A review of Cunaxidae (Acariformes, Trombidiformes): Histories and diagnoses of subfamilies and genera, keys to world species, and some new locality records

Abstract Cunaxidae are predaceous mites found in a variety of habitats. This work provides comprehensive keys to world subfamilies, genera, and species. Diagnoses and historical reviews are provided for subfamilies and genera. Cunaxa boneti, C. denmarki, C. exoterica, C. floridanus, C. lehmanae, C. lukoschusi, C. metzi, C. myabunderensis, C newyorkensis, C. rackae, C. reevesi, and C. reticulatus are moved to Rubroscirus and C. otiosus,﻿ C. valentis, and C. rasile are returned to Rubroscirus. Cunaxoides neopectinatus is moved to Pulaeus. Neocunaxoides pradhani and N. gilbertoi are transferred to Scutopalus. Pulaeus minutus and P. subterraneus are moved to Lupaeus. Pseudobonzia bakari, P. malookensis, and P. shamshadi are transferred to Neobonzia. Dactyloscirus bifidus is transferred to Armascirus. Scirula papillata is reported from the Western Hemisphere for the first time. Armascirus ozarkensis, A. primigenius, and Dactyloscirus dolichosetosus are reported from new localities.


Introduction
Cunaxidae ( Fig. 1) are common predatory mites that are present in forest systems, grasslands, agricultural fields, and anthropogenically disturbed areas. Surveys of mites in these habitats often report only family or generic-level identification. This is problematic because little is known about where cunaxid species occur, both regionally and in what habitats, and unfortunate because such reports are potentially very useful collectively if species were identified.
Part of the reason behind the lack of specific identification is the difficulty in reliably identifying cunaxids without extensive knowledge of the primary literature. Keys to cunaxid species are often regional, so of little use to researchers outside of that specific region, and scattered across countless journals. The last comprehensive attempt to present keys to world species was by . The number of described species since Smiley published his monograph has more than doubled (166 to 400+). Updated keys reflecting known diversity and current taxonomic opinion are therefore imperative if researchers are to identify individuals to the specific rather than generic or family level.
Both ambush and active hunting have evolved within the family, sometimes within the same subfamily. Within Cunaxinae, for instance, Armascirus and Dactyloscirus wait, sometimes for hours, to ambush prey (Walter and Proctor 1999), whereas Allocunaxa actively search for prey (Castro and Moraes 2010).
Cunaxids appear to be active year round. Den Heyer (1980a) collected all life stages of Neocunaxoides in the Transvaal Highveld during the summer (30 °C+) and winter (minimum 0 °C) months. Zaher et al. (1975b) collected cunaxids throughout the year and demonstrated a positive correlation between abundance and temperature; they also found a slight negative correlation between abundance and relative humidity.
Cunaxids have been reported to be found phoretically on bark beetles, though they were not identified to species (Penttinen et al. 2013).
Both sexual reproduction and thelytokous parthenogenesis have been reported in cunaxids Proctor 1999, Castro andMoraes 2010). Within Cunaxinae, Coleoscirinae, and Cunaxoidinae, precopulatory guarding of the quiescent tritonymphal female has been reported (Walter and Kaplan 1991). Dactyloscirus males possess a well-developed, sclerotized aedeagus; Armascirus and Rubroscirus males also possess an aedeagus, though less developed and sclerotized than in Dactyloscirus (Den Heyer 1978a, 1979a, 1981a. Castro and Moraes (2010) suggest that Cunaxatricha tarsospinosa may be cyclically or facultatively parthenogenetic -one population they studied consisted entirely of females while another population approximately 450 km distant contained males -and that parthenogenesis may be induced by cellular endosymbionts.
Cunaxids spin silk, which is used for a variety of purposes. Cunaxatricha tarsospinosa produces a webbing around eggs laid on leaves, but not branches; Castro and Moraes (2010) report that destruction of webbing may reduce viability of the eggs. Nymphal Armascirus taurus, Dactyloscirus inermis, Coleoscirus simplex, and an undescribed Pulaeus construct silken molting chambers (Alberti and Ehrnsberger 1977;Walter and Kaplan 1991); the breadth of this behavior suggests it may be widespread among cunaxids. Cunaxa setirostris constructs an irregular net of two silk varieties which is used during prey capture (Alberti and Ehrnsberger 1977). It has also been proposed that some species may be venomous, though this has not been confirmed (Den Heyer 1980a, Walter and Proctor 1999. Biogeography. Cunaxids have been found on every continent except Antarctica. South Africa and the Philippines have the most well-documented cunaxid diversity -68 and 57 species respectively -thanks to the efforts of Den Heyer and Corpuz-Raros (Den Heyer 2011a). South America was little studied until Castro and Den Heyer described 8 genera and 10 species from Brazil between 2008 and. Only two species are known from Australia, both reported by Womersley (1933), though  reported 5 undescribed species in 4 genera and Callan et al. (2011) reported another two species at the family level, suggesting many species await discovery there.
The cunaxid fauna of Europe and North America north of Mexico fall between these extremes. Most reports have been sporadic and span more than a century, beginning with Banks (1894) in the United States and Berlese (1887) in Europe. Robert L. Smiley, a well-known North American worker, never collected material. He instead worked on samples that were sent to him, often intercepted by the USDA at ports of entry, so rather than focusing on North American fauna he more generally worked on world species. This has led to a scattered understanding of the species and genera that occur in North America.

Methods
The diagnoses and keys presented are based on published descriptions and examination of available type specimens. However, for many species the types were not available for examination. The accuracy of the keys is therefore dependent upon the accuracy of the published descriptions. This also influenced which characters were chosen for couplets. Often a character that is potentially useful and informative (such as the presence or absence of a cheliceral seta) was not reported in the original description. Thus, unlike previous keys, characters such as setal counts of leg segments were often preferred. This may prove to be problematic as extra setae are sometimes reported on leg segments; however, examination of multiple specimens in a population should help overcome this.  transferred many species into different genera in the Bdelloidea database that is used by Species 2000 and ITIS Catalogue of Life (CoL). However, nomenclatural acts proposed within these databases are not considered valid under The International Code of Zoological Nomenclature as they do not conform to Article 8.4.2.2. This is intentional for a number of reasons, including avoiding circularity (e.g., a paper that cites CoL about a nomenclatural act, and CoL citing that paper) and time limitations in pursuing a publication that includes all nomenclatural acts proposed within the databases each year (Roskov and Bailly, 2 May 2014, pers. comm.).

