New taxa of mites associated with Australian termites (Acari: Mesostigmata)

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New taxa of mites associated with Australian termites(Acari: Mesostigmata)Robert B. Halliday aa CSIRO Entomology, GPO Box 1700, Canberra, ACT, 2601, Australia E-mail:

Version of record first published: 17 Mar 2009.

To cite this article: Robert B. Halliday (2006): New taxa of mites associated with Australian termites (Acari: Mesostigmata),International Journal of Acarology, 32:1, 27-38

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Vol. 32, No. 1 Internat. J. Acarol. 27

NEW TAXA OF MITES ASSOCIATED WITH AUSTRALIAN TERMITES (ACARI: M ESOSTIG MATA)

Robert B. Hal l iday CSIRO Entomology, GPO Box 1700, Canberra ACT 2601, Australia ([email protected]).

A B S T R A C T - New species, genera, and families of Mesostigmata are described from the nests of Austra- lian termites - Laelaptonyssus lacticolus n. sp. (Laelaptonyssidae) from Coptotermes lacteus; Teranyssus howardensis n. gen., n. sp., (Teranyssidae n. ram.) from Mastotermes darwiniensis; Uronyssus milleri n. gen., n. sp. (Uronyssidae n. ram.) from Nasutitermes graveolus, and U. watsoni n. sp. from Nasutitermes exitiosus. The ecological role of these species is not clear, but they may be parasites of termites. Key words - Acari, Mesostigmata, Laelaptonyssidae, Teranyssidae n. ram., Uronyssidae n. faro., Laelap- tonyssus, Teranyssus, Uronyssus, mites associated with termites.

INTRODUCTION

The nests of social insects provide accommodation for many groups of mites (reviewed by Eickwort, 1990). These include large numbers of species of mites that are simply saprophytic, and take advantage of the controlled climate and abundant food provided by their hosts. Some, such as Varroa, are clearly parasitic. Others, such as the genus Laelaptonyssus Womersley, have a more ambigu- ous relationship with their hosts. The morphological spe- cialisations of Laelaptonyssus, the fact that they are often found riding on the heads or bodies of termites, and the fact that they die soon after the death of their hosts (Wang et al., 2002), suggests an intimate relationship, but the na- ture of that relationship is not known. Krantz (2000) sug- gested that Laelaptonyssus are phoretic on termites rather than parasitic, and feed on nematodes in the termites' nest material.

The work described in this paper is a continuation of a previous study of termite-associated species of Laelaptonyssus in Australia (Halliday, 1987). Laelapto- nyssus was described by Womersley (1956) as the type genus of his new family Laelaptonyssidae. Lee (1970) and Halliday (1987) placed Laelaptonyssus in the Rhodacaridae, implying its placement in a subfamily Laelaptonyssinae, but Halliday (1998, 2001) and Krantz (2000) then reverted to Womersley's original usage of the family Laelaptonyssidae. The family currently includes the species L. mitis Womersley 1956 (Australia), L. chinensis (Sarn~infik 1964) (China), L. lenzi Halliday 1987 (Australia), L. darwiniensis Halliday 1987 (Austra- lia), L. hallidayi Krantz 2000 (Australia) and L. setosus

Krantz 2000 (USA). Wang et al. (2002) referred to L. setosus as Laelaptonyssus sp. Baker and Johnston (1959) described a new species as Laelaptonyssus phytoseioides, but Krantz and Khot (1962) transferred this species to their new genus Treatia in the family Otopheidomenidae.

Laelaptonvssus mitis was found in a laboratory culture of the housefly, while all other species are associated with termites. The known collections of L. mitis, L. chinensis, L. darwiniensis, and L. setosus are as previ- ously described (Halliday 1987; Krantz 2000). I here de- scribe a new species of Laelaptonyssus from Australia, and take the opportunity to document some recently collected specimens of L. hallidavi and L. lenzi.

The search for specimens of Laelaptonyssus also yielded some remarkable mites in apparently related groups. These cannot be accommodated in Laelapto- nyssus, so they are placed in two new genera - Uronyssus n. gen. and Teranvssus n. gen. These latter two genera are sufficiently distinctive to warrant the erection of two new families, Uronyssidae n. fam. and Teranyssidae n. fam.

M A T E R I A L S A N D M E T H O D S

Specimens were cleared in Nesbitt's solution and mounted in Hoyer's medium using methods described by Krantz (1978). Specimens are deposited in the Australian National Insect Collection, Canberra (ANIC) unless otherwise stated. The system of nomenclature for the dorsal shield setae is that of Lindquist and Evans (1965), and the leg chaetotaxy that of Evans (1963). All measurements are given in micrometers (~tm).

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F a m i l y L A E L A P T O N Y S S I D A E W o m e r s l e y

Laelaptonyssidae Womersley, 1956: 543. Laelaptonyssinae: Lee, 1970: 72. Laelaptonyssidae: Halliday, 1998:130; Krantz, 2000: 26.

