349/2yok-2sif

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

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

Döker I. *et al.*

Licensed under
Creative Commons CC-BY 4.0

<footnote><sup>a</sup>Cukurova University, Agricultural Faculty, Department of Plant Protection, Acarology Laboratory, 01330, Adana, Türkiye.</footnote>

<footnote><sup>b</sup>Florida Department of Agriculture and Consumer Services, 32608, Gainesville, FL, USA.</footnote>

<footnote><sup>c</sup>Guangdong Key Laboratory of Animal Conservation and Resource Utilization, Institute of Zoology, Guangdong Academy of Sciences, Guangzhou, 510260, Guangdong, China.</footnote>
species are known for their broad predatory habits, feeding on a variety of prey such as mites, thrips, whiteflies, and nematodes (McMurtry *et al.* 2013). Some species, including *A. andersoni* (Chant, 1957) and *A. swirskii* Athias-Henriot, 1962, have been successfully implemented in biological control and integrated pest management programs worldwide.

*Amblyseius cucurbitae* Rather, 1985, and *A. paraaerialis* Muma, 1967, were described from India approximately 40 and 60 years ago, respectively. However, both descriptions are brief, based on simplified illustrations, and lack key diagnostic features, making their reliable identification challenging. A review of the literature suggests that, likely due to these deficiencies, both species may not have been properly recognized and could have been described under different names in subsequent studies. Based on their original descriptions, several nominal species such as *A. amorphalae*, *A. bhadrakshae*, *A. coffeae*, *A. indirae*, *A. kundurukkae*, *A. mohanasundarami*, and *A. velayudhani*, appear morphologically similar to *A. cucurbitae*. Likewise, *A. hainanensis*, and *A. brachycalyx*, show strong affinities to *A. paraaerialis*. Given these similarities, we attempted to examine the type specimens of all these species in order to clarify their identities. While we were able to access and study the types of some species, others were unavailable; for those, detailed morphological comparisons were carried out based on the original descriptions.

The aim of this study is to redescribe *A. cucurbitae* and *A. paraaerialis* based on their type material and to assess the taxonomic status of the potentially synonymous species through direct comparison with the respective type specimens and/or a comprehensive literature review.

## Material and methods

Specimens were examined using a Zeiss Axio Imager.M2 compound microscope equipped with differential interference contrast (DIC) optics and an AxioCam 820 mono camera, as well as Zeiss Axioscope 5 TL compound microscopes with phase contrast optics and Canon EOS 80D and 90D cameras. All relevant morphological features commonly used in phytoseiid mite taxonomy were measured. Measurements were taken with ZEN 2012 software (version 8.0) and are reported in micrometers (µm). The dorsal shield length was measured along the midline from anterior to posterior margins. The ventrianal shield length, including the cribrum, was measured along the midline. Leg lengths were measured from the base of the coxa to the tip of the tarsus, excluding the ambulacrum.

The classification system follows Chant and McMurtry (2007). Dorsal idiosomal setal nomenclature is based on Lindquist and Evans (1965), as modified by Rowell *et al.* (1978) for phytoseiids. Ventral setal nomenclature follows Chant and Yoshida-Shaul (1991). Adenotaxy, poroidotaxy, and sigillotaxy were interpreted according to Athias-Henriot (1971, 1975), Evans and Fain (1995), and Tsolakis and Ragusa (2016). Leg chaetotaxy follows Evans (1963), and macrosetal terminology (e.g., *SgeIV*, *StiIV*) follows Athias-Henriot (1957). Following Athias-Henriot (1975), the term solenostomes used in this study refers to gland openings.

A paratype of *A. cucurbitae* and the holotype of *A. paraaerialis*, examined in this study, are deposited at the Florida State Collection of Arthropods, 1911 SW 34th St, Gainesville, FL 32608, USA. A paratype of *A. hainanensis* is deposited at the Guangdong Public Laboratory of Wild Animal Conservation and Utilization, Guangdong Institute of Applied Biological Resources, Guangzhou, Guangdong, China.

## Results

### *Amblyseius cucurbitae* Rather

*Amblyseius cucurbitae* Rather, 1985: 291.

