Lindquist, Evert E. ¹ and Moraza, Maria L. ²

¹Canadian National Collection of Insects, Arachnids, and Nematodes, Science & Technology Branch, Agriculture and Agri-Food Canada, Ottawa, ON, K1A 0C6, Canada.
²✉ Universidad de Navarra, Facultad de Ciencias, Departamento de Biología Ambiental, Campus Universitario, 31080 Pamplona, España.

2023 - Volume: 63 Issue: 4 pages: 998-1016

ZooBank LSID: CAD31ED5-B03E-4856-B9A4-FBF5F94E07CF

Original research

Keywords

Abstract

Introduction

The taxon Lasioseius (Endopodalius) was first proposed as a subgenus by Christian and Karg (2006) in their review of the predatory mite genus Lasioseius Berlese, 1916. Over forty years previously, the same taxon had been recognized and proposed as the ''Scutalis Group» of the genus Lasioseius in the unpublished thesis by Lindquist (1964), and thereafter by Lindquist and Evans (1965, p.54) who listed all of the then-named species in that group. The same species group was also referred to as the ''Lasioseius alter group» by Christian and Karg (2006, p. 135). More recently, the taxon named as Endopodalius has continued to be recognized as a subgenus of Lasioseius by various authors (Lindquist and Moraza, 2010, 2012; Moraza and Lindquist, 2011, 2018; Moraes et al., 2016; Abo-Shnaf et al., 2016; Argolo et al., 2018; Quintero-Gutiérrez et al., 2020; Moraza and Balanzategui, 2023). The relationship of Endopodalius to Lasioseius has been based on various morphological aspects, including the absence versus presence of a phytoseioid sperm access system (Alberti, 2002; Di Palma and Alberti, 2005), which latter is a basic autapomorphic characteristic of the superfamily Phytoseioidea sensu Lindquist et al. (2009; see below), to which Lasioseius (Blattisociidae) formally belongs. Loots (1980) particularly illustrated the spermatheca of Lasioseius mouchei Loots, 1980 (herein recognized as belonging in Endopodalius), as did Moraza and Balanzategui (2023) for L. (Endopodalius) ibericus Moraza & Balanzategui, 2023. However, their descriptions did not describe or distinguish these structures as of the phytoseioid-type, and their illustrations do not show any detailed sclerotized structures definitive of the phytoseioid-type, the latter characterized by a sclerotized calyx and associated embolus (Evans, 1992; Alberti and Coons, 1999; Alberti, 2002) from which a long, fine accessory duct leads to a secondary structure, the spermathecal capsule (Athias-Henriot, 1971), which sometimes is elaborated into species-specific forms (Lindquist, 1964; Walter and Lindquist, 1997). Otherwise, the sperm access system had not been illustrated or described in the female of any of the eight species of Lasioseius assigned to Endopodalius by Christian and Karg (2006), who overlooked assigning Lasioseius imitans (Berlese, 1910), L. americanus Chant, 1963, and L. mouchei Loots, 1980 to their subgenus. In addition, females of Lasioseius gabrielae Santos and Argolo, 2018 (in Argolo et al., 2018), and of a further, five undescribed Neotropical species of Endopodalius examined by one of us (E.E.L.) do not present sperm access structures of the phytoseioid-type.

Material and methods

Studied specimens were previously prepared, sometimes over a century ago (e.g., Berlese, 1910; Banks, 1914), mounted on microslides in a variety of media, e.g., either resin-based, permanent Canada balsam or water-miscible Hoyer's or a similar modification of Berlese's gum chloral medium (Singer 1967) and sealed with asphaltum or GLPT insulating varnish.

Morphological observations and illustrations were made using compound microscopes equipped with differential interference contrast (DIC) and phase contrast optical systems, drawing tubes and stage-calibrated eyepiece micrometers. Setal notation for the idiosoma follows that of Lindquist and Evans (1965) as modified slightly by Lindquist (1994). Notation for leg and palpal setation follows that of Evans (1963, 1964, 1969). Distinction of pore-like structures on the idiosomal integument as either poroids (lyrifissures) or glandular openings (solenostomes), as distinguished morphologically by Athias-Henriot (1969) and physiologically by Krantz and Redmond (1987), is applied. Notation for pore-like structures of the peritrematal region follows that of Johnston and Moraza (1991).

Observational notes were made (E.E.L.) from studies of primary type material of each of the following species accorded herein to Lasioseius (Endopodalius): Lasioseius (Zygoseius) alter Vitzthum, 1925, Zoologische Sammlung des Bayerischen Staates, Munich; Lasioseius americanus Chant, 1963, Canadian National Collection of Insects, Arachnids, and Nematodes (CNCI), Agriculture & Agri-Food Canada, Ottawa; Lasioseius convexus Krantz, 1962, Oregon State University, Corvallis; Lasioseius humberti Athias-Henriot, 1959, Laboratoire d'Acarologie de l′École Pratique des Hautes Études, Paris and Hochschule für Bodenkultur, Vienna; Lasioseius (Endopodalius) ibericus Moraza & Balanzategui, 2023, Universidad de Navarra; Lasioseius imitans Berlese, 1910, Research Centre for Plant Protection and Certification (CREA-DC), Florence; Hypoaspis scutalis Banks, 1914, Museum of Comparative Zoology, Harvard University, Cambridge; Lasioseius tectus Hyatt, 1964, British Museum (Natural History), London.

Background systematics

The systematic placement of Endopodalius had been problematical prior and subsequent to the description and illustration of the spermathecal structure of any of its species (e.g., by Loots, 1980, figure 28 for mouchei; by Esteca et al., 2020, figure 3F for prorsoperitrematus Abo-Shnaf et al., 2016; and by Moraza and Balanzategui, 2023, figure 2D for ibericus). What are its sister-group relationships, and in what family of the cohort or infraorder Gamasina should Endopodalius be placed? Using the key to families of the order Mesostigmata presented by Lindquist et al. (2009), the taxon Endopodalius readily reaches couplet 59, where it nearly entirely ′fits' 59b for Blattisociidae, except for one major aspect – the form of the female sperm access system not having any of the apomorphic aspects of the phytoseioid-type, as characterized by Evans (1992; see below). Alternatively, female Endopodalius do not share the peculiarities of couplet 59a for Melicharidae, particularly apomorphic aspects of the fixed and movable cheliceral digits (inflated form of pilus dentilis and mucronate midventral process, respectively), and form of the epigynal and opisthogastric shields.

If Endopodalius females have a sperm access system that has become degenerated, simplified, reverting as an atavism to a variant of the laelapoid-type form (see figures in Athias-Henriot, 1968), this would follow the contentious notion of Karg (1998, 2003, followed by Christian & Karg, 2006) that at a higher phylogenetic level, sperm access systems are insignificant because both types can be found within various families, and thus allowing for retention of Endopodalius in the Blattisociidae. However, Karg's notions regarding evolution of dermanyssine sperm-access systems were evaluated as unfounded by Alberti (2002).

It should be noted that two of the species currently assigned to Endopodalius, Lasioseius (Zygoseius) alter Vitzthum, 1925 and L. (Z.) vitzthumi Westerboer, 1963, were placed by Westerboer (1963) in Zygoseius Berlese, 1916, a previously monotypic genus based on Zygoseius furciger Berlese, 1916. Westerboer distinguished her placement of Zygoseius as a subset of Lasioseius on its species having smooth, pointed dorsal body setae; the sternal shield having large extensions between the bases of legs I/II and of legs II/III; very strongly developed endopodal plates extending beneath the metasternal platelets; and shortened hypostomatic setae C3 and C4 (hypostomatic setae hyp 2 and capitular setae c.s., respectively). To these attributes, there can be added that, unlike Lasioseius (other than Endopodalius), females have a sternal shield with convex posterior margin, and males have a holoventral shield. The systematic placement of Zygoseius remains problematical, as reviewed by Halliday (1997). However, Zygoseius is currently thought to be placed in the Pachylaelapidae, superfamily Eviphidoidea, not closely related to the families of Ascoidea (Lindquist et al., 2009), in part based on such attributes as a three-tined palptarsal claw, leg I femur with 13 setae (v4 present), dorsal shield divided in the deutonymph but entire in adults, and other aspects (Halliday, 1997; see Discussion, below).