Krantz (2000) drew attention to the extraordinary amount of variation in some morphological characters in Laelaptonvssus species, but identified the following features as diagnostic of the family: (1) elongate and attenuate chelicerae; (2) narrow acicular corniculae; (3) fusion of the palp tibia and tarsus; (4) loss of claws and peritremes; (5) elongate recurved spermatodactyli in the male.

Laelaptonyssus W o m e r s l e y

Laelaptonyssus Womersley, 1956: 543. Type species - Laelaptonyssus mitis Womersley, 1956, by original designation.

Puchihlungia Samginfik, 1964: 39. Type species - ] Puchihlungia sinensis Sam~infik, 1964: 39, by origi-

nal designation. Synonymy by Lee, 1970.

Laelaptonyssus hallidayi Krantz

Laetaptonyssus haltidayi Krantz, 2000:31.

Materia l examined - Holotype female, Australian Capital Territory, Brindabella Range, 31.v. 1978, M. Lenz coll., in nest of Porotermes adamsoni. Paratypes - 5 females, 6 males, 1 deutonymph, same data as holotype (ANIC).

New materia l e x a m i n e d - Australian Capital Terri- tory, 8 females, 6 males, 6 deutonymphs, Canberra, CSIRO Entomology, xi.1986, in laboratory colony of Porotermes adamsoni, origin Piccadilly Circus ACT, M, Lenz coll. (ANIC).

Notes -The new material examined here came from the same locality as the type specimens. The fact that the population ofL. hallidayi was still present eight years after the original collection shows that this is a long-term association between mite and termite, and not just a coin- cidence.

Laelaptonyssus lenzi Hal l iday

Laelaptonyssus lenzi Halliday, 1987: 86.

Material examined - Holotype female, New South Wales, Termeil, near Bateman's Bay, 16.xi.1983, M. Lenz coll., on Coptotermes lacteus. Paratypes - 2 females, 4 males, same data as holotype (ANIC).

New material e x a m i n e d - New South Wales, 1 female, 4 males, Gundaroo, 15.iv.1996, M. Lenz coll., in nest of Coptotermes lacteus; 3 females, Termeil,

19.i.1989, M. Lenz coll., in nest of Coptotermes lacteus (ANIC).

Notes - The new material from Termeil was from the same locality as the type specimens, six years after the original collection. The specimens from Gundaroo (30 km north of Canberra) represent a significant range extension for this species, since this site is 150 km inland from the type locality.

Laelaptonyssus lacticolus n. sp. (Figs. 1-7)

F E M A L E - D o r s u m (Fig. 1) - Dorsal shield divided completely in two by transverse suture at the level of coxa IV. Podonotal shield smooth and unornamented, posterolateral comers slightly produced; with a pair of irregularly shaped selerotised ridges between setae j5. Length 449-504, width 554-613 (n = 7). Shield with 16 pairs of setae; s3, s6, z6, j6 long (110-120), j l , j2, j3, j4, j5, z3, s4 minute (12-17 long), others intermediate (30-60 long), z5 vestigial, represented only by its insertion. Opisthonotal shield smooth and unornamented, with an irregular mass of heavy sclerotisation along the anterior margin; posterior margin eroded, emarginate. Length 273-386, width 529-559 (n = 5). Shield with 15 pairs of setae, length 75-150, homologies difficult to determine.

Venter (Fig. 2) - Tritostemum with an ovoid base and smooth laciniae. Sternal shield with four pairs of setae, two pairs of lyriform pores, and a pair of lateral in- cisions at the level of the first pair of pores. These inci- sions joined by a transverse line; faint polygonal ornamentation present posterior to this line, shield otherwise smooth to slightly granular, except for a heavily sclerotised transverse ridge between setae st3 and st4. Epigynial shield elongate, with indistinct polygonal ornamentation medially, and one pair of setae, posterior margin slightly concave. Ventrianal shield elongate, with indistinct polygonal ornamentation in the anterior half, four pairs of ventrianal setae, a pair of paranal setae, and a postanal seta, setae increasing in length posteriorly. Opisthosomal integument with a pair of elongate metapodal plates, a pair of setae inserted in the soft integument, and a pair of setae inserted on elongate platelets. Peritremes minute, confined to an area opposite coxa IV.

G n a t h o s o m a - Hypostomal setae hl and h3 and palp coxal setae subequal in length (ca. 40 long), hypostomal setae h2 about one-third this length. Hypostomal groove without denticles; corniculae slender and attenuate; internal malae broad, smooth (Fig. 3). Fixed digit of chelicera long and slender, with a single pointed retrorse tooth; movable digit long and slender, with downward-directed tip and two pointed retrorse teeth (Fig. 4); palp tibia and tarsus fused; chaetotaxy - trochanter 2, femur 4, genu 5; epistome not visible.

