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# Phytoseiid mites (Mesostigmata: Phytoseiidae) from Fuerteventura (Canary Islands): A taxonomic and biogeographical approach

Francisco Ferragut

► To cite this version:

Francisco Ferragut. Phytoseiid mites (Mesostigmata: Phytoseiidae) from Fuerteventura (Canary Islands): A taxonomic and biogeographical approach. Acarologia, 2025, 65 (3), pp.962-997. 10.24349/33af-ivt4. hal-05387022

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https://hal.science/hal-05387022v1

Submitted on 28 Nov 2025

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Acarologia

Open Science
in Acarology

# Phytoseiid mites (Mesostigmata: Phytoseiidae) from Fuerteventura (Canary Islands): A taxonomic and biogeographical approach

Francisco Ferragut IDª

## Original research

### ABSTRACT

The oceanic origin of the Canary Islands; that is, islands built from the deep oceanic crust that never were connected with the continent, provides special interest not only for knowing what species thrive there now but also for proposing hypotheses on their biogeographical origin. The phytoseiids (Mesostigmata, Phytoseiidae) from the Canaries have been studied over the past three decades, but the existing publications refer to the three central islands of the archipelago, while the fauna of the eastern and western islands remains unknown. This study presents the results of a survey on indigenous vegetation in Fuerteventura, the easternmost island located near the African coast. Five phytoseiid species were collected and one of them, *Neoseiulella extraseta* sp. nov., characterized by having an extra setal pair *J1* on the dorsum is proposed as new species. The remaining four species are fully re-described, and wherever possible all the motile developmental stages as well as the ontogenetic development of idiosomal and leg setation, pores, and poroids are described. The species richness is analysed from the perspective of the geo-environmental dynamics experienced in Fuerteventura, and some plausible hypotheses are suggested to explain the biogeographic origin of the species found.

**Keywords** Acari; taxonomy; new species; Macaronesia; Island biogeography
**Zoobank** http://zoobank.org/9201E22D-95EA-4759-B6C0-B2D84784B15E

### Introduction

Oceanic islands like the Canaries are considered important natural laboratories for the study of evolution and life's diversity and have been used as ecological models for understanding how their geological and environmental history has determined the existing biotas (Whittaker *et al.* 2023). The Canaries were built over oceanic crust hotspots on the African Plate and have never been connected with the African continent. The plants and animals that survive there arrived from the continent or near islands by dispersal or have evolved and differentiate as new species within the archipelago.

The phytoseiid mites (Mesostigmata, Phytoseiidae) of the Canary Islands have been studied since the late 1980's, and today 29 species are known from the entire archipelago, including eight Canarian endemisms (Evans, 1952; Ferragut *et al.* 1988; Ferragut and Peña-Estévez, 2003, 2007; Moraza and Peña-Estévez, 2005, 2006; Moraza *et al.* 2005; Pande *et al.* 1989). All these reports refer to collections made in the central islands, Gran Canaria, Tenerife and La Gomera, while the mite species from the eastern (Fuerteventura and Lanzarote) and western (La Palma and El Hierro) islands of the archipelago remain largely unexplored.

Received 25 July 2025
Accepted 15 September 2025
Published 26 September 2025

Corresponding author
Francisco Ferragut ID:
*fjferrag@eaf.upv.es*

Academic editor
Kreiter, Serge

https://doi.org/10.24349/33af-ivt4

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

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Ferragut F.

Licensed under
Creative Commons CC-BY 4.0

<footnote>ª Instituto Agroforestal Mediterráneo. Universidad Politécnica de Valencia. Camino de Vera, s/n. 46022 Valencia, Spain.</footnote>
In this study, I examine the phytoseiid fauna of Fuerteventura, the easternmost island of the archipelago, separated from the African coast by only 96 km and characterized by a semi-desertic climate that only allows the development of a sparse cover of shrubs and herbaceous plants with few trees but with a high level of plant endemism. This research has led to the discovery of a new phytoseiid species, *Neoseiulella extraseta* sp. nov. In addition, two species are recorded for the first time in the Canaries. The study also includes a detailed morphological description of all the collected immature stages, and provides a tentative biogeographical analysis of the mite fauna.

## Material and methods

The mites were collected from 25 to 30 May 2024 in Fuerteventura and the nearby islet of Lobos. Only natural and native vegetation was surveyed. Mites were extracted from plants by examining the leaves under a stereomicroscope or beating parts of the plant on a fine sieve. Specimens were mounted and clarified in Heinze-PVA medium and stored in an oven at 50 °C until total clarification. After clarification, the phytoseiids were examined using differential interference contrast (DIC) in a compound microscope Nikon Eclipse Ni-U (Nikon Corporation, Tokyo, Japan). Illustrations were performed on a graphic tablet using Concepts software (https://concepts.app). Images were captured with a digital camera Nikon DS-Fi3 (Nikon Corporation, Tokyo, Japan) and edited in Lightroom and Photoshop CC (© Adobe Systems Inc.). Measurements of morphological traits were obtained with the software NIS-Elements D 3.1 and are given in the text in micrometres (µm). In the description of the new species, values of measurements in the female holotype and the male allotype are represented in bold, followed by the ranges of all individuals studied (in parentheses). Measurements in the immatures of the new species and adults of already known species correspond to the range of all specimens examined. World distribution of each species is based on Demite *et al.* (2025).

The generic concepts are those proposed by Chant and McMurtry (1994, 2007). Idiosomal setal nomenclature follows that of Lindquist and Evans (1965) as adapted by Rowell *et al.* (1978) for the dorsal surface and Chant and Yoshida-Shaul (1991) for the ventral surface. Leg chaetotaxy follows Evans (1963) and Rowell and Chant (1979). Terminology for the morphology of the spermatodactyl is that of Beard (2001). Nomenclature for idiosomal solenostomes and poroids follows Athias-Henriot system (Athias-Henriot, 1975) recently modified by Moraza (2025). I adopted the modifications proposed by Moraza (2025) for being more practical and intuitive without losing the ontogenetic value of the notation system. The modifications introduced in the text affect the designation of the peritrematal gland pore *gd3* which is now *gp*, the peritrematal poroid *id3*, now *ip*, and the opisthonotal poroids *is1*, *idm1*, *idm5*, *idm6* and *idx*, renamed as *idl1*, *id3*, *idl5*, *idm5* and *idm3*, respectively. The remaining idiosomal pores and poroids maintain the same nomenclature.

## Results

**Subfamily Typhlodrominae Wainstein, 1962**

**Tribe Typhlodromini Wainstein, 1962**

**Genus Neoseiulella Muma, 1961**

*Neoseiulella extraseta* Ferragut sp. nov.

Zoobank: D5A739E3-E76F-4368-9299-8DB82D8AE4BB

### Type material and depository

Female holotype, seven female paratypes, one deutonymph on *Caroxylon tetrandrum* (Forsskål)
Akhani & Roalson (Amaranthaceae) and two females on *Arthrocaulon macrostachyum*
(Moricand) Piirainen & Kadereit (Amaranthaceae); Isla Lobos, 28°44'21"N, 13°49'21"W, 3 m a.s.l.; 28 May 2024. Three females on unidentified Amaranthaceae; El Cotillo, Aljibe de la Cueva beach, 28°39'53"N, 14°00'41"W, 18 m a.s.l.; 27 May 2024. Fourteen females, five males, one deutonymph, one protonymph on *Traganum moquinii* Webb ex Moquin-Tandon (Amaranthaceae) and one female on *Cakile maritima* Scopoli subsp. *maritima* (Brassicaceae); Dunas de Corralejo, 28°40'13"N, 13°50'00"W, 12 m a.s.l.; 27 May 2024. Female holotype, eleven paratype females, male allotype and one male paratype, and one deutonymph paratype were deposited in the Museo de Ciencias Naturales de Tenerife, Santa Cruz de Tenerife, Canary Islands. The remaining specimens are in the Acari collection, Instituto Agroforestal Mediterráneo, Universidad Politécnica de Valencia.

Diagnosis

Female dorsal shield thoroughly reticulated; midline cells between setae *j4–J2* elongate, oriented axially; cells between setae *J2–Z4* diverging from the midline and obliquely oriented; most of cells covered by longitudinal striae. Twenty pairs of short setae, the common dorsal setal pattern for *Neoseiulella* plus the pair *J1*, occurring on all the developmental stages and sexes (larvae were not examined); setae *j1* longer than *j3*, setae *J2* shorter than *J5*. Six pairs of dorsal solenostomes, lacking gland pore *gd4*. Peritremes ending anteriorly between setae *j1–j3*. Sternal shield with two pairs of setae. Ventrianal shield subpentagonal, transversally striated, with four pairs of preanal setae and without preanal pores. Spermathecal calyx bowl or dome-shaped, 3–5 deep. Fixed digit of chelicera with two subapical teeth; mobile digit edentate. Genua of legs II–IV with seven setae each; one macroseta on tarsus IV. Male dorsal shield elongate, similar in shape to that of female, not capturing the setal pairs *r3* and *R1*, which remain on the soft cuticle. Dorsal ornamentation as in female. Peritrematal shields merged to the dorsal shield at the level of *j3*. Ventrianal shield with seven pairs of preanal setae. Spermatodactyl L-shaped, with an inconspicuous heel and a short foot.

Description

**Female** (twenty-eight females examined, ten females measured; Figures 1 a–e, 5 a–c)

**Dorsal and lateral idiosoma** — (Figure 1a). Dorsal shield slender and elongate, 417 (377–420) long, 196 (175–196) wide, ratio L/W 2.1 (2.1–2.2). The lateral waist that separates podonotum from opisthonotum little pronounced. Surface evenly ornamented, consisting in lateral striae and closed cells, large cells pentagonal or elongate between the laterals and the midline, and smaller cells elongate, forming a peculiar axial pattern in the midline, between *j4* and *J2*; posterior to that point the cells are oriented transversally to both sides of the shield until the insertions of setae *Z4*. Most of cells, mostly on the opisthosoma, are furrowed by longitudinal striae (Figure 5 b–c). Twenty pairs of short, smooth, and pointed setae, except *Z4*, *Z5* blunt, and *Z5*, *J5* barbed. Lengths of *j1* 22 (20–22), *j3* 16 (14–17), *j4* 12 (12–14), *j5* 11 (11–14), *j6* 13 (13–14), *J1* 14 (12–16), *J2* 13 (13–15), *J5* 15 (15–18), *z2* 16 (12–16), *z3* 15 (13–16), *z4* 13 (12–17), *z5* 12 (12–15), *Z1* 13 (13–16), *Z4* 28 (24–30), *Z5* 30 (27–35), *s4* 16 (14–19), *s6* 20 (18–22), *S2* 21 (19–24), *S4* 21 (17–23), *S5* 22 (19–27). Sublateral setae *r3* 19 (19–21), *R1* 22 (20–25). Six pairs of dorsal solenostomes, with gland pore *gd4* absent; pore opening *gd1* slit-like, the remaining openings circular; pore *gd1* lateral or slightly posterolateral to sockets of setae *j3*; pore *gd8* anteroantiaxial to setae *Z4*; *gd9* anteroparaxial to *S5*. The complete set of sixteen poroids present. Peritrematal shield not fused anteriorly to dorsal shield; peritremes 202 (169–214) long, reaching the level of poroid *id1*, between setae *j1–j3*; internal groove with two rows of microvilli. Peritrematal shield with poroids *ip* and solenostome *gp* on the margin of a shield widening.

**Ventral idiosoma** — (Figure 1b). Tritosternum 97 (96–100) long, with two plumose laciniae 63 (51–63) long. Sternal shield with two pairs of setae, *st1–st2*, and two pairs of poroids *iv1–iv2*; anterior margin concave, posterior margin poorly defined, apparently straight; setae *st3–st4* on piriform platelets, poroids *iv3* not found; distance *st1–st3* 83 (70–83), *st2–st2*
<figure><img src="image_4.png" /><figcaption>**Figure 1** *Neoseiulella extraseta* sp. nov. female. a – dorsal and lateral idiosoma. b – ventral idiosoma. c – spermatheca. d – chelicera. e – genu, tibia and basitarsus IV.</figcaption></figure>
63 (57–64). Epigynal shield **131** (118–136) long, distance *st5-st5* **60** (55–65). Ventrianal shield subpentagonal, anterior margin convex between the insertions of *ZV2*, laterals concave between insertions of *ZV2–JV3*; **141** (126–143) long, **101** (87–103) wide at level of *ZV2*, **90** (81–91) wide at level of anus. Shield surface with a few transverse striae on the anterior part, striae more pronounced around the anus; bearing the preanals *JV1*, *JV2*, *JV3*, *ZV2*, without preanal pores, setae *JV2* anterior to the insertions of *JV3*. Four pairs of setae, *ZV1*, *ZV3*, *JV4*, *JV5*, and five pairs of poroids, four *ivo* and *ivp*, around the ventrianal shield. Posterior metapodal plate **32** (32–39) long. Setae *JV5* pointed, **35** (30–39).