Recently, Esteca et al. (2020, figure 3F) included a micrograph of the spermatheca of Lasioseius (Endopodalius) prorsoperitrematus, and Moraza and Balanzategui (2023, figure 2D) drew the spermathecal remnant of L. (E.) ibericus. These authors did not identify the type of spermatheca illustrated; however, unlike those of species of Lasioseius, the structures are not configured as of the phytoseoid-type spermathecal apparatus.

Furthermore, other distinctive attributes of adult Endopodalius are unlike those of Lasioseius: (1) dorsal shield humeral setae r3 are not apomorphically erect or modified in form; (2) nearly all dorsal shield setae are simple, none apomorphically tricarinate, although Z4 and/or Z5 may be penicillate; (3) gnathotectum with apomorphically elongate, apically split median branch; (4) female movable chela multidentate, apomorphically with four or more teeth (the movable chela among Lasioseius species generally has three teeth, though exceptionally eight teeth in Lasioseius ruehmi Hirschmann, 1972; see below); (5) female sternal shield with posterior margin apomorphically convex (other than its general convexity, the posterior margin may be rarely slightly concave medially); (6) tritosternum autapomorphically with small sclerite adjacent to its base; (7) endopodal plates beside coxae III and IV expansive, autapomorphically underlapping metasternal platelets; (8) male with holoventral shield. Concerning attributes (1) and (2), exceptionally, in adults of Lasioseius angustus Evans & Sheals, 1959, L. frontalis Evans & Sheals, 1959 and L. polydesmophilus Evans & Sheals, 1959, humeral setae r3 are not erect, and all of the dorsal shield setae are simple in form. These are secondary modifications derived in species that have become closely associated with millipedes — their adults conformed closely to the shapes of the spiracular despressions in which they are found on their hosts (Evans and Sheals, 1959).

In view of the above, the genus Endopodalius is so distinctive apomorphically to justify its own family grouping, even lacking information on the attributes of the immature instars except for a drawing of the larval dorsal idiosoma of E. araucariae (Hirschmann, 1972, figure 43RL; copied in Christian & Karg, 2006, figure 3.2.2). To be placed among the families and superfamilies of free-living Gamasina as currently recognized, the taxon Endopodalius or scutalis group of species presents attributes that currently distinguish the superfamily Ascoidea (see below). However, based on current classificatory concepts (Lindquist et al., 2009; Karg, 1993, 2006), distinctions between the superfamilies Ascoidea, Eviphidoidea and Dermanyssoidea are not discrete among the more derivative taxa associated with terrestrial insects and vertebrates, and Endopodalius mites associated with coleopterans are another example.

Whether sensu Lindquist et al. (2009) or Karg (1993, 2006), members of Ascoidea have a usually two-tined palptarsal apotele; adult males have a presternal genital aperture (Karg's concepts accept either a presternal or midsternal genital aperture); the adult male spermatodactyl arises from external face of movable digit, with an open groove, freely projecting distally (Karg's concepts accept presence or absence of a spermatodactyl); adult females have metasternal elements free from the sternal shield; adult males have posterior (caudal) margins of the dorsal and anal or ventrianal shields free from one another (Karg's concepts accept conjoining of those margins as they occur in Zerconidae, in Ascoidea); adults have microtrichia of cribrum confined to the postanal region of anal or ventrianal shield (not extended onto caudal margin of dorsal shield); tibia I maximally has 13 setae, lacking a fourth ventral seta (pv-2) (Karg's concepts allow 14 setae, including pv-2, as they occur in Zerconidae); genu IV maximally has nine setae, lacking pv (Karg's concepts allow ten setae, including pv, as they occur in Zerconidae).

The above paragraph leads us to consider an accommodation of the new taxon in the superfamily Ascoidea, based further in part on it having a plesiomorphic ''type A» laelapoid-type sperm access system rather than the apomorphically derivative phytoseioid-type system typical of the superfamily Phytoseioidea (see Alberti 2002). Based on cheliceral and leg chaetotactic attributes detailed by Lindquist et al. (2009), the new taxon is not readily accommodated among free-living and insecticolous taxa within the superfamilies Eviphidoidea and Dermanyssoidea, while the superfamily Ascoidea is characterized by a series of many attributes, as presented above, which readily accommodate Endopodalius.

Within the Ascoidea, however, the new taxon is not readily accommodated in any of the four recognized families as currently defined, Ascidae, Ameroseiidae, Melicharidae, and Antennochelidae, based on the following rationale. Unlike Ascidae, its adult females do not have the third pair of sternal poroids inserted plesiomorphically on the posterior margin of the sternal shield (they are on metasternal platelets), and its males do not have the endopodal strips beside coxae III-IV apomorphically free or at most narrowly connected to the sternitigenital shield (they are fully consolidated with that shield, as well as further consolidated into a holoventral shield).

Unlike Ameroseiidae, Endopodalius adults do not have the following apomorphic attributes: absence of the pair of caudal setae J5 (plesiomorphically present in Endopodalius); absence of R-series of marginal setae other than R1 (a full R-setal series present in Endopodalius); femur II with usually stable seta al-2 absent (Endopodalius femur II retains seta al-2 basal to the lyrifissure); and the basal row of deutosternal denticles is disrupted medially and widened laterally to extend beyond insertions of the capitular setae (basal row of deutosternal denticles not disrupted medially and not widened laterally in Endopodalius).

Unlike Melicharidae, in Endopodalius the corniculus lacks the small, apparently stiff, acuminate apomorphic projection arising paraxially from its base; the pilus dentilis of the fixed cheliceral digit is setiform, not apomorphically modified into a hyaline flap; the movable cheliceral digit lacks a pointed apomorphic projection, the mucro, on its mid-ventral face; the posterior margin of the adult female epigynal shield is strongly truncated, rather than being gently convex; the adult female has a well-developed ventrianal shield bearing at least four pairs of opisthogastric setae, rather than an ovate or elliptical anal shield bearing usually only the anal setae or rarely with the nearest one pair of opisthogastric setae; the adult male has a holoventral shield rather than a typical ventrianal shield abutting or coalesced with, but delineated from, a sternitigenital shield.

Adult Endopodalius resemble more those of blattisociids in having the peritrematal shield moderately broadly connected to the exopodal plate beside coxa IV, rather than being free from or narrowly connected to it, as in melicharids. The more plesiomorphic laelapoid form of the female sperm access system of Endopodalius, as illustrated by Abo-Shnaf et al., 2016, for prorsoperitrematus (their figure 50), by Loots, 1980, for mouchei (his figure 28), and herein for ibericus Moraza and Balanzategui, 2023 (Figure 2G), resembles that found in Melicharidae (e.g., as shown by Fain et al., 1977 and by Naskrecki and Colwell, 1998) and in Ascidae, but as also found in other families (Athias-Henriot, 1968), so it is not diagnostic in differing from the autapomorphic phytoseioid-type.

Unlike Antennochelidae Lindquist and Moraza, 2014, a taxon based on a single genus characterized mostly by apomorphic gnathosomatic attributes, the cheliceral shafts of adult and nymphal Endopodalius are not greatly elongated and projectable, body-length, beyond the gnathosoma; the subcapitulum of Endopodalius adults does not have the four pairs of hypostomatic setae aligned longitudinally (hyp2 and hyp3 are aligned transversely); the gnathotectum is not modified into an elongated triramous extension (only the mid-tine is elongated); the sternal and peritrematal shields are not broadly fused at the level of endopodal extensions between legs I and II (they are broadly adjacent, contiguous in Endopodalius); the intercoxal region of adults lacks a strongly sclerotized pair of invaginations associated with poroids iv2; and trochanter I retains seta ad, femur II retains al-2 basal to the lyrifissure, genu III retains pv-1 (these setae absent in Antennocheles).

