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A checklist and key to species ofEupodoidea (Acari: Prostigmata) fromAustralia and New Zealand and theirsubantarctic islandsTing‐Kui Qin a

a Landcare Research , Private Bag 92 170, Auckland, New ZealandE-mail:Published online: 30 Mar 2010.

To cite this article: Ting‐Kui Qin (1998) A checklist and key to species of Eupodoidea (Acari:Prostigmata) from Australia and New Zealand and their subantarctic islands, Journal of the RoyalSociety of New Zealand, 28:2, 295-307, DOI: 10.1080/03014223.1998.9517565

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© Journal of The Royal Society of New Zealand,Volume 28, Number 2, June 1998, pp 295-307

A checklist and key to species of Eupodoidea (Acari:Prostigmata) from Australia and New Zealand and theirsubantarctic islands

Ting-Kui Qin*

The Eupodoidea are reportedly represented in Australia and New Zealand (includingtheir subantarctic islands) by 5 families, 14 genera, and 67 species (37 not identified tonamed species), as follows. Eriorhynchidae: Eriorhynchus - 5 species; Eupodidae:Claveupodes - 1, Cocceupodes - 3, Eupodes - 16, Linopodes - 2; Penthaleidae:Chromotydaeus - 1, Halotydeus - 6, Linopenthalo ides - 1, and Penthaleus - 6;Penthalodidae: Penthalodes (generic determination need to be confirmed) - 1,Stereotydeus - 1 1 ; Rhagidiidae: Coccorhagidia - 3, Poecilophysis - 1, and Rhagidia -10. A checklist of all the reported species and a key to the 30 named species areprovided.

Keywords: Eupodoidea; Prostigmata; Acari; mites; Australia; New Zealand; subantarctic islands; key

INTRODUCTIONThe superfamily Eupodoidea (Acari: Prostigmata) currently includes six families:Eriorhynchidae, Eupodidae, Penthaleidae, Penthalodidae, Rhagidiidae and Strandtmanniidae.Members of first five families have been recorded from Australia and New Zealand, includingthe subantarctic Auckland Island, Campbell Island and Macquarie Island.

Until 1994, no comprehensive systematic study of Eupodoidea had been undertaken inthese countries, and early records were scattered in the literature (Womersley & Strandtmann1963; Strandtmann 1964, 1981; Zacharda 1980); and eupodoids were often listed asunidentified species (Wood 1964; Spain 1967; Styles 1967; Watson 1967; McMillan 1969;Luxton 1982a, 1982b, 1983a, 1983b; Martin 1983). Further systematic research on theEupodoidea in this region was stimulated by the work of Halliday (1991), who reviewed thetaxonomic literature on the redlegged earth mite Halotydeus destructor (Tucker), an importantpasture pest. He found that there was confusion about the identity of H. destructor andtherefore initiated a programme to study it and its relatives. The results are published in Qin(1994, 1996, 1997) and Qin & Halliday (1995, 1996a, 1996b, 1997).

To date, 67 species in 14 genera of Eupodoidea have been reported from Australia andNew Zealand, and their subantarctic islands, although more than half have not been formallynamed and/or described (see checklist below for details). Halotydeus destructor and Penthaleusmajor (Duges) are recognised pests, and P. falcatus Qin & Halliday is considered to be apotential pest (some early records of P. major are probably P. falcatus: see Qin & Halliday1996b). All other named species have been recorded from natural habitats.

This paper provides a checklist and key to all 30 named species of Eupodoidea fromAustralia, New Zealand, and their subantarctic islands. Thirty-seven unidentified and/or

*Landcare Research, Private Bag 92 170, Auckland, New Zealand; email: QinT@landcare.cri.nz

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unnamed species are not included in the key. The record ofRhagidia kerguelensis (Cambridge)from Macquarie Island by Spain & Luxton (1971) is probably a mistake, as first indicated byZacharda (1980).

CHECKLIST OF EUPODOID SPECIES OF AUSTRALIA AND NEW ZEALANDAND THEIR SUBANTARCTIC ISLANDS

ERIORHYNCHIDAE Qin & Halliday 1997Eriorhynchus australicus (Womersley)Penthalodes australicus Womersley, 1941: 294, fig. J-M.Eriorhynchus australicus: Qin & Halliday, 1997: 103, figs 1-14.Distribution: Australia (Victoria).

Eriorhynchus hades Qin & HallidayEriorhynchus hades Qin & Halliday, 1997: 106, figs 15-27.Distribution: Australia (Victoria).

Eriorhynchus ramosus Qin & HallidayEriorhynchus ramosus Qin & Halliday, 1997: 107, figs 28-37.Distribution: Australia (Tasmania).

Eriorhynchus walteri Qin & HallidayEriorhynchus walteri Qin & Halliday, 1997: 111, figs 38-47.Distribution: Australia (Queensland).

Eriorhynchus womersleyi Qin & HallidayEriorhynchus womersleyi Qin & Halliday, 1997: 111, figs 48-58.Distribution: Australia (New South Wales).

EUPODIDAE C. L. Koch

Claveupodes delicatus Strandtmann & PrasseClaveupodes delicatus Strandtmann & Prasse, 1977: 11, figs 42-50; Martin 1983: 15.Distribution: New Zealand (Nelson); Germany.