**Spermathecal apparatus** — (Figure 1c). Major duct tubular, of a uniform thickness; atrium simple; calyx short, bowl or dome-shaped, 3–5 deep, 17–20 wide.

**Gnathosoma** — (Figure 1d). Base of gnathosoma **56** (48–58) long, **91** (86–96) wide. Tectum subtriangular, anterior angle rounded, poorly defined. Hypostomal setae *h1* **26** (26–28), *h2* **20** (19–22), *h3* **24** (19–24), *pc* **25** (24–28). Length of first cheliceral segment **30** (29–32), of second segment **100** (93–102). Fixed digit of chelicera **35** (34–39) long, with two subapical teeth; mobile digit **32** (31–36) long, without teeth. Palp **121** (117–127) long; palp chaetotaxy (from trochanter to tarsus) 2-5-6-14-15; all setae smooth and simple except for femoral seta *al* thickened and distally spatulate and genual setae *al1-al2* thickened and distally knobbed and blunt, respectively; palp apotele 2-tined.

**Legs** — (Figure 1e). Lengths of leg I 326 (313–335), leg II 275 (261–278), leg III 271 (251–271), leg IV 367 (347–367). Ventral coxal glands of leg I hypertrophied forming a large and paired opening (about 12 long) in the form of eight. Chaetotaxy of leg segments I–IV: coxeae 2-2-2-1; trochanters 6-5-5-5; femora 12-10-6-6; genua 10-7-7-7; tibiae 10-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanters to tibiae as follows. Leg I: trochanter 1 0/1, 1/2 1; femur 2 3/1, 2/2 2; genu 2 2/1, 2/1 2; tibia 2 2/1, 2/1 2. Leg II: 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: 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: 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. One moderately long macroseta on tarsus IV (*pd3*), smooth and blunt apically, slightly curved at the base, **27** (25–31).

**Male** (five examined and measured; Figures 2 a–c).

**Dorsal and lateral idiosoma** — (Figure 2a). Dorsal shield elongate, ratio L/W 2.1–2.2, similar to that of female by the narrowing of the shield at the level of *r3* and *R1*, both setal pairs not captured by the dorsal shield; **327** (320–331) long, **158** (148–158) wide. Surface thoroughly reticulated; most of cells multistriated, following the female ornamental pattern. Peritrematal shield attached to the dorsal shield in unusual anterior position, at the level of *j3*. Twenty pairs of short, smooth and pointed dorsal setae excepting *Z5* and *J5* barbed. Lengths of *j1* **21** (20–21), *j3* **15** (15–16), *j4* **12** (10–13), *j5* **11** (11–12), *j6*, *J1*, *J2* **13** (12–13), *J5* **15** (13–15), *z2* **12**, *z3* **13** (12–14), *z4* **15** (13–15), *z5* **11** (11–12), *Z1* **14** (13–15), *Z4* **23** (23–24), *Z5* **25** (24–25), *s4* **15** (15–16), *s6* **18** (16–18), *S2* **20** (18–21), *S4* **17** (16–18), *S5* **19** (18–19). Sublateral setae *r3* **16** (16–19), *R1* **18** (17–18). Dorsal adenotaxy and poroidotaxy as in the female; gland opening *gp* on the peritrematal shield. Peritremes arriving anteriorly to the level of solenostome *gdl*, between setal insertions of *j3-z2*; length **151** (148–152).

**Ventral idiosoma** — (Figure 2b). Sternogenital shield with five pairs of setae (*st1-st5*) and three pairs of poroids (*iv1-iv3*), smooth, except on the posterior region covered by longitudinal faint striae; posterior margin of the shield lobate; posterolateral margins partially attached to the exopodal plates; distance *st1-st5* **135** (130–136), *st3-st3* **47** (46–49). Exopodal plates not fused with the ventrianal in the males examined; gland pore *gv2* on the soft cuticle. Ventrianal shield subtriangular attached to the peritrematal shield, anterior margin convex; **125** (121–127) long, **160** (150–163) wide. Surface evenly striate, transversal striae on the anterior part, concentric striae around the anal opening; some transversal striae can be connected forming closed cells. Seven pairs of preanal setae (*JV1*, *JV2*, *JV3*, *JV4*, *ZV1*, *ZV2*, *ZV3*) and four pairs of poroids (*iv5*, three *ivo*) on the shield. Soft integument around the shield bearing the pair *JV5*, **24** (23–25) long, and poroids *ivo* and *ivp*.
<figure><img src="image_5.png" /><figcaption>**Figure 2** *Neoseiulella extraseta* sp. nov. male. a – dorsal and lateral idiosoma. b – ventral idiosoma. c – spermatodactyl and chelicera.</figcaption></figure>
**Chelicerae and spermatodactyl** — (Figure 2c). Fixed digit of chelicera **27** (27–28) long, with two teeth distal to pilus dentilis; movable digit **25** (24–26), with one strong tooth. Spermatodactyl L-shaped, with a short foot and heel not developed; shaft **19** (19–21), foot **5** (4–6) long; one lateral and pointed projection facing upwards, near the distal end of foot. Vellum present.

**Legs** — Leg chaetotaxy similar to female; macroseta on tarsus IV smooth and blunt, **26** (24–28) long.

**Deutonymph** (two examined and measured; Figure 3 a–b).

**Dorsal idiosoma** — (Figure 3a). Dorsal shield 298–333 long, 140–154 wide; lateral margins partially incised at the level of gland pore *gd6*. Shield ornamentation similar to that of female, though less accentuated. Most of cells on the podosomal midline and throughout the opisthosoma multi-striated. Twenty pairs of smooth and pointed dorsal setae, except *Z5*, *J5* barbed. Lengths of *j1* 21–23, *j3* 17, *j4* 13–14, *j5* 12, *j6* 13–14, *J1* 12–13, *J2* 13–14, *J5* 13, *z2*, *z3* 14, *z4* 14–15, *z5* 12–14, *Z1* 13–14, *Z4* 37–38, *Z5* 33–36, *s4* 16–18, *s6* 17–19, *S2* 24–25, *S4* 22–25, *S5* 25–27. Sublateral setae *r3* 18, *R1* 18–19. Six pairs of solenostomes, without gland pore *gd4*, and sixteen pairs of poroids. Peritremes 145–157 long, reaching anteriorly the level of setae *z2*.

**Ventral idiosoma** — (Figure 3b). Base of tritosternum 12–13 wide, total length 94–97, laciniae 49–51. Sternogenital shield well-defined, elongate and smooth, with four pairs of setae (*st1–st4*) and two pairs of poroids (*iv1–iv2*); setae *st5* on the soft cuticle; distance *st1–st5* 150–156, *st3–st3* 52–54. Opisthogastric region with eight pairs of setae (*ZV1*, *ZV2*, *ZV3*, *JV1*, *JV2*, *JV3*, *JV4*, *JV5*) and five pairs of poroids (four *ivo* and *ivp*) surrounding the anal plate. Setae *JV5* smooth and pointed, 35–37.

**Gnathosoma** — Gnathosomal base 69–74 long, 78–83 wide. Deutosternum with seven rows of paired and lateral denticles. Hypostome with four pairs of setae, *h1* 24–26, *h2* 19–20, *h3* 19–22, *pc* 23. First cheliceral segment 25–26, second segment 82–86. Fixed digit of chelicera 30–32 long, with two teeth, movable digit 26–29, without teeth; cheliceral seta present, very short and subtle, about 2 ☐m. Palp 113–116 long; palp chaetotaxy (from trochanter to tarsus) 2-5-6-14-15; femoral seta *al* thickened and expanded apically, genual setae *all* pointed and longer than *al2*, which is blunt; apotele 2-tined.

**Legs** — Chaetotaxy of leg segments I–IV: coxae 2-2-2-1; trochanters 6-5-5-5; femora 12-10-6-6; genua 10-7-7-7; tibiae 10-7-7-6; basitarsi 0-4-4-4. Setal formulas as follows. Leg I: trochanter 1 0/1, 1/2 1; femur 2 3/1, 2/2 2; genu 2 2/1, 2/1 2; tibia 2 2/1, 2/1 2. Leg II: 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: 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: 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. Macroseta on tarsus IV (*pd3*) pointed, 30–33. Pore-like structures resembling gland openings on the distal margin of femur, genu and tibia, and on the proximal margin on tarsus of legs I–IV.

**Protonymph** (one examined and measured; Figure 4 a–b)

**Dorsal idiosoma** — (Figure 4a). Podonotal shield 147 long, 120 wide. Surface smooth, with nine pairs of setae (*j1*,*j3*,*j4*,*j5*,*j6*,*z2*,*z4*,*z5*,*s4*), poroids *id5*, *id2*, *id4*, and solenostomes *gd1* and *gd2*. Setae *r3* and *s6* on the soft integument. Mesonotum between the podonotal and pygidial shields bearing the setal pairs *J1*, *J2*, *Z1*, *S2*, and *R1*, the gland pore *gd6* and the poroids *idl1*, *idl2*, and *idm1*. Pygidial shield 59 long, 74 wide, reticulate, composed by elongated cells, pentagonal or hexagonal. Five pairs of setae (*J5*, *S4*, *S5*, *Z4*, *Z5*), four pairs of poroids (*idm4*, *idm5*, *idl4*, *idl5*) and gland openings *gd8*, *gd9*, on the shield. Setae *Z5* barbed, *J5* basally barbed. Lengths of *j1* 19, *j3* 15, *j4* 13, *j5* 12, *j6* 14, *J1* 11, *J2* 12, *J5* 11, *z2* 13, *z4* 16, *z5* 12, *Z1* 13, *Z4* 19, *Z5* 36, *s4* 19, *s6* 16, *S2* 19, *S4* 17, *S5* 19, *r3* 16, *R1* 15. Peritremes very short, 34 long.

**Ventral idiosoma** — (Figure 4b). Base of tritosternum 12 wide, total length 76, laciniae 39. Margins of sternal shield poorly defined, with three pairs of setae, *st1–st3*, and two pairs of poroids, *iv1*, *iv2*. Setal nubs of *st5* discernible. Setae *JV1*, *JV2*, *ZV2*, *JV5*, and poroids *ivo* and *ivp* around the anal plate. Setae *JV5* blunt, 19 long.
<figure><img src="image_6.png" /><figcaption>**Figure 3** *Neoseiulella extraseta* sp. nov. deutonymph. a – dorsal surface. b – ventral surface.</figcaption></figure>
<figure><img src="image_7.png" /><figcaption>**Figure 4** *Neoseiulella extraseta* sp. nov. protonymph. a – dorsal surface. b – ventral surface.</figcaption></figure>

**Gnathosoma** — Base of gnathosoma 48 long, 68 wide. Deutosternal groove with seven rows of denticles. Setae *h1* 17, *h2* 16, *h3* 19, *pc* 18. First cheliceral segment 19, second segment 64. Fixed digit 26, with two subapical teeth; mobile digit 23, without teeth. Palp 91 long; palp chaetotaxy (from trochanter to genu) 1-4-5; femoral setae *al* slightly expanded apically, setae *al* on genu blunt; apotele 2-tined.

**Legs** — Chaetotaxy of leg segments I–IV: coxae 2-2-2-1; trochanters 4-4-4-4; femora 10-7-5-4; genua 8-6-6-5; tibiae 8-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanter to tibia as follows. Leg I: trochanter 1 0/1, 0/1 1; femur 2 2/1, 2/1 2; genu 1 2/1, 2/1 1; tibia 1 2/1, 2/1 1. Leg II: trochanter 1 0/1, 0/1 1; femur 1 2/1, 2/0 1; genu 1 2/0, 2/0 1; tibia 1 1/1, 2/1 1. Leg III: trochanter 1 0/1, 0/1 1; femur 1 2/1, 1/0 0; genu 1 2/0, 2/0 1; tibia 1 1/1, 2/1 1. Leg IV: trochanter 1 1/1, 0/1 0; femur 1 2/0, 1/0 0; genu 1 2/0, 2/0 0; tibia 1 1/1, 2/0 1. Macroseta on tarsus IV 29 long.