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L e g s - Chaetotaxy - Leg I: coxa 0 0/1 0/1 0; trochanter 0 1/2 1/1 l; femur 1 2/1 3/2 1; genu 1 2/2 2/1 1; tibia 1 2/1 2/1 1; tarsus ca. 30 setae. Leg II: coxa 0 0/1 0/1 0; trochanter 0 1/2 0/1 1; femur 1 1/1 2/3 2; genu2 2/1 2/2 0; tibia 2 1/1 2/1 l; tarsus 3 4/1 4/1 1 + mv, md. Leg III: coxa 0 0/1 0/1 0; trochanter 0 1/2 0/1 1; femur 1 1/1 2/2 0; genu 1 2/1 2/1 1; tibia 1 1/1 2/1 1; tarsus 3 4/1 4/1 3 + my, md. Leg IV: coxa 0 0/0 0/1 0; trochanter 1 1/2 0/1 0; femur 1 2/1 1/1 1; genu 1 2/1 1/1 l; tibia 1 1/1 2/1 1; tarsus 3 4/1 4/1 3 + mv, md. Micro setae are femur I, pv2, pd2, genu I, pd2, tibia I, ad2, pd2, femur II, pd2, tibia II, pd2, femur III, pd2, genu III, ad2, tibia III, pd2, genu IV, pd2, tibia IV, pd2. Claws absent; pre-tarsi represented only by a membranous pad.

M A L E - D o r s a l i d i o s o m a - Length 651-659, width 420-437 (n = 4). Structure and chaetotaxy as for female.

V e n t e r (Fig. 5) - Tritosternum as for female. Sternogenital shield smooth, with five pairs of setae and two pairs of pores, anterior margin weakly defined, genital opening near anterior margin of shield. Ventrianal shield with indistinct polygonal ornamentation medially, five pairs of ventrianal setae, a pair of paranal setae, and heavy postanal seta. Opisthosomal integument with a pair of elongate metapodal platelets, and two pairs of small platelets each bearing a heavy seta. Peritremes as for female.

Gnathosoma - As for female except movable digit of chelicera long and edentate, with a terminal hook; spermatodactyl very robust, recurved, tip pointed; fixed digit long, attenuate, edentate (Fig. 6).

L e g s - As for female except one ventral seta on femur II modified into a heavy finger-like spur (Fig. 7).

M a t e r i a l e x a m i n e d - Holotype female, Australia, New South Wales, Tallaganda State Forest (near Cap- tain's Flat, 35°38'S 149°33'E), 10.viii.1989, M. Lenz coll., in nest of Coptotermes lacteus. Paratypes - 6 females, 4 males, same data as holotype (ANIC)

Etymology - The name of this species refers to its association with its host Coptotermes lacteus.

R e m a r k s - In the key of Krantz (2000), L. lacticolus keys out to L. lenzi (palp trochanter with two setae, podonotum with sclerotised bands between setae j5/j6). It is interesting to note that these two species of Laelaptonyssus occur with the same species of host, and have overlapping geographic distributions (populations ofL. lenzi at Gundaroo and L. lacticolus at Tallaganda are separated by only 60 kin). However, L. tacticolus may easily be distinguished from L. tenzi on at least two crite- ria. In the soft ventral skin behind the metapodal plates, L. lenzi has a pair of large setae inserted on small sclerotised platelets, and two pairs of short setae inserted in the soft skin. Laelaptonyssus lacticolus has the corresponding large setae inserted in platelets, but instead of two pairs of short setae, it has a single pair of long setae inserted in the soft skin. In L. lenzi, the posterior margin of the podonotal shield carries two pairs of long setae and two pairs of

short setae. In L. lacticolus, the posterior margin of the podonotal shield carries three pairs of long setae.

Laelaptonyssus lacticolus continues the trend towards bizarre aberrations in morphology observed in this genus. The opisthonotal shield has an irregular band of heavy sclerotisation along its anterior margin, and an eroded and emarginate posterior margin; and a slightly hypertrichous chaetotaxy. The leg chaetotaxy also reflects the irregular patterns observed in other species of Laelaptonyssus. Many setae have migrated towards the distal end of their segments, so their precise position is subject to differences in interpretation, as noted by Krantz (2000). There is an extraordinary combination of macro- and micro-setae on some segments. Laelaptonyssus lacticolus also shows some reductions in chaetotaxy that have not been seen before - 9 setae on genu I, 8 on tibia I, and 8 on tibia II. This species does show the sclerotised nodules on the podonotal shield between setae j5, which are remininscent of those found in the Rhodacaridae. These nodules, along with 4 pairs of setae on the sternal shield, and a divided dorsal shield, once again support the placement of the Laelaptonyssidae in the Rhodacaroidea.