*Amblyseius (Amblyseius) indirae* Gupta, 1985: 209. **New synonym.**

*Amblyseius (Amblyseius) kundurukkae* Anithalatha & Ramani, 2004: 58. **New synonym.**
*Amblyseius (Amblyseius) mohanasundarami* Anithalatha, 2005: 81. **New synonym.**

*Amblyseius (Amblyseius) amorphalae* Sadanandan & Ramani, 2006: 2267. **New synonym.**

*Amblyseius (Amblyseius) bhadrakshae* Sadanandan & Ramani, 2006: 2267. **New synonym.**

*Amblyseius (Amblyseius) coffeae* Sadanandan, 2006: 39. **New synonym.**

*Amblyseius velayudhani* Santhosh & Sadanandan, 2015: 67. **New synonym.**

**Material examined**

Paratype female, *Amblyseius cucurbitae* on *Cucurbita pepo* L. (Cucurbitaceae), 14 August 1977, Srinagar, Jammu and Kashmir, India, coll. A. Q. Rather.

**Diagnosis**

Idiosomal setal pattern 10A: 9B/14 *JV-3: ZV* (*r3* and *R1* off shield). Dorsal shield smooth with seven pairs of solenostomes (*gd1*, *gd2*, *gd4*, *gd5*, *gd6*, *gd8*, and *gd9*). Dorsal setae smooth, except *Z5* with a few barbs and *J5* with one barb. Dorsal setae *Z4*, *Z5*, and *s4* greatly elongated; *j1*, and *j3* moderately elongated; remaining setae short/minute. Peritremes long, extending bases of setae *j1*. Solenostome *gd3* present on peritrematal shield. Sternal shield smooth with three pairs of setae (*st1-3*). Genital shield smooth wider than ventrianal shield. Ventrianal shield elongated, vase-shaped, smooth, with three pairs of pre-anal setae (*JV1*, *JV2*, and *ZV2*); one pair of large crescentic solenostomes. Spermatheca with bell-shaped calyx with nodular and forked atrium attached to calyx, neck absent. Fixed digit of chelicera with 10 teeth and movable digit with three teeth. Trochanter I with five (1 0/1 0/2 1) and genu II with seven setae (2 2/0 2/0 1). Macrosetae present on all legs. Tarsus I with two erect setae. *SgeI* (*pd1*), *SgeII* (*pd1*), *SgeIII* (*ad1*), *StIII* (*ad*), *StIII* (*ad3*), *SgeIV* (*ad1*), *StIV* (*ad*), and *StIV* (*pd3*) present.

**Re-description**

**Female (n = 1) (Figures 1-3)**

*Dorsal idiosoma* – (Figure 1A). Dorsal setal pattern 10A: 9B (*r3* and *R1* off shield). Dorsal shield sclerotized, smooth; with seven pairs of solenostomes (*gd1*, *gd2*, *gd4*, *gd5*, *gd6*, *gd8*, and *gd9*), *gd9* conspicuously larger than others; 11 pairs of visible poroids (*id1-id6*, *idm2*, *idm3*, *idm6*, *is1*, *idl1*, and *idl3*). Muscle marks (sigilla) visible on podosoma. Length of dorsal shield 355, width at level of *s4* 210, width at level of *S2* 250. All dorsal setae smooth except *Z5* with a few barbs and *J5* with one barb. Measurements of dorsal setae as follows: *j1* 33, *j3* 38, *j4* 8, *j5* 8, *j6* 8, *J2* 11, *J5* 10, *z2* 13, *z4* 13, *z5* 8, *Z1* 12, *Z4* 115, *Z5* 238, *s4* 105, *S2* 15, *S4* 13, *S5* 8, *r3* 19, and *R1* 11.

*Peritrematal shield* – Peritrematal shield fused with dorsal shield at level of seta *j1*. Peritremes begin at the stigmatic opening with four to five lines of microvilli, continuing with two lines anteriorly; extending bases of setae *j1*. Solenostomes *gd3* and *gd10*, and poroid *id3* visible. Solenostome *gdp*, large, inserted posterior to stigmatic opening.

*Ventral idiosoma* – (Figure 1B). Ventral setal pattern 14: *JV-3: ZV*. Sternal shield sclerotized, smooth; with three pairs of setae (*st1-3*) and two pair of elliptical poroids (*iv1* and *iv2*); distance between *st1-st3* 69, distance between setae *st2* 71; metasternal setae *st4* and poroid *iv3* on metasternal platelets. Genital shield smooth, wider than ventrianal shield, with one pair of setae *st5*, and three pairs of sigilla (*gs1-3*); 115 in length, 73 width at level of *st5*. Ventrianal shield vase-shaped, smooth; with three pairs of pre-anal setae (*JV1*, *JV2*, and *ZV2*), one pair of para-anal setae *PA*, unpaired post-anal seta *PST*, and a pair of large crescentic pre-anal solenostomes *gv3* located near base of setae *JV2*, distance between solenostomes 28. Length of ventrianal shield 110, width at level of *ZV2* 60, width at level of paraanal setae 68, width at level of waist 45. Four pairs of caudoventral setae (*ZV1*, *ZV3*, *JV4*, and *JV5*) on soft cuticle surrounding ventrianal shield. Setae *JV5* smooth inserted on small platelets, 60 in length.