**Larva.** Not found

**Etymology**

The specific name *extraseta* is derived from the Latin terms “extra» meaning above or beyond the common or ordinary and “seta”, in reference to the additional pair *J1* occurring in this species, which represents a novel dorsal setal pattern in the genus *Neoseiulella*.
Comparative diagnosis and remarks

*Neoseiulella extraseta* sp. nov. features the new dorsal setal pattern 12A:10B in the genus by the addition of setae *J1* on the opisthosoma. The occurrence of this setal pair has been previously mentioned by some authors in another member of *Neoseiulella*. Rivnay and Swirski (1980) described *N. montforti* (Rivnay & Swirski) from Israel mentioning and depicting a member of the pair *J1* on the dorsal shield of one specimen. Denmark and Rather (1984) examined a female paratype and reported that the specimen had the two members of setae *J1*. Later, Chant and Yoshida-Shaul (1989) agree with the original description by studying an additional paratype and illustrate a single seta *J1* considering that this character is "an aberration" in the normal dorsal chaetotaxy. By contrast, all the specimens of *N. extraseta* sp. nov. examined in this study plus an abundant set of specimens collected in the nearby island of Lanzarote (unpublished data) have the complete pair *J1*, indicating that this setal pair is stable for the species. Furthermore, *N. montforti* differs from the new species by additional features, such as the presence of preanal solenostomes on the ventral shield, the absence of gland pores *gd5* and the presence of *gd4*, the cup-shaped calyx of the spermatheca, and the longer dorsal setae with ratios *J2/J5* equal to 2.8 instead of 0.9 in *N. extraseta* sp. nov. In the original description of *N. montforti* it is said that the ventral shield lacks solenostomes, but the drawing that accompanies it (Fig. 15 in Rivnay and Swirski, 1980) shows a pair of small punctiform solenostomes. Likewise, the later redescriptions of the species that examined type material (Chant and Yoshida-Shaul, 1989; Denmark and Rather, 1984, 1996; Kanouh et al. 2012) confirm the presence of these pores and include them in their illustrations. The existence of this morphological character is also indicated in the revision of the phytosciids of Israel (Swirski et al. 1998). Table 1 lists the *Neoseiulella* species related to *N. extraseta* sp. nov., all of them native to the Mediterranean basin and the Macaronesian islands. *Neoseiulella extraseta* sp. nov. is the only one with a flattened spermathecal calyx, and the only one, alongside *N. elongata* Ferragut & Peña-Estévez, in having short dorsal setae; however, the latter taxon only has four pairs of dorsal solenostomes, the dorsal ornamentation is distinct, and the peritremes are shorter, with the internal groove striate. The new species also differs from the others by the greater length of setae *j1* and *J5* in relation to setae *j3* and *J2*, respectively. Differences between the new species and *N. litoralis* (Swirski & Amitai) are discussed below. *Neoseiulella paralias* Stathakis, Kapaxidi & Papadoulis (not included in the table) also resembles the new species, but it can easily be differentiated by having three macrosetae on leg IV, cup-shaped calyx of the spermatheca, and six pairs of preanal setae on the male ventral shield (Stathakis et al. 2016).

*Neoseiulella extraseta* sp. nov. exhibits some very original morphological traits in both, females and males. The female and male dorsal ornamentation is made of striated cells resembling scales, which form characteristic patterns comprised of cells of different size and shape arranged in a characteristic way on the shield. This suggests that a detailed description of the dorsal sculpture may provide useful insights as well as additional characters for species diagnosis in the genus. The female peritrematal plate is not fused anteriorly with the dorsal shield, often it is fully separated or at most attached to the dorsal shield by a bridge of soft integument. This feature has never been mentioned in the literature for *Neoseiulella* species, but can also be observed in at least *N. litoralis* (see Figures 5a and 6a).

The dorsal aspect of the male is eye-catching as the lateral sides of the dorsal shield are roughly parallel, with the podonotal region about as wide as the opisthonotal rather than wider than the opisthonotal, as is the norm in the males of Phytoseiidae. Because of this narrowing, the setal pairs *r3* and *R1*, commonly found on the shield, are in the males of this species placed on the soft integument, like in the females. Furthermore, the peritrematal shield is not merged with the dorsal plate at the level of setae *r3* as usual in other species, but in a very anterior position, near the insertion of setae *j3*, as in females.

In the ontogenetic development, dorsal setae *z3*, dorsal solenostomes *gd1* and *gd5*, and ventral setae *st5* (visible as incipient setal nubs), *JV3*, *JV4*, *ZV1*, *ZV3* are added to the
deutonymph. Additionally, some leg setae are incorporated in the moult to deutonymphal stage. On leg I, setae *ad1* and *pv2* of trochanter; *ad3* and *pv2* of femur; *al2*, *pl2* of both genu and tibia are added. On leg II seta *pv2* of trochanter; setae *al2*, *ad3*, *pv1* of femur; *al2* of genu are incorporated. On leg III, *ad1* of trochanter, *pl1* of femur, and *av1* of genu are added. On leg IV, *pv2* of trochanter; *av1*, *pl1* of both femur and genu first appeared in the deutonymph. No differences in leg chaetotaxy were observed between the deutonymphal and adult stages.

## Neoseiulella litoralis (Swirski & Amitai, 1984)

*Typhloctonus litoralis* Swirski & Amitai, 1984: 73.

*Neoseiulella* (*Typhloctona*) *litoralis* Denmark & Rather, 1996: 71.

*Neoseiulella litoralis* Swirski & Amitai, 1997: 37.

## Specimens examined and depository

Ten females, three males, four deutonymphs, one protonymph on *Ononis hesperia* (Maire) H. Förther & Podlech (Fabaceae) and two females on *Cakile maritima* (Brassicaceae); Dunas de Corralejo, 28°40'13"N, 13°50'00"W, 12 m a.s.l.; 27 May 2024. Three females on *Suaeda vera* Forsskål ex J.F. Gmelin (Amaranthaceae) and three females on *Suaeda vermiculata* Forsskål ex J.F. Gmelin (Amaranthaceae); Isla Lobos, 28°45'13"N, 13°49'35"W, 15 m a.s.l.; 28 May 2024. One female on *Suaeda vermiculata* (Amaranthaceae); Betancuria, mirador Morro Velosa, 28°26'68"N, 14°02'59"W, 663 m a.s.l.; 29 May 2024. Twelve females, six males, two deutonymphs, two protonymphs on *Caroxylon tetrandrum* (Forsskål) Akhani & Roalson (Amaranthaceae); Morro Jable. El Saladar, 28°03'03"N, 14°19'34"W, 2 m a.s.l.; 30 May 2024.

Thirteen females, one male, and three deutonymphs were deposited in the Museo de Ciencias Naturales de Tenerife, Santa Cruz de Tenerife, Canary Islands. The remaining specimens are in the Acari collection, Instituto Agroforestal Mediterráneo, Universidad Politécnica de Valencia.

## Geographical distribution

Mediterranean and Macaronesian species, distributed from Israel in the east to the African Atlantic islands of Cape Verde in the west. Usually found on plants near the coast. This is the first report in the Canaries.

<table><caption>**Table 1** *Neoseiulella* species with dorsal shield reticulate, four pairs of preanal setae, one macroseta on leg IV and without preanal pores. Differences between *N. extraseta* sp.nov. and related species. Symbol “-” refers to the absence of data in the literature. Data retrieved from: 1. Rivnay & Swirski, 1980; Chant & Yoshida-Shaul, 1989: Denmark & Rather, 1984; Kanouh *et al.* 2012. 2. Ferragut & Peña-Estévez, 2003; Kanouh *et al.* 2012. 3. Döker, 2018. 4. Swirski & Amitai, 1984; Chant & Yoshida-Shaul, 1989: Denmark & Rather, 1996; Ferragut *et al.* 2010; Kanouh *et al.* 2012. 5. Athias-Henriot, 1960; Chant & Yoshida-Shaul, 1989: Denmark & Rather, 1984; Kanouh *et al.* 2012. 6. Ferragut & Peña-Estévez, 2003; Kanouh *et al.* 2012* [*] Kanouh *et al.* (2012) consider that the integumentary organ between setae s4-s6 illustrated in the original description of *N. splendida* is the solenostome *gd4*, but that structure is, in fact, the poroid *id4*.</caption><thead><tr><th></th><th>N. extraseta sp. nov.</th><th>N. carmeli (Rivnay & Swirski) <sup>1</sup></th><th>N. elongata Ferragut & Peña-Estévez <sup>2</sup></th><th>N. kazaki Döker <sup>3</sup></th><th>N. litoralis (Swirski & Amitai) <sup>4</sup></th><th>N. perforata (Athias-Henriot) <sup>5</sup></th><th>N. splendida Ferragut & Peña-Estévez <sup>6</sup></th></tr></thead><tbody><tr><td>Calyx spermatheca</td><td>shallow, bowl-shaped, 3-5 deep</td><td>cup or bell-shaped, 12 deep</td><td>cup or bell-shaped, 12 deep</td><td>cup or bell-shaped, 10-11 deep</td><td>cup or bell-shaped, 10-12 deep</td><td>cup or bell-shaped, 11 deep</td><td>cup or bell-shaped, 12 deep</td></tr><tr><td>Setae genu II</td><td>7</td><td>6</td><td>7</td><td>8</td><td>7</td><td>8</td><td>7</td></tr><tr><td>Dorsal solenostomes (pairs)</td><td>6 (without gd4)</td><td>6 (without gd5)</td><td>4 (without gd1, gd4, gd5)</td><td>4 (without gd4, gd5, gd8)</td><td>6 (without gd4)</td><td>6 (without gd4)</td><td>6 (without gd4)</td></tr><tr><td>Sternal setae (pairs)</td><td>2</td><td>3</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td></tr><tr><td>Ratio J1/J3</td><td>1.4</td><td>0.8</td><td>1.3</td><td>0.6</td><td>0.9</td><td>1</td><td>0.9</td></tr><tr><td>Ratio J2/J5</td><td>0.9</td><td>2.3</td><td>1.5</td><td>2.2</td><td>1.3</td><td>1.3</td><td>1.1</td></tr><tr><td>Length S2</td><td>19-24</td><td>35-40</td><td>18-21</td><td>41-42</td><td>35-38</td><td>27-30</td><td>42-49</td></tr><tr><td>Length Z5</td><td>27-35</td><td>51-56</td><td>26-36</td><td>50-52</td><td>42-66</td><td>50-53</td><td>70-80</td></tr><tr><td>Male preanal setae (pairs)</td><td>7 pairs</td><td>-</td><td>-</td><td>-</td><td>6 pairs</td><td>6 pairs</td><td>7 pairs</td></tr><tr><td>Distribution</td><td>Canary Islands</td><td>Israel</td><td>Canary Islands</td><td>Türkiye</td><td>Mediterranean, Macaronesia</td><td>West Mediterranean</td><td>Canary Islands</td></tr></tbody></table>
<figure><img src="image_8.png" /><figcaption>**Figure 5** *Neoseiulella extraseta* sp. nov. female. a – anterior part of dorsal shield showing the free peritrematal shield and the position of seta z2. b – striate pattern on the dorsal cells between setae J2–Z4 (1000x). c – ventrianal shield.</figcaption></figure>

Diagnosis

Female dorsal shield suboval, thoroughly reticulate; cells on the dorsal midline between setae *j5–J2* narrow, elongate and oriented axially, which posteriorly become larger and divergent heading towards solenostomes *gd8* and insertions of *Z4*; cells smooth, not striated. Nineteen pairs of dorsal setae; setae *j1* similar in length to *j3*; *z2* short, near setae *z3* and in some females distinctly paraxial to the line *j3–z3*; setae *z3* twice as long as *z2*; setae *J2* longer than *J5*. Six pairs of dorsal solenostomes, gland pore *gd4* absent. Peritremes ending between *j3–z2*; peritrematal shield not fused anteriorly with dorsal shield. Sternal shield with two pairs of setae. Ventrianal shield transversally striate, with four pairs of preanal setae and without preanal pores; margin anterior to setae *ZV2* broadly convex. Spermathecal calyx cup-shaped, 9–10 deep. Fixed digit of chelicera with three subapical teeth, only two visible in profile; movable digit with 0–1 tooth. Genua of legs II–IV with seven setae each; one macroseta on leg IV. Male peritrematal shield fused to the dorsal shield at the level of *j3*; setae *r3* on the soft cuticle, *R1* on the dorsal shield. Ventrianal shield with six pairs of preanal setae, without solenostomes. Spermatodactyl L-shaped; heel rounded and reduced in size; foot short; lateral projection short, pointed, at the end of foot.
<figure><img src="image_9.png" /><figcaption>**Figure 6** *Neoseiulella litoralis* female. a – anterior part of dorsal shield with the peritrematal shield and the position of seta z2. b – ornamentation pattern on the dorsal shield between setae *j6–Z4* (600x). c – ventrianal shield.</figcaption></figure>