Laelaptonyssus lacticolus also continues the trend of association between species of Laelaptonyssus and termites - L. setosus Krantz with Reticulotermes flavipes in USA, L. lenzi Halliday with Coptotermes lacteus in Aus- tralia, L. hattidayi Krantz with Porotermes adamsoni in Australia, L. darwiniensis Halliday with C. acinaciformis in Australia, and L. chinensis (Samginfik) in China. The occurrence of L. mitis Womersley in a laboratory culture of housefly at the University of Western Australia (Womersley, 1956) is anomalous. It is possible that L. mitis may be found in association with Coptotermes in Western Australia. To test this idea I examined 30 collections of Coptotermes from the southern half of Western Australia. Seven of these samples contained acarid hypopodes and one contained a single pygmephorid, but none contained any Mesostigmata. It should also be re- membered that there is precedent for Womersley's published collection records to be found to be inaccurate (Southcott, 1986).

T E R A N Y S S I D A E N. F A M .

D i a g n o s i s - Rhodacaroidea with an entire dorsal shield, without sclerotised nodules; female with an anal shield with one pair of paranal setae and a postanal seta; epigynial shield rounded posteriorly, with one pair of setae; tarsi without claws; pretarsus I much bigger than III-IV; legs I much thicker than II-IV; peritremes very reduced; palp tibia and tarsus not fused, palp tarsal claw 2-tined; cheliceral digits attenuate, edentate. Male with straight spermatodactyl; legs without spurs.

T y p e g e n u s - Teranyssus n. gen., here designated. Etymology - The name of the family is based on the

type genus.

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32 Halliday 2006

IV

Fig. 15. Teranyssus howardensis n. sp. (female) - Legs I-IV. For each leg the upper figure shows the ventral and lateral setae, the lower figure shows the dorsal setae (scale bar = 100 ~tm).

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Vol. 32, No. 1 Intemat. J. Acarol. 33

Teranyssus n. gen .

D i a g n o s i s - Teranyssidae with dorsal shield almost circular, posteriorly truncate, with 24 pairs of setae; anal shield much wider than long, partly visible dorsally; coxa I with seta pv short, thick, rounded; seta av on coxae III-IV short, thick, spine-like; femur, genu and tibia IV with several very long and heavy dorsal macrosetae; tritosternum with very elongate base and short laciniae.

T y p e s p e c i e s - Teranyssus howardensis n. sp., here designated.

E t y m o l o g y - The name of this genus is derived from teras (Gr., monster) and nysso (Gr., prick), referring to its fanciful resemblance to a spiky monster, caused by the appearance of the leg setae.

Teranyssus howardensis n. sp. ( F i g s . 8 - 1 5 )

F E M A L E - D o r s u m (Fig. 8) - Dorsal shield almost circular (length 601-647, width 575-617, n = 10), slightly truncate and sometimes eroded posteriorly; surface smooth, lacking sclerotised nodules, but with distinct sigillae between j l/j2 and adjacent to j4. Shield with 24 pairs of setae, all smooth and pointed, increasing in length posteriorly; j 1 fine, all others robust. Anal shield extending onto dorsal surface, with robust post-anal seta and circular cribrum.

V e n t r a l i d i o s o m a (Fig. 9) - Tritostemum with elongate base and short brush-like laciniae. Sternal shield smooth, with a heavily sclerotized strip across the anterior margin, and extending into the endopodal area between coxae I/II/III. Posterior comers of shield fused with endopodal plates embracing coxae IV. Shield with four pairs of setae and two pairs of lyriform pores. Epigynial shield drop-shaped, smooth, with one pair of setae. Anal shield very wide, with a pair of paranal setae on the ventral surface, postanal seta and cribrum on a dorsal extension of the shield. Peritremes extending from opposite coxa IV to the midlevel of coxa III, peritrematal plates with a lightly sclerotized posterior extension embracing coxae IV. Opisthogastric integument without setae or metapodal plates.

G n a t h o s o m a - Palp coxal seta long (ca. 47 long), hypostomal seta h2 short (ca. 16 long), hypostomal setae hl and h3 intermediate (ca. 32 long). Hypostomal groove without denticles; corniculae short, slender, attenuate; internal malae conspicuous, smooth (Fig. 10). Fixed digit of chelicera straight, pointed, edentate except for a few blunt serrations proximally; movable digit edentate, curved, with a blunt tip (Fig. 11). Epistome not visible. Palp with 5 free segments, tibia and tarsus not fused; chaetotaxy: trochanter 2, femur 4, genu 6, tibia 5, tarsus ca. 15; palp tarsal claw 2-tined, one tine pointed, the other spatulate (Fig. 12).

