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1 Acarologia A quarterly journal of acarology, since 1959 Publishing on all aspects of the Acari All information: acarologia@supagro.inra.fr Acarologia is proudly non-profit, with no page charges and free open access Please help us maintain this system by encouraging your institutes to subscribe to the print version of the journal and by sending us your high quality research on the Acari. Subscriptions: Year 2018 (Volume 58): Previous volumes ( ): 250 / year (4 issues) Acarologia, CBGP, CS 30016, MONTFERRIER-sur-LEZ Cedex, France The digitalization of Acarologia papers prior to 2000 was supported by Agropolis Fondation under the reference ID through the «Investissements d avenir» programme (Labex Agro: ANR-10-LABX ) Acarologia is under free license and distributed under the terms of the Creative Commons-BY-NC-ND which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original author and source are credited.

2 1 1 A REVIEW OF THE GENERA OF THE F AiVIIL Y MACROCHELIDAE VITZTHUM 1930 (ACARINA : MESOSTIGMATA) 1 BY G. w. KRANTZ 2 INTRODUCTION. The family Macrochelidae has been the subject of a number of descriptive and revisionary studies in recent years. BERLESE (1903, 1910, 1918) supplied much of the needed basic information for subsequent work on the group, describing the the majority of genera as well as many species. In his most extensive contribution on the family (1918), BERLESE established several supraspecific categories primarily on the strength of differences in sternal shield sculpture. The shape of the ventral shields and the extent of shield fusion also were considered to be characters of supraspecific significance. Unfortunately, BERLESE's work was of a nature which has made recognition of many 'of his species difficult or impossible without referring to type material. HuLL (1918, 1925) used dor~al chaetotaxy as a taxonomie character in addition to those proposed by BERLESE but, as pointed out by EvANS an_d BROWNING (1956), HuLL's descriptions and figures are inadequate and often inaccurate. Little more had been learned by the time that VITZTHUM (1930) had established the family Macrochelidae. OuDEMANS (1931) emphasized setal number on the ventrianal shield in his key to genera, establishing the genus M acrholaspis to contain those macrochelids with two pairs of preanal setae. Subsequent generic separations (EvANS and BROvVNING 1956; EvANS 1956) are based in part on OuDEMANs' scheme. EvANs (1956) discusses TRA.GÂDH's attempts to formulate a subfamilial classification in the Macrochelidae (1949, 1952). In 1952, TRA.GÂRDH described the genus Areolaspis, and proposed the following subfamilial divisions : 1. Approved as Technical Paper No. 1441, Oregon Agricultural Experiment Station, Cor. vallis, Oregon. 2. Associate Professer of Entomology, Oregon State University, Corvallis, Oregon. Acarologia, t. IV, fasc. 2, IO

3 «r. Metasternal shi~lds connected with the sternal shield through a narrow bridge.. Protoholaspinae Tragiirdh, Metasternal shields free Peritrematic shields not fused with the exopodal shields.... Macrochelinae Tragiirdh, Peri trema tic shields fused with the exopodal shields.. Areolaspinae subfam. nov. >> Protoholaspis subsequently was removed from the family Macrochelidae' (EvANS and BROWNING. 1956), and the separation of the Macrochelinae and Areolaspinae was found to be based on secondary characters (EVANS 1956) 0 Another separation was proposed by EvANS, in which those macrochelids.with looped peritremes and lateral accéssory sclerites on the epigynial shield were separated from those which did not possess these characters. The following key was given by Evans : «, r. Peritreme with a U-shaped loop and leaving stigma posteriori y ; genital shield with lateral sclerites Macrochelinae Tragii.rdh. - Peritreme straight and leaving stigma anteriorly; genital shield without lateral sclerites...: Parholaspinae subfam. nov.» KRANTZ (1g6o) redefined the Parholaspinae, raising it to family status. This resulted in the suppression of subfamilial categories in the Macrochelidae and in a clearer definition of its taxonomie limits. The Macrochelidae, as presently conceived, appears to be a natural group which is evolving toward a parasitic existence. Many species are free-living predators with large well-sclerotized dorsal and ventral shields, while others have been found to be phoretic or paraphagous on insects, or dependent in some other way on particular insect species for maintenance (PEREIRA and CASTRO 1945 ; FILIPPONI 1955 ; 1959; FILIPPONI and CERVONE 1957). Evolution toward parasitism in sorne genera apparently results in the progressive loss of sclerotization, a factor which supplies some rather attractive supraspecific characters but which also confuses natural affinities between species groups. Other genera appear to be following the opposite trend, exhibiting secondary expansion and fusion of ventral shields. Generic concepts in the Macrochelidae, therefore, have been somewhat tenuous, and have tended to stress specifie rather than supraspecific structures with the result that the limit? of several macrochelid genera are difficult to establish. EvANS (rg56) proposed a key to macrochelid genera in which severa! genera were provisionally relegated to synonymy under Macrocheles Latr., the type genus for the family. These groups were placed in synonymy in a review of British Macrochelinae (Macrochelidae) which was published the same year (EvANS and BROWNING rg56). To date, thirteen genera are recognized in the family Macrochelidae, including two genera recently described by KRANTZ (1961). The monotypic genus Trichocelaeno Berlese, of which_ only the male is lo:iown, is omitted from further consideration in the following discussion. It is the purpose of this review to bring together the available information on