**Complementary description**

**Female** (Thirty-one females examined, ten females measured; Figures 6 a–c)

**Dorsal and lateral idiosoma** — (Figures 6 a–b). Dorsal shield suboval, lateral margins with a wide concave waist between setae *s6* and poroids *idl2*; 347–398 long, 155–189 wide, ratio L/W = 2.1–2.3; surface reticulate, except on the anterolateral margins, which are striate-reticulate and the region between the insertions of *Z4*, with transversal striae which can form irregular cells; ornamental pattern with midline cells (*j5*-beyond *J2*) narrow, elongate, and axially oriented, posterior cells becoming larger and diverging towards gland pores *gd8*; cells smooth, without striae. Nineteen pairs of dorsal setae, all smooth and pointed except *Z5* barbed
and blunt and *J5* basally barbed. Lengths of *j1* 20–25, *j3* 19–25, *j4*, *j5* 12–14, *j6* 15–18, *J2* 16–21, *J5* 13–15, *z2* 10–13, *z3* 19–24, *z4* 17–21, *z5* 12–14, *Z1* 15–19, *Z4* 32–45, *Z5* 42–57, *s4* 21–26, *s6* 21–30, *S2* 27–40, *S4* 21–28, *S5* 21–27. Sublateral setae *r3* 19–24, *R1* 20–25. Six pairs of dorsal solenostomes, *gd4* absent; all the gland openings conspicuous and circular. Sixteen pairs of dorsal poroids. Peritremes arriving anteriorly at the level of pore *gd1*, between insertions of *j3* and *z2*; 138–165 long. Peritrematal shield usually extends anteriorly through a narrow and sclerotized strip, but without merging with the dorsal shield.

**Ventral idiosoma** — (Figure 6c). Tritosternum 80–85 long, with two plumose laciniae 43–46 long. Sternal shield with two pairs of setae and two pairs of poroids, poroids *iv2* just on the posterior margin; lateral margins straight, posterior margin concave and irregular; distance *st1-st3* 66–73, *st2-st2* 54–61. Setae *st3* and *st4* on platelets. Epigynal shield narrower than the ventrianal; 112–131 long, distance *st5-st5* 51–70. Ventrianal shield transversally striate, with striae bending around the anal opening (Figure 6c); 119–139 long, 89–102 wide at level of *ZV2*, 76–88 wide at level of anus; anterior margin of the shield sharply convex, laterals straight; with four pairs of preanal setae and without preanal pores. Opistogastric region around the ventrianal plate with setal pairs *ZV1*, *ZV3*, *JV4*, *JV5*, and poroids *ivo* (four pairs) and *ivp*. Setae *JV5* smooth, on small platelets, 36–45 long. Posterior metapodal shield bacillar, with laterals straight, 29–42 long.

**Spermatheca** — Major duct thin and tubular, 25–27 long; atrium simple; calyx cup-shaped, with thin walls; 9–10 deep, 14–16 wide.

**Gnathosoma** — Gnathosomal base 40–43 long, 83–88 wide. Tectum subtriangular, anterior angle rounded. Hypostomal setae *h1* 22–25, *h2* 19–20, *h3* 23–26, *pc* 24–28. First cheliceral segment 23–28, second segment 80–88 long. Fixed digit on chelicerae 27–36 long, with three subapical teeth, almost parallel, commonly only two visible in lateral view; movable digit 26–34 long, usually with one tooth, but some females without teeth. Palp 111–116 long: anterolateral setae (*al*) on palpfemur thicker than the others on the segment, but scarcely spatulated distally; on the palpgenu setae *all* longer than *al2*, setae *all* pointed, *al2* blunt.

**Legs** — Ventral coxal glands of leg I hypertrophied with an opening in the form of eight (about 10 long). Chaetotaxy of leg segments I–IV: coxea 2-2-2-1; trochanters 6-5-5-5; femora 12-10-6-6; genua 10-7-7-7; tibiae 10-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanters to tibiae as follows. Leg I: trochanter 1 0/1, 1/2 1; femur 2 3/1, 2/2 2; genu 2 2/1, 2/1 2; tibia 2 2/1, 2/1 2. Leg II: 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: 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: 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. One macroseta on tarsus IV (*pd3*), smooth and blunt, 32–36 long.

Male (nine examined, five measured; Figure 7 a–b)

**Dorsal and lateral idiosoma** — Dorsal shield suboval, 302–330 long, 147–175 wide; weakly reticulate, ornamental pattern resembling that of female but with midline cells broader and more irregular in shape. Twenty pairs of smooth and acuminate dorsal setae, except *Z4*, *Z5*, blunt, and *J5* basally barbed. Lengths of *j1* 19–22, *j3* 19–21, *j4* 11–13, *j5* 10–13, *j6* 13–14, *J2* 13–16, *J5* 12–14, *z2* 10–12, *z3* 17–21, *z4* 17–19, *z5* 11–14, *Z1* 12–16, *Z4* 30–40, *Z5* 37–43, *s4* 19–21, *s6* 21–26, *S2* 23–31, *S4* 16–21, *S5* 13–21, *r3* 17–18, *R1* 17–19. Setae *r3* always on the soft integument, *R1* on the shield. Six pairs of dorsal solenostomes (*gd4* absent), openings with the same morphology as on females. Peritremes ending at the level of gland pores *gd1* and *gva*, between setae *j3-z2*; 128–132 long. Peritrematal shield merged with dorsal shield at the level of *j3* insertions or even between *j1-j3*.

**Ventral idiosoma** — (7a). Sternogenital shield mostly smooth, with a few lateral striae; posterior margin convex, lobate; distance *st1-st5* 121–137, *st3-st3* 47–61. Exopodal plates not attached to the ventrianal shield. Ventrianal shield subtriangular, broadly convex anteriorly, covered by transversal striae, 122–130 long, 152–179 wide; with six pairs of preanal setae (*JV1*, *JV2*, *JV3*, *ZV1*, *ZV2*, *ZV3*) and without preanal solenostomes. Setae *JV5* 24–29, blunt and smooth, on small platelets.
<figure><img src="image_10.png" /><figcaption>**Figure 7** *Neoseiulella litoralis* male. a – ventrianal shield. b – spermatodactyl and chelicera.</figcaption></figure>

**Chelicera and spermatodactyl** — (Figure 7b). Cheliceral fixed digit 24–28 long, with two subapical teeth, the distal tooth larger than the proximal; movable digit with a strong tooth, 22–24 long. Spermatodactyl L-shaped; shaft 17–18 long; heel rounded, slightly developed; foot short, 5–6; lateral projection near the toe, acute, directed upwards; vellum present.

**Legs** — Ventral coxal gland on leg I similar to that of female. Leg chaetotaxy as in females. One macroseta on tarsus IV (pd3), smooth and blunt, 28–34 long.

**Deutonymph** (six examined and measured; Figure 8 a–b)

**Dorsal and lateral idiosoma** — (Figure 8a). Dorsal shield 291–308 long, 124–142 wide, lateral margins widely invaginated at the level of gland pores gd6 and poroids id11, both cuticular organs on the posterior margin of invagination. Surface of the shield weakly reticulate, cells more marked on the opisthonotum. Nineteen pairs of smooth and pointed dorsal setae, except for Z5, J5, barbed. Lengths of j1 19–21, j3 18–20, j4, j5 11–14, j6 14–16, J2 13–16, J5 12–13, z2 12–15, z3 15–18, z4 14–17, z5 12–14, Z1 14–19, Z4 39–46, Z5 42–45, s4 20–23, s6 21–24, S2 25–31, S4 19–25, S5 21–25. Sublateral setae r3 17–19, R1 16–21. Six pairs of dorsal solenostomes, without gland pore gd4, and sixteen pairs of poroids. Peritremes 130–135 long, reaching the level of setae z2.

**Ventral idiosoma** — (Figure 8b). Sternogenital shield poorly sclerotized, with irregular margins, anterior margin deeply concave; setae st1–st4 on the shield, st5 on the soft cuticle; distance st1–st5 137–144, st3–st3 50–58. Eight pairs of setae (ZV1, ZV2, ZV3, JV1, JV2, JV3, JV4, JV5) and five pairs of poroids (four ivo and ivp) around the anal plate. Setae JV5 smooth and blunt, 31–36.

**Gnathosoma** — Hypostomal setae h1 21–23, h2 15–17, h3 19–21, pc 21–24. Fixed digit of chelicera 25–27, with two subapical teeth; movable digit 23–25, with one small tooth. Palp 93–101 long; tip of seta al on femur slightly expanded, all on genu acuminate, al2 blunt.

**Legs** — Chaetotaxy of leg segments I–IV: coxae 2-2-2-1; trochanters 6-5-5-5; femora 12-10-6-6; genua 10-7-7-7; tibiae 10-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanter to tibia as follows. Leg I: trochanter 1 1/1, 0/2 1; femur 2 3/1, 2/2 2; genu 2 2/1, 2/1 2; tibia 2
<figure><img src="image_11.png" /><figcaption>**Figure 8** *Neoseiulella litoralis* deutonymph. a – dorsal surface. b – ventral surface.</figcaption></figure>
<figure><img src="image_12.png" /><figcaption>**Figure 9** *Neoseiulella litoralis* protonymph. a – dorsal surface. b – ventral surface.</figcaption></figure>

2/1, 2/1 2. Leg II: 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: 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: 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. Macroseta on tarsus IV (*pd3*), distally blunt, 33–42.

**Protonymph** (three examined and measured; Figure 9 a–b)
**Dorsal and lateral idiosoma** — (Figure 9a). Podonotal shield 135–149 long, 113–125 wide, with nine pairs of setae (j1, j3, j4, j5, j6, z2, z4, z5, s4), four pairs of poroids (id5, id2, id4, id6) and solenostomes gd1 and gd2; gland opening gd1 minute, on the shield margin, at the level of the insertions of setae j3. Surface of the shield with faint striae. Setae r3 and s6 on the soft cuticle. Mesonotum with five pairs of sclerotized scutella, setal pairs J2, Z1, S2, R1, solenostomes gd6, and poroids id3, idl1, idl2, idm1, idm2. Pygidial shield reticulate, anterior margin lobate, 59–65 long, 77–89 wide, bearing the setae S4, S5, Z4, Z5, J5, the gland pores gd8, gd9, and the poroids idm3, idm4, idm5, idl3, idl4, idl5. All the dorsal setae smooth and pointed, except Z4, Z5 barbed and J5 basally barbed. Lengths of j1 17–21, j3 15–16, j4 10–12, j5 9–12, j6 13, J2 11–12, J5 9, z2 11–12, z4 15–16, z5 10–11, Z1 12–14, Z4 37–49, Z5 32–38, s4 19–22, s6 16–20, S2 21–25, S4 16, S5 14–18, r3 11–14, R1 9–12. Peritremes 27–33 long, extending to between s4–s6.

**Ventral idiosoma** — (Figure 9b). Sternogenital shield poorly sclerotized, bearing the setal pairs st1–st3 and the poroids iv1, iv2; anterior margin concave, posterior margin subtriangular; distance st1–st3 69–81, distance st2–st2 48–57. Setal nubs of st5 and setal pairs JV1, JV2, ZV2, JV5 on the integument anterior to the anal plate. Setae JV5 smooth and blunt, 16–19 long.

**Gnathosoma** — Cheliceral fixed digit 13–25 long, with two subapical teeth; movable digit 19–22, without dentition. Setae h1 15–17, h2 13–14, h3 17–18, pc 18–20. Palp 78–95 long; anterolateral seta on femur distally expanded, on genu thicken and pointed.

**Legs** — Chaetotaxy of leg segments I–IV: coxae 2-2-2-1; trochanters 4-4-4-4; femora 10-7-5-4; genua 8-6-6-5; tibiae 8-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanter to tibia as follows. Leg I: trochanter 1 0/1, 0/1 1; femur 2 2/1, 2/1 2; genu 1 2/1, 2/1 1; tibia 1 2/1, 2/1 1. Leg II: trochanter 1 0/1, 0/1 1; femur 1 2/1, 2/0 1; genu 1 2/0, 2/0 1; tibia 1 1/1, 2/1 1. Leg III: trochanter 1 0/1, 0/1 1; femur 1 2/1, 1/0 0; genu 1 2/0, 2/0 1; tibia 1 1/1, 2/1 1. Leg IV: trochanter 1 1/1, 0/1 0; femur 1 1/0, 1/0 1; genu 1 2/0, 2/0 0; tibia 1 1/1, 2/0 1. Macroseta on tarsus IV (pd3), blunt, 28–37 long.