L e g s (Fig. 15) - Legs I and IV longer and thicker than II and III. Chaetotaxy: Leg I: coxa 0 0/1 0/1 0; trochanter 1 0/1 0/2 1; femur 2 2/2 2/2 0; genu 2 2/2 2/2 0; tibia 2 2/2 2/2 0; tarsus ca. 30 setae. Leg It: coxa 0 0/1 0/1 0; trochanter 1 0/1 0/2 1; femur 1 2/2 3/1 0; genu2 2/1 2/1 2; tibia 2 2/1 2/1 2; tarsus 3 4/1 4/1 3 + mv, rod. Leg III: coxa 0 0/1 0/1 0; trochanter 1 0/1 0/2 1; femur 1 2/2 2/1 0; genu 2 2/1 2/1 0; tibia 1 2/1 2/1 2; tarsus 3 4/1 4/1 3 + mv, md. Leg IV: coxa 0 0/0 0/1 0; trochanter 1 0/1 0/2 1; femur 1 2/1 3/1 1; genu 2 2/1 3/1 2; tibia 2 2/1 2/1 2; tarsus 3 4/1 4/1 3 + my, md. Pre-tarsi without claws, pretarsus I much bigger than II-IV.

M A L E - Dorsum - Dorsal shield length 563-580, width 508-559 (n = 6); structure and chaetotaxy as for female.

V e n t e r (Fig. 13) - Tritostemum as for female. Stereo-genital shield smooth, with a heavily sclerotised marginal ridge extending along anterior and lateral edges; shield with five pairs of setae and two pairs of pores; genital opening at anterior margin of shield. Ventrianal shield smooth, with one pair ofventrianal setae, a pair ofparanal setae, and a long heavy postanal seta, cribrum not visible ventrally. Opisthogastric integument smooth, without setae or metapodal plates. Peritreme extending from midlevel of coxa IV to midlevel of coxa III.

G n a t h o s o m a - As for female except movable digit of chelicera narrow, curved; spermatodactyl completely fused with digit except for a slight distal protuberance; fixed digit straight, with blunt tip (Fig. 14).

L e g s - As for female, without spurs or modified setae.

M a t e r i a l e x a m i n e d - Holotype female, Australia, Northern Territory, Howard Springs, 21 km SE of Dar- win, 7.vii.1985, M. Lenz coll., in nest of Mastotermes darwiniensis. Paratypes - 14 females, 6 males, same data as holotype; 1 male, Berrimah Research Farm, 21.iv.1999, M. Geyer coll., ex body of Mastotermes darwiniensis; 1 female, 1 male, Kakadu National Park, Kapalga Billabong, 28.xi. 1990, L. Miller coll., in nest of Mastotermes darwiniensis. (ANIC).

E t y m o l o g y - The name of this species is derived from the type locality.

N o t e s - The fusion of the sternal and metastemal shields in the female to form a stemal/metastemal shield with 4 pairs of setae would place Teranyssus in the Rhodacaroidea sensu Krantz (1978). However, this is un- comfortable due to its possession of an anal rather than ventri-anal shield. Within the Rhodacaroidea, Teranyssus cannot be placed in the Rhodacaridae, Digamasellidae, or Laelaptonyssidae, because these families have a divided dorsal shield. Teranyssus has many features in common with Laelaptonyssus, but in addition to its entire dorsal shield, it differs from Laelaptonvssus in having a straight spermatodactyl, clearly separate palp tibia and tarsus, and strongly developed endopodal extensions of the sternal shield. The peritrematal plate of Teranyssus is larger than

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34 Halliday 2006

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\ 18 19

21 .... 2

Figs. 16-21. Uronyssus milleri n. sp., 16-19, female - 16. dorsal idiosoma; 17. ventral idiosoma; 18. hypostome; 19. chelicera. 20-21, male - 20. ventral idiosoma; 21. chelicera (scale bar = 100 ~tm for 16, 17; 50 gm for 18, 19, 21).

that of Laelaptonyssus, and has a long posterior extension that embraces the posterior edge of coxa IV, and there are no metapodal plates. The male of Teranyssus lacks the ventral spur on femur II that is typical of the males of Laelaptonyssus. Teranyssus is similar to the Ologama- sidae in having an entire dorsal shield, no scleronoduli, and the anterior portion of the sternal shield not differenti- ated from the remainder of the shield. However, it has a

series of specializations not found in the Ologamasidae, such as attenuate cheliceral digits, reduced peritreme, anal shield, and highly modified tritostemum and leg chaetotaxy. It may be that Teranyssus is a highly derived relative of the Ologamasidae, but the distinctive characters of this genus make it necessary to create a new family to accom- modate it.

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Another feature that Teranyssus has in common with Laelaptonyssus is its aberrant leg chaetotaxy. In many segments the setae have migrated to the distal end of the segment, which makes recognition of the chaetotaxic pattern difficult in some cases, and potentially subject to differences in interpretation. It also shares with Laelaptonyssus a mixture of macro- and microsetae on some segments. Setapv on coxa I of Teranvssus is a thick peg with a rounded end. Some ventral setae on the femur, genu and tibia of legs II-IV, are short, thick and spine-like, and many dorsal setae are very long and thick, with blunt tips, especially those at the distal end of segments.