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5 - I46- the genera of the family Macrochelidae, and to attempt to show relationships between generic groups on the basis of primary morphological characters. Family 'MACROCHELn=iAE Vitzthum. Macrochelidae Vitzthum, H. G. (1930). Zool. ]ahr. Syst. Bd Dorsal shield entire, variously ornamented and with twenty-eight to thirty pairs of setae. Sternal shield wit4 three pairs of setae ; metasternal shields free, absent, or fused into sternal-endopodal complex. Peritremes looped posteriorly and joining the stigma posteriorly. Epigynial shield with a pair of lateral accessory~ sclerites ; with a ventrianal or anal shield. Male with a separa te ventrianal or anal shield, or with a holoventral shield ; genital orifice presternal. Gnathosoma well-developed; with salivary styli and distinct corniculi; tectum tripartite or unipartite; palpi with five free segments, palpai claw three-tined. Chelicerae chelate-dentate and with one or two setal brushes at the base of the digitus mobilis; male with strong spermatophoral process. Legs I without ambulacra in either sex; legs II-IV with claws and pulvilli; legs II and IV of -male, and rarely legs II of female spurred. Laterodistal elements of pretarsi membranous, broad basally, and divided or acuminate distally. Free-living in soil or vegetable debris, or paraphages of insects. The laterodistal elements of the pretarsi (fig. 32) show definite variations at the generic level in the M~crochelidae, but their value as a generic character is tempered by the fact that the structure is difficult to observe in most specimens. Usually it appears as a more or less setate appendage which is strongly acuminate distally. When viewed laterally, however, it is seen to be a broad membrane, the distal edge of which may be entire or divided into a number of spinose projections. In the following generic diagnoses, the condition of the laterodistal pretarsal elements is included where known, but the structure has not been used in key.separation of the genera. REYIEW OF GENERA. 1. M acrochelès La treille. Macrocheles LATREILLE, P. Règne anim. (CuviER), Ed. 2(4) : 282. Macrholaspis ÜUDEMANS, A. C. (1931). Ent. Ber. 8(18o) : 272, new synonymy. Andrholaspis TURK, F. A. (1948). Froc. Zool. Soc. London ns : 103, new synonymy. Coprholaspis BERLESE, A. {1918). Redia 13 : 146. Nothrhçlaspis BE~LESE, A. (1918). Redia 13 : 169 (in part). Dissoloncha FALCONER, W. (1923). Naturalist, London : 151. Monoplites Huu, J. 'E. {1925). Ann. Mag. Nat. Hist. 15, ser. 9 : 215. '. J, - ::

6 . ~ With twenty-eight to thirty pairs of dorsal shield setae ; dorsal shield without distinct anterior protuberance, serrated or smooth laterally. Sternal shield with three pairs of setae ; metasternal shields free. Genital shield with a pair of setae and accessory sclerites. Ventrianal shield with two or three pairs of ventrianal setae. Peritrematal shield not fused to exopodal shields. Male with sterr:itigeniti-ventral shield and separate ventrianal or with holoventral shield ; genital orifice presternal. Hypostome with well-developed corniculi and salivary styli. Palps with five free segments ; palpai claw tridentate. Tectum trifid (except in lvi. superbus Hull). Chelicera with a strong brush of setae ventrolaterally; movable digit of male with a large spermatophoral process. Legs I without ambulacra ; legs II and often legs IV spurred in the male ; pretarsal elements divided distally. TYPE SPECIES. - Acarus marginatus Hermann 1804 (= Acarus m;uscae domesticae Scopoli 1772).. iviacrocheles sup er b~ts Hull may represent a supraspecific taxon on the basis of its tectal structure, which is unipartite rather than tripartite as in other species of Macrocheles (fig. 12). The structure of the tectum and the arrangement of hypostomal setae in lvi. superbus, as well as the general facies of the gnathosoma, point to a possible relationship with the genus Geholaspis Berlese. Until further stud y of characters can be made, however, M. super b~ts is retained in lvi acrocheles. ivi acrocheles is by far the largest genus in the family Macrocheliçlae, with weil over roo species having been described. Many of these forms are free-living in decaying organic material or sail, where they prey on mites and other small arthropods. Others are found in phoretic association with insects. The genus is world-wide in distribution. In their diagnosis of the genus iviacrocheles, Evans and Browning (1956) list the following genera as synonyms of lvi acrocheles Coprholaspis Berlese Northrholaspis Berlese Dissoloncha Falconer Monoplites Hull C oprholaspis. This group was based primarily on the type of sternal shield sculpture exhibited by those species which were included in it. BERLESE (r9r8) also emphasized the sculpture of the ventrianal and epigynial shields in his diagnosis of the subgenus. He named ten phalanges in Coprholaspis, each possessing a different combinaiton of sternal shield characters. SELLNICK (1940) synonymized Coprholaspis with ivi acrocheles s. str., but the former name persisted in the literature until sorne years lat~r. The great variation in dorsal and ventral shield sculpture bath within.

7 and betwee:ri genera in the family Macrochelidae makes the use of such characters extrerriely difficult above species level, particularly when supporting characters are not evident. N othrholaspis. WüMERSLEY (1942) gives the following diagnosis for Nothrholaspis : "Sternum variously and densely rugose, rugosities not reduced, transverse median line sqmetimes obsolete; porose areas sometimes present. Body and legs with scaly secretions. Dorsal setae penicillate. " WüMERSLEY's diagnosis, derived from BERLESE (rgr8), is based on characters which have since proven to be unreliable as generic indicators. Dissoloncha. Dissoloncha wac; established as a subgenus by FALCONER (1923) '.vho based his diagnosis on the following characters : "Apex of the epistome forked on a forked lamellar base. Body elongate pentagonal or oval. Dorsal scutum in two parts of diverse texture. The femora of the second and fourth pairs of legs in the male spurred... " Falconer named M acrocheles super bus Hull as the type species of Dissolonchti, and included M acrocheles plumiventris Hull rgzs ( Glyptholaspis fimicola Berlese r88z) as a second species. FALCONER's description of tectal structure in Dissoloncha apparently was based on inadequate observations, inasmuch as M. superbus has a unipartite tectum which is forked distally, rather than the tripartite type common.to other species of M acrocheles. Monoplites. HuLL (1925) named Monoplites as a subgenus of Macrocheles, describing the males as having spurs on the second pair of legs only, and the female as having a more or less cordiform ventrianal shield which is well separated from the. epigynial shield. These characters are considered by the writer to be secondary, and not of subgeneric or generic importance. The following discussion is relative to proposais of new synonymies in Macro- cheles. Citations of these synonymies are listed above. M acrholaspis. In their review of the subfarnily Macrochelinae, EvANS and BROWNING (1956) give the following diagnosis for the genus M acrholaspis : " Dorsal shield strongly attenuated posteiiorly and bearing twenty-nine pairsl of plumose setae 1. Lateral margins of the shield markedly serrated. Sternal shield with three pairs of setae ; metasternals free. Genital shield with acces BALOGH (r958) has since described Nlucrholaspis evansi, which has smooth D 1, D 1, D 1, and M 1 setae.