Linking Endopodalius as a sister-group to any of the above families is problematical. However, the following attributes argue for some commonality with the morphologically more highly derived Antennochelidae: (1) Female sternal shield with a somewhat convex posterior margin, more strongly so in Endopodalius: other than in the genus Zygoseius (see below), this attribute is rarely found among other gamasines of the hyporder or subcohort Dermanyssiae. (2) Adult endopodal sternal anterolateral extensions broadly projecting and fused or contiguous with exopodal-peritrematal plates between legs I and II: these sternal extensions are generally more narrowly projecting and free from or narrowly contiguous with exopodal-peritrematal plates in other gamasines. (3) Adult endopodal plates strongly developed between legs III and IV; in Antennocheles (and in Zygoseius) these plates are nearly as strongly developed as in Endopodalius, although they do not extend under the metapodal platelets. (4) Gnathotectum with elongated tines; in Endopodalius only the mid-tine is elongated. (5) Male with holoventral shield: this attribute is not found elsewhere among taxa of Ascidae, although this is characteristic of the genus Zygoseius, sometimes having been related with Lasioseius (Westerboer, 1963) but provisionally considered generally to be a member of the family Pachylaelapidae (Lindquist and Evans, 1965; Krantz and Ainscough, 1990; Halliday, 1997; Lindquist et al., 2009; see Discussion, below). The placement of Zygoseius in Pachylaelapidae was not accepted by Mašán and Halliday, 2014; yet they were unable to argue or opt for its placement in any other family. (6) Dorsal shield with one or two pairs of the Z-setal series differentiated in thickened form from adjacent setae; thickened Z3 are tapered in Antennocheles, while Z4 and/or Z5 are untapered and penicillate in Endopodalius. (7) Peritremes of adults conspicuously bent or elbowed subapically at level of setae s1. (8) Adult mites are closely associated as females phoretic on tropical insects, all coleopterans in families of the infraorder Cucujiformia, suborder Polyphaga: Antennocheles with chrysomelid hispine beetles; Endopodalius with beetles of various other families, i.e., E. tectus with Hololepra humilis Payk (sic for Hololepta humilis Paykull, 1811) (Histeridae) in Venezuela (Hyatt, 1964) and with Scyphophorus acupunctatus Gyllenhaal, 1838 (Dryophthoridae) in southern Spain (Moraza and Balanzategui, 2023); E. americanus with a scarabaeid and with a curculionioid Metamasius in Brasil and Costa Rica, respectively (Lindquist, 1964); E. prorsoperitrematus Abo-Shnaf et al. 2016 with Sphenophorus levis Vaurie, 1978 (Curculionidae) in Brasil (Esteca et al., 2020); E. scutalis (Banks, 1914) with Scarabaeidae in Brasil; E. imitans (Berlese, 1910) with Oryctes rhynoceros (Linnaeus, 1758) (Scarabaeidae) in India; and an undescribed species of Endopodalius with a curculionid in Costa Rica (E.E.L., personal observation).

The above arguments for a sister relationship between Antennocheles and Endopodalius are tenuous, but perhaps could be tested by other morphological aspects (apart from eventual molecular analyses). For example, in both the protonymph and deutonymph of Antennocheles, the anal shield is expanded, much wider than long, enough so as to bear setae JV5 in the deutonymph. This poses a test for the nymphs of Endopodalius, which as yet remain unknown. A sister relationship may argue for either of two perspectives – either the placement of Endopodalius in a separate family, or as a subfamily as a less-derived subset basal to the morphologically and behaviourly more derived Antennocheles, whose family group name has seniority over any such name proposed for Endopodalius. In this regard, we are persuaded to opt for a separate family, to avoid a revised, expanded concept of Antennochelidae when so little is known about what other taxa phylogenetically related to Endopodalius and Antennocheles may remain unaccounted for, among the tropical diversity of insect-associated gamasine mites.

Family Endopodaliidae fam. nov.

Type genus: Endopodalius Christian and Karg, 2006, new status, based on subgenus Lasioseius (Endopodalius) Christian and Karg, 2006.

Family based on adult females representing 13 described species, keyed below, and a further two undescribed species (E.E.L., both from Costa Rica), and males of two and the larva of one of those species. Quintero-Gutiérrez et al. (2020) assigned three other species of Lasioseius to Endopodalius, L. frontalis, L. polydesmophilus, and perhaps L. angustus (the latter based only on the male), without rationale. Based on arguments presented above, those species are retained in the genus Lasioseius, as diagnosed in our other studies (e.g., Moraza and Lindquist, 2018).

(Figures 1–4)

Diagnosis

Adults are apomorphic among members of the superfamily Ascoidea in the females having a sternal shield with (1) posterior margin strongly convex, such that its posteromedial margin reaches same transverse level as laterally adjacent, free metapodal platelets, and (2) anterior margin extending laterally into broadly projecting endopodal extensions between legs I and II where connecting with endopodal-peritrematal plates; (3) females with endopodal plates between legs III and IV uniquely enlarged, extending under metasternal platelets; (4) adults uniquely with a small unpaired platelet at base of tritosternum; (5) female movable cheliceral digit multidentate, with four to nine teeth in addition to apical hook. Other diagnostic attributes include: female sperm access system of laelapoid type, simplified to a short major duct (small section of tubular piece discernibly sclerotized, opening at the base of coxae III), lacking a sclerotized calyx and adjacent atrium with elongated minor duct; fixed cheliceral digit more multidenticulate than movable digit, and with setiform pilus dentilis; movable digit ventral face without a mucronate projection; gnathotectum anterior margin three-tined, with median tine elongated, fringed apically; dorsal shield setae simple, none tricarinate; humeral setae r3 of adult not differentiated by perpendicular projection; male with holoventral shield.

Description

Idiosomal dorsum — Adult female (Figure 1A). Dorsal shield entire, without lateral incisions, well sclerotized and well ornamented, sometimes complexly so (Westerboer, 1963, figure 153), without a delineated lateral rim alongside the lateral series of r-S setae; surrounding soft cuticle smoothly striate. Dorsal shield holotrichous, with a complement of usually 37 pairs of setae, including usually 22 podonotal (j1-j6, z1-z6, s1-s6, r2-r5) (r5 sometimes off shield) and 15 opisthonotal (J1-J5, Z1-Z5, S1-S5); dorsal setae nearly all simple, none tricarinate, though Z4 and/or Z5 bushy, penicillate distally and J5 minute, spiculate basally; humeral setae r3 not differentiated by more perpendicular projection. Dorsal shield with complement of 23 pairs of discernible pore-like structures (9 podonotal, 14 opisthonotal), of which seven pairs (four podonotal, three opisthonotal) superficially appear secretory (gland pores) and 16 pairs (five podonotal, 11 opisthonotal) non-secretory (poroids). Soft lateral cuticle with seta r6, six or seven pairs of marginal R-setae, several pairs of submarginal UR- setae, and pair of marginal poroids idRp. Peritrematal plates broadly uniting with dorsal shield anteriorly at level of setae z2-s1; peritremes well developed, reaching to level of, but well laterad, setae j1; peritremes conspicuously bent or elbowed subapically at level of setae s1.

Adult male. Dorsal shield similar in size, most ornamentation and setation to that of female, except its lateral margins slightly wider to bear setae r5 and variably some of R and UR setae; form and relative lengths of shield setae as in female. Extent and form of peritremes as in female.