Terminology
An effort is made to utilize terminology that is broadly applicable and well-accepted across mite taxa, despite conventions used among bdelloid researchers. Some terms widely used by bdelloid researchers are either inaccurate or outdated, and others are misleading. Therefore, we follow the suggestions outlined by Fisher et al. (2011), which are elaborated upon below.
Subcapitulum. The part of the gnathosoma that bears the palps and chelicerae has been variously termed by researchers of Bdelloidea. One such term -hypostome -more properly refers to the area of the subcapitulum anterior to the oral opening (Evans 1992;Krantz and Walter 2009), and therefore its use in reference to the entire subcapitulum is incorrect. The other term -hypognathum -is synonymous with subcapitulum, and is therefore not inaccurate, but also not broadly used across mite taxa. Thus, we reject the use of hypognathum in favor of subcapitulum and reserve the use of hypostome to the region of the subcapitulum anterior to the oral opening.
Body segmentation. The terminology associated with the acariform idiosoma remains controversial. Classically, these regions have been most widely called the propodosoma and hysterosoma. However, Grandjean (1970) proposed an alternate view of acariform idiosomal organization based on a segmentation hypothesis of van der Hammen (1963). Grandjean postulated that the podosoma is dorsally overtaken by the gnathosoma and the opisthosoma and termed the outgrowth of the gnathosoma that obscures the propodosoma the 'aspidosoma'. Under this hypothesis, referring to the anterio-dorsal half of the idiosoma as the propodosoma is inaccurate, while referring to posterio-dorsal idiosoma as the hysterosoma (opisthosoma + metapodosma) is more inclusive than necessary and should instead be denoted simply as the opisthosoma. This hypothesis has gained popularity and 'aspidosoma' is currently used across disparate acariform taxa (e.g., Caeculidae : Coineau 1974;Erythraeidae: Mąkol 2010;Penthalodidae: Jesionowska 2010;Tydeidae: Kazmierski 2008). Contrary to this, Weigmann (2001) pointed out there is neither evidence for the dorsal overgrowth of the gnathosoma obscuring the propodosoma, nor for the overgrowth of the opisthosoma obscuring the metapodosoma. Further, he provided good evidence for retaining 'propodosoma' and 'hysterosoma'. Ultimately, this matter will not be resolved without detailed investigation into developmental biology. Thomas (2012, 2013) investigated the embryology of an oribatid (Archegozetes longisetosus Aoki, 1965) and demonstrated the opisthosoma of that mite comprises only two segments. Unfortunately, their investigations are as yet unable to resolve the problem of the dorsal podosoma. Fisher et al. (2011) proposed avoiding hypothesis-dependent terminology pending further evidence for a given hypothesis. Thus, they retained 'hysterosoma' to refer to the idiosoma posterior to the sejugal furrow and implemented 'proterosoma' for the anterior idiosoma. Both terms were considered hypothesis-independent, but suffered from being more inclusive than necessary. Regardless, 'hysterosoma' is already used by many authors to refer to the dorsum posterior to the sejugal furrow, therefore its implementation is uncontroversial. Conversely, 'proterosoma' is not widely used to refer to the anterior idiosoma. Thus, referring to those setae as 'proterosomal setae' is novel, and therefore less preferred. However, recent investigations provide some support for implementing 'proterosoma' -this is discussed below.
Phylogenetic analyses of large datasets that include molecular data has corroborated previous suspicions of the non-monophyly of "Acari" and provided substantial support for a clade that combines camel spiders with acariforms called Poecilophysidea (Dabert et al. 2010, Pepato et al. 2010). In addition to characteristics of the reproductive system that have been previously noted (Alberti 1980a, b, 2000, Alberti and Peretti 2002, Klann et al. 2009), Dunlop et al. (2012) suggested that the sejugal furrow of Acariformes is homologous to a similar body division in Solifugae, lending another potential synapomorphy for this clade. Because of this, the sejugal furrow was elevated as a key morphological trait among both camel spiders and acariforms, which now makes it possible to construct terminology founded in a well-supported hypothesis. This renders terms that are denoted relative to the sejugal furrow (like 'proterosoma' and 'hysterosoma') as hypothesis-dependent, which is only preferred over hypothesisindependent terminology when the hypothesis is well-supported.
Therefore, we continue with the suggestions of Fisher et al (2011) in using 'proterosoma' and 'hysterosoma' for two reasons: 1) they are hypothesis-independent with respect to Grandjean's 'aspidosoma' and Weigmann's 'propodosoma'; and 2) since 2011, they have been found to be hypothesis-dependent, but on well-supported hypotheses. Obviously, as future research resolves the issue of the acariform idiosomal dorsum (i.e. Grandjean vs. Weigmann), we suggest that new terminology based on those hypotheses should be adopted.
Idiosomal setae. For hysterosomal setae, we follow the notation of Grandjean (1939Grandjean ( , 1947 that has been widely adopted by acarologists (e.g., van der Hammen 1970;Lindquist 1976Lindquist , 1977Kethley 1990;Swift 1996). However, proterosomal setae remain problematic. Historically, proterosomal chaetotaxy followed Grandjean (1939Grandjean ( , 1947, which identified internal/external verticals (vi and ve) and internal/external scapulars (sci and sce). This notation has always been cumbersome for groups like Bdelloidea which have sci always external to sce. Given that homology has not been determined for these setae across mite taxa, some authors suggested simply switching the designations of sci and sce to reflect their position (Den Heyer and Castro 2008a, b, c;Den Heyer 2011c). As a result, frustratingly, the literature now has both sci and sce referring to each set of setae.
Therefore, we reject the suggestion of Den Heyer and Castro (2008) and follow the suggestion of Fisher et al. (2011), which resorts to a modified version of Atyeo (1960) when referring to proterosomal setae: anterior/posterior trichobothria (at/pt), and lateral/median proterosomal setae (lps/mps). Obviously, once homology of these setae can be determined across mite taxa, we suggest revising the terminology accordingly.
Illustrations were produced using the methods outlined by Fisher and Dowling (2010).
for those mites in the family Bdellidae (sensu Dugés) that have pedipalps with a curved terminal segment and movable chela only (= Cunaxidae sensu Thor). Thor (1902) erected Cunaxidae as a family separate from Bdellidae. Oudemans (1906) disregarded Thor's (1902) erection of Cunaxidae and kept Cunaxinae as a subfamily within Bdellidae. Van der Hammen (1972) erected the superfamily Cunaxoidea over Bdelloidea, disregarding the priority of Bdella Latreille (1795) over Cunaxa Von Heyden (1826). Den Heyer (1977b) erected Bonziinae for Bonzia and Parabonzia. Den Heyer (1978a) preserved the name Cunaxinae, but limited its concept to those cunaxids possessing 5-segmented pedipalps which extend past the subcapitulum by at least the distal two segments. Den Heyer (1978b) erected Coleoscirinae. Den Heyer (1980c) erected the monobasic Scirulinae and recognized the priority of Bdelloidea over Cunaxoidea. Bu and Li (1987a) erected Orangescirulinae.  erected Denheyernaxoidinae, Neobonzinae, and Paracunaxoidinae as monotypic subfamilies and monographed and provided keys to known species. Den    (Figs 3-6). Pedipalps 3-, 4-, or 5-segmented and end in a strong claw (except in Pseudobonzia). They may be shorter than, equal to, or extend beyond the distal end of the subcapitulum. Femora of 5-segmented pedipalps divided into basi-and telofemora, though may be secondarily fused; a dark line often indicates the previous articulation ( Fig. 5a, b illustrate a fully divided femur and Fig.  6a, b illustrate a secondarily fused femur. This is for illustration purposes only, i.e., cunaxids with long and short 5-segmented pedipalps may have either fully divided or secondarily fused femora). Telofemora and genua are uniquely fused in Allocunaxa, though the basifemoral/telofemoral articulation is present. Apophyses present or not on the telofemora, adjoining the genua and tibiotarsi, or on the tibiotarsi. Subcapitulum wedge-shaped and may be patterned with random dots or papillae, dots or papillae forming lines, a single row of cells on the posterior edge, or reticulations forming polygonal cells. Subcapitulum with up to 6 pairs of setae are present: hg 1-4 and 2 pairs of adoral setae. Seta hg 1 usually straight, but geniculate in Bonziinae and may be curved in Neoscirula; hg 4 often longest pair of subcapitular setae. Chelicerae with or without seta near the cheliceral digit.
Idiosoma, dorsal (Fig. 7a). Idiosoma diamond-shaped. Dorsal proterosoma covered with a sclerotized shield that bears 2 pairs of setae (lps and mps) and 2 pairs of setose sensilla (at and pt); rarely one pair of setae or sensillae absent. Dorsal hysterosoma complemented with 0-2 large shields or plates and 0-4 pairs of platelets. These plates and platelets may capture one or more pairs of setae. Up to 8 pairs of dorsal hysterosomal setae present (c 1 -h 1 , c 2 , f 2 , and h 2 ); h 2 may occur ventrally. Setae may occur on small platelets that are barely larger than the setal socket. Integument not covered in shields, plates, or Figures 3-6. a. dorsal. b. ventral. 3 3-segmented pedipalp (Cunaxoidinae) 4 4-segmented pedipalp (Scirulinae) 5 5-segmented pedipalp that does not extend beyond the subcapitulum by more than the distal half of the genua (Bonziinae, Coleoscirinae, and Orangescirulinae) 6 5-segmented pedipalp that reaches beyond the subcapitulum by at least the distal half of the genua (Cunaxinae). platelets is striated. Cupule im present, usually laterad and slightly posterior to e 1 . Dorsal idiosomal shields and plates smooth or patterned with random dots or papillae, dots or papillae forming lines, reticulations forming polygonal cells, or cells which form rows. Idiosoma,ventral (Fig. 7b) Ventral idiosoma may be complemented with 1 or a few small platelets in addition to the coxae. Coxae fused to body and form plates. Coxae I-II are often fused in adults and may coalesce medially to form a sternal shield. Coxae III-IV are often fused in adults and may extend caudally beyond the genital plates. Each coxa complemented with 0-4 setae; in addition, extensive coxae or sternal shields may capture setae normally on the integument and therefore have more. Coxae may be plain or patterned with random dots or papillae, dots or papillae forming lines, or reticulations forming polygonal cells. Genital plates (sometimes called anal valves) present in adults and bear 3 (rarely) or 4 (usually) setae, except in Parabonzia which have up to 9 pairs of setae. 2 pairs of genital papillae visible underneath the plates. Anal plates (sometimes called anal valves) bear 1-2 setae (ps 1-2 ). Setae ps 2 may occur off the anal plates. Legs 6-segmented in larvae, 7-segmented in nymphs and adults. In adults these segments are coxa, trochanter, baifemur, telofemur, genu, tibia, and tarsus, however, the coxae are often treated separately from the other leg articles. Femora undivided in larvae. Trichobothrium present on leg tibia IV. Ambulacral claws present on either side of a 4-rayed empodium.