8 -149-.sory sclerites and a pair of genital setae. Ventrianal sh'ield with two pairs of pre.anal setae. Other characters as in lvi acrocheles. 1 An additional seta may be present in sorne species (cf. M acrholaspis dentattts :sp. n.. ) 1, -.. The genus M acrholaspis has been separated from species of 1\II acrocheles primarily by the presence, in the latter group, of three pairs of pre-anal setae and usually only twenty-eight. pairs of dorsal shield setae. Recent examina tian of undescribed material tentative! y assigned to Jl![ acrocheles on the basis of ventrianal shleld setation were found to possess twenty-nine pairs. of dorsal shield setae and marginal dorsal shield serrations, as found in described species of 11 acrholaspis. Further comparisons with species of bath genera revealed that most of the dia _gnostic features for either group are not constant, and that characteristics assigned to one group may be found to occur in the ether. It was felt, therefore, that a re-evaluation of M acrholaspis should be attempted so as to establish the true relationship of this assemblage to the genus 11 acrocheles. A study of the ventrianal shields of 111_~crholaspis and lvi acrholaspis-like species with two and with three pairs of preanal setae on the shield shows. that this charader is dependent only on the relative anterior development of the ventrianal shield. In those species where anterior clevelopment is strongly retarded (or reduced), the first pair of preanals are free in the integument (fig. 3, PR 1 ), while in those species where the ventrianal shield is more completely expanded, preanals I are included on the shield (fig. 2). The question arase, then, as to the signifi Cance of this difference as a generic character. Examina tian of ventrianal shield development of various speci~s of 111 acro.cheles s. str. revealed wide intrageneric variations in size and shape, and in relative position of insertion or preanals I. For example, the ventrianal shield of ~Macro cheles mttscaedomesticae (Scopoli) is a well-developed structure, extending anteriorly so that it abuts the epigynial shield along a more or less transverse juncture (fig. r). Three pairs of pre anal setae are inserted on the shield. 111 acrocheles carinatus (C. L. Koch) ~Iso has the typical three pairs of preanal setae on the ventrianal shielcl, but the shield itself is so abbreviated anteriorly that preanals I are marginally inserted. The condition of the shield allows it to attain a more or less roundecl aspect anteriorly, the shield being weil separated from the posterior margin of the epigynial shield. Similar reduction is seen in ether members of the genus (NI. tardtts (C. L. Koch) and M. cattcasictts Bregetova and Koroleva, for example), with placement of. preanals I being marginal. Identicat' development occurs in the M acrholaspis-like species referred to previously. Ventrianal shield reduction in either group commonly is correlatecl with the presence of a series of small sclerotized platelets lying in the integument between the epigynial and ventrianal shields (figs. 2, 3). The wide variation in ventrianal shield development in the genus Macrocheles (Bregetova and Koroleva rg6o), and the fact that the shield structure in species of ]1.1 acrocheles resembles that of species which more properly fit the diagnosis of M acrholaspis leads the writer to believe that ventrianal shielcl

9 ,, ----;--~ ~------IOOU------~ - --._- / ~ - ---=-- 8 ~ U ,---, /_,_,-~, /.,--- ' 1 \ , -\ / 1 ',!~\ '-' 9 -=.:...!_-, ~ U T \ / 1 \ '\ 10 <é u ----~ Fig. 5 Holocelaeno 1'otuncla Berl. Anterior portion of dorsal shield of female. - Fig. 6. Holocelaeno sp. (Argentina) Anterior portion of dorsal shield of female. - Fig. 7 Ettholocelae-no bursiformis Berl. Venter of female. - Fig. 8. Ettholocelaeno btwsiformis Berl. Chelicera of female.- Fig. g. Holocelaeno rotttncla Berl. Ventrianal shield of female. -Fig. 10. Holocelaeno sp. (Argentina) Ventrianal shield of female. -Fig. 11. Holocelaeno sp. (Costa Rica) Ventria:iJ.al shield of female.

10 - rsrvariation of this magnitude should not in it5elf be considered as having diagnostic value at the generic level in the family Macrochelidae. Examination of undescribed 1VIacrholaspis-like species also reveals that the dorsal shield usually is attenuated posteriorly, and that lateral serrations may or may not be evident. Lateral serrations on the dorsal shield occur in several cases in the genus MacrocJ:teles (M. superb~ts Hull, M. penicilliger Berl. and others), and posterior attenuation of the dorsal shield in this group is not unknown (M. ivanovi Breg. and Korol.). The importance of these characters, therefore, appears to be less thau previously supposed. Plumosity of dorsal setae also is a feature which is shared by members of both genera. lvi acrholaspis evansi Balogh has several pairs of smooth dorsal setae and lacks distinct marginal serration, differing fromm acrocheles s. str. orîly in preanal setal insertions. '. Comparative studies involving the following additional characters showed no definitive differences when applied to Macrocheles, Macrholaspis, and Machrolaspis-like species : r. Tectostracum present, commonly obscuring dorsal setal condition; dorsum appears rugose., 2. Setae D 1 strongly plumose over most of their length. 3 Dorsal setae in well-defined raised insertions. 4 With stout articuladed spines on genu and tibia I. 5 Laterodistal elements of pretarsi divided distally. It was found that those species which resemble J1,1acrholaspis, as well as those which agree with the diagnosis of the genus cited earlier in this discussion, have an extra pair of dorsal (D) setae inserted between set~e D 7 ar:id D 8 An additional D seta occurs, in sorne cases, in the same general area. In one undescribed species, the additional seta is paired, resulting in two extra pairs of D setae. Three or four platelets may be found between the epigynial and ventrianal shields of most of the species possessing thes_e characteristics, and all have an attenuated dorsal shield. It also has been observed that various species which more closely resemble J11acrocheles may possess extra D setae (J1,1. neomontamts Ryke and Meyer), and ventral platelets are present in others (M. carinatus (C. L. Koch), M. s~tbrnotus Falconer). Plumose D 1 setae occur in several Macrocheles species (EvANS and BROWNING 1956; BREGETOVA and KüLOLEVA rg6o). The strong similarities between J11acrocheles and J11acrholaspis, and the rather unsubstantial bases on which definitive separation may be achieved point to an intra-rather thau an intergeneric relationship between the two assemblages. It is possible that J11acrholaspis represents a subgeneric group, but the writer has been unable to find a reliable character on which subgeneric separation may be based. It is herein proposed that the genus M acrholaspis be placed in synonymy with L11 acrocheles.