*Gnathosoma* – (Figures 1C, 2). Subcapitulum sclerotized, with three pairs of hypostomal setae *h1*, *h2*, and *h3* each 26 in length, and one pair of palp coxal setae *pc* 30 in length; internal
<figure><img src="image_3.png" /><figcaption>**Figure 1** *Amblyseius cucurbitae* Rather, 1985, a paratype female. A – Dorsal idiosoma; B – Ventral idiosoma; C – Chelicera; D – Spermatheca.</figcaption></figure>
<figure><img src="image_4.png" /><figcaption>**Figure 2** *Amblyseius cucurbitae* Rather, 1985, subcapitulum of a paratype female.</figcaption></figure>

malae paired, not extending tip of corniculi; deutosternal groove with seven transverse rows of denticles, each with two lateral denticles. Second segment of chelicera 90 in length. Antiaxial poroid and external arthrodial brush visible. Fixed digit 31 long, with 10 teeth, and pilus dentilis; movable digit 34 long, with three teeth.

*Spermatheca* – (Figure 1D). Calyx, bell-shaped, widening apically, 14 long, 9 wide at mid-point; atrium nodular incorporated with calyx without neck; major duct visible, minor duct not visible.

*Legs* – (Figures 3A–E). Length of legs I–IV 405, 334, 340, and 427, respectively. Chaetotaxy of legs as follows; leg I: coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/2 2, genu 2 2/1 2/1 2, tibia 2 2/1 2/1 2. Leg II: coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/1 1, genu 2 2/0 2/0 1, tibia 1 1/1 2/1 1. Leg III: coxa 0 0/1 0/1 0, trochanter 1 1/1 0/1 1, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/1 1. Leg IV: coxa 0 0/1 0/0 0, trochanter 1 1/1 0/1 1, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/0 1. Chaetotaxy of tarsi II–IV typical for Phytoseiidae and bears 18 setae 3 3/2 3/2 3 + *mv* and *md*. Macrosetae present in all legs. Tarsus I with two erect setae, both sharp pointed and 46–49 in length. Measurements of macrosetae as follows: *SgeI* (*pd1*) 41, *SgeII* (*pd1*) 35, *SgeIII* (*ad1*) 42, *StIII* (*ad*) 42, *StIII* (*ad3*) 28, *SgeIV* (*ad1*) 120, *StIV* (*ad*) 85, and *StIV* (*pd3*) 73.

## Remarks

*Amblyseius cucurbitae* was originally described by Rather (1985) based on specimens collected from various host plants in Srinagar, Jammu and Kashmir, India. The original description is limited, relying on simplified illustrations and providing only dorsal setal measurements. It lacks several critical diagnostic characters that are essential for distinguishing phytoseiid species, such as number of dorsal solenostomes, complete leg chaetotaxy, and measurements of macrosetae on the first three legs. Although Denmark and Muma (1989) and Gupta (2003) later redescribed the species, they repeated the same setal measurements, which were most likely taken directly from the original description. To address these limitations, we redescribed a
<figure><img src="image_5.png" /></figure>

**Figure 3** *Amblyseius cucurbitae* Rather, 1985, right legs of a paratype female. A – Leg I (trochanter-tibia); B – leg II (trochanter-basitarsus); C – Leg III (trochanter-basitarsus); D – Leg IV (trochanter-femur) E – Leg IV (genu-basitarsus). Macrosetae drawn in solid black for clarity.
female paratype of *A. cucurbitae* and provide a more comprehensive and reliable morphological diagnosis for future studies.

We examined the original descriptions of several species, all of them described from India. These species include, *A. amorphalae*, *A. bhadrakshae*, *A. coffeae*, *A. indirae*, *A. kundurukkae*, *A. mohanasundarami*, and *A. velayudhani*. Except for *A. indirae*, which was described from a somewhat more distant location (approximately 300–350 km away), all other species were described based on specimens collected within a restricted area of about 50–80 km in Kerala. In fact, *A. kundurukkae* and *A. bhadrakshae* were even collected from the same location (the Ayurvedic Herbal Garden of Kottakal Arya Vaidya Sala in Malappuram District), but from different host plants. Considering the close proximity of these collection sites, it seems unlikely that these specimens represent truly distinct species. Nevertheless, the authors described them as separate taxa without providing any differential diagnoses or comparisons, either among themselves or with the valid species proposed in this study, *A. cucurbitae*.