U R O N Y S S I D A E N. F A M .

D i a g n o s i s (Female) - Dermanyssoidea with mode - rately to strongly hypertrichous dorsal shield; metasternal plates and setae absent; peritremes and hypostomal denticles absent; corniculae very reduced and attenuate; cheliceral digits elongate, attenuate. Paranal setae and postanal seta hypertrophied, very long and thick; anal shield with strongly concave anterior margin; epigynial shield elongate, triangular.

Type genus - Uronvssus n. gen., here designated. E t y m o l o g y - The name of the family is based on the

type genus.

Uronyssus n. gen .

D i a g n o s i s (Female) - Uronyssidae with at least three pairs of conspicuous elongate setae on caudal margin of dorsal shield; setae in posterior half of dorsal shield long enough to exceed insertions of next most posterior setae; sternal shield with three pairs of setae and two pairs of pores; epigynial shield with three or four pairs of setae; ventri-anal shield with one pair of ventrianal setae, one pair of paranal setae and an unpaired postanal seta; opisthogastric integument with a pair of elongate metapodal platelets; movable digit of chelicera with rounded ventral lobe, palp tibia and tarsus fused; chaetotaxy - trochanter 1, femur 4, genu 5.

Type species - Uronyssus watsoni n. sp., here designated.

Etymology - The name of this genus is derived from oura (Gr., tail) and nvsso (Gr., prick), referring to the exceptional development of the para-anal and postanal setae to form a three-pointed tail.

Uronyssus milleri n. sp. ( F i g s . 1 6 - 2 1 )

F E M A L E - Dorsum (Fig. 16) - Surface of dorsal shield smooth, without sclerotised nodules. Length 400-437, width 200-231. Shield with ca. 180 smooth, pointed setae, pairwise arrangement not discernable; setae

increasing in length posteriorly, 3 caudal pairs noticeably longer (ca. 75 long).

V e n t e r (Fig. 17) - Tritosternum with rectangular base and lightly pilose laciniae. Sternal shield with weak polygonal ornamentation laterally, smooth medially; with three pairs of setae and two pairs of lyriform pores; fused with endopodal plates between coxae II/III. Metasternal plates and metasternal setae absent. Metasternal pores lo- cated in soft integument mediad of endopodal plates IV. Epigynial shield triangular, with very faint linear ornamentation and four pairs of long pointed setae. Ventri- anal shield heart-shaped, with a pair of heavy setae inserted in anterolateral horns; paranal setae very long and heavy (125 long), curved outwards, postanal seta shorter, straight. Opisthogastric integument with a pair of elongate metapodal plates. Peritrematal plates extending from behind coxa IV to midlevel ofcoxa I, peritremes absent.

G n a t h o s o m a - Palp coxal seta and hypostomal setae hl and h3 subequal in length (ca. 25 long), hypostomal seta h2 less than half this length. Hypostomal groove without denticles; corniculae short, slender, attenuate; internal malae long and conspicuous (Fig. 18). Palp tarsal claw 2-tined. Fixed digit of chelicera narrow, curved, edentate, with a bifid tip, dorsal seta very long and conspicuous; movable digit edentate with a rounded ventral lobe (Fig. 19).

L e g s - Chaetotaxy - Leg I: coxa 0 0/1 0/1 0; trochanter 1 0/1 0/2 1; femur 1 3/2 2/1 1; genu 2 2/1 2/1 1; tibia 2 2/1 2/1 1; tarsus ca. 30 setae. Leg II: coxa 0 0/1 0/1 0; trochanter 1 1/2 0/1 0; femur 1 2/2 2/1 1; genu 2 2/1 2/1 1; tibia 2 2/1 2/1 1; tarsus 3 4/1 4/1 3 + mv, md. Leg III:coxa 0 0/1 0/1 0; trochanter 1 1/2 0/1 0; femur 1 2/2 1/1 1; genu 2 2/1 2/1 1; tibia 2 1/1 2/1 1; tarsus 3 4/1 4/1 3 + mv, rod. Leg IV: coxa 0 0/1 0/0 0; trochanter 1 1/2 0/1 0; femur 1 2/2 1/1 1; genu2 2/1 2/1 1; tibia 2 1/1 2/1 1; tarsus 3 4/1 4/1 3 + mv, md. Setae adl ,pdl on femur I and II and adl on femur IV thickened, all other setae fine, smooth, pointed. Pretarsus I much bigger than II-IV, claws absent.

M A L E - Dorsum - Dorsal shield length 366-376, width 195-200 (n = 2), structure and chaetotaxy as for female.