11 Andrholaspis. TURK (1_948) presents the following diagnosis for Andrholaspis " Witn the characters of the genus Coprholaspis Berlese, but with the first and se~ond pairs of legs of the female with one or more segments markedly crassate and spurred, and with the coxae of the third and fourth pairs of legs with one or more spurs. Epistôme an apically bifurcate, simple lamellar base with a single, pencillate central piece with two simple acute bifurcations apically. Dorsum with a few faint reticulations and sorne long harply pointed spine-like hairs. " Andrholaspis is separated from Macrocheles only by the presence, in the former, of spurs or spines on legs I and II. Such a variation in itself is felt by the writer to be more of specifie than generic value. It is proposed that Andrholaspis be synonymized with Macrocheles. Several species in the genus Macrocheles were originally described as belonging to the laelaptid ge1ms Holostaspis, a hypoaspidine group associated with insects. Several of the resulting synonymies are listed by EvANS and BROWNING (rgs6). 2. Glyptholaspis Filipponi and Pegazzano. Glyptholaspis FILIPPONI, A. and F. PEGAZZANO (rg6o). Redia 45 : 136. Macrocheles (Macrocheles) BERLESE, A. (rgr8). Redia 13: 172 (non LATREILLE r8zg). Nothrholaspis BERLESE, A. (rgr8), Redia 13 : r6g (in part). Macrocheles Latreille, P. (r8zg). EVANS, G. 0. and E. BROWNING (1956). Bul. Brit. M'us. (Nat. Hist.) Zool. 4: ro (in part). With the characters of M acrocheles except for the following : Dorsal shield strongly crenulate-reticulate; 'dehtate laterally and posteriorly. Sternal shield with raised polygonal design and extending laterally and posteriorly arou~1d the epigynial shield to the level of the posterior angles of coxae III (fig. 4) ; posferiot cornets abutting more or less rounded metasternal shields. Epigynial and ventrianal shields crenula.te-reticulate ; ventrianàl shield transversely subelliptical. Dorsal seta of fixed cheliceral digit denticulate. TYPE SPECIES. - Gamaszts tard us Berlese r882., FrLIPPONI and PEGAZZANO (rg6o) erected the genus Glyptholaspis to accomodate four species <;>f macrochelids originally ascribed to BERLESE's subgenus Macrocheles (rgr8), plus an additional species de'scribed at the time of generic designation. FrLIPPONI and PEGAZZANO demonstrated quite conclusively that those species assigned to Glyptholaspis are intimately related and thus merit a separate taxon. Inasmuch as their separation from M acrocheles was based in large part on variations in dorsal and sternal shield sculpture patterns, the elevation of Glyptholaspis

12 -153- to generic rather than to subgeneric rank is debatable. FILIPPONI (personal correspondence) has reported that additional characters being used in current unpublished research further indicate thé valid.ty of generic rank for Glyptholaspis. The lateral extensions of the sternal shield (fig. 4), combined with the crenulate reticulation of the dorsal shield, are considered to be definitive characters in this rev1ew. The extensive ventral sclerotization in Glyptholaspis may be indicative of an evolutionary trend toward consolidation and fusion of ventral shields in this group. A similar situation is seen in the genus Synaphaspis, which will be discussed in a later section. 3 H olocelaeno Berlese. Holocelaeno BERLESE, A. (r9;ro). Redia 6 : 249. Neoholaspis Turk, F. A. (1948). Froc. Zool. Soc. London n8 99, new synonymy. The genus H olocelaeno may be characterized as follows : Dorsal shield rounded posteriorly and bearing twenty eight pairs of unadornecl setae (setae D 8 may be plumbse) ; shielcl with or without an anterior projection on which setae D 1 are inserted (fig. 5, 6). Dorsocentral setae (D 4 -D 7 ) typically shorter than 6ther dorsal setae. Sternal shield with three pairs of setae. With a single pair of epigynial se1:ae and with accessory sclerites ;. metasternal shields free. Peritrematal shields not frised to exopodal shields. Ventrianal shield with two, th:tee or four pairs of preanal setae. Cheliceral brush filamentous and extencling at least to the distal end of the digitus mobilis. Laterodistal elements of pretarsi broad, distally divided or acuminate. Other characters as in M acrocheles Latr. TYPE SPECIES. - Holocelaeno mitis Berlese EvANS (1956) indicates in his review of the M acrochelidae that H olocelaeno may be synonymous with 1\.facrocheles. He does not, however, list Holocelaeno as a provisional synonym of Macrocheles. TuRK (1948) erected the genus N eoholaspis on the basis of two species of macrochelicl mites, one of which was describecl as new. Examination of the cheliceral brush of the type species, N. corpophihts, leaves little doubt that this form is a species of H olocelaeno. The second species assigne cl to N eoholaspis by TuRK was N. cooremani (Cooreman), which originally was described as, 111 acrocheles sp. " by CooREMAN ( 1943). This species agrees with the diagnosis of M acrocheles rather than with H olocelaeno on the basis of the generic attributes cited above. The genus N eoholaspis is herein synonymized with H olocelaeno and N eoholaspis cooremani (Cooreman) is assigned to the genus 111 acrocheles. Examination of several species of Holocelaeno reveals at least one consistent character which points to the generic validity of this group the presence of a long

13 14~~\i~. Tc li } l~!t ~' \ 12 "" - :'"'> ~~)' /~1., \f( 1 ir\, 'r 1 rl(j ;;~1 ~,~~:~~djf/ «---200).J --~ oou----~ ~--- IOOU----" Fig. 12. Macrocheles dinzidiatus Berl. Tectum of female. - Fig. 13. /j.feolaspis sp. (Georgia, U.S.A.) Lateral view of peritœmatal exopodal fusion in female. -Fig. 14. Areolaspis sp. (Kentucky, U.S.A.) Dorsum of female. - Fig. 15. Geholaspis (Longicheles) mandibularis (BerL) Tectum of female. - Fig. 16. Holostaspella sp. (India) Dorsum of female. - Fig. 17. Gelwlaspis (Longicheles) mandibularis (Berl.) Chelicera of female. - Fig. 18. Geholaspis (Geholaspis) longispinosus (Kramer) Ventral aspect d chelicera of female. - Fig. 19. Holostaspella sp. (India) Chelicera of female. {..