**Larva.** Not found

**Remarks**

The most distinctive feature in the females is the shape of the ventrianal shield. This shield is broad, and the margin of the anterior third, above the insertions of setae ZV2, is remarkably rounded, without anterior corners (Figure 6c). On the dorsal shield, setae z2 are short, about half as long as j3 and z3, placed near z3, and with their insertions paraxial to the line j3–z3 (Figure 6a). Furthermore, as pointed out by Ferragut and Baumann (2021), the peritrematal shield is not fused with the dorsal shield but simply extends forward over a narrow bridge (Figure 6a). The number of teeth on the movable cheliceral digit is variable, from zero to one tooth depending on the source and depending on the females, as observed in the sample of Fuerteventura. The male dorsal shield never contains the setal pair r3, which is located on the interscutal integument, and the peritremal shield is attached to the dorsal shield between setae j1–j3. The number of male preanal setae varies according to the literature. Although it is considered that the males have six pairs of setae as originally described by Swirski and Amitai (1984), the corresponding figure in the original description shows six setae on the left side of the shield and five on the right side (figure 7, page 75). Later, Denmark and Rather (1996) examined one paratype male and concluded (and depicted) that it has only five pairs of preanal setae. Subsequent redescriptions (Kanouh et al. 2012; Stathakis et al. 2016) emphasized that the males have six pairs of setae, but Ferragut and Baumann (2021) observed seven pairs on the three males collected in Cape Verde archipelago, with the addition of setae JV4.

This species is common and widespread in the coastal regions of Fuerteventura and Lobos Island and it can coexist on the same plants with *N. extraseta* sp. nov. Females of *N. litoralis* can be easily differentiated from those of the new species by: 1) the cup-like calyx of the spermatheca, 2) the absence of dorsal setal pair J1, 3) the dorsal ornamentation, made of smooth cells, never lineate or striate, and 4) the longer dorsal setae (about 10–66% longer).
The ontogenetic development of dorsal and ventral setation and adenotactic pattern, although incomplete by the absence of larvae, seems identical to that of *N. extraseta* sp. nov., with setae z3, dorsal solenostomes gd1, gd5, and ventral setae st5, JV3, JV4, ZV1, ZV3 incorporated to the deutonymph. Likewise, development of leg chaetotaxy seems similar to that of the new species. On leg I, setae ad1 and pv2 of trochanter; ad3 and pv2 of femur; al2, pl2 of both genu and tibia are added. On leg II seta pv2 of trochanter; setae al2, ad3, pv1 of femur; al2 of genu are incorporated. On leg III, ad1 of trochanter, pl1 of femur, and av1 of genu are added. On leg IV, pv2 of trochanter; av1, pl1 of both femur and genu are added to the deutonymphal complement. There were no differences in leg chaetotaxy between the deutonymphal and adult life stages.

**Genus Typhlodromus Scheuten, 1857**

**Subgenus Anthoseius De Leon, 1959**

**Typhlodromus (Anthoseius) charactus Ueckermann**

*Typhlodromus charactus* Ueckermann, 1996: 21.

*Typhlodromus (Anthoseius) charactus* – Moraes *et al.*, 2004: 318.

*Amblydromella charactus* – Denmark & Welbourn, 2002: (although these authors incorporate most of species of the subgenus *Anthoseius* within the genus *Amblydromella* Muma, they never explicitly mentioned this species. Had they done, they should have named it *Amblydromella* [*Lindquistoseia* characta).

**Specimens examined**

Three females on *Heliotropium ramosissimum* (Lehmann) Sieber ex de Candolle (Boraginaceae); Tindaya, Barranco de Esquinzo, 28°37′47″N, 14°00′18″W, 59 m a.s.l.; 26 May 2024.

**Geographical distribution**

Previously known from Yemen. The new record in Fuerteventura proves it has a disjunct distribution.

**Diagnosis (female)**

Dorsal shield deeply incised laterally at the level of sublateral setae R1; shield striate-reticulate on the anterior and lateral margins, almost smooth between setae j4 and j6, longitudinally striate between j6–J2, with some large cells, transversal striae on the posterior part. Eighteen pairs of short, smooth, and pointed dorsal setae; all shorter than 20 µm. Five pairs of dorsal solenostomes (gd2, gd4, gd6, gd8, gd9). Peritremes extending to between setae j1–j3. Sternal shield with two pairs of setae, setae st3 on the soft cuticle; posterior margin of the shield lobate. Ventranial shield longer than wide; anterior margins straight or slightly convex, laterals concave; surface on the anterior half smooth, on the posterior half striate-reticulate; three pairs of preanal setae (JV1, JV2, ZV2) and preanal pores present, posteroantiaxial to JV2. Major duct of spermatheca broad, atrium fissured and conical, calyx short cup-shaped. Fixed digit of chelicera with three subapical teeth, dentition of movable digit not discernible (four teeth in the original description). Genua of legs II–IV with eight, seven, and eight setae, respectively. Without macrosetae on leg IV.

**Complementary description**

**Female** (Three females examined and measured; Figures 10 a–c)

**Dorsal and lateral idiosoma** — (Figures 10a). Dorsal shield piriform, 359–378 long, wider at the level of opisthosoma (S4, 221–226) than at the level of podosoma (179–192); with a characteristic lateral incision near the insertion of setae R1, deep and extending beyond poroid
<figure><img src="image_13.png" /><figcaption>**Figure 10** *Typhlodromus (Anthoseius) charactus* female. a –dorsal shield showing the lateral incision. b – ventricular shield. c – spermatheca.</figcaption></figure>
*id1* or even poroid *id3*. Dorsal ornamentation consisting of elongate and narrow cells covering the margins of the shield; midline between setae *j4-j6* smooth, between *j6-J2* reticulate, with large cells; posterior region between the insertions of *Z4* transversally striate. Eighteen pairs of short, smooth, and pointed dorsal setae, only *J5* barbed basally. Lengths of *j1* 11–12, *j3* 14–15, *j4* 9–13, *j5* 9–11, *j6* 8–13, *J2* 9–13, *J5* 9–11, *z2* 13–14, *z3* 13–15, *z4* 13–16, *z5* 10–11, *Z4* 12–14, *Z5* 16–17, *s4* 14–18, *s6* 16–19, *S2* 16–18, *S4*, *S5* 14–16. Sublateral setae *r3* and *R1* 14–16, placed on small platelets. Five pairs of dorsal gland pores, *gd2*, *gd4*, *gd6*, *gd8*, *gd9*; gland pore *gd8* anteroantiaxial to *Z4*, pore *gd9* paraxial and aligned with the insertions of *S5*; sixteen pairs of poroids on the shield. Peritremes reaching the level of *j1* or between *j1-j3*, 202–205 long; peritrematal shield narrow, attached to the dorsal shield above the poroid *id1*.

**Ventral idiosoma** — (Figure 10b). Tritosternum relatively short (69–72), plumose throughout except at the base, divided into two laciniae, 34–36 long. Sternal shield smooth, with a few lateral striae; anterior margin concave, posterior margin trilobate, posterolateral lobes penetrating deeply into the shield, up to the level of setae *st2*, lateral margins divergent; poroid *iv2* just on the posterior margin of lateral lobes. Shield with two pair of setae (*st1-st2*) and poroids *iv1*, *iv2*. Setae *st3* placed directly on the soft integument; *st4* on piriform metasternal plates, with poroid *iv3* in anterior position. Distance *st1-st3* 68–79, *st2-st2* 62–66. Epigynal shield narrower than the ventranal, 111–124 long, distance *st5-st5* 59–66. Ventranal shield elongate, almost vase-shaped (Figure 10b); anterior margin slightly convex, laterals between *ZV2* and the anal opening deeply concave; 122–137 long, 88–95 wide at the level of *ZV2*, 74–80 at the level of anus; ratio L/W = 1.4–1.6. Anterior half of the shield almost smooth, posterior half irregularly striate, some striae forming closed cells. Three pairs of setae *JV1*, *JV2*, *ZV2*, and solenostomes *gv3* on the shield; setae *JV2* in posterior position, transversally aligned with the greatest shield narrowing; preanal pores punctiform, posteroantiaxial to the sockets of *JV2*. Setae *ZV1*, *ZV3*, *JV4*, *JV5*, four pairs of poroids *ivo* and one pair *ivp* surrounding the shield. Setae *JV5* smooth and pointed, 17–19 long, on platelets. Metapodal shields bacillar; posterior metapodal 25–33 long.

**Spermatheca** — (Figure 10c). Major duct short, vacuolated near the atrium; atrium well developed, almost as long as calyx; calyx short, subconical, 4–6 deep, 8–9 wide at the junction with the vesicle.

**Gnathosoma** — Base of gnathosoma 27–30) long, 72–76 wide. Tectum dome shaped. Deutosternal groove with seven rows of two denticles. Hypostomal setae *h1* 19–20, *h2* 14–15, *h3* 16, *pc* 18–19. Length of first cheliceral segment 14–15, of second segment 69–76. Fixed digit of chelicera 24–28 long, with three subapical teeth; movable digit 19–14 long, dentition not visible. Palp 92–96 long; femoral seta *al* thickened and distally enlarged, genual setae *al1-al2* thickened and distally blunt; apotele 2-tined.

**Legs** — Lengths of leg I 250–262, leg II 220–224, leg III 206–213, leg IV 264–269. Distal margin of coxeae on all the legs dentate, with 8–10 acute spine-like teeth. All the setae on the legs short and pointed. Chaetotaxy of leg segments I–IV: coxeae 2-2-2-1; trochanters 6-5-5-5; femora 12-10-6-6; genua 10-8-7-8; tibiae 10-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanter to tibia as follows. Leg I: trochanter 1 1/1, 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: trochanter 1 0/1, 0/2 1; femur 2 3/1, 2/1 1; genu 2 2/1, 2/0 1; tibia 1 1/1, 2/1 1. Leg III: 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: trochanter 1 1/1, 0/2 0; femur 1 2/1, 1/0 1; genu 1 2/1, 2/0 2; tibia 1 1/1, 2/0 1. Without leg macrosetae.

**Remarks**

*Typhlodromus (A.) charactus* is an atypical member of the subgenus *Anthoseius* in that the lateral margins of the female dorsal shield are incised in such a way that it resembles that shield in the phytoseiid deutonymphal stage. Unfortunately, no males or immatures of this or other related species are known. This feature is shared by three species, *T. (A.) charactus*, *T. (A) kutabus* Schicha & Corpuz-Raros from the Philippines, and *T. (A.) eremicus* Ueckermann from Northern Cape Province in South Africa. The three taxa share, in addition, an elongate ventranal shield
wider at the level of anterior corners than at the level of anus, a subconical calyx and enlarged atrium in the spermatheca, and only three pairs of preanal setae. Despite the limited number of species, there is much confusion about their taxonomical identity. Ueckermann (in Meyer and Ueckermann, 1989) described *T. (A.) eremicus* from the Kalahari Gemsbok National Park (now incorporated into the Kgalagadi Transfrontier Park). The description was rather complete, indicating that genua II and IV have eight setae. Later, Schicha and Corpuz-Raros (1992) described superficially *T. (A.) kutabus*, with an account of the most relevant characters and some measures but without references to the leg chaetotaxy, although their illustrations are clear and the genu IV bears eight setae. Nevertheless, there are no comments on this work to the previously described *T. (A.) eremicus*. Finally, Ueckermann (1996) described *T. (A.) charactus*. The morphological description and pictures are pretty informative, but again he did not include either *T. (A.) eremicus* and *T. (A.) kutabus* in the comparison to justify the status of the new species.

According to the information appearing in the mentioned papers, *T. (A.) eremicus* seems to be distinct by the shortness of the peritremes, which extend to between the lateral setae *z2-z3*, and the size of the spermathecal calyx, at least twice the length of this structure in the other two species. Ueckermann *et al.* (2008) indicated that there are five pairs of dorsal solenostomes, but they did not specify which ones, and their Figure 17A only shows the gland pores *gd2*, *gd4*, and *gd9*. *Typhlodromus* (*A.*) *kutabus* and *T. (A.) charactus* are similar; the former differs by having three instead of two sternal setae, and the posterior margin of sternal shield is concave rather than lobate. Even though there is no indication of the dorsal adenotaxy for any of the two species, the respective drawings seem to show five pairs of pores in *T. (A.) kutabus* (without openings *gd1* and *gd5*) and six pairs in *T. (A.) charactus* (without *gd5*), but this interpretation is subjective and it is based on the confidence that the drawings accurately reflect mite morphology.