Idiosomal venter — Adult female (Figure 2A). Tritosternum with laciniae free or fused for at most half their length, without basal elaborations. Presternal region with median sclerotized strip just anterior to base of tritosternum and one pair of relatively large platelets (Figure 2A), these sometimes split in two (Figure 2B) or with several small platelets adjacent to anterior margin of sternal shield (e.g., figure 47 of Lasioseius prorsoperitrematus Abo-Shnaf et al., 2016). Sternal shield entire, its anterior margin with a transverse strip of heavier, darker sclerotization behind first pair of sternal setae and poroids which blends into strongly developed, wide endopodal extensions between coxae I and II, contiguous with endopodal-peritrematal plate, and lacking gland pore (gbv) in that area; sternal shield with three pairs of setae, two pairs of poroids, strong endopodal extensions between coxae II and III, and with distinctively convex posterior margin, such that its posteromedial margin at same transverse level as laterally adjacent, free metasternal platelets. Poroids iv3 together with setae st4 on small metasternal platelets. Endopodal strips between coxae III and IV strongly developed, free, with anterior extremities extending under metasternal platelets and touching sternal shield posterolateral margin, and posterior tips nearly touching parapodal elements which bear gland pores gv2. Epigynal shield with rounded hyaline anterior margin nearly abutting posterior edge of sternal shield, with posterior margin truncate; setae st5 on shield's posteriorly widening lateral margins, paragenital poroids iv5 on soft cuticle; postgenital furrow with transverse row of four small platelets. Opisthosomatic venter with single pair of freely-standing metapodal platelets. Ventrianal shield well developed, slightly wider than long, with anterior margin abutting posterior edge of epigynal shield where flanked by setae ZV1 on soft cuticle, and with usually four to sometimes six pairs of opisthogastric setae and two pairs of poroids; paranal setae inserted at mid-level of anus, postanal seta slightly longer than paranal setae; anal opening not enlarged; shield with cribrum formed as a narrow band between pair of prominent gland pores gv3 along posterior margin; four or five pairs of other opisthogastric setae (JV4, JV5, ZV3-ZV5) and two pairs of poroids, flanked by posterior pairs of marginal and submarginal setae, on soft cuticle. Peritrematal shield narrowly separated from exopodal strip from midlevel of coxae II to midlevel of coxae IV, and fused to that strip more anteriorly, and posterior to coxa IV; peritrematal shield with two poroids (ip3, ist) and one gland pore (gp3) in area behind stigma, and with poroid ip2 and gland pore gp2 along mid-length of shield; poroid ip1 and gland pore gp1 not visible dorsolateral to peritreme at level of coxa II. Spermathecal apparatus weakly developed, of laelapoid type, with tubular piece (major duct) discernibly associated with posterior surface of coxae III (Figure 2G).

Adult male. Form of presternal area, including a median sclerotized strip just anterior to base of tritosternum, as in female. Sternitigenital, ventrianal and podal-peritrematal shields consolidated to form holoventral shield, with transverse line of fusion delineating connection between sternitigenital and ventrianal portions (Westerboer, 1963, figure 158); anterior margin of shield, at level of genital opening, prominent, broadly widened as endopodal extensions between legs I and II as in female but not darkened; shield with five pairs of setae and three pairs of poroids in sternitigenital region, and fully united with endopodal strips between coxae II and III, III and IV; shield ventrianal region with six to eight pairs of opisthogastric setae (usually JV1-JV4, ZV1, ZV2, sometimes also ZV3, JV5) in addition to circumanal setae; metapodal sigilla incorporated in shield; posterior margin of shield broadly rounded, with cribral band as in female; sternitigenital and opisthogastric setae smooth, similar in shape and length as in female. Exopodal and peritrematal shielding and peritremes as in female, except peritrematal union with dorsal shield broader anteriorly.

Gnathosoma — Female and male. Gnathotectum with convex anterior margin bearing three projections, median one longer and split terminally (Figure 1C). Subcapitular platform elongate, narrow (Figure 1B). Chelicerae capable of complete retraction, yet most of length projectable anteriorly free from body; chelicerae elongate, slender (Figure 2C), without any conspicuous process along antiaxial or paraxial lateral surfaces basal to digits, where hyaline rim smooth. Fixed cheliceral digit fully developed, with or without a mucronate process distally, with bifid apical hook (Figures 2D, E), apposed to apical hook of movable digit, with typically short setiform pilus dentilis, and with a continuous ridge of seven to 13 similarly-sized teeth along masticatory surface and few small teeth at base of digit, flanked paraxially by a hyaline multidenticate rim (Figure 2E, from Abo-Shnaf et al., 2016, figure 52a); movable digit of female with four to nine teeth (Figures 2C, F), that of male with one large tooth, and with spermatodactyl tubular, directed anteriorly, bent or hooked distally (vide Westerboer, 1963, p. 285; Abo-Shnaf et al., 2016, figure 59). Corniculi of female moderately long, slender, parallel (Figure 1B), those of adult male slightly more widely spaced, with more conspicuous internal projection from their bases (Westerboer, 1963, figure 157a); internal malae elongate, extending well beyond tips of corniculi. Apex of labrum extending to or slightly beyond tips of corniculi. Salivary styli not reaching tips of corniculi. Subcapitular setae smooth, pair hp3 nearly transversely aligned with hp2 and usually longer than pairs hp1 and c.s. Deutosternum with seven moderately wide rows of denticles, similarly multidenticulate, free (Figure 1B) or flanked by lateral margins (Westerboer, 1963, figure 154b). Palpi (Figure 1D) with holotrichous setation as described for Gamasina by Evans (1964); palpfemoral seta al slightly lanceolate and palpgenual setae al-1 and al-2 clearly spatulate; palptarsal apotele two-tined, with spatulate tips; palptrochanter with inner seta not markedly elongated.

Legs — Female and male. Legs of moderate length, I and IV no longer than dorsal shield. Legs I slightly longest and thinner, and II slightly thicker, than legs III and IV (Figures 4A–D). Distal margins of coxae I–IV without serrated ridges or spur-like processes (Figures 4E–H). Legs I to IV with pretarsi bearing paired claws of moderate size, inconspicuous paradactyli, and rounded pulvilli (Figures 3A, E). Leg I tarsus with 45 setae (Figure 3B) and distinct pedicel bearing pretarsal structures (Figures 3A, C). Tarsus I with sensilla s inconspicuous, not noticeably lanceolate distally (Figure 3D). Legs II to IV with tarsus (excluding pretarsus) about twice as long as tibia. Tarsi II–IV with apical setal processes ad-1, pd-1 short, less than half as long as pretarsi (Figures 3E, F), and with rounded triangular apical process ventrally. Setation and their ontogeny on segments of legs I to IV with full complement of Ascidae (as presented by Lindquist and Evans 1965 for Ascini), coxae, 2-2-2-1; trochanters, 6-5-5-5; femora, 12 (2 3/1 2/2 2) –11 (2 3/1 2/2 1) – 6 (1 2/1 1/0 1) – 6 (1 2/1 1/0 1); genua, 13 (2 3/2 3/1 2) – 11 (2 3/1 2/1 2) – 9 (2 2/1 2/1 1) – 9 (2 2/1 3/0 1), genu IV lacking pv-1; tibiae, 13 (2 3/2 3/1 2) – 10 (2 2/1 2/1 2) – 8 (2 1/1 2/1 1) – 10 (2 1/1 3/1 2). Tarsi II–IV with seta ad-2 somewhat elongated but not extending to base of pretarsi (Figure 3E); paired ventral setae (v-1) and (v-2) somewhat spine-like and claws strong (Figure 3F); other leg setae simple, not strongly modified on female. Male leg setal dimorphism not notably developed (Westerboer, 1963).

Genus Endopodalius Christian and Karg, 2006

Subgenus Lasioseius (Endopodalius) Christian and Karg, 2006

Type species: Lasioseius (Zygoseius) alter Vitzthum, 1925, original designation.