Despite repeated efforts to contact the authors and their collaborators, we were unable to obtain any verifiable access to the type specimens of *A. amorphalae*, *A. bhadrakshae*, *A. coffeae*, *A. kundurukkae*, *A. mohanasundarami*, and *A. velayudhani*. Dr. Anithalatha-Sadanandan, who authored all of these taxa, stated that the materials were deposited at the Zoological Survey of India, Calicut, but could not provide any documentation or accession information, referring us instead to colleagues who did not respond to our queries. According to Dr. Ramani, her former Ph.D. advisor, the type specimens were initially kept in their university lab but were later taken by her students. Following her retirement, the acarology laboratory was reassigned to new faculty members with entirely different specializations, and as a result, all taxonomic material, literature, and equipment were lost. No new acarology personnel were appointed, and currently, the department lacks successors in the field. We also attempted to contact directly the Zoological Survey of India in Calicut over a period of two years regarding these types, but received no response. Therefore, we must rely on the original descriptions and make our conclusions based on the morphological characteristics and other information provided therein.

In nearly all cases, morphological differences were neither consistent nor reliable; most were vague, involving slight variations in setal lengths or, in a few instances, questionable observations of movable digit dentition, likely due to limitations of the optic equipment used at the time. For instance, the observed differences between *A. mohanasundarami*, *A. kundurukkae*, and *A. cucurbitae*, specifically in the number of teeth on the movable digit of the chelicera (one, none, and three, respectively) and in certain dorsal setal and macrosetal lengths (e.g., *j3*, and *SgeIV*) show less than 20% variation and are considered to fall within the range of intraspecific variation observed among different populations of the same species (Tixier 2012). Therefore, in the absence of accessible type materials and given the vague, overlapping, and inconsistent morphological characters provided in the original descriptions, we consider these species to be junior synonyms of *A. cucurbitae*. Although these synonymies are strongly supported by the evidence presented above, they should be reconsidered if the type material becomes accessible in the future.

Similarly, based on our examination of the original descriptions, the only difference between *A. cucurbitae* and *A. indirae* is the number of teeth on the movable digit of the chelicera: three in the former and none in the latter. Considering the overall morphological similarity and the fact that members of the *largoensis* species group typically possess three or four teeth on the movable digit (Zannou *et al.* 2007; Liao *et al.* 2020; Döker *et al.* 2022a, b), we consider the toothless state in *A. indirae* to be a potential error in the original description from earlier optics. Furthermore, as *A. cucurbitae* was published one month earlier (August 1985) than *A. indirae* (September 1985), it has priority. Therefore, we consider *A. indirae* to be a junior synonym of *A. cucurbitae*.

Amblyseius paraaerialis Muma

*Amblyseius paraaerialis* Muma, 1967: 270.
*Amblyseius (Amblyseius) hainanensis* Wu, in Wu and Qian 1983: 264. **New synonym.**

*Amblyseius brachycalyx* Karmakar, Bowmik and Sherpa, 2017: 44. **New synonym.**

**Material examined**

Holotype female, *Amblyseius paraaerialis*, on *Citrus* sp. (Rutaceae), 12 December 1962, Palghat, Kerala, India, coll. V. P. Rao. A paratype female, *Amblyseius hainanensis*, on *Citrus* sp. (Rutaceae), 3 October 1980, Xinglong, Hainan, China, coll. X. Qian.

**Diagnosis**

Idiosomal setal pattern 10A: 9B/14 *JV-3: ZV* (*r3* and *R1* off shield). Dorsal shield smooth with seven pairs of solenostomes (*gd1*, *gd2*, *gd4*, *gd5*, *gd6*, *gd8* and *gd9*). Dorsal setae smooth, except *Z5* with a few barbs and *J5* with one barb. Dorsal setae *Z4*, *Z5*, and *s4* greatly elongated; *j1*, and *j3* moderately elongated; remaining setae short/minute. Peritremes long, extending bases of setae *j1*. Solenostome *gd3* and *gd10* present on peritrematal shield. Sternal shield smooth with three pairs of setae (*st1-3*). Genital shield smooth as wide as ventrianal shield. Ventrianal shield, pentagonal, smooth, with three pairs of pre-anal setae (*JV1*, *JV2*, and *ZV2*); one pair of large crescentic solenostomes. Spermatheca with calyx cup-shaped with very small atrium attached to base of calyx, neck absent, with broad major duct. Fixed digit of chelicera with nine teeth and movable digit with three teeth. Trochanter I with five (1 0/1 0/2 1) and genu II with seven setae (2 2/0 2/0 1). Macrosetae present on all legs. Tarsus I with two erect setae, both sharp pointed and 26-28 in length. *SgeI* (*pd1*), *SgeII* (*pd1*), *SgeIII* (*ad1*), *StIIII* (*ad*), *SgeIV* (*ad1*), *StIV* (*ad*), and *StIV* (*ad3*) present.