Taking into account all these uncertainties, I opted to identify provisionally the specimens collected as *T. (A.) charactus* for the following reasons: 1) some differential features in this species are coincident with the material collected in Fuerteventura; e.g., the number of pairs of sternal setae and the posterior trilobate margin of the sternal shield, and 2) the biogeographical proximity of this species compared to the Philippine *T. (A.) kutabus*. Further detailed examination of the respective types and genetic studies would be necessary to finally elucidate the taxonomic validity of these taxa and their phylogenetic relationships.

**Typhlodromus (Anthoseius) maspalomensis Ferragut & Peña-Estévez, 2003**

*Typhlodromus (Anthoseius) maspalomensis* Ferragut & Peña-Estévez, 2003: 158.

**Specimens examined**

Thirteen females, four males on *Tamarix canariensis* Willdenow (Tamaricaceae); Puerto del Rosario, 28°30'02"N, 13°50'53"W, 5 m a.s.l.; 26 May 2024. One female on *Dittrichia viscosa* (L.) Greuter (Asteraceae) and one female on *Suaeda vermiculata* (Amaranthaceae); Betancuria, mirador Morro Velosa, 28°26'68"N, 14°02'59"W, 663 m a.s.l.; 29 May 2024. Three females, one male on *Tamarix canariensis* (Tamaricaceae) and one male on *Salsola divaricata* Masson ex Link s. l. (Amaranthaceae); ravine near Vega del Rio Palmas, 28°23'25"N, 14°05'41"W, 213 m a.s.l.; 29 May 2024.

**Geographical distribution**

Canary Islands, Greece, and Morocco. It also occurs in the Iberian Peninsula (unpublished data).

**Diagnosis**

Female dorsal shield mostly reticulate, with the midline on the opisthonotum smooth. Dorsal setae smooth and pointed; setae *j3* longer than *j1*; *S5* longer than *S2* and *S4*. Five pairs of dorsal
<figure><img src="image_14.png" /><figcaption>**Figure 11** *Typhlodromus (Anthoseius) maspalomensis* female. a – anterior part of dorsal shield with pronounced muscle marks. b – cavities on the posterior part of dorsal shield.</figcaption></figure>

gland pores, *gd2*, *gd4*, *gd6*, *gd8*, *gd9*; *gd8* anteroantiaxial to *Z4*, *gd9* distant from *S5*. Peritremes extending to level between *j1–j3*. Sternal shield with two pairs of setae and posterior margin trilobate. Ventrianal shield subpentagonal, with four pairs of setae *JV1*, *JV2*, *JV3*, *ZV2*, and without preanal pores. Spermathecal calyx bell-shaped, with a short neck between the base of calyx and the atrium. Cheliceral fixed digit with two–three subapical teeth, movable digit with one tooth. Genua of legs II–IV with seven setae. Three macrosetae on leg IV. Male dorsal shield with six pairs of solenostomes; gland pore *gp* on the shield, marginal to the insertions of *s4*. Ventrianal shield with four pairs of setae. Spermatodactyl almost straight, a bit curved, not being able to distinguish where the foot begins; distal toe bifurcate; vellum present.

**Complementary description**

**Female** (eighteen examined, eight measured. Figures 11 a–b)

**Dorsal and lateral idiosoma** — (Figures 11 a–b). Dorsal shield with lateral waist slightly developed. Margins of the shield striate-reticulate, inner region of podonotum without reticulation, with fine furrows and crevices; opisthonotum striate-reticulate, cells very long, obliquely oriented; very noticeable muscle marks throughout the shield, many of them scattered between podonotal setae *j3–j4* and around setae *z5* (Figure 11a), and four large marks between gland pore *gd6* and insertions of *Z4* (Figure 11b). Eighteen pairs of smooth and acute dorsal
setae, except Z5 barely barbed on its distal half and J5 basally barbed. Lengths of j1 18–19, j3 22–24, j4 13–14, j5 12–15, j6 14–17, J2 14–19, J5 6–8, z2 13–16, z3 and z4 17–18, z5 13–16, Z4 27–29, Z5 41–43, s4 17–19, s6 20–23, S2 22–26, S4 21–25, S5 24–29. Sublateral setae r3 17–19, R1 16–17, placed on small platelets. Five pairs of dorsal gland pores, gd2, gd4, gd6, gd8, gd9; gland pore gd8 transversally aligned with poroid idm4, anteroantiaxial to the insertions of Z4; gd9 separated from and anteroparaxial to S5. Fifteen pairs of poroids on the shield; poroid idl2 placed on the lateral integument, near the level of S2. Peritremes ending between the setae j1–j3, detached from the dorsal shield, 170–207 long; internal groove with two continuous rows of microvilli.

**Ventral idiosoma** — Tritosternum 76–80 long, with two laciniae 55–57 long. Sternal shield subquadrate, with two pairs of setae and two pairs of poroids; posterior margin lobate; distance st1–st3 62–68, st2–st2 54–58; setae st3 and st4 on sclerotized platelets, that of st3 larger. Epigynal shield 104–117 long, distance st5–st5 62–66. Ventrianal shield pentagonal; 99–109 long, 87–94 wide at the level of ZV2, 64–67 wide at the level of anus. Anterior margin between insertions of JV1 straight, laterals between ZV2 and anus strongly concave, anterior corners between JV1–ZV2 sometimes eroded. Shield surface smooth, with four pairs of setae and without preanal solenostomes (Figure 16c). Setae ZV1, ZV3, JV4, JV5 and five pairs of poroids surrounding the ventrianal shield; setae JV5 33–36. Posterior metapodal shield 23–28 long, elliptical.

**Spermatheca** — Major duct short, proximal part near the sperm induction pore wide, apparently membranous, distal part near the atrium tubular, narrow and sclerotized; atrium nodular, a short neck between atrium and calyx; calyx bell-shaped, relatively small, 7–9 deep, 10–12 wide close to the vesicle.

**Gnathosoma** — Base of gnathosoma 26–30 long, 69–74 wide. Deutosternal groove with seven rows of paired and acute denticles. Hypostomal setae h1 22–24, h2 18–19, h3 20–22, pc 20–24. Length of first cheliceral segment 16–20, of second segment 76–83. Fixed digit of chelicera 26–29 long, with two or three subapical teeth; movable digit 22–25 long, with one tooth. Palp 98–105 long; femoral seta al thickened and distally spatulate, 14–15 long, genual setae al1–al2 thicken and blunt, similar in size, 8–9 long; 2-tined apotele.

**Legs** — Length of leg I 250–261, leg II 218–226, leg III 220–229, leg IV 293–299. Chaetotaxy of leg segments I–IV: coxae 2-2-2-1; trochanters 6-5-5-5; femora 12-10-6-6; genua 10-8-7-8; tibiae 10-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanter to tibia as follows. Leg I: trochanter 1 0/1, 1/2 1; femur 2 3/1, 2/2 2; genu 2 2/1, 2/1 2; tibia 2 2/1, 2/1 2. Leg II: 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: 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: 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. Three macroseta on leg IV; those of genu and tibia blunt or very scarcely knobbed, 24–27 and 23–28 long, respectively; that on basitarsus knobbed, 45–49 long.

### Male (six examined and measured)

**Dorsal and lateral idiosoma** — Dorsal shield reticulate, except on the midline of the podonotum, which is smooth; ornamentation on the opisthonotum as in female, but cells are smaller. Shield inversely oval, wider at the level of s4 (169–180) than at level of S4 (135–148), 252–290 long. Lengths of j1 16–18, j3 22–23, j4 11–13, j5 11–12, j6 11–15, J2 13–14, J5 6–7, z2 13–14, z3 13–15, z4 15–17, z5 12–14, Z4 25–27, Z5 36–37, s4 16–17, s6 17–18, S2 18–20, S4 17–20, S5 22–24, r3 15–19, R1 13–14. Six pairs of dorsal solenostomes, gd2, gd4, gd6, gd8, gd9, gp; gland pore gp very near the shield margin, hypertrophied, opening fundibuliform, placed in line with the insertions of setae s4. Poroid idl2 off the shield, as in females. Peritremes extending forward to setae z2 or between j3–z2, 135–140 long; peritrematal shield fused with the dorsal shield at the level of r3 insertions.

**Ventral idiosoma** — Sternogenital shield with five pairs of setae (st1–st5) and three pairs of poroids (iv1–iv3); distances st1–st5 107–119, st3–st3 54–57; anterior margin indiscernible, posterior margin gently convex; shield surface mostly smooth, with marginal striae. Ventrianal shield strongly convex anteriorly, subtriangular, transversally striated, 93–102 long, 137–153
wide; setae *JV1*, *JV2*, *JV3*, *ZV2* and four pairs of poroids (*iv5*, *ivo1-3*) on the shield; poroid *ivo2* in marginal position, *ivo3* slightly posteroparaxial to *ivo2* and equidistant from *ivo2* and setae *ZV2*. A fifth poroid *ivo*, poroids *ivp* and setae *JV5* off the shield; *JV5* smooth and pointed, 20–24 long.

**Chelicera and spermatodactyl** — Fixed digit of chelicera with two subapical teeth, 19–22 long; movable digit with one tooth, 18–21 long. Spermatodactyl almost straight, describing a smooth curve in which the foot is a continuation of the shaft; combined length shaft plus foot 29–33; foot ending in a bifid toe. Vellum present on the shaft.

**Legs** — Leg chaetotaxy and setal formulas like females. Leg IV with three macrosetae on genu 19–22, tibia 20–23, and basitarsus 37–39.

**Remarks**

*Typhlodromus* (*A.*) *maspalomensis* is very close to *T. (A.)* *henanoides* and both species may coexist on the same plants. The latter species can be easily distinguished by the following attributes: 1) calyx of spermatheca long tubular, about 20 micrometres, 2) occurrence of preanal pores, 3) only one macroseta on leg IV (at least the Canarian specimens), 4) setae Z4 long enough to reach the insertions of S5, 5) gland pore *gd9* adjacent to S5, instead of clearly separated from it, and 6) male spermatodactyl L-shaped rather than slightly curved in *T. (A.)* *maspalomensis*.

As stated by previous authors (Ferragut and Peña-Estévez, 2003; Stathakis *et al.*, 2016; Tixier *et al.* 2016) the species appears to have a close relationship with plants of the genus *Tamarix* (Tamaricaceae) on which it has been collected across its geographical range. Other host plants reported in Fuerteventura, e.g. Amaranthaceae, always were close to tamarisks.

**Typhlodromus (Anthoseius) rhenanoides Athias-Henriot, 1960**

*Typhlodromus rhenanoides* Athias-Henriot (1960): 85.

*Neoseiulus rhenanoides* – Schuster & Pritchard (1963): 205.

*Anthoseius rhenanoides* – Charlet & McMurtry (1977): 186.

*Amblydromella rhenanoides* – Tenorio *et al.* (1985): 303.

*Typhlodromus (Anthoseius) rhenanoides* – Ueckermann & Loots (1988): 60.

*Amblydromella (Aphanoseia) rhenanoides* – Denmark & Welbourn (2002): 308.

**Specimens examined**

Seven females and two males on unidentified Amaranthaceae, three females on *Suaeda vermiculata* (Amaranthaceae), one male on *Cakile maritima* (Brassicaceae); Dunas de Corralejo, 28°40'13"N, 13°50'00"W, 12 m a.s.l.; 27 May 2024. One female on *Suaeda* sp. (Amaranthaceae); Beach of Aljibe de la Cueva, El Cotillo, 28°39'53"N, 14°00'41"W, 18 m a.s.l.; 27 May 2024. Two females, two males and one deutonymph on *Limonium* sp. (Plumbaginaceae), two females on *Suaeda vera* (Amaranthaceae), two females, one male, one deutonymph on *Salsola divaricata* (Amaranthaceae), three females and two deutonymphs on *Caroxylon tetrandrum* (Amaranthaceae); Isla Lobos, 28°44'21"N, 13°49'21"W, 3 m a.s.l. Three females and one deutonymph on *Suaeda vermiculata* (Amaranthaceae); Isla Lobos, 28°45'13"N, 13°49'35"W, 15 m a.s.l.; 28 May 2024. Nineteen females, two males, one deutonymph, two protonymphs on *Andryala pinnatifida* Aiton subsp. *cuneifolia* (Schultz Bipontinus) M.Z. Ferreira (Asteraceae), one female on *Echium bonetti* Coincy (Boraginaceae); Betancuria, Mirador de Guise y Ayose, 28°26'27"N, 14°03'22"W, 694 m a.s.l.; 29 May 2024. One female on *Dittrichia viscosa* L. Greuter (Asteraceae); Betancuria, mirador Morro Velosa, 28°26'68"N, 14°02'59"W, 663 m a.s.l.; 29 May 2024. Three females on *Echium decaisnei* Webb & Berthelot subsp. *purpuriensis* Bramwell (Boraginaceae); Betancuria, 28°25'31"N, 14°03'24"W, 390 m a.s.l.; 29 May 2024. Four females, one male, one deutonymph, one protonymph on *Salsola divaricata* (Masson ex Link) Akhani (Amaranthaceae), one female on *Pulicaria* sp. (Asteraceae); La Pared, Pueblo del Mar, 28°12'28"N, 14°13'27"W, 69 m a.s.l.; 30 May 2024.
**Geographical distribution**

Mediterranean basin countries, Madeira, Canary Islands, Belgium. Also reported from California and Hawaii, although these records due to their biogeographical distance and age should be revised.