Diagnosis — Adults of Endopodalius share with other genera included in the superfamily Ascoidea the series of attributes given above for diagnosis of that superfamily. As the only genus of Endopodaliidae, they are distinguished from those of other taxa of Ascoidea by the following characteristics, some of which are apomorphic, as indicated below: Dorsal shield with (1) humeral setae r3 not erect or differentiated in form from adjacent setae, and with (2) nearly all setae simple, none tricarinate, but (3) usually with setae Z4 and/or Z5 brushy, penicillate distally (apomorphy); (4) adult sternal shield anterior margin with more heavily sclerotized strip continuing as broadly widened extensions to unite with endopodal-peritrematal shield on either side (apomorphy); (5) female sternal shield posterior margin strongly convex, such that its posteromedian margin reaches same transverse level as adjacent, free metasternal plates (autapomorphy); (6) female endopodal plates between legs III-IV enlarged, extending under metasternal platelets (autapomorphy); (7) adult presternal region with small unpaired platelet at base of tritosternum (autapomorphy), and (8) with one to several paired platelets; (9) movable cheliceral digit with usually five or more teeth (apomorphy); (10) gnathotectum with elongated median process, fringed apically (apomorphy); (11) male with holoventral shield. Attributes 1, 2, 5-7 and 9-11 are repeated from the family diagnosis as distinctions from genera in closely related families.

Key to species of Endopodalius, adult females

The following key has been elaborated from the original descriptions and illustrations of the previously described species and from our observations of type or reference material of some of them, as noted above. Several of those species are so inadequately described that further studies of type material may indicate other synonymies, especially some of those species among couplets 10 through 12. On the other hand, several other species inadequately described under the genus and subgenus Lasioseius may belong to Endopodalius (as we have determined for Lasioseius americanus Chant, 1963, which was inadvertently placed in the inguinalis species complex of the subgenus Lasioseius by Christan and Karg, 2006). For example, the adult female of Lasioseius ruehmi, Hirschmann, 1972, needs further study, as it has a similarly multidentate (6–7 teeth) movable chela; none of its dorsal shield setae are tricarinate, though Z5 is penicillate; and its sternal shield posterior margin is partially convex.

1. Dorsal shield with all setae simple, smooth, tapered; sternal shield coarsely ornamented with large, scattered fovea; metapodal plates encompassed into anterolateral corners of ventrianal shield
...... 2

— Dorsal shield with one or two of setae Z4, Z5 untapered, thickened distally; sternal shield more finely ornamented with puncta, striae; metapodal plates free from ventrianal shield
...... 3

2. Ventrianal shield shown with six pairs of opisthogastric setae, ZV1 and JV4 on shield; two pairs of presternal platelets
...... alter (Vitzthum, 1925)

— Ventrianal shield shown with four pairs of opisthogastric setae, ZV1 and JV4 off shield; one pair of presternal platelets
...... hirschmanni Christian and Karg, 2006 (= alter sensu Bhattacharyya, 1969)

3. Dorsal shield with both of setae Z4, Z5 untapered, penicillate distally
...... 4

— Dorsal shield with only one of either Z4 or Z5 untapered, thickened distally
...... 5

4. Sternal shield anterior margin with wide, rectangular, median notch, posterior margin slightly concave medially (other than its general convexity); dorsal shield setae of J-series shorter than intervals between their bases
...... humberti (Athias-Henriot, 1959)

— Sternal shield anterior margin not notched, posterior margin entirely convex; dorsal shield setae of J-series slightly longer than intervals between their bases
...... araucariae (Hirschmann, 1972)

5. Fixed chela with mucronate process distally (Figure 2E); movable chela with 8–10 teeth
...... scutalis (Banks, 1914) (=prorsoperitrematus Abo-Shnaf et al. 2016, new synonym)

— Fixed chela without mucronate process distally; movable chela with 4–7 teeth
...... 6

6. Dorsal shield setae Z4 untapered, stout, blunt-tipped, penicillate distally, Z5 simple
...... 7

— Dorsal shield setae Z4 slender, pointed, smooth, but Z5 untapered, thickened distally
...... 9

7. Ventrianal shield with five or six pairs of opisthogastric setae, including JV5, sometimes JV4; presternal region with two pairs of well-separated platelets
...... vitzthumi (Westerboer, 1963)

— Ventrianal shield with four pairs of opisthogastric setae, JV4, JV5 on adjacent soft cuticle; presternal region with one pair of platelets divided into two contiguous pairs (Figure 2B)
...... 8

8. Dorsal shield with prominent, transverse punctated line/groove across entire face behind setae J4 and Z4; fixed cheliceral digit with ca 13 teeth
...... tectus (Hyatt, 1964)

— Dorsal shield without transverse punctated line across entire face at any level; fixed cheliceral digit with seven to nine teeth
...... ibericus Moraza and Balanzategui, 2023

9. Distally thickened setae Z5 abruptly curved distally, about 1.7x longer than setae Z4
...... 10

— Distally thickened setae Z5, straight, similar in length or longer than setae Z4
...... 11

10. Setae Z5 serrated, penicillate distally; ventrianal shield setae JV3 over twice as long as other opisthogastric setae; movable cheliceral digit with four teeth
...... americanus (Chant, 1963)

— Setae Z5 knobbed, smooth distally; ventrianal shield setae JV3 scarcely longer than other opisthogastric setae; movable cheliceral digit with seven teeth
...... gabrielae (Santos and Argolo, 2018)

11. Dorsal shield with setae J1 to J4 slightly longer than intervals between their bases, J4 as long as Z5; presternal platelets divided but its parts contiguous
...... convexus (Krantz, 1962)

— Dorsal shield with setae J1 to J4 ca half to nearly as long as intervals between their bases, J4 ca half to 0.7 as long as Z5; presternal platelets entire, subtriangular
...... 12

12. Dorsal shield with only setae j5, z6 shorter (about 0.7 as long) than adjacent setae (E.E.L., personal observation)
...... imitans (Berlese, 1910)

— Dorsal shield with setae j4 and hexagonal area setae j5, z5, j6 short, about half as long as adjacent j3, z6
...... mouchei (Loots, 1980; possible synonym of imitans)

Distribution and habitats

Endopodalius is known at present from species found in Seychelles islands east of Africa (north of Madagascar), India, Sumatra, South and Central America (Brasil, Dominican Republic, and Costa Rica), and new records noted below from Micronesia. Among the thirteen named described species, males of only two species, E. prorsoperitrematus (Abo-Shnaf, Sánchez and Moraes, 2016) and E. vitzthumi (Westerboer, 1963), and the larva of only E. araucariae (Hirschmann, 1972) have been described, while nymphal instars remain unknown so far. Several described species have been reported associated with insects, all coleopterans (see details above).

Discussion

Cheliceral structures — The chelicerae of Endopodalius are rather ordinary in various morphological respects, in comparison with many of the gamasine groups of Mesostigmata (Karg, 1965; Evans, 1992; Alberti and Coons, 1999). However, compared with forms of chelicerae within the hyperdiverse genus Lasioseius, with which Endopodalius has been confused, the multidentate yet not elongate form of the movable digit, with more than three teeth, is relatively unique and perhaps relevant to a more particular feeding behavior, of which nothing is known.

Although occasionally illustrated (e.g., Lindquist, 1971and subsequent papers; Abo-Shnaf et al., 2006) little has been stated about the form and function of the hyaline structure with a denticulated rim on the paraxial surface of the chelicerae at the level of the movable chela's articulation in various blattisociid and melicharid mites (shown in dorsoventral aspect in Figure 2E). This structure is not considered in the major morphological treatises by either Evans (1992) or Alberti and Coons (1999). As a pattern, lateral view drawings of gamasine chelicerae are often from the outer side, showing the antiaxial lyrifissure, i.e., across from the paraxial side where the hyaline structure is located. Evident among various blattisociid and melicharid mites, these structures have not been illustrated or noted among other gamasine taxa such as the ascid subfamilies Ascinae, Arctoseiine and the blattisociid subfamily Platyseiinae (E.E.L. personal observations). These structures may be part of a sieving or straining system when functioning together between the chelicerae.