**Re-description**

**Female** (n = 1) (Figures 4–6)

*Dorsal idiosoma* – (Figure 4A). Dorsal setal pattern 10A:9B (*r3* and *R1* off shield). Dorsal shield sclerotized, smooth; with seven pairs of solenostomes (*gd1*, *gd2*, *gd4*, *gd5*, *gd6*, *gd8*, and *gd9*), *gd9* conspicuously larger than others; 16 pairs of visible poroids (*id1-id6*, *idm1-6*, *is1*, *idl1*, *idl3*, and *idl4*). Muscle marks (sigilla) visible on podosoma. Length of dorsal shield 329, width at level of *s4* 202, width at level of *S2* 228. All dorsal setae smooth except *Z5* with a few barbs and *J5* with one barb. Measurements of dorsal setae as follows: *j1* 28, *j3* 46, *j4* 7, *j5* 7, *j6* broken, *J2* 7, *J5* 7, *z2* 13, *z4* 9, *z5* broken, *Z1* 9, *Z4* 67, *Z5* 129, *s4* 60, *S2* 14, *S4* 9, *S5* 9, *r3* 18, and *R1* 12.

*Peritrematal shield* – Peritrematal shield fused with dorsal shield at level of seta *j1*. Peritremes begin at the stigmatic opening with three to four lines of microvilli, continuing with two lines anteriorly; extending bases of setae *j1*. Solenostomes *gd3* and *gd10*, and poroid *id3* visible. Solenostome *gdp* large inserted posterior to stigmatic opening.

*Ventral idiosoma* – (Figure 4B). Ventral setal pattern 14: *JV-3: ZV*. Sternal shield sclerotized, smooth; with three pairs of setae (*st1-3*) and two pairs of elliptical poroids (*iv1* and *iv2*); distance between *st1-st3* 60, distance between setae *st2* 66; metasternal setae *st4* and poroid *iv3* on metasternal platelets. Genital shield smooth, as wide as ventrianal shield, with one pair of setae *st5* and three pairs of sigilla (*gs1-3*); 120 in length, 74 width at level of *st5*. Two pairs of genital sigilla (*gs* 4–5) inserted on soft cuticle between genital and ventrianal shields and a pair of genital sigilla (*sgpa*) inserted posterior to setae *ZV1*. Ventrianal shield pentagonal, smooth; with three pairs of pre-anal setae (*JV1*, *JV2*, and *ZV2*), one pair of para-anal setae *PA*, unpaired post-anal seta *PST*, and a pair of large crescentic pre-anal solenostomes *gv3*, located posteromesad to setae *JV2*, distance between solenostomes 25. Length of ventrianal shield 113, width at level of *ZV2* 87, width at level of paraanal setae 74. Four pairs of caudoventral setae (*ZV1*, *ZV3*, *JV4*, and *JV5*) and five pairs of poroids (four pairs of *ivo*, and *ivp*) on soft cuticle surrounding ventrianal shield. Setae *JV5* smooth, 53 in length.

*Gnathosoma* – (Figures 4C, 5). Subcapitulum sclerotized, with three pairs of hypostomal setae *h1*, *h2*, and *h3* each 26 in length, and with one pair of palp coxal setae *pc* 30 in length;
<figure><img src="image_6.png" /><figcaption>**Figure 4** *Amblyseius paraaerialis* Muma, 1967, the holotype female. A – Dorsal idiosoma; B – Ventral idiosoma; C – Chelicera; D – Spermathecae.</figcaption></figure>
<figure><img src="image_7.png" /><figcaption>**Figure 5** *Amblyseius paraaerialis* Muma, 1967, subcapitulum of the holotype female.</figcaption></figure>

internal malae paired, not extending tip of corniculi; deutosternal groove with seven transverse rows of denticles, each with two lateral denticles, except two basal rows with two vertical teeth on each side. Second segment of chelicera 82 in length. Antiaxial poroid and external arthrodial brush not visible. Fixed digit 29 long, with 9 teeth, and pilus dentilis; movable digit 30 long, with three teeth.