**Diagnosis**

Medium sized phytoseiid mite with relative short legs. Female dorsal shield evenly reticulated except on the striate anterolateral margins. Dorsal setae simple, smooth and pointed, only Z4, Z5, J5 with a few barbs. Setae *j1* and *j3* subequal in length; *z3* equal to *z4*; *z2* 2/3 the length of *z3* and *z4*; *s4* equal to *s6*; Z4 reaching the S5 sockets; dimensions of Z5 variable, 47–60. Five pairs of noticeable dorsal solenostomes (*gd2*, *gd4*, *gd6*, *gd8*, *gd9*). Poroid *idl2* detached from the shield. Peritremes ending near the level of *j1*; peritrematal channel not stippled, with an internal strip through all of its length. Sternal shield with two pairs of setae and posterior median lobe (inversely triangular). Ventrianal shield with four pairs of preanal setae; preanal pores small, punctiform, posteroparaxial to *JV2*. Calyx of spermatheca tubular, about 20 µm, atrium inserted into the base of calyx. Fixed digit of chelicera with four teeth, three subapical and smaller and one posterior to pilus dentilis and larger; movable digit with two teeth. Trochanter I with six setae, genua II–IV with seven setae. One slightly knobbed macrosetae on tarsus IV (other references mention three on leg IV, see Remarks). Male dorsal ornamentation as in female. Setae *r3*, *R1* on the dorsal shield. Six pairs of dorsal solenostomes, *gd2*, *gd4*, *gd6*, *gd8*, *gd9*, *gp*; gland opening *gp* on the margin of the dorsal shield, posterior to the level of *s4*. Cheliceral fixed digit with three teeth distal and one strong tooth proximal to pilus dentilis; movable digit with one large tooth. Spermatodactyl L-shaped with a short foot.

**Complementary description**

**Female** (52 examined, eight measured; Figure 12 a–c)

**Dorsal and lateral idiosoma** — (Figure 12a). Dorsal shield slender and elongate, 317–343 long, 145–166 wide; completely reticulate, except on the anterolateral margins, mainly striated; reticulation more extensive and marked on the opisthonotum. Eighteen pairs of setae, most of them smooth and pointed, Z4 with a few barbs, Z5 scarcely barbed, J5 basally barbed. Lengths of *j1* 23–26, *j3* 22–28, *j4* 13–18, *j5* 16–18, *j6* 18–21, *J2* 20–23, *J5* 9–10, *z2* 14–16, *z3* 20–25, *z4* 20–23, *z5* 16–19, *Z4* 32–41, *Z5* 47–60, *s4* 22–27, *s6* 25–28, *S2* 28–33, *S4* 32–38, *S5* 17–20. Sublateral setae *r3* 23–25, *R1* 22–27, on small basal platelets. Five pairs of very conspicuous solenostomes, *gd2*, *gd4*, *gd6*, *gd8*, *gd9*; gland opening *gd2* buttonhole-like, the remaining horseshoe-shaped; *gd9* adjacent to the insertions of setae *S5*. Fifteen pairs of poroids on the shield; poroid *idl2* always on the soft integument, posterior to setae *R1*. Peritremes extending anteriorly to almost the level of *j1*, 154–190 long; peritrematal groove without papillae or with just a few microvilli around the stigmata, with a continuous and internal strip (Figure 12a).

**Ventral idiosoma** — (Figure 12b). Tritosternum 77–84 long, split into two laciniae, 43–51 long. Sternal shield slightly sclerotized, almost smooth, with parallel sides and posterior margin with a median subtriangular lobe; bearing setae *st1–st2* and poroids *iv1*, *iv2*, the latter poroid just on its posterior margin. Each seta *st3–st4* on platelets, poroid *iv3* free on the unsclerotized cuticle, between the metasternal platelets. Distance *st1–st3* 65–70, *st2–st2* 51–56. Epigynal shield 104–111 long, distance *st5–st5* 57–64. Ventrianal shield broadly pentagonal (Figure 12b); anterior margin between setae *JV1* straight, laterals between *ZV2–JV3* distinctly concave. Surface of the shield almost smooth, with a few and weak transversal striae; with four pairs of setae (*JV1*, *JV2*, *JV3*, *ZV2*) and preanal solenostomes, punctiform and widely separated (distance between them 34–40), placed posteroparaxial to *JV2* and almost at the same level as the insertions of *JV3*. Elongate and narrow platelets on the soft cuticle, near the anterolateral corners of the shield. Posterior metapodal shield 22–28 long. Setae *ZV1*, *ZV3*, *JV4* and *JV5* around the ventrianal shield; *JV5* smooth and pointed, on platelets, 43–50 long.
<figure><img src="image_15.png" /><figcaption>**Figure 12** *Typhlodromus (Anthoseius) rhenanoides* female. a – peritreme showing the lineate peritrematal groove. b – ventrianal shield. c – spermatheca.</figcaption></figure>
**Spermatheca** — (Figure 12c). Major duct short, narrow and cylindrical; atrium c-shaped just on the base of calyx; calyx tubular, 19–22 long, lateral walls almost parallel, flaring distally, 7–11 wide at the junction with the vesicle. The proximal portion of calyx near the atrium is less sclerotized and may appear folded.

**Gnathosoma** — Base of gnathosoma 40–54 long, 75–84 wide. Tectum dome-shaped, without a clear anterior angle. Deutosternal groove with seven rows of two lateral denticles. Hypostomal setae *h1* 23–25, *h2* 19–21, *h3* 24–25, *pc* 26–28. First cheliceral segment 20–25 long, second segment 81–85. Fixed digit of chelicera 27–30 long, with four teeth, three anterior and one larger and triangular (in profile) posterior to pilus dentilis; movable digit 24–26, with two teeth, the proximal being larger than the distal. Palp 113–118 long; femoral seta *al* 15–16 long, flattened and expanded at the tip; on the genu *al1* 9–10, *al2* 7–8 long, both thick and pointed.

**Legs** — Five-six calciciform glands and a large presumably gland opening on ventral side of coxa I; the opening elongate, piriform, in anteroventral position. Length of leg I 255–282, leg II 213–228, leg III 220–228, leg IV 297–305. Chaetotaxy of leg segments I–IV: coxae 2-2-2-1; trochanters 6-5-5-5; femora 12-10-6-6; genua 10-8-7-8; tibiae 10-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanter to tibia as follows. Leg I: trochanter 1 0/1, 1/2 1; femur 2 3/1, 2/2 2; genu 2 2/1, 2/1 2; tibia 2 2/1, 2/1 2. Leg II: 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: 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: 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. One macroseta on tarsus IV, slightly knobbed, 38–51 long.

**Male** (nine examined, five measured)

**Dorsal idiosoma** — Dorsal shield 257–264 long, wider at the podosoma (147–153) than at the opisthosoma (135–139); evenly reticulate except for the striated anterolateral margins, cells elongate and oriented axially between setae *j5-j6*, pentagonal and regular in size between setae *j6-J2*. Twenty pairs of dorsal setae morphologically similar to those of female. Lengths of *j1* 17–18, *j3* 19–22, *j4* 12–14, *j5* 11–14, *j6* 13–16, *J2* 15–17, *J5* 8–9, *z2* 12–17, *z3* 15–16, *z4* 14–20, *z5* 11–14, *Z4* 28–33, *Z5* 35–40, *s4* 16–22, *s6* 19–24, *S2* 21–26, *S4* 20–26, *S5* 14–18, *r3* 16–17, *R1* 15–18. Six pairs of dorsal solenostomes, without gland pores *gd1* and *gd5*, but with pore *gp* placed just on the shield margin, a bit posterior to the level of *s4*; all the gland openings more developed than on the female, horseshoe-shaped, except *gp*, oval in shape. Peritrematal shield attached to the dorsal shield at the level of *r3*; peritremes 127–131 long, ending anteriorly between *j3-z2*; peritrematal groove with a few papillae in the proximal section, near the stigmata, the remaining lineate, without microvilli, as in females.

**Ventral idiosoma** — Sternogenital shield mostly smooth; distance *st1-st5* 108–113, *st3-st3* 49–54; posterior margin between setae *st5* smoothly convex. Ventranal shield subtriangular, 101–108 long, 139–143 wide; surface transversally striate, with some closed cells anterior to preanal pores; four pairs of setae (*JV1*, *JV2*, *JV3*, *ZV2*), and four pairs of poroids (*iv5* and three pairs *ivo*) all anterior to preanal solenostomes; preanal pores *gv3* punctiform and posterior to the insertions of setae *JV2*, distance between them 23–28. Setae *JV5* smooth and pointed, 23–26 long.

**Chelicera and spermatodactyl** — Fixed digit of chelicera 22–25 long, with three teeth distal and one strong tooth proximal to pilus dentilis; movable digit 18–22 long, with one large tooth. Spermatodactyl L-shaped; shaft 17–19 long, a bit curved; heel rounded, little marked; foot short (about 5–6 µm), directed upwards, ending in small toe.

**Legs** — Same leg chaetotaxy as in female. Macroseta on tarsus IV blunt or knobbed, 31–33 long.

**Deutonymph** (seven examined, five measured. Figure 13 a–b)

**Dorsal and lateral idiosoma** — (Figure 13a). Elongate dorsal shield, 248–268 long, 110–114 wide; laterals incised between the insertions of setae *s6* and *S2*, incision reaching the gland opening *gd6*. Surface mostly reticulate, cells more abundant and pronounced on the opisthonotum. Eighteen pairs of simple and pointed setae, except *Z4*, *Z5*, finely barbed and *J5* basally barbed. Lengths of *j1* 19–20, *j3* 16–17, *j4* 10–12, *j5* 12–13, *j6* and *J2* 14–16, *J5* 8–9,
<figure><img src="image_16.png" /><figcaption>**Figure 13** *Typhlodromus (Anthoseius) rhenanoides* deutonymph. a – dorsal surface. b – ventral surface.</figcaption></figure>

z2 11–12, z3 11–14, z4 13–14, z5 11–13, Z4 35–38, Z5 40–46, s4 17–19, s6 17–20, S2 23–24, S4 24–26, S5 15–16. Sublaterals r3 18–20, R1 16–17. Same adenotaxy (absence of gland pores gd1, gd5) and poroidotaxy (complete set of sixteen poroids) than on female. Peritremes extending to the level between j3–z2, 124–141 long; peritrematal groove with an incomplete row of microvilli.
**Ventral idiosoma** — (Figure 13b). Tritosternum 77–82, with two laciniae 48–51 long. Sternogenital shield with four pairs of setae and two pairs of poroids, setae *st5* off the plate; distance *st1–st5* 124–136, *st3–st3* 48–50. Caudoventral soft cuticle surrounding the anal plate with eight pairs of setae (*JV1*, *JV2*, *JV3*, *JV4*, *JV5*, *ZV1*, *ZV2*, *ZV3*); pair *ZV3* half as long as companions. Setae *JV5* smooth and pointed, on platelets, 29–33 long.

**Gnathosoma** — Gnathosomal base 34–38 long, 66–70 wide. Deutosternal channel with seven rows of paired denticles. Hypostomal setae h1 21–22, h2 18, h3 19–20, pc 21–23. First cheliceral segment 21–22, second segment 70–73. Fixed digit of chelicera 21–24 long, with four teeth, three anterior and one posterior to pilus dentilis; movable digit 19–22, with two small teeth. Palp 97–102; anterolateral seta on femur spatulated, al1, al2 on genu thickened.