Diagnosis.
Gnathosoma. Pedipalps 5-segmented and reach beyond the subcapitulum by at most the distal half of the tibiae. Apophyses absent. A multi-branched seta present dorsally on the telofemora. Tibiotarsi terminate in a stout claw or two strong setae. 2 pairs of adoral setae present or absent. Subcapitulum with 4 pairs of setae (hg 1-4 ) present in Bonzia; up to 6 pairs of subcapitular setae (hg 1-4 + additional setae) present in Parabonzia. Idiosoma, dorsal. Proterosoma bears a shield complemented with 2 pairs of setae (at and pt) and 2 pairs of setose sensillae (lps and mps). Dorsal hysterosoma may bear a shield; if a shield is present it may be complemented with a variable number of setae depending on the extent of the shield. Setae c 1 -h 1 , c 2 , f 2 and h 2 present and are smooth or spiculate. Cupule im present laterad and caudally of e 1 . Integument that does not bear shields or plates is striated.
Idiosoma, ventral. Coxae I-II fused or not and coxae III-IV fused or not. Genital plates bear 4-9 setae; 2 pairs of genital papillae visible underneath the plates. Up to 4 pairs of setae present on the anal plates. Up to 9 pairs of setae present on the integument between coxae II and the anal plates. Legs. Trichobothrium present on leg tibia IV. The ambulacral claws occur on either side of a 4-rayed empodium.
Idiosoma, dorsal. proterosoma bears a shield complemented with 2 pairs of setae (at and pt) and 2 pairs of setose sensillae (lps and mps). The dorsal hysterosoma bears a shield that may be complemented with a variable number of setae depending on the extent of the shield. Setae c 1 -h 1 , c 2 , f 2 and h 2 present, and are smooth or spiculate. Cupule im present laterad and caudally of e 1 . Integument that does not bear shields or plates is striated.
Coxae I-II fused and coxae III-IV fused. Genital plates bear 4 setae; 2 pairs of genital papillae visible underneath the plates. 4 pairs of setae present on the anal plates. Trichobothrium on leg tibia IV present. Ambulacral claws occur on either side of a 4-rayed empodium.

Diagnosis.
Gnathosoma. Pedipalps 5-segmented and reach beyond the subcapitulum by at most the distal half of the tibiae. Apophyses absent. A multi-branched seta present dorsally on the telofemora. Tibiotarsi terminate in two strong setae. 2 pairs of adoral setae present or absent. Subcapitulum with up to 8 pairs of setae present.
Idiosoma, dorsal. Proterosoma bears a shield complemented with 2 pairs of setae (at and pt) and 2 pairs of setose sensillae (lps and mps). Dorsal hysterosoma may bear a shield; if a shield is present it may be complemented with a variable number of setae depending on the extent of the shield. Setae c 1 -h 1 , c 2 , f 2 and h 2 present and smooth. Cupule im is present laterad and caudally of e 1 . Integument that does not bear shields or plates is striated.
Idiosoma, ventral. Coxae I-II fused or not and coxae III-IV fused or not. Genital plates with up to 9 pairs of setae; 2 pairs of genital papillae visible underneath the plates. Up to 4 pairs of setae present on the anal plates. Up to 9 pairs of setae on the integument between coxae II and the anal plates. Legs. Trichobothrium on leg tibia IV present. The ambulacral claws occur on either side of a 4-rayed empodium. Historical review. Koch (1838) established Eupalus and described the first mite belonging to Cunaxoidinae, Eupalus croceus. Baker and Hoffmann (1948) proposed Cunaxoides to replace Eupalus Koch as the name was preoccupied (a fact that acarolo-gists had missed for 100 years) by Eupalus Gistl; they also redescribed and reillustrated a number of known species. Radford (1950) proposed Haleupalus to replace Eupalus, though this name is invalid because it is predated by Cunaxoides.  erected Neocunaxoides and reviewed Cunaxoides. Both genera were assigned to the newly established Cunaxoidinae by Den Heyer (1978c). Pulaeus was established by Den Heyer (1979b); the name is an anagram and nod to Eupalus. Den Heyer (1979c) erected Scutopalus for those cunaxoidines with well-demarcated dorsal and ventral plates.  synonymized Scutopalus with Neocunaxoides and Haleupalus with Cunaxoides; he also erected Denheyernaxoides and Paracunaxoides as monotypic genera in two new subfamilies, Denheyernaxoidinae and Paracunaxoidinae respectively. Castro and Den Heyer (2009) split a new genus, Lupaeus, from Pulaeus based on the number of setae on basifemora IV (1 and 2, respectively) and the number of pointed processes on the pedipalpal tibiotarsi (2 and 1, respectively). Den  split Bunaxella, Dunaxeus, Funaxopsis, and Qunaxella from Cunaxoides; they also moved Denheyernaxoides and Paracunaxoides to Cunaxoidinae, thus disregarding Denheyernaxoidinae and Paracunaxoidinae as valid subfamilies. Diagnosis. Gnathosoma. Pedipalps 3-segmented: a trochanter which lacks setae, fused femurogenu (femur + genu) which is complemented with 5 or 6 setae, and tibiotarsus (tibia + tarsus) which is complemented with 5 or 6 setae. Tibiotarsi may be complemented with a bladder-or bulb-like apophysis. Pedipalps do not reach beyond the subcapitulum by more than the distal half of the tibiotarsi. Chelicera with or with- Parabonzia key illustrations. 12a Unbranched pedipalp telofemoral seta 12b Multibranched pedipalp telofemoral seta 13 Lightly barbed pedipalp tibiotarsal sigmoid seta 14 Spur-like process on femora III. out seta near the cheliceral digit. Subcapitulum with 4 pairs of setae (hg 1-4 ) are present; setae hg 4 is often the longest. 2 pairs of adoral setae are present or absent.

Key to adult female
Idiosoma, dorsal. Female with proterosomal shield (absent in Cunaxoides ulcerosus) which is complemented with two pairs of setae (lps and mps) and two pairs of setose sensillae (at and pt) and may bear a hysterosomal plate complemented with a varying number of setae; when present the dorsal hysterosomal plate may be fused with the proterosomal shield. Dorsal plates well demarcated or not. Dorsal setae c 1 -h 1 are present; c 2 , f 2 and h 2 may also be present. If f 2 is present, f 1 and f 2 may be located together on a small platelet. Setae not on larger plates may be born on small platelets barely larger than the setal socket. Cupule im present laterad and posterior of e 1 . Integument that is not covered in shields or plates is striated Idiosoma, ventral. Coxae of female vary in size, from being restricted to the trochantral bases to being extensive and nearly forming a holoventral shield. Coxae may or may not be well demarcated. Coxae I-II fused (usually) or not, coxae III-IV fused (usually) or not. Coxae I-II may coalesce medially to form a sternal shield. The genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath the plates. The anal plates bear one pair of setae (ps 1 ); one pair of setae is present ventrally on the integument near the anal plates (either ps 2 or pa). Cupule ih is present ventrally laterad the integumental setae associated with the anal plates. The integument that is not covered in shields or plates is striated. Legs. Tarsi never constricted apically so as to end in lobes. Trichobothrium on leg tibia IV present. Ambulacral claws are rippled and occur on either side of a 4-rayed empodium. Diagnosis. Gnathosoma. Pedipalps 3-segmented. Femurogenua are at least twice as long as wide and complemented with 5 setae. Tibiotarsi at least twice as long as wide and usually complemented with 6 setae. A small apophysis present basally and a pointed process occurs near the terminal tip; a ridge present between the apophysis and pointed process. Subcapitulum with 6 pairs of setae (hg 1-4 and 2 pairs of adoral setae) present; setae hg 4 is often the longest. Chelicera without seta.