*Spermatheca* – (Figure 4D). Calyx, cup-shaped, widening apically, 6 long, 4 wide at widest point; atrium very small nodular incorporated with calyx without neck; major duct well-developed and broad, minor duct not visible.

*Legs* – (Figures 6A–E). Length of legs I–IV 310, 264, 255, and 352, respectively. Chaetotaxy of legs as follows; leg I: coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/2 2, genu 2 2/1 2/1 2, tibia 2 2/1 2/1 2. Leg II: coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/1 1, genu 2 2/0 2/0 1, tibia 1 1/1 2/1 1. Leg III: coxa 0 0/1 0/1 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/1 1. Leg IV: coxa 0 0/1 0/0 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/0 1. Chaetotaxy of tarsi II–IV typical for Phytoseiidae and bears 18 setae 3 3/2 3/2 3 + *mv* and *md*. Macrosetae present in all legs. Tarsus I with two erect setae. Measurements of macrosetae as follows: *SgeI* (*pd1*) 29, *SgeII* (*pd1*) 30, *SgeIII* (*ad1*) 34, *StIII* (*ad*) 25, *StIII* (*ad3*) 28, *SgeIV* (*ad1*) 72, *StIV* (*ad*) 45, and *StIV* (*ad3*) 59.

**Remarks**

*Amblyseius paraaerialis* was originally described by Muma (1967) based on specimens collected from *Citrus* sp. in two different locations in India. The original description is limited, relying on simplified illustrations and lacking several critical diagnostic characters that are essential for distinguishing phytoseiid species today, such as measurements of dorsal setae and macrosetae, number of dorsal solenostomes, and complete leg chaetotaxy. Although Denmark and Muma (1989) later provided a redescription with more details, their redescription still lacks several important morphological features.

Due to its morphological similarity with two other species, *A. brachycalyx* (described from
<figure><img src="image_8.png" /><figcaption>**Figure 6** *Amblyseius paraaerialis* Muma, 1967, left legs of the holotype female. A – Leg I (trochanter-tibia); B – leg II (trochanter-basitarsus); C – Leg III (trochanter-basitarsus); D – Leg IV (trochanter-femur) E – Leg IV (genu-basitarsus). Macrosetae drawn in solid black for clarity.</figcaption></figure>
India) and *A. hainanensis* (described from China), we examined a paratype specimen of *A. hainanensis* and confirmed that it is a junior synonym of *A. paraaerialis*. Unfortunately, we were unable to access the type material of *A. brachycalyx*. The only observed difference between *A. paraaerialis* and *A. brachycalyx* is the length of seta Z5 (129 vs. 150–155). However, this variation is likely due to intraspecific variation between geographically distinct populations and may be correlated with overall body size, as similarly reported for *A. paraaerialis* specimens from Thailand by Ehara & Bhandhufalck (1977) and Oliveira *et al*. (2012).

The holotype specimen of *A. hainanensis* could not be located in the respective collection during our study. Nevertheless, it should be noted that the setal measurements provided in the original description of *A. hainanensis*, specifically for *j1*, *j3*, *Z4*, *Z5*, *s4*, *JV5*, *SgeIV*, *StiIV*, and *StIV*, differ from those observed in the paratype specimen examined in this study. The measurements obtained from the paratype are as follows: 27, 43, 73, 148, 58, 54, 74, 47, and 59, respectively. These measurements are consistent with those observed in *A. paraaerialis*. The discrepancy may reflect variation among type specimens or differences in measurement methods.

## Acknowledgements

We are grateful to Drs. Serge Kreiter (France), Peterson Demite (Brazil), and Emilie Demard (USA) for sharing soft copies of some papers. This study was supported by the Cukurova University Scientific Projects Foundation Units, project number FAY-2022-14495. We are grateful to three anonymous reviewers for their constructive comments on an early version of the manuscript.

## ORCID

Ismail Döker https://orcid.org/0000-0002-1412-1554

Isabelle Atchia https://orcid.org/0000-0001-5407-8370

Jude Gasparro-Brewer https://orcid.org/0009-0009-6396-9349

Xiao-Duan Fang https://orcid.org/0000-0001-8286-4432

Samuel J. Bolton https://orcid.org/0000-0002-5807-8519

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