14 filamentous setal brush (fig. 8) at the base of the digitus mobilis. It is of interest to note that preanal set al variation similar ta that found in lvi acrocheles also exists in Holocelaeno (figs. g, 10, II). Holocelaeno species are almost all paraphagous on insects, with few free-living forms having been described. 4 Euholocelaeno Berlese. Euholocelaeno BERLESE, A. (rg18). Redia 13 : 176. With the characters of Holocelaeno, except that the peritrematal and,exopodal shields are fused and extend posteriorly around coxae IV, and the membranous laterodistal elements of the pretarsi are not divided distally. TYPE SPECIES. - Holocelaeno bursiformis Berlese The generic status of Euholocelaeno is dependent on the value of the peritrematal exopodal fusion as a generic character (fig. 7). For purposes of this review, the writer considers Euholocelaeno a valid genus.. 5 Areolaspis Tragârdh. Areolaspis TRA.GÂRDH, I. (1952). Ark. Zool. Bd. 4, no. 3 : 6o. TRAGÂRDH erected the genus Areolaspis ta contain a new species which exhibited the following characters : Vertical setae inserted on prow-like anterior projection of dorsal shield; with a row of aval, deeply punctured depressed areas along the lateral margins of the shield. Peritrematal and exopodal shields fused. With sclerotized ridges on legs II. Membranous laterodistal pretarsal elements divided distally. TYPE SPECIES. - Areolaspis bifoliàtus Tragârdh TRA.GÂRDH also erected a new subfamily for the genus, the Areolaspinae, which subsequently was suppressed (see Introduction). Areolaspis is primarily a subtropical or tropical group of insect paraphages which obvious affinities with Holostaspella Berl., as will be described below. Emphasis on the dorsal shield projectio~ character as a primary generic separation (fig. 14) has resulted in the association of Areolaspis with the E-uholocelaeno-Holocelaeno complex. Such an association is unrealistic on the basis of ~heliceral (fig. 8) and setal characters, and in light of the fact that anterior shield projections do not occur in sorne Holocelaeno species (fig. 6). Lateral depressed areas on the dorsal shield occur ta a grea ter or lesser extent in all species of Areolaspis examined., Similar depressions are evident in H olostaspella, which is described belo w. ' \

15 Holostaspella Berlese. Holostaspella BERLESE, A. (1903). Redia 1 : 241. Prholaspina BERLESE, A, (1918). Redia 13 : 174, new synonymy. Dorsal shield witl). twenty-eight pairs of setae: many of which surpass the bases of the setae immediately posterior to them. Vertical setae situated on an out-' growth of the dorsal shield; with well-defmed depressions on the lateral mârgins of the shield. Sternal shield with three pairs of setae ; metasternal shields free. Ventrianal shield with three or four pairs of preanal. setae. Gnathosoma normal. Leg II 1 in the femaje armed with stout spurs. Laterodistal element 0f pretarsï divided distally. Other characters as in 111 acrocheles Latr. TYPE SPECIES. - Holostaspis (Holostàspella) sculpta Berlese BERLESE (1918) described Prholaspina as a subgenus of Holostaspella. The group is considered by EvANS (1956) to be a valid genus, differing from Holostaspella in the number of preanal setae on the ventrianal shield. Prholaspina has only three pairs rather than four as ïs found in Holostaspella. The diagnosis of Holostaspella offered by RYIŒ and MEYER (1958) actually includes Prholaspina, defining Holostaspella as having three or four pairs of preanal sètae. This variation in setal number is s~milar to that encountered in separation of M acrocheles and. Macrholaspis, where the number of preanal s~tae on the shield depends on slight variation in shape or size of the shield. Inasmuchas Holostaspella and Prholaspina appear to agree closely on features cited in the diagnosis given for the former group, it is felt that Prholaspina should not be retained as a valid genus. It is, therefore, herein reduced to synonymy under Holostaspella. As mentioned earlier, Holostaspella shows strong affinities to Areolaspis. The presence of armature on legs II of the female, in conjunction with the lateral depressions and anterior extension of the dorsal shield (fig. r6), indicate such affinities. Holostaspella and Areolaspis maintain separate generic status primarily because of the presence or lack of fusion of the peritrematal and exopodal shields (fig. 13), as well as the apparently consistent differences in dorsal shield setallength (figs. 14, r6).. Holostaspella is found in temperate and tropical regions and may be free-living or paraphagous on insects. It appears probable that Areolaspis is an offshoot of this group. I. In their diagnosis of Holostaspella, EVANS and BROWNING (:igs6) mention that leg I of the female, rather tf1an leg n, is armed 'with stout spurs. This is undoubtedly a printer's error.

16 Geholaspis Berlese. Geholaspis BERLESE, A. (rgr8). Redia 13 : 145. The -genus Geholaspis has been reviewed by VALLE (1953) and by EvANS and BROWNING (1956). The following diagnosis is that of EVANS and BROIYNING : " Dorsal shield with twenty-eight pairs of setae and twenty-two pairs of, pores ". Sternal shield with three pairs of setae; metasternal shields free or fused with the endopodal. shields. Ventrianal shield in the female with five pairs of preanal setae. Chelicerae normal length with a few teeth or conspicuously elongate and multidentate. Tectum with median process only. Other characters as in Macro- cheles La tr. " TYPE SPECIES. - Gamas~ts longispinosus Kramer r876. Geholaspis is believed to be most primitive genus in the Màcrochelidae, and may be a more or less intermediate form between the more a9-vanced macrochelid genera and other closely related families (KRANTZ rg6o). VALLE (I9S3) has divided the genus into three subgenera, using the comparative lengths of setae Mv cheliceral teeth, and fusion of metasternal and endopodal shields as differentiai features. BALOGH (r958) described two species of Geholaspis (Longicheles) which do not possess the dorsal setal character attributed to the subgenus by Valle. The cheliceral,character is retained in this review as the primary subgeneric feature of Longiclieles, although further study of the genus may reveal that M 1 setallength provides a more reliable basis for subgeneric definition. The following key is adapted from V ALLE. r. Chelicerae of normal dimensions (fig. r8) ;. digitus mo'bilis with less th an five tee th. 2 - Chelicerae long and narrow (fig. 17) ; digitus mobilis with more than five teeth... Longicheles 2. lvietasternal shields fused with endopoclal shields Oyrtocheles - Mestasternal shields free..... Geholaspis 8. Neopodocinum Oudemans. Neopodocinmn ÜUDEMANS, A. C. (rgoz). Tijds. v. Ent. 45 :. 24. Neopodocinwm originally vvas described as a member of the Laelaptinae, and was distinguished by its lack of ambulacra on legs I (OuDEMANS rgoz). Its subsequent inclusion in the Macrochelidae as a separate genus was justified on the basis of the presence of an anal rather than a ventrianal shield as in other macrochelid ger~era (fig. zr). The genus is dêfined.as follows : Dorsal shield broad, covering most of the dorsum ; with twenty-eight pairof setae, most of which are smooth, setae D 1 and l\1 1 commonly ornamented. Ster-