Key to adult female Cunaxoidinae
Idiosoma, dorsal. Proterosoma bears an ill-defined and weakly sclerotized shield which is complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). The dorsal hysterosoma may or may not bear a plate; if a plate is present it is ill-defined and weakly sclerotized, may be complemented with a variable number of setae, and may or may not be fused with the proterosomal shield. Setae c 1 -h 1 , c 2 , and h 2 are present. Seta c 2 plumose or fan-shaped. Cupule im is present laterad and posterior of e 1 . Integument that is not covered in shields or plates is striated. Idiosoma, ventral. Coxae are weakly sclerotized and ill-defined; they can be recognized by possessing somewhat denser striations than the surrounding integument. Coxae I-II may be fused and may coalesce medially to form a sternal shield. Coxae III-IV fused or not. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath the plates. Anal plates bear one pair of setae; one pair of setae is present ventrally on the integument near the anal plates. Up to 7 pairs of setae present on the integument between the coxal and genital plates. Cupule ih present ventrally laterad the integumental setae associated with the anal plates. Integument that is not covered in shields or plates is striated. Legs. Tarsi are never constricted apically so as to end in lobes. Depression for the famulus on tarsus I is absent. Tibia III complemented with 1 bsl, 5 sts. Tibia IV is complemented with 4 sts and lacks a trichobothrium. Ambulacral claws occur on either side of a 4-rayed empodium.   Garman (1948) reported E. biscutum from apple trees in Connecticut. Baker and Hoffmann (1948) recognized that the name Eupalus was preoccupied and erected Cunaxoides to replace it; they transferred all known Eupalus to the new genus and figured each species. Haleupalus oliveri was described by Schruft (1971).  synonymized C. vitellinus with C. croceus and provided a translation of Thor and Willmann's (1941) description of C. croceus. Den Heyer (1978c) placed Cunaxoides as the type genus in the newly erected Cunaxoidinae; he also redescribed the genus and redescribed and designated a neotype for C. croceus. Kuznetzov and Livshitz (1979) described C. ulcerosus, C. longistriatus, C. fidus and C. desertus and reported and figured C. biscutum, and C. parvus from Russia. Gupta and Ghosh (1980) described C. nicobarensis. C. kielczewskii was described by Michocka (1982).   Diagnosis. Gnathosoma. Pedipalps 3-segmented. Femurogenua at least twice as long as wide and complemented with 5 setae. Tibiotarsi at least twice as long as wide and usually complemented with 6 setae. A small apophysis present basally and a pointed process present near the terminal tip; a ridge present between the apophysis and pointed process. Subcapitulum with 6 pairs of setae (hg 1-4 and 2 pairs of adoral setae) are present; setae hg 4 longest. Chelicera without seta.
Idiosoma, dorsal. Proterosoma bears an ill-defined and weakly sclerotized shield which is complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). The dorsal hysterosoma may or may not bear a plate; if a plate is present it is ill-defined and weakly sclerotized, may be complemented with a variable number of setae, and may or may not be fused with the proterosomal shield. Setae c 1 -h 1 , c 2 , and h 2 are present. Cupule im present laterad and posterior of e 1 . Integument that is not covered in shields or plates is striated.
Idiosoma, ventral. Coxae weakly sclerotized and ill-defined; they can be recognized by possessing somewhat denser striations than the surrounding integument. Coxae I-II may be fused and may coalesce medially to form a sternal shield. Coxae III-IV may be fused. Each coxa is complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath the plates. Anal plates bear one pair of setae; one pair of setae present ventrally on the integument near the anal plates. Up to 7 pairs of setae present on the integument between the coxal and genital plates. Cupule ih present ventrally laterad the integumental setae associated with the anal plates. Integument that is not covered in shields or plates is striated. Legs. Tarsi never constricted apically so as to end in lobes. Trichobothrium present on leg tibia IV. Ambulacral claws are rippled and occur on either side of a 4-rayed empodium.

Key to adult female Cunaxoides
The following species have not been included because the original descriptions and subsequent papers describing them (Thor and Willmann 1941;Baker and Hoffmann 1948) are not in English; known illustrations do not contain enough detail; and the types were not examined: C. minima (Trägårdh, 1910), C. minutissimus (Koch, 1938), C. vitellinus (Koch, 1941 Diagnosis. Gnathosoma. Pedipalps 3-segmented. Femurogenua at least twice as long as wide, complemented with 5 setae. Tibiotarsi at least twice as long as wide, usually complemented with 6 setae. A small apophysis occurs basally and a pointed process occurs near the terminal tip; a ridge runs between the apophysis and pointed process. Subcapitulum with 4 pairs of setae (hg 1-4 ); setae hg 4 often the longest. Adoral setae absent. Chelicera without seta.
Idiosoma, dorsal. Proterosoma lacks a shield, complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma lacks a plate. Setae c 1 -h 1 , c 2 , and f 2 , h 2 present. Cupule im present laterad and posterior of e 1 . Integument not covered in shields or plates is striated.
Idiosoma, ventral. Coxae I-II connected by small apodemes. Coxae III-IV fused. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae; 1 pair of setae present ventrally on the integument near the anal plates. 5 pairs of setae present on the integument between the coxal and genital plates. Cupule ih present ventrally laterad the integumental setae associated with the anal plates. Integument not covered in shields or plates is striated.
Legs. Femora I and II not divided. Trichobothrium on tibia IV absent. Tarsi never constricted apically so as to end in lobes. Ambulacral claws on either side of a 4-rayed empodium present. Diagnosis. Gnathosoma. Pedipalps 3-segmented. Femurogenua at least twice as long as wide, complemented with 5 setae. Tibiotarsi at least twice as long as wide, usually complemented with 6 setae. A small apophysis occurs basally and a pointed process occurs near the terminal tip; a ridge runs between the apophysis and pointed process. Subcapitulum with 4 pairs of setae (hg 1-4 and 2 pairs of adoral setae); setae hg 4 is often the longest. Chelicera without seta.
Idiosoma, dorsal. Proterosoma bears an ill-defined and weakly sclerotized shield which is complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma may or may not bear a plate; if a plate is present it is ill-defined and weakly sclerotized, may be complemented with a variable number of setae, and may or may not be fused with the proterosomal shield. Setae c 1 -h 1 , c 2 , and h 2 are present. Cupule im is present laterad and posterior of e 1 . The integument that is not covered in shields or plates is striated.
Idiosoma, ventral. Coxae weakly sclerotized and ill-defined; they can be recognized by possessing somewhat denser striations than the surrounding integument. Coxae I-II may be fused and may coalesce medially to form a sternal shield. Coxae III-IV fused. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath plates. Anal plates bear 1 pair of setae; 1 pair of setae present ventrally on the integument near the anal plates. Up to 7 pairs of setae present on the integument between the coxal and genital plates. Cupule ih present ventrally laterad the integumental setae associated with the anal plates. Integument not covered in shields or plates is striated. Legs. Tarsi never constricted apically so as to end in lobes. Tibia III complemented with 5 sts (4 short, 1 long). Tibia IV complemented with 5 sts (4 short, 1 long), and lacks a trichobothrium. Ambulacral claws on either side of a 4-rayed empodium present. Diagnosis. Gnathosoma. Pedipalps 3-segmented. Femurogenua at least twice as long as wide, complemented with 5 setae. Tibiotarsi at least twice as long as wide, usually complemented with 6 setae. A small apophysis occurs basally and a pointed process occurs near the terminal tip; a ridge runs between the apophysis and pointed process. Subcapitulum with 6 pairs of setae (hg 1-4 and 2 pairs of adoral setae); setae hg 4 is often longest. Chelicera without seta.

Key to adult female
Idiosoma, dorsal. Proterosoma bears an ill-defined and weakly sclerotized shield complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma may or may not bear a plate; if plate present, it is ill-defined and weakly sclerotized, may be complemented with a variable number of setae, and may or may not be fused with the proterosomal shield. Setae c 1 -h 1 , c 2 , and h 2 present. Cupule im present laterad and posterior e 1 . Integument not covered in shields or plates striated.
Idiosoma, ventral. Coxae weakly sclerotized and ill-defined; they can be recognized by possessing somewhat denser striations than the surrounding integument. Coxae I-II may be fused and may coalesce medially to form a sternal shield. Coxae III-IV may be fused. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae; 1 pair of setae present ventrally on the integument near the anal plates. Up to 7 pairs of setae present on the integument between the coxal and genital plates. Cupule ih present ventrally laterad integumental setae associated with the anal plates. Integument not covered in shields or plates striated. Legs. Tibia III complemented with 1 bsl and 3, 4, or 5 sts. Tibia IV complemented with 3 sts (2 short, 1 long) and lacks a trichobothrium. Tarsi never constricted apically so as to end in lobes. Ambulacral claws on either side of a 4-rayed empodium present. Historical review. Berlese (1916) described Eupalus subterraneus. Thor and Willmann (1941) redescribed E. subterraneus. Baker and Hoffmann (1948) erected Cunaxoides in place of Eupalus as Eupalus was preoccupied; they also described C. minutus and redescribed and illustrated C. subterraneus. Den Heyer (1979b) erected Pulaeus, moving those species with f 2 present and setae present on basifemora IV to the new genus from Cunaxoides; he also described P. martini and P. clarae and placed Pulaeus into the subfamily Cunaxoidinae. Pulaeus platygnathus was described by Bu and Li (1991). Corpuz-Raros (1996b) described P. dentatus, P. lenis, P. longisetus, P. villacarlosae, and P. filipinus from the Philippines. Hu (1997) reported P. platygnathus from China. Lin and Zhang (2000) reported P. platygnathus from China. Lin and Zhang (2003) reported P. minutus from China. Corpuz-Raros (2007)  Diagnosis. Gnathosoma. Pedipalps 3-segmented. Femurogenua at least twice as long as wide, complemented with 6 setae. Tibiotarsi at least twice as long as wide, usually complemented with 6 setae; they possess 2 or 3 pointed processes and may possess a bladder-or knob-like apophysis (Fig. 24a). Subcapitulum with 6 pairs of setae (hg 1-4 and 2 pairs of adoral setae); setae hg 4 often the longest. Chelicera with seta present.