Sternal shield aspects — Within the highly diverse and speciose genus Lasioseius (over 150 species described; Christian and Karg, 2006; Moraes et al., 2016), adult females of the scutalis species group now recognized as Endopodalius are distinctive in having their sternal shield's anterior margin strongly thickened and darkened as a transverse band. The dark, thickened aspect of this shield was not illustrated in previous species descriptions (other than for E. ibericus by Moraza and Balanzategui, 2023); however, personal observations (E.E.L.) of several species suffice to conclude that this thickening is prevalent or ubiquitous in the genus. Interestingly, a similar such sternal shield band (the shield's posterior margin also somewhat convex) was noted and illustrated by Bhattacharyya (1972) for a melicharid, Proctolaelaps sternalis Bhattacharyya, found under bark of a dead tree – a habitat similar to those of Endopodalius. A somewhat similar but broader transverse band of thickened sternal shield cuticle is evident in deutonymphs of the genus Poecilochirus G. & R. Canestrini, 1882, family Parasitidae. Such anterior thickenings of the sternal shield may be correlated with phoretic behavior, which is restricted to the instar and sex for which this thickened structure is found.

The unusually convex form of the female sternal shield's posterior margin, along with strongly developed endopodal plates between legs III and IV in species of Endopodalius are also found among those of Zygoseius, a taxon confused with Endopodalius, in relationships with Lasioseius (Westerboer, 1963; Halliday, 1997). Differences among other adjacent coxisternal structures (particularly in Endopodalius, incorporation of the metasternal platelets with setae st4 and poroids iv3 into the expanded endopodal plates alongside legs III-IV, with those plates extended and connected anteriorly to the sternal shield endopodal extensions between the bases of legs II and III) indicate that these similarities are morphological convergences. In Endopodalius, the posteromedial margin of the sternal shield extends to the same transverse level as the small metasternal platelets (bearing setae st4 and poroids iv3), which are freely standing on soft cuticle and underlapped by the endopodal plates which are devoid of other structures. In Zygoseius, the sternal shield's posteromedial margin does not extend so far posteriorly as in transverse level between the metasternal setae and poroids; and the latter structures are inserted on even more enlarged endopodal plates which connect with the sternal shield extensions between legs II and III.

Peritrematic aspects — Abo-Shnaf et al. (2016) drew attention to the anterior ends of the peritremes being distinctively bent forward at the level of the paravertical setae z1 in Lasioseius prorsoperitrematus, stated to be in distinction to those of L. americanus Chant, 1963 and L. araucariae Hirschmann, 1972. While this aspect was not noted or illustrated among previous descriptions and illustrations of other species of this genus, our observations (E.E.L.) of type or reference material indicated that the same, anteriorly bent form is present in L. americanus, L. scutalis, L. tectus, but not in material identified as near L. araucariae and L. imitans, and only slightly so in material of another undescribed species.

Lectotype designation — Observations by one of us (Lindquist, 1964) of Nathan Banks' microslide preparation of Hypoaspis scutalis (deposited in the Museum of Comparative Zoology, Harvard University), with the word ''type» written on the label, found it to include twelve adult female mites, four of which were identified as scutalis and are considered to be syntypes. The specimen most centrally located is here designated as the lectotype: from the orientation of its legs, this specimen is obviously the one upon which Banks based his original illustration, and it can be readily distinguished also by being the only specimen with setae Z5 and all legs intact. Evans and Hyatt (1960) erroneously anticipated Hypoaspis scutulis (sic) Banks, 1914 to belong to the genus Sejus Koch, 1843 (subsequently Cheiroseius Berlese, 1916, sensu Evans and Till, 1979); however, they had not examined Banks' type material of Hypoaspis scutalis.

Life history — Although the life cycle instars have not been observed for any species of Endopodalius, a presumably free-standing larva of L. (E.) araucariae Hirschmann was illustrated by Hirschmann (1972). An adult female identified as L. (E.) imitans at hand (E.E.L.) contains an egg with fully formed larval structures, indicating probable larviparity.

Coexistent or polymorphic forms — Further observations (E.E.L. notes, 1964) of Banks' type slide preparation of Hypoaspis scutalis determined that among the twelve adult females (of which only four were identifiable as scutalis), there are two other closely related forms, both belonging to Endopodalius. Of these, two females were at that time identifiable as Lasioseius americanus Chant, 1963 (original type material from Guatemala and Ecuador), and three other females as a species closely similar to Lasioseius tectus (Hyatt, 1964) (original type material from Venezuela), which was not yet described when those notes were first taken. Other than the scutalis females, those of the two other closely related forms were noted (E.E.L. personal observations) as lacking the apical mucronate process on the fixed chela. Three of the specimens were not adequately preserved for more specific identification. The coexistence of three of these forms may indicate a possible adult female polymorphism; however, we are not aware of any other example among gamasine mites where representatives of more than one of a closely related group of species are phoretically coexistent on an arthropod host. The example of Antennoseius janus Lindquist and Walter, 1989 exhibiting adult female dimorphism was shown to be phase morphism, involving phoretic and non-phoretic forms (Lindquist and Walter, 1989). Curiously, Hirschmann (1972) described closely similar adult females of two species, Lasioseius araucariae and Lasioseius ruehmi, which may possibly be another example of polymorphism, or if not, then further studies of material of L. ruehmi are needed to confirm its placement in Endopodalius. Both forms were collected in association with Araucaria trees, L. araucariae from phloem infested by numerous insects, and L. ruehmi from insect frass galleries in Araucaria and phoretic on bark beetles, Ptelobius valdivianus (Eggers). While female L. araucariae presents all of the attributes typical of Endopodalius (form of presternal structures and sternal shield; penicillate dorsal shield setae Z4, Z5, multidentate five teeth movable chela, median tine of gnathotectum elongate, apically split), female L. ruehmi has only setae Z5 penicillate and a similarly multidentate, six or seven toothed movable chela, median gnathotectal tine less distinctive in form from lateral tines, and lacking presternal platelets and not distinctively convex posterior margin of sternal shield.

Additional collection records of Endopodalius species — The Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, has one slide preparation with four females of Lasioseius (Endopodalius) scutalis (Banks) from Peru (no locale indicated), ex Brazil nuts; one female of L. (E.) americanus (Chant) from Guatemala (no locale) ex Metamasius sericeus Kuschel, (Dryophthoridae); one female of L. (E.) imitans (Berlese) from Bangalore, India, ex. Oryctes rhinoceros (Linnaeus), (Scarabaeideae); one female of L. (E.) undescribed sp. 1 from Jokaj Islet, Pohnpei (formerly named Ponape) Island, Micronesia, ex. decaying crown of dead coconut; one slide with two females of L. (E.) undescribed sp. 2 from Pohnpei Island, Micronesia, Mt. Nanlaud, elev. 1000 ft. (305 m), ex. dead palm petiole;and four slides each with one female of different described species of Endopodalius from La Selva Biological Station, Heredia, Costa Rica: one L.(E.) tectus, ex fogged Vitex foliage; one L.(E.) scutalis, ex fogged unidentified foliage; one, near L.(E.) araucariae, ex experimental swamp; one undescribed species, near L.(E). americanus ex Curculionidae.

Acknowledgments

Our thanks to Jerry W. Krantz (Oregon State University, Corvallis) and, many years ago, to Herbert W. Levi (Museum of Comparative Zoology, Harvard University, Cambridge) for the loans of type material of Lasioseius convexus Krantz, 1962, and Hypoaspis scutalis Banks, 1914, respectively; to Reham I.A. Abo-Shnaf (University of San Paulo, Piracicaba, Brazil) for permission to reproduce an illustration of Lasioseius prorsoperitrematus Abo-Shnaf et al. 2016; to Serge Laplante (Agriculture & Agri-Food Canada, Ottawa) for bibliographic support; and to Fred Beaulieu (CNCI-Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture & Agri-Food Canada, Ottawa) and Bruce Halliday (CSIRO Division of Entomology, Canberra) for their useful and persuasive reviews of the manuscript.