V e n t e r (Fig. 20) - Tritosternum as for female. Sterniti-genital shield smooth, with eight pairs of smooth pointed setae; fused posteriorly with ventrianal shield. Ventrianal shield with a pair of robust setae in anterolateral corners, a pair of very long, thick, curved setae (110 long) at posterolateral comers, and a straight postanal seta. Opisthogastric integument with a pair of elongate metapodal plates. Stigmata at level of coxa IV; peritrema - tal plates extending from behind coxa IV to midlevel of coxa I, peritremes absent.

G n a t h o s o m a - Fixed digit of chelicera sinuous, edentate, with rounded tip; dorsal seta short. Movable digit straight, pointed, edentate; spermatodactyl straight, fused with movable digit for most of its length, projecting

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24

Figs. 22-25. Uronyssus watsoni n. sp., 22-27, female - 22. dorsal idiosoma; 23. ventral idiosoma; 24. hypostome; 25. chelicera. 26-27, male - 26. ventral idiosoma; 27. chelicera (scale bar = 100 ~m for 22, 23, 26; 50 ~tm for 24, 25, 27).

slightly beyond movable digit (Fig. 21). Other features of gnathosoma as for female.

Legs - Unmodified, structure and chaetotaxy as for female.

M a t e r i a l e x a m i n e d - Holotype female, Northern Territory, Darwin, May 1986, L. Miller, in nest of Nasutitermes graveolus. Paratypes - 3 females, 2 males, same data as holotype. (ANIC).

Etymology - This species is named in honour of Leigh Miller, who collected the type specimens.

N o t e s - The presence of three pairs of setae on the sternal shield of the female, and the pointed posterior end of the epigynial shield, suggest dermanyssoid affinities for Uronyssus. However, Uronyssus does not agree with any known family of Dermanyssoidea, except perhaps for

a very loosely conceived family Laelapidae. The lack of deutosternal denticles, the very reduced corniculae, two ventral setae on genu IV, and the absence of peritremes, are not typical for the Laelapidae, so it is necessary to create a new family for this genus.

Uronyssus watsoni n. sp. (Figs . 22 -27 )

F E M A L E - D o r s u m (Fig. 22) - Dorsal shield length 281-297, width 208-224 (n = 10), surface smooth, truncate posteriorly. Shield with ca. 120 setae, increasing in length posteriorly; four pairs of long setae (75 long) on posterior margin of shield; two pairs of long setae inserted in soft integument posterior to shield; opisthonotal region

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Vol. 32, No. 1 Tnternat. J. Acarol. 37

hypertrichous, with many unpaired and asymmetrical setae.

r e n t e r (Fig. 23) - Tritostemum with rectangular base and lightly pilose laciniae. Stemal shield smooth, with three pairs of setae and two pairs of lyriform pores. Metastemal plates and setae absent; metastemal pores in soft integument level with coxae IV. Epigynial shield smooth, triangular, with three pairs of setae. Anal shield heart-shaped, with a pair of robust setae in the anterolateral comers, a pair of very long (125) heavy curved para-anal setae on the posterolateral comers, and a straight post-anal seta. Opisthogastric integument with a pair of setae on small platelets near posterior end of epigynial shield, and a pair of elongate metapodal platelets.

G n a t h o s o m a - Palp coxal seta and hypostomal setae hl and h3 subequal in length (ca. 10 long), hypostomal seta h2 shorter (ca. 5 long). Hypostomal groove and denticles absent; comiculae apparently absent; intemal malae short and broad (Fig. 24). Palp tarsal claw 2-tined. Fixed digit of chelicera edentate, sinuous, with blunt tip; dorsal seta long and conspicuous; movable digit edentate, shorter than fixed digit, with a prominent ventral keel (Fig. 25).

L e g s - Leg I much thicker than II-IV. Chaetotaxy as for U. milleri. Some setae conspicuously long and thick: femur I: adl, pdl; femur II, adl, pdl; femur III, adl; genu III: adl, pdl; femur IV: adl; genu IV: adl, pdl. Pre-tarsus I much bigger than II-IV, claws absent.

M A L E - D o r s u m - Dorsal shield length 234-239, width 171-176 (n = 2). Structure and chaetotaxy as for female.

V e u t e r (Fig. 26) - Tritostemum as for female. Stemiti-genital shield smooth, with eight pairs of setae; shield fused with ventrianal shield; ventrianal shield with one pair of long and thick setae inserted on anterior corners of shield. Paranal setae and postanal seta very long and thick. Opisthogastric integument with a pair of elongate metapodal plates and a pair of long thick setae laterad of the anus. Stigmata at level of coxa IV; peritrematal plates extending from behind coxa IV to midlevel of coxa I; peritremes absent.