Key to adult female
Idiosoma, dorsal. Proterosoma bears a well-sclerotized shield complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma bears a sclerotized plate that is variable in size and fused with the proterosomal shield; it may be complemented with a variable number of setae depending on the size of the plate. Setae c 1 -h 1 , c 2 , f 2 , and h 2 present. Cupule im present laterad and posterior of e 1 . Integument not covered in shields or plates is striated.

Idiosoma, ventral.
Coxae sclerotized and well-defined. Coxae I-II may be fused and may coalesce medially to form a sternal shield. Coxae III-IV may be fused. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ). Setae g 1,2,4 usually occur in a straight line near the midline and setae g 3 occur near the edge of the genital plates (Fig. 24b). 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae; 1 pair of setae present ventrally on the integument near the anal plates. Cupule ih present ventrally laterad; the integumental setae associated with the anal plates. Integument not covered in shields or plates striated. Legs. Tarsi never constricted apically so as to end in lobes. Trichobothrium on leg tibia IV present. Ba sifemora setal formula 4-6-3-1. Depression of the famulus occurs on distal half of tarsus I. Tibiae I-II possess striated blunt solenidia. Ambulacral claws rippled and occur on either side of a 4-rayed empodium.

Key to adult female Lupaeus
Lupaeus longisetus is known only from the male and is not included in the key. It can be recognized by the following characters: small platelet between the edges of a divided sternal shield absent, basifemora I with 3 sts, and setae e 1 elongate and barbed (Fig. 25a).
Lupaeus polilloensis is only known from the male and is not included in the key. It can be recoginized by the following characters: small platelet between the edges of a divided sternal shield absent; basifemora I-II setal formula 4-6; platelets complemented with setae f 1 , f 2 with fused medially into one plate; and the dorsal shield densely granulate (Fig. 25b).
Idiosoma, dorsal. Proterosoma bears a well-sclerotized shield which is complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma bears a sclerotized plate which is variable in size and fused with the proterosomal shield; it may be complemented with a variable number of setae depending on the size of the plate. Setae c 1 -h 1 , c 2 , and h 2 present. Setae f 2 absent. Cupule im present laterad and posterior of e 1 . The integument not covered in shields or plates is striated.
Idiosoma, ventral. Coxae sclerotized and well-defined. Coxae I-II may be fused and may coalesce medially for form a sternal shield. Coxae III-IV may be fused. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ), which are usually in a straight now; 2 pairs of genital papillae visible underneath the plates. Anal plates bear one pair of setae; one pair of setae is present ventrally on the integument near the anal plates. Cupule ih present ventrally laterad the integumental setae associated with the anal plates. Integument not covered in shields or plates is striated. Legs. Tarsi never constricted apically so as to end in lobes. Trichobothrium on leg tibia IV present. Basifemora setal formula 3-5-2-0. Ambulacral claws rippled and occur on either side of a 4-rayed empodium.

Key to adult female Neocunaxoides
Cunaxoides philippinensis (Corpuz-Raros, 2007) is regarded as belonging to Neocunaxoides because it has 6 seatae on the femurogenu and lacks setae f 2 . Neocunaxoides makapalus, N. philippinensis (Corpuz-Raros, 1996c), N. unguianalis, and N. rugosus are regarded as belonging to Scutopalus as they possess 5 sts on pedipalp femurogenu and extensive dorsal shields. They have therefore not been included in the following key.
Neocunaxoides biramus is not included in the key because it is only known from the male. It can be distinguished from all other Neocunaxoides, and indeed all de- Neocunaxoides metwallyi is not included in the key as, despite the best efforts of the authors and the University of Arkansas Interlibrary Loan Department, the description could not be obtained.
Idiosoma, dorsal. Proterosoma complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). A pair of oval shields formed by flat, bacilluslike striae present between the sensillae. Setae c 1 -h 1 , c 2 , and h 2 present. Setae f 2 absent. Integument not covered in shields or plates is striated.
Idiosoma, ventral. Coxae sclerotized and well-defined. Coxae I-II thinly connected. Coxae III-IV more broadly connected. Genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae; 1 pair of setae present ventrally on the integument near the anal plates. Integument not covered in shields or plates is striated. Legs. Trichobothrium on tibia IV present.
Idiosoma, dorsal. Proterosoma bears a well-sclerotized shield, complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma bears a sclerotized plate which is variable in size and fused with the proterosomal shield; it may be complemented with a variable number of setae depending on the size of the plate. Setae c 1 -h 1 , c 2 , f 2 , and h 2 and present. Cupule im present laterad and posterior of e 1 . Integument not covered in shields or plates striated.
Idiosoma, ventral. Coxae sclerotized and well-defined. Coxae I-II may be fused and may coalesce medially to form a sternal shield. Coxae III-IV may be fused. Each coxa complemented with 2-4 setae. Genital plates each bear 4 setae (g 1-4 ), which are usually in a straight row; 2 pairs of genital papillae visible underneath the plates. Anal plates bear one pair of setae; 1 pair of setae present ventrally on the integument near the anal plates. Cupule ih present ventrally laterad the integumental setae associated with the anal plates. The integument not covered in shields or plates striated. Legs. Tarsi never constricted apically so as to end in lobes. Trichobothrium on leg tibia IV present. Depression of the famulus occurs on proximal half of tarsus I. Tibiae I-II possess non-striated blunt solenidia. Ambulacral claws rippled and occur on either side of a 4-rayed empodium.

Key to adult female Pulaeus
P. ardeola was not included in the key because the original text is in Cyrillic script and the illustrations do not provide enough characters to differentiate it from other species. N. cinctus is moved from Neocunaxoides to Pulaeus based on features given in the original description, namely that f 2 is present and basifemora IV are complemented with 2 sts.
The following were species assigned to Pulaeus before Lupaeus was erected. The characters that divide the two genera are not given in the original species descriptions and types have not been viewed. These indeterminable species are therefore not included in either generic key, but instead characters are given for each species that will serve to identify them.
P. parapatzuarensis (Shiba, 1978) -This species has a divided sternal plate, lacks a sclerotized area anterior to the genital plates, and does not have f 1,2 located on platelets. In addition it has 6 pairs of setae on the integument between coxal and genital plates.
P. patzcuarensis (Baker & Hoffmann, 1948) -This species can be recognized by the sternal plates being connected anteriorly and divided in a v-shape posteriorly.
P. pseudominutus (Shiba, 1978) -Setae e 1 being 3 times the length of c 1 and d 1 distinguishes this species.
P. payatopalpus (Corpuz-Raros, 1996) -The hypostome is 2/3 the length of the gnathosoma and the pedipalps are extremely long and slender, at least 8 times longer than wide. In addition the tibiotarsus is complemented with a seta that is longer than the segment.
Idiosoma, dorsal. Proterosoma with weakly defined shield present which is complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma lacks a plate. Setae c 1 -h 1 , c 2 , and h 2 present. Setae c 1 -f 1 finely setose and c 2 , h 1 , and h 2 smooth. Setae f 2 absent. Integument not covered in shields or plates striated.
Idiosoma, dorsal. Proterosoma with a well-defined shield present, complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma with a well-defined plate fused to the proterosomal plate. Small platelets may be present laterad and posterior to the dorsal shield. Setae c 1 -h 1 , c 2 , and h 2 present. Setae f 2 absent. Integument not covered in shields or plates striated.
Key to female Scutopalus (modified from Rocha et al. 2013).
As suggested by Den Heyer (2011b) Neocunaxoides pradhani (Gupta and Ghosh 1980) and N. gilbertoi (Bashir and Afzal 2004) are transferred to Scutopalus as they posses 5 setae on the femurogenu instead of 6 as in Neocunaxoides and have well-demarcated plates. 1 Coxae I-II faintly or totally divided (Fig. 40a, b)  Coxae I-II faintly divided (Fig. 40a)  At least 2 pairs of thick rod-like setae on the dorsum (Fig. 41)
Idiosoma, dorsal. Proterosoma covered in a plate which bears 4 pairs of setae: 2 pairs of simple setae (lps and mps) and 2 pairs of setose sensilla (at and pt). Dorsal hysterosoma may or may not be complemented with a plate. 6 dorsal setae, c 1 -h 1 , c 2 present. Cupule im present.
Idiosoma, ventral. Coxae I-IV fused, resulting in a complete shield covering the ventral idiosoma. Genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. Cupule ih present. Anal plates bear 2 pairs of setae (ps 1 and ps 2 ); 1 pair of setae born on integument next to anal plates.