References

1. Abo-Shnaf, R.I., Sánchez, L., Moraes, G.J.D. 2016. Plant inhabiting Gamasina mites (Acari: Mesostigmata) from the Dominican Republic, with descriptions of four new species of Lasioseius (Blattisociidae) and complementary descriptions of other species. Systematic and Applied Acarology, 21(5): 607-646. https://doi.org/10.11158/saa.21.5.5
2. Alberti G. 2002. Ultrastructural investigations of sperm and genital systems in Gamasida (Acari: Anactinotrichida): Current state and perspectives for future research. Acarologia, 42: 107-126.
3. Alberti G., Coons L.B. 1999. Acari: Mites. In: Harrison F.W., Foelix R.F. (Eds.) Microscopic anatomy of invertebrates, Vol. 8C, Chapter 6, Chelicerate Arthropoda. Wiley-Liss, New York, pp. 515-1265.
4. Argolo P.S., Santos J.C., Oliveira A.R., de Moraes G.J. 2018. Two new species of Lasioseius Berlese (Acari: Blattisociidae) from Brazil, and a key for separation of the Brazilian species of the genus. Systematic & Applied Acarology, 23(8): 1567-1577 https://doi.org/10.11158/saa.23.8.7
5. Athias-Henriot C. 1959. Phytoseiidae et Aceosejidae (Acarina, Gamasina) d'Algérie. III Contribution aux Aceosejinae. Bulletin de la Société d'Histoire Naturelle de L'Afrique du Nord, 50: 158-195.
6. Athias-Henriot C. 1968. L'appareil d'insémination laelapoïde (Acariens anactinotriches: Laelapoidea). Premières observations. Possibilité d'emploi à des fins taxonomiques. Bulletin Scientifique de Bourgogne, 25: 229-274.
7. Athias-Henriot C. 1969. Les organes cuticulaires sensoriels et glandulaires des Gamasides. Poroïdotaxie et adénotaxie. Bulletin de la Société Zoologique du France, 94: 485-492.
8. Athias-Henriot C. 1971. Nouvelles notes sur les Amblyseiini (Gamasides Podospermiques, Phytoseiidae) 1. La dépilation des génuaux et tibias des pattes. Acarologia, 13: 4-15.
9. Banks N. 1914. The Stanford Expedition to Brazil, 1911. Acarians from Brazil. Psyche 21(5): 160-162. https://doi.org/10.1155/1914/61017
10. Berlese A. 1910. Brevi diagnosi di generi e specie nuovi di Acari. Redia, 6: 346-388.
11. Berlese A. 1916. Centuria prima di Acari nuovi. Redia, 12: 19-67.
12. Bhattacharyya S.K. 1969. Studies on Indian mites (Acarina: Mesostigmata). 7. Six species found under bark in North East Frontier Agency. Zoologische Jahrbuch, Systematische, 96: 69-80.
13. Bhattacharyya S.K. 1972. Studies on Indian mites (Acarina: Mesostigmata). 9. Additional species found under bark in North East Frontier Agency. Acarologia, 13: 272-279.
14. Canestrini G., Canestrini R. 1882. Gamasi Italiani. Atti della Societa Veneto-Trentina di Scienze Naturali, 8: 1-80.
15. Chant D.A. 1963. The subfamily Blattisocinae Garman (=Aceosejinae Evans) (Acarina: Blattisocidae Garman) (=Aceosejidae Baker and Wharton) in North America, with descriptions of new species. Canadian Journal of Zoology, 41: 243-305. https://doi.org/10.1139/z63-025
16. Christian A., Karg W. 2006. The predatory mite genus Lasioseius Berlese, 1916 (Acari, Gamasina). Abhandlungen und Berichten aus den Naturkundemuseum Görlitz, 77: 99-250.
17. Di Palma A., Alberti G. 2005. Functional morphology of accessory structures involved in reproduction among gamasid mites (Acari, Anactinotrichida): a review. Atti dell′Accademia Nazionale Italiana di Entomologia Rendiconti, 53: 361-378.
18. Esteca F.C.N., Borges V., Santos J.C., Neves L.S., Moraes G.J. 2020. Report of the mite Lasioseius prorsoperitrematus Abo-Shnaf, Sánchez & Moraes, 2016 (Acari: Blattisociidae) in Brazil associated with the insect Sphenophorus levis Vaurie, 1978 (Coleoptera: Curculionidae). Entomological Communications, 2, 2020: ec02028 https://doi.org/10.37486/2675-1305.ec02028
19. Evans G.O. 1963. Observations on the chaetotaxy of the legs in the free-living Gamasina (Acari: Mesostigmata). Bulletin of the British Museum (Natural History) Zoology, 10: 275-303. https://doi.org/10.5962/bhl.part.20528
20. Evans G.O. 1964. Some observations on the chaetotaxy of the pedipalps in the Mesostigmata (Acari). Annals and Magazine of Natural History, Series 13, 6: 513-527. https://doi.org/10.1080/00222936308651393
21. Evans G.O. 1969. Observations on the ontogenetic development of the chaetotaxy of the tarsi of legs II-IV in the Mesostigmata (Acari). In: Evans G.O. (Ed.) Proceedings of the 2^nd International Congress of Acarology, 1967, Akadémiai Kiadó, Budapest, pp. 195-200.
22. Evans G.O. 1992. Principles of Acarology. CAB International, Wallingford, xviii+563 pp. https://doi.org/10.1079/9780851988221.0000
23. Evans G.O., Hyatt K.H. 1960. A revision of the Platyseiinae (Mesostigmata: Aceosejidae) based on material in the collections of the British Museum (Natural History). Bulletin of the British Museum (Natural History), Zoology 6(2): 27-101. https://doi.org/10.5962/bhl.part.26844
24. Evans G.O., Sheals J.G. 1959. Three new mesostigmatic mites associated with Millepedes in Indonesia. Entomologische Berichten, 19: 107-111.
25. Evans G.O., Till W.M. 1979. Mesostigmatic mites of Britain and Ireland (Chelicerata: Acari-Parasitiformes). An introduction to their external morphology and classification. Transactions, Zoological Society of London, 35: 139-270. https://doi.org/10.1111/j.1096-3642.1979.tb00059.x
26. Fain A., Hyland K.E., Aitken T.H.G. 1977. Flower mites of the family Ascidae phoretic in nasal cavities of birds (Acarina: Mesostigmata). Acta Zoologica Pathologica Antverp, 69: 99-154.
27. Gyllenhaal L. 1838. In: Schönherr C.J., Gyllenhaal L., Boheman C.H. Genera et species Curculionidum, cum synonymia hujus familae species novae. Roret, Paris, Vol. 4, Part 2, pp. 855-859.
28. Halliday R.B. 1997. Revision of the genus Zygoseius Berlese (Acarina: Pachylaelapidae). Acarologia, 38: 3-20.
29. Hirschmann W. 1972. Von Dr. W. Rühm während seiner Tätigkeit an der Univ. Austral de Chile (Valdevia) gesammelte Araukarien-Milben aus Südchile u. Südbrasilien. Acarologie, Gangsystematik der Parasitiformes, Teil 4, 17: 29-33, 9 plates.
30. Hyatt K.H. 1964. A collection of Mesostigmata (Acari) associated with Coleoptera and Hemiptera in Venezuela. Bulletin of the British Museum (Natural History), Zoology, 11(7): 465-509. https://doi.org/10.5962/bhl.part.4723
31. Johnston D.E., Moraza M.L. 1991. The idiosomal adenotaxy and poroidotaxy of Zerconidae (Mesostigmata: Zerconina). In: Dusbábek F., Bukva V.(Eds.) Modern Acarology, Academia, Prague, Vol. 2, pp. 349-356.
32. Karg W. 1965. Larvalsystematische und phylogenetische Untersuchung sowie Revision des Systems der Gamasina Leach, 1915 (Acarina, Parasitiformes). Mitteilungen aus dem Zoologischen Museum in Berlin 41: 193-341. https://doi.org/10.1002/mmnz.19650410207