17 T }~ ~ l '...,_.) H Ul O:J 20 Fig. 20. Neopodocinum jaspersi (Oud.) Dorsum àf female. - Fig. 21. Neopodocinum jaspersi (Oud.) Venter of female.

18 ) nal shield transversely narrowed medially, with three pairs of setae; metastemal shields free. With simple anal shield. Tectum unipartite (fig. 24). Lateral elements of pretarsi short, broad and distally acuminate. Other characters as in M acrocheles Latr. TYPE SPECIES. - Neopodocinum jaspersi Oudemans The dorsal shield of N eopodocinum is of interest in that an extra dorsal se!a may be present (fig. 20). Recently, specimens of Neopodicinum were e~amined which have lateral ridges that resemble those of Holostaspella and Areolaspis. The generic importance of the first character cited is questionable, but it may be that the dorsal shield configuration points to an intergeneric relationship between the above mentioned genera. The genus Lordocheles Krantz, to be referred to below, shows the same type of configuration as well as other similarities which will be described. Members of the genus N eopodocinum are primarily tropical or subtropical, and are found in association with scarabeid, trogid, or geotrupid beetles. The mites usually are confined to the region of the mouthparts, and may congregate in great numbers between the man di bles and under the labrum of the beetle " hosts ". g. Lordocheles Krantz. Lordocheles KRANTZ, G. W. (rg6r). Instit. Parcs Nat. Congo, Fasc. 24 (r) : 3 The following diagnosis of the genus Lordocheles is from the original description by KRANTZ (rg6r) : \ " Dorsal shield strongly sculptured, scalloped laterally ; with twenty-eight pairs of dorsal setae and a single unpaired seta D 7 ; medially with a longitudinal depression bordered posteriori y by a series of large " pores ". Stemal shield reduced ; endopodal shields free in the integument. Metastemal shields absent. Anal shield small, without pre-anal setae. Chelicerae strongly bent dorsad, variously toothed. Laterodistal elements of pretarsi considerably longer than claws, undivided distally. " TYPE SPECIES. - Lordocheles desaegeri Krantz rg6r. Like N eopodocimtm, Lordocheles has an anal rather than a ventrianal shield. Reduction in the stemal and epigynial shields (fig. 23), is another feature shared by both genera. The presence of lateral dorsal shield depressions in Lordocheles (fig. 22) may, as mentioned earlier, indicate a common line of intrafamilial descent for Lordocheles, Holostaspella, Areolaspis and Neopodocinum. Lordocheles is based upon two species of mites taken in the Congo.!lcarologia, t. IV,.fasc. 2, x962. II ' '

19 500U. 1,r. /. ' 1 1 ~ ':. Ill/// ", 1 ~ ~" H 0\ 0 -<"... ~... 1 \. Fig. 22. Loràocheles rykei Krantz Dorsum of female. Fig. 23. Lordocheles desaegeri Krantz Venter of female (From Krantz rg6r).

20 Trigonholaspis Vitzthum. Trigonholaspis VITZTHUM, H. G. {1930). Zool. jahr. Syst. Bd. 59 : 300. The genus Trigonholaspis, as originally described, included macrochelid mites with the following characters : Dorsal shield smooth or reticulate ; dorsal setae short, either simple, pilose, spinose, or cylindrical; lateral margins of shield smooth or serrulate. Sternal shield excavated antermedially, with three pairs of setae ; metasternal shields free. Ventri'anal shield with three pairs of preanal or with eight pairs of circumanal setae. Tectum tripartite. TYPE SPECIES. - Trigonholaspis salti Vitzthum Although the writer has not had the opportunity to examine specimens of Trigonholaspis, it is apparent from the descriptions of species (VITZTHUM 1930) that the type species differs in several important respects from the remaining three forms. T. salti has a posteriorly truncate abbreviated dorsal shield which is unornamented and adorned with short or minute setae -(fig. 25)- The lateral and posterior margins of the shield are without serrations, and the ventral setae are extremely minute. The three' remaining species in the genus (T. columbiana, T. trigonarum, and. T. amaltheae) differ from T. salti in the following respects : r. The dorsal shield is not truncate posteriorly, and many of the dorsal setae are cylindrical and blunt. 2. The lateral and posterior margins of the dorsal shield are strongly serrulate, and the surface of the shield is distinctly sculptured. 3 The ventral setae are of normal dimensions. The primary generic character of Trigonholasp-is as conceived by VITZTHUi\1 is the excavated anteromedian aspect of the sternal shield (fig. z6). Inasmuch as similar concavities may be found in other macrochelid genera (Macrocheles, N eopododnmn), the value of this character in generiè diagnoses is doubtful. The dorsal setation of the. three species of Trigonholaspis other than the type species, and the marginal condition of the dorsal shield (fig. z8), provide a basis for generic separation of these forms from T. salti, It is proposed that the genus Trigonholaspis be reduced to a monotypic genus, and that the species columbiana, trigonar1:tm, and amaltheae be placed 'in a separa te genus. The genus Trigonholaspis is re-defined as follows Dorsal shield unornamented, truncate and strongly abbreviated posteriori y ; lateral margins smooth. Dorsal setae short or minute. Sternal, epigyni<ll and