G n a t h o s o m a - Fixed digit of chelicera straight, edentate, with truncate tip; movable digit straight, edentate, with pointed tip; spermatodactyl completely fused with movable digit (Fig. 27). Other features of gnathosoma as for female.

L e g s - Structure and chaetotaxy as for female. M a t e r i a l e x a m i n e d - Holotype female, New South

Wales, Urimbirra, 20 km north of Canberra, 11.ii.1986, J. A. L. Watson and H. Abbey coll., in nest of Nasutitermes exitiosus. Paratypes - 15 females, 2 males, same data as holotype (ANIC).

Etymology - This species is named in honour of the late Tony Watson, who was partly responsible for collect- ing the type specimens.

N o t e s - The two species of Uronyssus described here may be distinguished by the greater degree of hypertrichy of the dorsal shield in U. milleri, and by the fact that the fourth pair of epigynial setae are on the epigynial shield in U. milteri, and on separate platelets in U. watsoni.

DISCUSSION

The morphological diversity of the mites described here offers considerable challenges to their classification. It could be argued that all these taxa should belong to the family Laelaptonyssidae. They are united by several morphological specializations - comiculae very reduced; peritremes reduced or absent; chelicerae attenuate and pointed; leg chaetotaxy highly modified. However, this would create a very heterogeneous family Laelaptonyssi- dae, with variation in some important characters: dorsal shield divided or undivided; female with anal or ventrianal shield; palp tibia and tarsus either fused or not fused; metastemal setae either present or absent; metapodal plates present or absent; podonotal shield with me- dial sclerotised bands or sigillary zone present or absent; femur II of male with or without spurs; spermatodactyl either straight or recurved. This family Laelaptonyssidae would be radically different from the one that is presently conceived (Krantz, 2000).

Furthermore, some of these morphological speciali- sations also occur in other quite diverse insect-associated groups of Mesostigmata - reduced peritremes, comiculae and cheliceral digits in Myrmonyssus (Laelapidae) (Hunter and Hunter, 1963); reduced peritremes, cheliceral digits and comiculae, and modified leg chaetotaxy, in Canestriniphis (Eviphididae) (Potter and Johnston, 1976); reduced peritremes and chelicerae in Blattisocius (Ascidae) (Haines, 1979); and reduced peritremes, comiculae and cheliceral digits in Nabiseius (Otopheidomeni- dae) (Halliday, 1994). It should therefore not be assumed that these character states are synapomorphies for the taxa described here. Further light will undoubtedly be thrown on the relationships among these taxa when further collections are made, perhaps from different species of hosts and different geographical areas. Also, the concepts of Teranyssus, Teranyssidae, Uronyssus and Uronyssidae developed here must be regarded as provisional, and subject to revision when further species in these groups are discovered.

ACKNOWLEDGEMENTS

I am very grateful to Leigh Miller and Michael Lenz for drawing my attention to these specimens, and Jerry Krantz for his stimulating discussions on the classification of these remarkable mites. I also thank Wendy Whitby for help with access to the ANIC termite collection.

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REFERENCES

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Haines, C. P. 1979. A revision of the genus Blattisocius Keegan (Mesostigmata: Ascidae) with especial ref- erence to B. tarsalis (Berlese) and the description of a new species. Acarologia 20: 19-38.

Halliday, R. B. 1987. Two new species of Laelaptonyssus Womersley (Acarina : Rhodacaridae) from the nests of Australian termites. J. Aust. Ent. Soc. 26: 85-91.

Halliday, R. B. 1994. First record of the family Oto- pheidomenidae (Acarina: Mesostigmata) in Austra- lia, with description of Nabiseius melinae sp. n. J. Aust. Ent. Soc. 33: 347-350.

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Krantz, G. W. andN. S. Khot. 1962. A review of the family Otopheidomenidae Treat 1955 (Acarina : Mesostigmata). Acarologia 4: 532-542.

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Lindquist, E. E. and G. O. Evans. 1965. Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Mem. Ent. Soc. Can. 47: 1-64.

Potter, D. A. and D. E. Johnston. 1976. Canestriniphis megalodacne n. g., n. sp. (Acari: Eviphididae) from a pleasing fungus beetle, Megalodacne heros. Ann. Ent. Soc. Am. 69: 494-496.

Samgin~ik, K. 1964. Termitophile Milben aus der VR China. I. Mesostigmata. Ent. Abh. Staat. Mus. Tierk. Dresden 32: 33-52.

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Wang, C., J. E. Powell and B. M. OConnor. 2002. Mites and nematodes associated with three subterranean termite species (Isoptera: Rhinotermitidae). Fla. Ent. 85: 499-506.

Womersley, H. 1956. On some new Acarina- Mesostig- mata from Australia, New Zealand and New Guinea. J. Linn. Soc. (Zool.) 42: 505-599.

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Documents

New taxa of mites associated with Australian termites (Acari: Mesostigmata)