33. Karg W. 1993. Acari (Acarina), Milben Parasitiformes (Anactinochaeta) Cohors Gamasina Leach. Raubmilben. Die Tierwelt Deutschlands Teil 59: 523 pp.
34. Karg W. 1998. Zur Kenntnis der Eugamasides Karg mit neuen Arten aus den Regenwäldern von Ecuador (Acarina, Parasitiformes). Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische Reihe, 74: 185- 214. https://doi.org/10.1002/mmnz.4850740204
35. Karg W. 2003. Neue Raubmilbenarten aus dem tropischen Regenwald von Ecuador mit einem kritischen Beitrag zur Merkmalsevolution bei Gamasina (Acarina, Parasitiformes). Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische Reihe, 79: 229 - 251. https://doi.org/10.1002/mmnz.4850790203
36. Karg W. 2006. The systematics of Parasitiformes, especially of Gamasina Leach (Acarina), with new species from Ecuador. Mitteilungen aus dem Museum für Naturkunde in Berlin, Zoologische Reihe, 82: 140-169. https://doi.org/10.1002/mmnz.200600002
37. Koch C.L. 1843. Übersicht des Arachnidensystems. Heft 3. C. H. Zeh'schen Buchhandlung, Nürnberg, 131 pp., 13 plates.
38. Krantz G.W. 1962. Acari. Free-living Mesostigmata. II. The family Aceosejidae. Parc National De La Garamba, Mission H. De Saeger, 34: 3-29.
39. Krantz G.W., Ainscough B.D. 1990. Acarina: Mesostigmata (Gamasida). In: Dindal, D.L. (Ed.) Soil Biology Guide, John Wiley & Sons, New York, pp. 583-665.
40. Krantz G.W., Redmond B.L. 1987. Identification of glandular and poroidal idiosomal systems in Macrocheles perglaber F. & P. (Acari: Macrochelidae). Experimental & Applied Acarology, 3: 243-253. https://doi.org/10.1007/BF01270460
41. Lindquist E.E. 1964. A revision of mites of the subfamily Blattisocinae (Acarina: Blattisocidae) in America north of Mexico. Ph.D. dissertation, University of California, Berkeley, University Microfilms, Inc., Ann Arbor, Michigan, 64-2093. iii + 413 pp.
42. Lindquist E.E. 1971. New species of Ascidae (Acarina: Mesostigmata) associated with forest insect pests. Canadian Entomologist, 103: 919-942. https://doi.org/10.4039/Ent103919-7
43. Lindquist E.E. 1994. Some observations on the chaetotaxy of the caudal body region of gamasine mites (Acari: Mesostigmata), with a modified notation for some ventrolateral body setae. Acarologia, 35: 323-326.
44. Lindquist E.E., Evans G.O. 1965. Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the Entomological Society of Canada, 47: 1-64. https://doi.org/10.4039/entm9747fv
45. Lindquist E.E., Moraza M.L. 2010. Revised diagnosis of the family Blattisociidae (Acari: Mesostigmata: Phytoseioidea), with a key to its genera and description of a new fungus-inhabiting genus from Costa Rica. Zootaxa, 2479: 1-21. https://doi.org/10.11646/zootaxa.2479.1.1
46. Lindquist E.E., Moraza M.L. 2012. A new genus of fungus-inhabiting mites of the family Blattisociidae (Acari Mesostigmata Phytoseioidea) from Costa Rica, with an updated key to genera of the subfamily Blattisociinae. Redia, 95: 9-19.
47. Lindquist E.E., Moraza M.L. 2014. Mites coexistent with neotropical hispine beetles in unfurled leaves of Heliconia: a new genus and family of the Ascoidea (Acari: Mesostigmata: Gamasina). Journal of Natural History, 48(27-28): 1611-1651. https://doi.org/10.1080/00222933.2013.877995
48. Lindquist E.E., Walter D.E. 1989. Antennoseius (Vitzthumia) janus n. sp. (Acari: Ascidae), a mesostigmatic mite exhibiting adult female dimorphism. Canadian Journal of Zoology, 67(11): 2797-2813. https://doi.org/10.1139/z89-184
49. Lindquist E.E., Walter D.E., Krantz G.W. 2009. Order Mesostigmata. In: Krantz G.W, Walter D.E. (Eds.) A Manual of Acarology, Third Edition, Texas Tech University Press, Lubbock, pp. 124-232.
50. Linnaeus C. 1758. Systema Naturae, 10^th ed. L. Salvii, Stockholm.
51. Loots G.C. 1980. Contributions à l'étude de la faune terrestre des îles granitiques de l'archipel des Séchelles (Mission P.L.G.Benoit - J.J. Van Mol 1972). Freeliving Gamasina (Mesostigmata - Acarina) - Revue de zoologie africaine, 94: 745-772.
52. Mašán P., Halliday B. 2014. Review of the mite family Pachylaelapidae (Acari: Mesostigmata). Zootaxa, 3776(1): 1-66. https://doi.org/10.11646/zootaxa.3776.1.1
53. Moraes G.J. de, Britto E.P.J., Mineiro J.L. de C., Halliday B. 2016. Catalogue of the mite families Ascidae Voigts & Oudemans, Blattisociidae Garman and Melicharidae Hirschmann (Acarina: Mesostigmata). Zootaxa, 4112(1): 1-299. https://doi.org/10.11646/zootaxa.4112.1.1
54. Moraza M.L., Balanzategui I. 2023. A new species of Lasioseius (Endopodalius) (Acari: Mesostigmata: Blattisociidae) coexistent with the invasive agave weevil (Coleoptera: Dryophthoridae) in the southeast of the Iberian Peninsula. Acarologia, 63: 605-614. https://doi.org/10.24349/vfv7-8tut
55. Moraza M.L., Lindquist E.E. 2018. A new species-group with new species of the genus Lasioseius (Acari: Mesostigmata: Blattisociidae) associated with neotropical hispine beetles in furled leaves of Heliconia. Acarologia, 58(1): 62-98. https://doi.org/10.24349/acarologia/20184227
56. Naskreski P., Colwell R.K. 1998. Systematics and host plant affiliations of hummingbird flower mites of the genera Tropicoseius Baker & Yunker and Rhinoseius Baker & Yunker (Acari: Mesostigmata: Ascidae). Monographs, Thomas Say Publications in Entomology, iii+185 pp. https://doi.org/10.4182/QYFA2793
57. Paykull G de. 1811. Monographia Histeroidum. Stenhammar & Palmblad, Uppsala. 123 pp + 13 plates.
58. Quintero-Gutiérrez E.J., Sandmann D., Cómbita-Heredia O., Klarner B., Widyastuti R., Scheu S. 2020. A new species of the genus Lasioseius (Acari: Blattisociidae) inhabiting litter of secondary rainforest in Sumatra, Indonesia. Acarologia, 60(2): 338-352. https://doi.org/10.24349/acarologia/20204371
59. Singer G. 1967. A comparison between different mounting techniques commonly employed in acarology. Acarologia, 9: 475-484.
60. Vaurie P. 1978. Revision of the genus Sphenophorus in South America. Bulletin of the American Museum of Natural History, 2656: 1-30.
61. Vitzthum H. 1925. Fauna sumatrensis (Beitrag Nr. 5). Acarinae. Supplementa Entomologica Berlin, 11: 1-79.
62. Walter D.E., Lindquist E.E. 1997. Australian species of Lasioseius (Acari: Mesostigmata: Ascidae) the porulosus group and other species from rainforest canopies. Invertebrate Taxonomy, 11: 525-547. https://doi.org/10.1071/IT96003
63. Westerboer I. 1963. Die Familie Podocinidae Berlese, 1916. In: Stammer H.J. (Ed.), Beiträge zur Systematik und Ökologie mitteleuropäischer Acarina, Band II, Mesostigmata 1. Akademie Verlagsgesellschaft, Leipzig, pp.179-450.

instructions