21 r6z r 1 1 f 27 ' Fig. 24. Neopodocinum J aspersi (Oud.) Tectum of female. -Fig. 25. Trigonholaspis salti Vitzthum Dorsum of female (From Vitzthum 1930). - Fig. 26. Trigonholaspis salti Vitzthum Sternal shield of female (From Vitzthum 1930). - Fig. 27. Grafia columbiana (Vitzthum) Ventrianal shield of female (From Vitzthum 1930). - Fig. 28. Grafia columbiana (Vitzthum).Dorsal shield of female (From Vitzthum 1930).. \

22 -!63- ventrianal setae minute. Sternal shield excavated anteromedially ; metasternal shields free. Tectum tripartite. Other. characters as in M acrocheles. rr. Grafia n. gen. Trigonholaspis (in part) VITZTHUM, H. G. (1930). Zool. jahr. Syst. Bd. 59 : 300. Dorsal shield reticulate, rounded posteriorly, and with distinct serrulations laterally and posteriorly. With several pairs of blunt cylindrical dorsal shield setae, sorne of which are strongly abbreviated. Setae D 1 pilose. Sternal, epigynial and ventrianal setae of normal dimensions ; ventrianal shield with three pairs of preanal or with èight pairs pf circumanal setae. Sternal shield excavated ànteromedially, with three pairs of setae ; m~tasternal shields free. Other characters as in Macrocheles (Latr.). TYPE SPECIES. - Grafia trigonarum (Vitzthum) ' ' The inclusion of eight pairs of circumanal setae on the expanded ventrianal shield of G. columbiana (fig. 27) l~ads to further speculation on the comparative value of dorsal shield characters and ventrianal characters as generic indicators. While it is possible that such extreme expansion of the anal or ventrianal shield, and the many. setal insertions afforded by such development, may offer a useful means of generic classification and separation, it is questionable whether differences such as these should be used vvithout supporting factors. Geholaspis is readily separable from other genera on the basis of ventrianal setal number, but the unipartite tectal condition and the development of the corniculi are considered by the writer to be more definitive characters for separation of the genus. Lordochel'es and N eopodocinum are distinctive because of the reduction, or limited development, of the shield surrounding the anal opening. The two groups, however, are, aqually separable on cheliceral and tectal characters. _ The strong dorsal shield similàrities between G. columbiana and the remaining two species of the genus are felt to offer a sound generic basis for the group, ventrianal shield differences notwithstanding. 12. Synaphaspis Krantz. Synaphaspis KRANTZ, G. W. (rg6r). Instit. Parcs Nat. Congo, 24 (r) : ro. The following diagnosis oj the fenus Synaphapis is from the original description by KRANTZ (rg6r) : " Dorsal shield strongly pitted (fig. 29) ; with twenty-eight pairs of setae. :Metasternal shields fused with sternal, podal and ventrianal shields; discrete metapodal shields absent. Peritrematal shields present, not fused to parapodals

23 29 \ \ <'----loo u Fig. 29. Synaphaspis congoensis Krantz Dorsum of female (From Krantz 1961). - Fig. 30. Synafthaspis congoensis Krantz Chelicera of female (From Krantz 1961).- Fig. 31. Synaphaspis congoensis Krantz Tectum of female (From Krantz 1961). - Fig. 32. Synaphàspis congoensis Krantz Tarsus II of female showing lateral pretarsal elements (From Krantz 1961).

24 , posterior to coxae II. Legs without ridges or spurs ; laterodistal elements of pretarsi divided distally. Tectum (fig. 31) trifid; lateral elements free. " TYPE SPECIES. - Synaphaspis congoensis Krantz Synaphapis differs from all other macrochelid genera in having fused sternal metasternal and ventral shields (fig. 33). Its affinites are somewhat clouded, due to its resemblance to bath ad vanced ànd unspecialized forms. Synaphaspis exhibits most of the generalized macrochelid characters founcl. in J.V! acrocheles, and possesses the type of shield consolidation which is less évident in the genus Glyptholaspis Filipponi and Pegazzano. Assuming the genus M acrocheles to be more or less typical of the family, the consolidation of ventral sclerites in Synaphaspis might be considered either a primitive or an advanced condition. If shield reductio~ is to be considered the only criterion for phylogenetic advancement vvithin the family, then Synaphaspis must be relegated a position well below that of Mqcrocheles. The fact that Synaphaspis has been found on b~etles indicates that it is phoretic rather than free-living, which indicates an advanced mode of life. Lack of distinct teeth on the chelicerae (fig. 30) also signifies advanced development. It appears likely, therefore, that the consolidation of ventral scierites in Synaphaspis is a result of expansion and subsequent fusion of shields, and that the genus is a morphologically specialized offshoot of J.V! acrocheles. 13. Ancistrocheles n. gen. Dorsal shield not reduced, with an indeterminate number of simple setae on on the shield. With three pairs of sternal setae; metasternal shields absent. Ventrianal shield oblong, rounded anteriorly and with only one pair of preanal setae. Fixed digit of chelicera abbreviated, giving the digitus mobilis a hook-like aspect ; dorsal seta of fixed digit simple. Corniculi long, narrow and yointed, internally notched. Tectum trifid. Pé}lpi with five free segments; palpai claw with three tines. Pre tarsal elements not seen. Other characters as in NI acrocheles Latr. TYPE SPECIES. - Ancistrocheles bregetovae n. sp. Ancistrocheles bregetovae n. sp. FEMALE (figs ). Length of idiosoma 1050-II09 [1. ; width between external angles of coxae IV fl. With the characters of the genus. Setae D 1 simple, well separated ; other dorsal setae simple or expanded distally. Dorsal shield (fig. 34) patterned with fine punctations and distinctly reticulate; lateral and posterior margins rough and uneven. Tritostemum biramous. Sternal,

25 - r U l ' Fig. 33 Synaphaspis congoensis Krantz Dorsum of female (From Krantz 1961).