1
Hysterosomal shield present (Fig. 45a) Vitzthum (1931) raised Dactyloscirus to full generic status but later (1940-43) treated it as a subgenus. Thor and Willmann (1941) again elevated Dactyloscirus to generic status and designated Dactyloscirus eupaloides as the type specimen. Baker and Hoffmann (1948) regarded Dactyloscirus as a senior syno-nym of Cunaxa.  synonymized Rosenhofia with Dactyloscirus. Den Heyer (1978a) preserved the name Cunaxinae, but limited its concept to those cunaxids possessing 5-segmented pedipalps that extend past the subcapitulum by at least the distal two segments; he also erected Armascirus. Den Heyer (1979d) erected Rubroscirus for R. africanus. Gupta and Ghosh (1980) erected Indocunaxa.  synonymizedRubroscirus with Cunaxa but failed to give his reasoning for doing so. Den Heyer (2006) erected Riscus for a species known only from Thailand. Castro and Den Heyer (2008) erected Cunaxatricha and provided a key to the genera of Cunaxinae. Den Heyer and Castro (2008) erected Allocunaxa for a Neotropical species, synonymized Indocunaxa with Armascirus, and provided the most up-to-date key to world genera of Cunaxinae. Diagnosis. Gnathosoma. Pedipalps 5-segmented and extend beyond the subcapitulum by at least the distal half of the tibiae. Basifemora and telofemora fused but often dark line remains to indicate the division between the segments; telofemora and genua also fused in this manner in Allocunaxa. Apophyses may be present on the telofemora and between the genua and tibiotarsi. Tibiotarsi end in a strong claw. Chelicera with or without seta. Subcapitulum with up to 6 pairs of setae; setae hg 1-4 always present, 2 pairs of adoral setae present or absent. Setae hg 4 longest. In species with pedipalpal apophyses, the apophyses of the males shorter.
Idiosoma, dorsal. Female proterosoma bears a shield complemented with 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Dorsal hysterosoma may bear any combination of a median plate and lateral platelets (i.e., median plate and platelets absent, only median plate present, only lateral platelets present, or both median plate and lateral platelets present). Median plate, if present, may be complemented with 0-6 pairs of dorsal setae; lateral platelets, if present, may bear setae c 2 . Setae not born on plates or platelets may be born on tiny platelets barely larger than the setal socket. Integument that does not bear plates or platelets striated. Males differ in that the dorsal shields often more extensive and may be holodorsal.
Idiosoma, ventral. Coxae I-II fused or divided and may coalesce medially to form a sternal shield; coxae III-IV fused or divided and may extend caudally past the genital plates. Coxae each complemented 0-3 setae. Genital plates each bear 4 setae (g 1-4 ); 2 pairs of genital papillae visible underneath the plates. Anal plates complemented with at least one pair of setae, ps 1 . Setae ps 2 present or absent, either on the anal plates or on the integument adjacent to the anal plates. Setae h 2 present ventrally on the integument adjacent to the anal plates. Cupule ih present laterad of h 2 . Integument that does not bear plates striated. Legs. Tarsi constricted apically so as to end in lobes. A trichobothrium on tibia IV present or absent. Pedipalpal basifemora with simple seta (Fig. 46f); coxae II-IV setal formula usually 1-3-3 (male) or 2-3-3 (female); famulus normal; pedipalpal apophyses (when present) usually long in females and short in males, and with pointed apices (Fig. 46f)  Pedipalpal basifemora with spine-like seta (Fig. 46g); coxae II-IV setal formula usually 3-3-3; famulus large, broad based with tri-pronged tip; pedipalpal apophyses (when present) usually equal length in females and males, and with bulbous apices (Fig. 46g)  Diagnosis. Gnathosoma. Pedipalps 5-segmented, end in a strong claw, and extend beyond the subcapitulum by at least the last segment. Pedipalpal apophyses absent. Basifemora complemented with a long simple seta and telofemora with a short simple seta; these two segments fused, although a line remains visible and they can thus be differentiated. Telofemora and genu nearly fused, although a line remains visible and they can thus be differentiated. Subcapitulum complemented with 6 pairs of setae (hg 1-4 and 2 pairs of adoral setae) and covered by integumental papillae.
Idiosoma, ventral. Coxae I and II fused. Coxae III and IV fused. Genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. Integument between plates striated and bears 4 pairs of additional setae. Legs shorter than the body. Leg 4 longest. Famulus on tarsi I normally shaped. Tarsi constricted apically, resulting in large tarsal lobes. Trichobothrium on leg tibia IV present. Ambulacral claws on either side of a 4-rayed empodium present.

Armascirus Den Heyer, 1978
Historical review. The first Armascirus was described by Kramer (1881) as Scirus taurus. Berlese (1888) described S. taurus var. bison. Banks (1894) described S. quadripilis. Thor (1902) transferred S. taurus to Cunaxa. Banks (1914) described C. armata. Miller (1925) reported S. quadripilis from Ohio. Womersley (1933) reported C. taurus from Australia. Thor and Willmann (1941) transferred S. taurus var. bison to Cunaxa and raised it to full species status, viz. C. bison and transferred S. quadripilis to Cunaxa; they also redescribed and figured C. armata, C. bison, C. quadripilis, and C. taurus. Baker and Hoffmann (1948) synonymized S. quadripilis and C. armata with C. taurus; they followed Thor and Willmann (1941) in placing C. taurus var. bison in Cunaxa but declined to recognize it as a species and instead kept it as a variety or subspecies of C. taurus. Zaher et al. (1975b) collected C. taurus in Egypt. Chaudhri (1977) described Dactyloscirus ebrius and D. fuscus from Pakistan. Den Heyer (1978a) split Armascirus from Dactyloscirus and Cunaxa and raised the subfamily Cunaxinae to accommodate them, thus refining the definitions of all three genera; he transferred C. taurus and C. bison to the new genus Armascirus; and described A. huyssteeni, A. lebowensis, A. limpopoensis, and A. albiziae. Kuznetzov and Livshitz (1979) redescribed and figured C. taurus and C. bison from Russia, either disagreeing with or being unaware of Den Heyer's 1978 publication. Tseng (1980) reported A. taurus from Taiwan. Chaudhri (1980) described D. fixus from Pakistan. Den Heyer (1980c)  made Dactyloscirus and Armascirus the sole representatives. Gupta and Ghosh (1980) erected Indocunaxa, a monotypic genus with I. smileyi as the type species. Liang (1983) reported A. taurus from China. Shiba (1986) described A. hastus and A. multioculus. Michocka (1987) described D. rafalskii from Poland. A. mactator and A. pluri were described by Muhammad and Chaudhri (1991b).   Diagnosis. Gnathosoma. Pedipalps 5-segmented, end in a strong claw, and extend beyond the subcapitulum by at least the last segment. Apophysis between the genua and tibiotarsi, which tapers to a point, usually present; this apophysis shorter in males than in females. Basifemora complemented with a simple seta; telofemora with a spine-like seta. These two segments fused, although a line remains visible and they can thus be differentiated. Subcapitulum complemented with 6 pairs of setae (hg 1-4 and 2 pairs of adoral setae). It can be covered by integumental papillae which are either randomly distributed or form a polygonal, reticulated pattern.
Idiosoma, dorsal. Female dorsal idiosoma with at least one sclerotized plate that bears 2 pairs of setose sensillae (at and pt) and 2 pairs of simple setae (lps and mps). 0-4 other major plates and platelets may also be present. All plates, if present, covered by integumental papillae that form a reticulated pattern. Integument between the plates is striated. 7 pairs of setae, c 1-2 , d 1 -h 1 , present. Each seta, when not on a major plate or platelet, surrounded by a minute platelet that is only slightly larger than the setal socket. Cupule im present, usually laterad or in the proximity of e 1 . Dorsal idiosoma of males is similar except a single large plate complemented with c 1-2 , d 1 -e 1 present.
Idiosoma, ventral. Coxae reticulated in the same manner as the dorsal plates. Coxae I-II often fused; Coxae III-IV often fused. Setal formula of coxae I-IV in males 3-1-3-3 (including the paracoxal seta), in females 3-2-3-3 (including the paracoxal seta). Genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae (ps 1 ). 2 pairs of setae (ps 2 and h 2 ) associated with but do not occur on the anal plates. Cupule ih present in close proximity to h 2 . Integument between plates striated and bears 5-7 pairs of additional setae. The ventral idiosoma of males similar except the coxae are much more extensive. A sclerotized aedeagus is often visible in association with the genital plates. Legs comparatively long, at least ¾ the length, and often longer than the body. Famulus on tarsi I normally shaped. Tarsi are constricted apically, resulting in large tarsal lobes. Trichobothrium on leg tibia IV present. Ambulacral claws occur on either side of a 4-rayed empodium.