**Legs** — Coxa I with eight calciform glands arranged into two groups of four; anteroventral opening present. Same chaetotaxy and setal formulas as the female. Only one macroseta on basitarsus IV, usually knobbed, 41–46 long.

**Protonymph** (three examined and measured. Figures 14 a–b)

**Dorsal and lateral idiosoma** — (Figure 14a). Podonotal shield smooth, weakly sclerotized, 125–136 long, 104–115 wide. Solenostomes and poroids not discernible on the shield. Nine pairs of setae on the shield; length of *j1* 17–18, *j3* 14, *j4* 11–12, *j5* 10–11, *j6* 14–15, *z2* 12–13, *z4* 13–14, *z5* 10–11, *s4* 17. Setae *s6*, *r3*, *R1* off the shield; *s6* 16–17, *r3* 17–18, *R1* 12–13. Setal pairs *J2* and *S2*, and six pairs of mesonotal scutella, one of the platelets bearing the gland pore *gd6* between podonotal and pygidial shields. Setae *J2* 13–14, *S2* 21–22. Pygidial shield finely reticulated, anterior margin convex, 53–57 long, 66–68 wide. Setae *S4*, *S5*, *Z4*, *Z5*, *J5*, and poroids *idl3*, *idl4*, *idl5*, *idm5* on the shield; setae *Z4* and *Z5* scarcely barbed, *J5* basally barbed. Lengths of *J5* 8–9, *Z4* 39–44, *Z5* 34–36, *S4* 19–21, *S5* 14. Peritremes 31–37 long, almost reaching the level of setae *s4*; peritrematal channel with papillae or microvilli.

**Ventral idiosoma** — (Figure 14b). Tritosternum 67–68 long, laciniae 46–48. Sternogenital shield poorly sclerotized, with setae *st1–st3* and poroids *iv1–iv2*; distance *st1–st3* 71–76, *st2–st2* 44–46. Opisthoventral soft integument with setae *JV1*, *JV2*, *ZV2*, and *JV5* around the anal plate; solenostomes *gv3* on the plate, posterior to *JV2*, punctiform, minute. Setae *JV5* 20–21 long.

**Gnathosoma** — Base of gnathosoma 45–47 long, 59–63 wide. Deutosternum with seven rows of denticles. Setae *h1* 18, *h2* 14–15, *h3* 16–18, *pc* 19–20. First cheliceral segment 14–16, second segment 56–59 long. Cheliceral fixed digit 20–21 long, with three teeth anterior and one tooth posterior to pilus dentilis; movable digit 17–18 long, with two small denticles. Palp 83–89 long; setae *al* on femur 12–13 long, distally expanded; setae *al* on genu 8–9, spiniform.

**Legs** — Three calciform glands and one small and punctiform pore on the ventral surface of coxa I. Chaetotaxy of leg segments I–IV: coxeae 2-2-2-1; trochanters 4-4-4-4; femora 10-7-5-4; genua 8-6-6-5; tibiae 8-7-7-6; basitarsi 0-4-4-4. Setal formulas from trochanter to tibia as follows. Leg I: trochanter 1 0/1, 0/1 1; femur 2 2/1, 2/1 2; genu 1 2/1, 2/1 1; tibia 1 2/1, 2/1 1. Leg II: trochanter 1 0/1, 0/1 1; femur 1 2/1, 2/0 1; genu 1 2/0, 2/0 1; tibia 1 1/1, 2/1 1. Leg III: trochanter 1 0/1, 0/1 1; femur 1 2/1, 1/0 0; genu 1 2/0, 2/0 1; tibia 1 1/1, 2/1 1. Leg IV: trochanter 0 0/2, 1/1 0; femur 1 2/0, 1/0 0; genu 1 2/0, 2/0 0; tibia 1 1/1, 2/0 1. Macrosetae on genu (*ad1*) and tibia (*ad*) IV differentiate; lengths of macroseta on genu 15–17, on tibia 18–19, on tarsus 40–42.

**Larva.** Not found.

## Remarks

Athias-Henriot (1960) pointed out that this species was the most abundant and widespread in Algeria, in both coastal and semi-arid inland landscapes. Likewise, according to Tixier *et al.* (2016), it appears to be very common and abundant in Morocco. Although the phytoseiid fauna in the Canaries is still incompletely known, the available information suggests that *T.* (*A.*) *rhenanoides* is likely the most abundant and frequent phytoseiid throughout the archipelago (unpublished data).
<figure><img src="image_17.png" /><figcaption>**Figure 14** *Typhlodromus (Anthoseius) rhenanoides* protonymph. a – dorsal surface. b – ventral surface.</figcaption></figure>
Since its original description it has been redescribed several times, although some important
morphological features were overlooked by previous authors and the morphology of immature
stages was completely unknown. There is no mention in the literature to the peculiar peritrematal
groove, which is devoid of microvilli or papillae and is instead furrowed by a continuous strip
from the stigmata to the end of the peritreme (Figure 12a). In some females there are papillae
around the stigmatal opening and at the beginning of the channel. All the previous illustrations
in the literature show a "normal" and stippled peritreme, but the excellent photograph by Döker
et al. (2022) (their Figure 5a) reveals the lineate structure on females collected in Belgium. It
would be desirable to check this character in mites from other geographical locations to confirm
whether it is universal in the species or restricted to certain populations.

Another trait that raises questions is the shape of the sternal posterior margin in females. Athias-Henriot neither mentioned or illustrated this character. In Ueckermann and Loots (1988) and Moraes et al. (1999) the margin is described as straight or concave, whereas according to Papadoulis et al. (2009), Ferragut et al. (2010), Faraji et al. (2011), and Tixier et al. (2016) it is lobate. Again, this character should be mentioned in future redescriptions, and ideally revisited, in particular in specimens collected far from the supposed centre of geographical distribution, the western Mediterranean and North of Africa. Similarly, some reports indicated the absence of preanal pores (e.g., Charlet and McMurtry, 1977; Faraji et al., 2011), although the occurrence of this trait is evident in other redescriptions and it was highlighted in Athias-Henriot's original description, where she depicted them and mentioned their remarkable separation. Finally, the number of macrosetae on leg IV is variable in the literature. The original description includes measures of three macrosetae, and this number has been reported by Schuster and Pritchard (1963), Ueckermann and Loots (1988), Faraji et al. (2011), Tixier et al. (2016), Döker et al. (2022), and it occurs in females from the Iberian Peninsula (per. observation). However, other references report a single macroseta (Papadoulis et al., 2009). These discrepancies may result more from differing criteria for identifying a macroseta than from a real morphological variation. Nevertheless, the females from Fuerteventura do not have the dorsal setae *ad1* on genu and tibia IV differentiate as macrosetae. The length of those setae in the females is only 2-3 micrometres longer than the remaining setae on the segment; on the contrary, in the protonymph these setae are clearly longer. Thus, the absence of three macrosetae seems to be characteristic of the Canarian populations, as females from Gran Canaria and Tenerife Islands only have one macroseta. Accordingly, the mentioned morphological variability should be traced using molecular techniques on specimens from different locations to determine whether they represent a single species or, conversely, they hide a complex of closely related taxa.

Geo-environmental dynamism reveals patterns of island
colonization

A biogeographical approach to better understand the occurrence of phytoseiid species in the Canaries must be based on the geo-environmental dynamics of the islands since their emergence from the Atlantic Ocean and on the origin and dispersal pattern of host plants on which the mites thrive. The seven main islands in the archipelago are, more or less, arranged in a linear age sequence from the continental mainland, decreasing in age with greater distance from Africa. Fuerteventura lies barely 96 km from the African littoral and it is the oldest island, estimated to have emerged above sea level between 20-24 Ma (million years ago) (Carracedo and Troll, 2021). Geologically, each oceanic island undergoes a life cycle or ontogeny with several stages: since its emergence breaking the sea surface, to further growth to great heights increasing its topographic complexity, followed by erosion and lengthy island subsidence, and eventually, both processes reducing the island to a seamount again (Whittaker et al., 2024). Hence, Fuerteventura is an "old oceanic island" on which the high volcanos have long since disappeared due to erosion, though these structures are still evident in the landscape. There are no high altitudes, and as a result, clouds from the ocean do not release rain, rendering the island semi-desertic, almost devoid of trees, and similar to the nearby African coast.
As this short review illustrates, Fuerteventura is under extreme environmental conditions, particularly in terms of low humidity, scarce water resources, and limited habitat complexity, and the mites living there must thrive in these harsh conditions. Unfortunately, the phytoseiid fauna on the whole archipelago and the near African coast is only partially known or completely unknown and, referring to the species' geographic origin, I can only offer tentative explanations, aiming to provide at least the foundations for further and improved predictive models of both species richness and insular colonization patterns.

## Species richness

Even though different habitats were surveyed over several days, only five species were collected. This limited diversity contrast with Fuerteventura's land surface (about 1,670 km²) which is second only to Tenerife in the archipelago, and the expected species-area relationship, which predicts a rise in species richness with increased land surface. However, in this case, other interlinked factors have a greater predictable power, as for example the island age, elevation, climate and habitat diversity. The mean altitude of the island is only 44 m, and most of the landscape is dominated by plains and lowlands, with just one southern mountain range reaching 800 m a.s.l. This topographical simplicity is a consequence of the island age, which in turn influences the other factors. Low oceanic islands tend to have dry climates, and islands that undergo a long period of subsidence and erosion have limited habitats and microclimates for future colonizers (Leihy et al. 2018). Surely, Fuerteventura once hosted a richer mite fauna, but the geo-environmental changes over time have greatly reduced ecological opportunities and driven most of species to extinction.

## Biogeographical origin of species

The origin of the species collected can be inferred from their currently known geographic ranges and the distribution of their host plants. *Typhlodromus* (A.) *rhenanoides*, *T. (A.) maspalomensis* and *N. litoralis* have wide distribution ranges that span the Mediterranean basin and are common and locally abundant in Northwest Africa and Southwest Europe. Hence, they are strong candidates for having colonized the islands via long-distance dispersal from the African continent. Phytoseiids disperse mostly by aerial means aided by their small size (Sabelis and Dicke, 1985), and the Canaries are influenced by constant northeast to southwest trade winds that enable the transport of small animals and plant propagules from the continent. Both *T. (A.) rhenanoides* and *N. litoralis* are commonly found on a wide range of host plants from different plant families, but *T. (A.) maspalomensis* is more plant-specific, mainly retrieved from *Tamarix* (Tamaricaceae) species. Three tamarisks occur in the Canaries, *T. africana*, *T. canariensis* and *T. boveana*, all of which also inhabit Southwest Europe, Morocco, and other Macaronesian islands (Sauebier et al. 2023). The distribution of the preferred host-plant may not only explain mite distribution, but also play a crucial role in providing ecological opportunities for colonizing mites when they arrived to new environments already furnished with suitable habitats.

*Typhlodromus* (A.) *charactus* exhibits a curious disjunct distribution. It was described from Yemen and this remains the only known record of the species. This distribution bridging the Arabic peninsula and the Canary Islands via northern Africa is congruent with the biogeographical concept of “Rand flora”; that is, the disjunct distribution in many plant lineages between Macaronesia–Northern Africa, Southwest Asia, and East–South Africa (Sanmartín et al. 2010; Pokorni et al. 2015). Several hypotheses have been proposed to explain the evolutionary origin of this flora, but what is relevant here is that phytoseiid mites are carried by plants and it is not possible to understand mite movements accurately without considering the long-distance displacements of their host plants. A plausible explanation for their arrival in the Canaries involves a route across the huge Sahara Desert. In the current scenario, the Sahara is a barrier to the species dispersal, but its desert character is relatively recent in geological terms (it began forming 7 My ago) and the desert was once a verdant land that sustained rich and abundant live communities enabling the free movement of plants and animals across
northern Africa (the desert experienced several intervals of wet and dry climates, but current aridity in the Sahara began only 4500–4000 years ago. See Williams (2023) for a readable and comprehensive account). Finally, *N. extraseta* sp. nov. is likely a Canarian endemic. However, based on current knowledge, we cannot safely report that. The narrow coastal strip in southwest Morocco, which harbour vegetation quite similar to that of the eastern Canary Islands, remains unexplored. This taxon could have originated in situ on the islands by diversification from a former *Neoseiulella* species, or could have conspecific populations thriving in the African littoral or, even, being a relicual form once present in the continent but now extinct on the mainland following island colonization.

## Acknowledgements

The author was provided with a permit to collect plants by the Cabildo de Fuerteventura (Resolución 1087/2024).

## ORCID

Francisco Ferragut ID https://orcid.org/0000-0003-1545-6011

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