26 ~ ' ' epigynial and veritrianal shields weakly reticulate; ventral setae simple. Metasternal setae inserted laterad of epigynial shield. Ventrianal shield (fig. 36) slightly broader anteriorly than posteriorly and with one pair of preanal setae inserted mediolaterally. Peritrematal shields free; peritreme extending anteriorly to the middle of coxa I. Ventral integument 'striate-reticulate. Hypostome with five rows of denticles ; gnathosoma punctate dorsally. Cornic~tli (fig. 36) narrow,. extending to a point level with the proximal porton iof the palpai femur ; variously notched internally. Techtm trifid distally, triangular basally (fig. 35). Digitus mobilis of chelicera (fig. 37) reduced, truncate dis~ally ; dorsal seta simple ; fixed digit with a large tooth and a terminal hook. Legs with simple setae, not ridged or spurred. Type material- Two females with the following data: Canon, Arizona, January 2, 1937; ex Cochliomyia macellaria mouthparts (USNM ); collected by C. E. DEONIER. H olotype female will be deposited in the collection of 'the U. S. National Museum, Washington, D. C. The paratype female will be retained in the collection at Oregon State University, Corvallis, Oregon. On the basis of shield reduction, condition of the corniculi and tectal structure, Ancistrocheles shows strong a:ffinities to the genus Lordocheles. The basal tectal structure, as well as the general facies of A. bregetovae, also points to a possible relationship with the genus Neopodocinum. As mentioned earlier in the discussion on Neopodocinum, the latter genus may be related to Lordocheles, which irt turn appears to be allied to the Areolaspis-H olostaspella group. The group of genera to which Ancistrocheles appears to be related, therefore, may consitute a primary developmental branch with highly advanced forms showing progressive shield reduction (Lordocheles, Neopodocinum, Ancistrocheles) while the Areolaspis H olostaspella group represents a specialized offshoot of the primary branch of development. KEY TO GENERA OF THE FAMILY MACROCHELIDAE. Females.. r. Ventrianal shield fused with podal shields; metasternal shields fused in sternal- '. K endopodal complex Synaphaspzs rantz - With free. ventrianal or anal shield; metasternal shields free, fused to the endopal shields, or ~bsent :... 2,;.; 2. With severa! pairs of blunt cylindrical dorsal shield setae, sorne of which are strongly abbreviated, setae D 8 pilose; dorsal shield serrulate marginally; in bee nests.... Grafta n. gen. - Without the above combination of characters Setal brush of digitus mobilis filamentous, extendirig to or beyond the distal end of the digit; dorsal setae simple (D 8 may be pilose), dorsocentral setae (D 4 -D 7 ) typically shorter than other dorsal setae, spine-like; ventrianal shield with two, three or four pairs of preanal setae

27 Fig. 34 Ancistrocheles bregetovae n. gen. n. sp. Dorsum of.female. - Fig. 35 Ancistrocheles bregetovae n. gen. n. sp. Tectum of female. _:_ Fig. 36. Ancistrocheles bregetovae n. gen. n. sp. Venter of female. -Fig. 37 Ancistrocheles bregetovae 11. gen. 11. sp. Chelicera of female. 36 H 0'\ 00 ~}~ê~,,~~)~~ r.=_...j. f~ L -~" -... ~ ~ 9''. \. \ ir. ~~- ~~~.~~'~f ~....~ /l;c Î~r~~~:~~~ \: \ ;,r. 1 \,,.,,.,\ ""' ~ jlî. 35 ~~2::~' J") :.,.,:~,~;:):(('...:. ~. _. : ~ _ ':... : i:i,.-ir~.,j. l'... 3~'f\t~ 500).1 \ _, ~ ~ 37 ~ 06) ~

28 - r6g- - Setal brush o'f digit us mobilis brush-like, not filamentous, never extending -beyond digit ; dorsal setae variable ; with one to five pairs of preanal setae on the ventrianal shield, or with anal shield... :... ' Peritrematal shields fused with exposai shields and extending posteriorly with them around coxae IV Ettholocelaeno Berl. - Peritrematal shields free Holocelaeno Berl. 5 Ventrianal shield with two or more pairs of preanal setae; free-living or insect paraphages.... : :- Ventrianal shield with one pair of preanal setae, or witlï simple anal shield; insect paraphages Tectum tripartite (except NI. superbus Hull) ; with two to four pairs of preanal setae on the ventrianal shield ; metasternal shields free. : Tectum unipartite; ventrianal shield with five pairs of preanal serae; metasternal shields free or fused to endopodal shields, free-living Geholaspis Berl. 7 With anal shield; chelicerae not reduced; tectum unipartite With one pair of preanal setae on the ventrianal shield ; fixed cheliceral digit abbreviated, giving the digitus mobilis a hook-like appearance; lateral elements of tectum fused proximally, -trifid distally Ancistrocheles n. gen. 8. Chelicerae distinctly curved dorsad; metasternal shields absent; tritosternum reduced. Lordocheles Krantz - Chelicerae not curved as above ; metasternal shields present ; tritosternum normal - for family N eopodocinum Oud. 9 Vertical setae inserted on prow-like projection of dorsal shield ; femora of legs H with sclerotized ridges or spurs Io - Without anterior projection of dorsal shield; femora of legs II rarely with sclerotized ridges ors purs II TO. Peritrematal and exopodal shields fused ; ventrianal shield with three pairs of preanal setae... ;... :..... Areolaspis Tragdh... - Peritrematal and exopodal shields not fused ; ventrianal shield with three or four pairs of preanal setae H olostaspella Berl. II. Dorsal shield truncate and abbreviated posteriorly ; sternal, epigynial and ventrianal setae minute, smooth ; in bee nests Trigonholaspis Vitzthum - Dorsal shield rounded or attenuated posteriorly; ventral setae variable but never minute ; with two or three pairs of preanal setae on the ventrianal shield ; para~ phages of Coleoptera or Diptera, or free-living IZ TZ. Dorsal shield strongly crenulate-reticulate ; sternal shield extending posterolaterally around epigynial shield to the level of the posterior angles of coxae III and abutt~ng rounded -metasternal shields... Glypthlaspis Filipponi and Pegazzano - Dorsal shield smooth, punctate, reticulate, or areolate ; sternal shield not extending posterolaterally to the level of the posterior angles of coxae III ; metasternal shields rounded or elongate, separated from, or abutting, posterolateral angles of sternal shield :.. lvi acrocheles Latr.