Key to adult female Armascirus (modified from Kalúz and Vrabec 2013)
Dactyloscirus bifidus Corpuz-Raros, 2008 is transferred to Armascirus as it posessess a spine-like seta on the pedipalpal basifemora.
Armascirus gojraensis and A. sabrii appear to be nymphs based on the leg setal counts given in the original descriptions. Having not seen the type material, however, they are retained within the key. Caution should be exercised if these species are reached.
Idiosoma, dorsal. Proterosoma bears a shield that is complemented with 2 pairs of setae (at and pt) and 2 pairs of setose sensillae (lps and mps). Dorsal hysterosoma may bear a shield; if a shield is present, it may bear up to 4 pairs of setae. Dorsal shields may be smooth or patterned with random dots, but never reticulated. Lateral platelets (as in Armascirus and Dactyloscirus) absent. Setae c 1 -h 1 , and c 2 present. Setae not born on the median plate may be born on small platelets that are barely larger than the setal socket. Cupule im present laterad and caudally of e 1 . Integument not bearing the proterosomal shield and median plate (if present) striated. These striations smooth or lobed but never papillated.
Idiosoma, ventral. Coxae I-II may be fused and coxae III-IV may be fused. Coxae II-IV setal formula 1-3-2. Genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae (ps 1 ). 1 pair of setae (h 2 ) associated with, but do not occur on, the anal plates. Cupule ih present in close proximity to h 2 . Integument between plates striated and bears up to 7 pairs of additional setae. Legs. Tarsi long and slender. Tarsi constricted distally but the tarsal lobes are small and not conspicuous as in Armascirus and Dactyloscirus. A trichobothrium on tibia IV present. Ambulacral claws on either side of a 4-rayed empodium present.

Key to adult female Cunaxa
Cunaxa bochkovi is not included in the key because the original description is in Cyrillic and the illustration does not contain enough detail or diagnostic characteristics. Den Heyer (pers. comm., Jan. 13, 2014) indicated that Cunaxa setirostris var. plurisetosa and C. setirostris var. diversa were described in "Mihelčič, F. 1958" but did not have the entire citation and had not seen the original description. The authors have also not been able to locate such a publication after extensive searching and so have not included the taxa here.
Idiosoma, ventral. Coxae I and II fused, as are coxae III and IV. 6 pairs of setae present between and posterior to the coxae. Genital plates each bear 4 setae; 2 pairs of genital papillae not visible underneath the plates. Integument between plates striated and bears 4 pairs of additional setae. Legs shorter than the body. Leg 4 longest. Famulus on tarsi I normally shaped and set in a deep depression. Tarsi slightly constricted apically, resulting in small tarsal lobes. Basifemora and telofemora of legs I and II partially fused. A trichobothrium on leg tibia IV absent. Ambulacral claws on either side of a 4-rayed empodium present.
Idiosoma, dorsal. Female dorsal idiosoma has at least one sclerotized plate that bears 2 pairs of setose sensillae (at and pt) and 2 pairs of simple setae (lps and mps). 0-4 other major plates and platelets present. All plates, if present, covered by integumental papillae that form a reticulated pattern. Integument between plates striated. 7 pairs of setae (c 1-2 , d 1 -h 1 ) present. Each seta, when not on a major plate or platelet, surrounded by a minute platelet only slightly larger than the setal socket. Cupule im present, usually laterad or in the proximity of e 1 . Dorsal idiosoma of males similar except a single large plate complemented with c 1-2 , d 1 -e 1 present.
Idiosoma, ventral. Coxae I and II often fused; coxae III and IV often fused. Setal formula for coxae I-IV 3-3-3-3 (including paracoxal seta). Genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae (ps 1 ). 2 pairs of setae (ps 2 and h 2 ) associated with, but do not occur on, anal plates. Cupule ih present in close proximity to h 2 . Integument between plates striated and bears 5-7 pairs of additional setae. Ventral idiosoma of males similar except the coxae much more extensive. A sclerotized aedeagus often visible in association with the genital plates. Legs comparatively short, generally not exceeding ¾ the length of the body. Famulus on tarsi I enlarged and ends in a tri-tipped prong. Tarsi constricted apically, resulting in large tarsal lobes. Trichobothrium on leg tibia IV present. Ambulacral claws occur on either side of a 4-rayed empodium.
Key to adult female Dactyloscirus (modified from Skvarla and Dowling 2012)  transferred Cunaxoides nicobarensis to Dactyloscirus as D. nicobarensis (Gupta & Ghosh, 1980). However, later in the same work he attributes the same holotype (No. 3146/17) and same description (viz. Gupta and Ghosh 1980:191) to Cunaxoides nicobarensis Gupta & Ghosh, 1980. The original description and illustration by Gupta and Ghosh clearly state the species in question has three pedipalpal segments, which precludes it from being assigned to Dactyloscirus. Smiley illustrated a Dactyloscirus with 5-segmented pedipalp "after Gupta and Ghosh 1980" when discussing D. nicobarensis, though it looks like nothing in the publication. Because of this Dactyloscirus nicobarensis (Gupta and Ghosh 1980) is declared nomen dubium. Pedipalpal telofemora with 1 apophysis (Figs 65a, b)  (6) ......... Apophysis adjoining pedipalpal genua and telofemora shorter than length of genu, blunt distally (Fig. 61a); median shield absent (Fig. 63d) ...8 -Apophysis adjoining pedipalpal genua and telofemora as long or longer than length of genu, blunt or pointed distally (Fig 61 c); median shield present or absent (Figs 63b, c) (Fig. 64b); apophysis adjacent to pedipalpal genua and tibiotarsi blunt distally (Fig. 61c) Historical review. Gupta and Ghosh (1980)  apophysis, which is about as long as the width of the telofemur 65b Pedipalp telofemur with one apophysis, which is shorter than the width of the telofemur 65c, d Pedipalp telofemur with two apophyses, one apical and one basal which is flattened and disc-shaped.

Diagnosis.
Gnathosoma. Pedipalps 5-segmented, extend beyond the subcapitulum by at least the last segment, and end in a strong claw; apophysis absent. Basifemora and telofemora complemented with simple setae; these two segments fused, although a line remains visible and they can thus be differentiated. Subcapitulum complemented with 6 pairs of setae (hg 1-4 and 2 pairs of adoral setae). Setae hg 3 and hg 4 both near the coxal bases of the pedipalps.
Idiosoma, dorsal. Female dorsal idiosoma has a sclerotized plate that bears 2 pairs of setose sensillae (at and pt) and 2 pairs of simple setae (lps and mps). Idiosomal shield covered by integumental papillae that form a reticulated pattern. Hysterosoma lacks a plate and bears 7 pairs of setae (c 1-2 , d 1 -h 1 ). Cupule im present, usually laterad or in the proximity of e 1 .

Key to adult female
Idiosoma, dorsal. Proterosoma bears a shield, complemented with 2 pairs of setose sensillae (at and pt) and 2 pairs of setae (lps and mps). Sensillae at and pt not as densely pilose as in Allocunaxa, Cunaxatricha, and Riscus. Proterosomal shield reticulated. Hysterosomal shield absent in females. Lateral platelets (as in Armascirus and Dactyloscirus) absent. Setae c 1 -h 1 , and c 2 present. Cupule im present laterad and caudally of e 1 . Integument not bearing the l shield striated. Striations papillated, not smooth or lobed as in Cunaxa. Idiosoma, ventral. Coxae I-II may be fused; coxae III-IV may be fused. Coxae II-IV setal formula 1-3-1. Genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. Anal plates bear 1 pair of setae (ps 1 ). 1 pair of setae (h 2 ) associated with, but do not occur on, the anal plates. Cupule ih present in close proximity to h 2 . Integument between plates striated and bears up to 7 pairs of additional setae. Legs. Tarsi long and slender, and constricted distally but tarsal lobes small and not conspicuous as in Armascirus and Dactyloscirus. A trichobothrium on tibia IV present. Ambulacral claws either side of a 4-rayed empodium present.
Idiosoma, dorsal. Dorsal idiosoma heavily sclerotized and the plates well-demarcated. A single dorsal shield present; it may range in size from terminating anteriorly to cupule im to being holodorsal. No papillated line or other marking indicates the separation of the proterosomal and hysterosomal shields. 2 pairs of setae and 2 pairs of setose sensillae present on the proterosomal. Setae c 1 -h 1 , c 2 , and f 2 and cupule im present dorsally. Dorsolateral plates ( such as present in Scutascirus) absent.
Idiosoma, ventral. Coxae I-II fused and coalesce medially to form a sternal shield which often has a prominent apex caudally. Sternal plate complemented with 5-7 pairs of setae. Coxae III-IV fused and may extend laterally and caudally past the genital plates. Genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. Anal plates bear two pairs of setae (ps 1 and ps 2 ). Seta h 2 located ventrally near the anal plates. Cupule ih present in close proximity to h 2 . Legs shorter than the idiosoma, never constricted apically so as to end in lobes. The apices of solenidia, especially on tarsi I, may be swollen. Trichobothrium on leg tibia IV present. Ambulacral claws on either side of a four-rayed empodium present.
Males similar, except up to three shields or plates may occur on the dorsal idiosoma (that is the proterosomal shield may not be fused to a hysterosomal plate and up to two hysterosomal plates may be present) and coxae I-IV may be fused into a holoventral shield.

Key to adult female Coleoscirus
Coleoscirus brevicornis (Berlese) has been excluded from the key as the original publication (Berlese 1904) and subsequent publication detailing the species (Thor and Willmann 1941) are in Italian and German and the accompanying illustrations provide too little detail. Den Heyer (1978b) is the last author to mention the species, but only indicates that it belongs to the genus Coleoscirus.
Coleoscirus carex, C. kifayati, and C. mardi have been excluded from the key as the authors did not provide enough information in the original descriptions to include them.
Coleoscirus zaherii is not included in the key as, despite the best efforts of the authors and the University of Arkansas Interlibrary Loan Department, the description could not be obtained.  telofemora. Pedipalp tibiotarsi long and S-shaped (as opposed to short and cylindrical as in Neoscirula). Subcapitulum with 4 pairs of setae (hg 1-4 ). 2 pairs of adoral setae present. Chelicera with seta usually present. Extensive reticulated pattern absent from the gnathosoma, though a row of single cells may be present caudally.
Idiosoma, dorsal. Plates lightly sclerotized and may not be well defined or demarcated. Proterosomal plate bears 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Extensive reticulated pattern absent, although a pair of rows of up to 6 cells may be present. Proterosomal plate may be covered with random dots or papillae. Hysterosomal plate absent. Setae c 1 -h 1 , and usually c 2 and f 2 present dorsally; h 2 present or absent. Cupules im present laterad and sometimes caudally of e 1 . Integument striated.
Idiosoma, ventral. Coxae usually restricted to the trochantral bases, though sometimes coxae I-II may nearly touch medially. Coxae I-II fused. Coxae III-IV fused. All coxae lightly sclerotized and may be ill-defined. Extensive reticulated pattern absent from the coxae, though a row of cells or reticulated pattern may be present near the edges. Coxae may be covered with random dots or papillae. Coxae I-IV usually have the simple setal formula 3-3-3-3 (N. parilis is the exception with 2-2-3-2). Genital plates each bear 3-4 setae; 2 pairs of genital papillae visible underneath the plates. 2 pairs of setae (ps 1-2 ) usually occur on the anal plates and 1 pair of setae (pa) occurs on the integument near the anal plates. However, at least one species (N. clavata) has 3 pairs of setae present on the anal plates and 0 pairs of setae on the integument. Cupules ih present ventrally near the anal plates. Legs. Tarsi never constricted apically so as to end in lobes. The apices of solenidia cylindrical, not swollen as in Coleoscirus and Scutascirus. Trichobothrium on leg tibia IV present. Ambulacral claws rippled and occur on either side of a 4-rayed empodium.
Neobonzia parvirostris (Berlese, 1910) is known only from the male and so is not included in the key. N. breviscuta (Luxton, 1982) is not included in the key as an insufficient number of characters are given in the original description.
Idiosoma, ventral. Coxae I-II separate or fused medially into a single sternal shield. Coxae III-IV contiguous on either side, restricted to area around trochantral bases. Dorsal cupules im present laterad to e 1 ; ventral cupules ih present near h 2 , anal plates. Legs shorter than body. Tarsi never constricted apically so as to end in lobes. Apices of solenidia cylindrical, not swollen as in Coleoscirus and Scutascirus. Trichobothrium on leg tibia IV present. Ambulacral claws smooth and occur on either side of a 4-rayed empodium.

Key to adult female Neoscirula.
Neoscirula hoffmannae Mejía-Recamier & Palacios-Vargas, 2007 is excluded from the following key as it is only known from the male. Pedipalp tibiotarsal claw a tooth present, giving bifid appearance (Fig. 82a) .. 8

Figures 75, 76.
Neobonzia key illustrations 75a Sensilla at and pt clavate, short, length less than the width of the proterosomal shield 75b Sensilla at and pt clavate, long, length greater than the width of the proterosomal shield 75c Sensilla at and pt normal, not clavate 76a Coxae I-II nearly touching medially 76b Coxae I-II widely separated medially.
pairs of adoral setae present. Chelicera with seta present (usually) or absent. Extensive reticulated pattern present on the gnathosoma. Idiosoma, dorsal. Plates lightly sclerotized and not be well defined or demarcated. The proterosomal plate bears 2 pairs of setae (lps and mps) and 2 pairs of setose sensillae (at and pt). Extensive reticulated pattern present. Hysterosomal plate absent. Setae c 1 -h 1 present; setae c 2 , f 2 , and h 2 present or absent. Cupules im present laterad and caudally of e 1 . Integument striated.
Idiosoma, ventral. Coxae restricted to the trochantral bases. Coxae I-II fused. Coxae III-IV fused. All coxae lightly sclerotized and may be ill-defined. Coxae with extensive reticulated pattern. Coxae I-IV usually have setal formula 3-3-3-3. Genital plates each bear 3-4 setae; 2 pairs of genital papillae visible underneath the plates. 2 pairs of setae (ps 1-2 ) occur on the anal plates and 1 pair of setae (pa) occurs on the integument near the anal plates. Cupules ih present ventrally near the anal plates. Legs. Basal leg podomeres with reticulated pattern present or absent. Tarsi never constricted apically so as to end in lobes. Apices of solenidia cylindrical, not swollen as in Coleoscirus and Scutascirus. Trichobothrium on leg tibia IV present. Ambulacral claws are rippled and occur on either side of a 4-rayed empodium.

Key to adult female Pseudobonzia (modified from Den Heyer and Castro 2008)
1 Pedipalp basifemora and telofemora with similar setae, either spine-like or simple (Fig. 88a, (Fig. 89c) Diagnosis. Gnathosoma. Pedipalps 5-segmented and reach beyond the subcapitulum by at most the distal half of the tibiotarsi. Basifemora and telofemora fused but retain a dark line. The tibiotarsi complemented with a tubercle and a dorsodistal solenidion. Pedipalps end in a stout claw. Chelicera with seta present or absent. Subcapitulum bears 6 pairs of setae: 2 pairs of adoral setae and 4 pairs of subcapitular setae (hg 1-4 ). Setae hg 4 often the longest.
Diagnosis. Gnathosoma. Pedipalps 5-segmented and reach beyond the subcapitulum by at most the distal half of the tibiotarsi. Basifemoral seta simple or spine-like. Telofemoral seta spine-like. Pedipalps end in a stout claw. Subcapitulum bears 6 pairs of setae: 2 pairs of adoral setae and 4 pairs of subcapitular setae (hg 1-4 ). Setae hg 1 long and bent.
Idiosoma, dorsal. Proterosoma covered in a shield which bears 4 pairs of setae: 2 pairs of simple setae (lps and mps) and 2 pairs of setose sensilla (at and pt). Dorsal hysterosoma median plate present, fused to proterosomal shield; 1 to 5 pairs of dorsolateral plates present. Plates and shields smooth or reticulated. Seven pairs of setae present on the dorsal hysterosoma (c 1 -f 1 , c 2 , h 2 ). Unsclerotized integument striated.
Idiosoma, ventral. Coxae I-II fused, coxae III-IV fused; coxae may coalesce medially for form a sternal shield. Each pair of coxae complemented with 3 pairs of setae. The genital plates each bear 4 setae; 2 pairs of genital papillae visible underneath the plates. 4-9 pairs of setae present on the integument between coxae II and the genital plates. Anal plates complemented with 2 pairs of setae (ps 1-2 ). Two pairs of setae (h 2 , pa) located on the integument near the anal plates. Cupule ih present in close proximity to h 2 . Legs shorter than idiosoma; they are never constricted apically so as to end in lobes. Trichobothrium on leg tibia IV present. Ambulacral claws on either side of a four-rayed empodium present.