Cocceupodes sp.Cocceupodes sp. (nrparadoxus Weis-Fogh), Martin, 1983: 15.Distribution: New Zealand (Nelson).

Cocceupodes sp.Cocceupodes sp., Luxton, 1982b: 302Distribution: New Zealand (Waikato).Cocceupodes sp.Cocceupodes sp., Luxton, 1983b: 140Distribution: New Zealand (Waikato).Eupodes longisetatus StrandtmannEupodes longisetatus Strandtmann, 1964: 148, fig. 1; Shiba, 1969: 66.

Distribution: New Zealand (Campbell Island); Japan.

Eupodes minutus (Strandtmann)Protereunetes minutus Strandtmann, 1967: 55, fig. 1.Eupodes minutus (Strandtmann), Strandtmann, 1970: 89.Distribution: Australia (Macquarie Island); New Zealand (South Island); Antarctic Peninsula.Eupodes sp.Eupodes sp. a, Martin, 1983: 15.Distribution: New Zealand (Nelson).

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Eupodes sp.Eupodes sp. (nr crozetensis Strandtmann & Davies), Martin, 1983: 15.Distribution: New Zealand (Nelson).Eupodes sp.Protereunetus sp.n., Martin, 1983: 15.Distribution: New Zealand (Nelson).Eupodes sp.Eupodidae sp., Kinnear, 1991: 278.Distribution: Australia (Western Australia).Note: I confirm that the specimens of Kinnear (1991) belong to Eupodes.

Eupodes sp.Eupodes sp., McMillan, 1969: 386, 393, 399, 402.Distribution: New Zealand (Hurt Valley).Eupodes sp.Eupodes sp., Spain, 1967: 67.Distribution: New Zealand (Steward Island).

Eupodes sp.Eupodes sp., Luxton, 1982b: 302Distribution: New Zealand (Waikato).

Eupodes sp.Eupodes sp., Luxton, 1982a: 562Distribution: New Zealand (Waikato).

Eupodes sp.Eupodes sp., Luxton, 1983a: 97Distribution: New Zealand (Waikato).

Eupodes sp.Eupodes sp., Luxton, 1983b: 140Distribution: New Zealand (Waikato).

Eupodes sp.Protereunetes sp., Wood, 1964: 39.Distribution: New Zealand.

Eupodes sp.Eupodes sp. nov., Watson, 1967: 51.Distribution: Australia (Macquarie Island)

Eupodes sp.Protereunetes sp., Watson, 1967: 52.Distribution: Australia (Macquarie Island)

Eupodid sp.Eupodid sp., Spain, 1967: 67.Distribution: New Zealand (Magister Ridge, South Island).

Linopodes sp.Linopodes sp., Wood, 1964: 39.Distribution: New Zealand.

Linopodes sp.Linopodes sp., Styles, 1967: 1060.Distribution: New Zealand.

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PENTHALEIDAE Oudemans

Chromotydaeus quartus Qin & HallidayChromotydaeus quartus Qin & Halliday, 1996b: 1834, figs 1-12.Distribution: Australia (South Australia, Western Australia).Halotydeus bakerae Qin & HallidayHalotydeus bakerae Qin & Halliday, 1996a: 445, figs 5-15.Distribution: Australia (South Australia, Victoria, Western Australia).

Halotydeus castellus Qin & HallidayHalotydeus castellus Qin & Halliday, 1996a: 445, figs 16-24.Distribution: Australia (New South Wales, Victoria).

Halotydeus destructor (Tucker)Penthaleus destructor Tucker, 1925: 23.Halotydeus destructor. Womersley, 1933: 108; Baker, 1995: 263, figs 1-61; Qin & Halliday,1996a: 447, figs 3 ^ .Distribution: Australia, New Zealand, South Africa.Halotydeus spectatus Qin & HallidayHalotydeus spectatus Qin & Halliday, 1996a: 449, figs 25-33.Distribution: Australia (New South Wales, South Australia).

Halotydeus sp.Halotydeus sp., Watson, 1967: 52.Distribution: Australia (Macquarie Island)Halotydeus sp.Halotydeus sp., Luxton, 1983b: 140Distribution: New Zealand (Waikato).Linopenthaloides novazealandicus StrandtmannLinopenthaloides novazealandicus Strandtmann, 1981: 392, figs 1-14.Distribution: New Zealand (Charleston)

Penthaleus falcatus Qin & HallidayPenthaleus falcatus Qin & Halliday, 1996b: 1837, figs 13-20.Distribution: Australia.

Penthaleus major (Duges)Tetranychus majeur Duges, 1834: 57.Rhyncholophus major (Dug[es]), Murray, 1877: 126.Notophallus bicolor Froggatt, 1921: 33.Penthaleus bicolor (Froggatt), Womersley, 1933: 110.Penthaleus major (Duges), Thor, 1930: 83; Womersley, 1935a: 163; 1941: 292; Thor &Willmann, 1941: 80; Qin & Halliday, 1996b, 1840, figs 21-27.Distribution: Australia; New Zealand; North America; South America; South Africa; Asia.

Note: some records of P. major in the literature may actually be misidentifications (e.g.Narayan 1962).

Penthaleus minor (Canestrini)Notophallus minor Canestrini, 1886: 705; Berlese, 1892: no. 5.Penthaleus minor (Canestrini): Oudemans, 1905: 241; Womersley, 1941: 292; Thor &Willmann, 1941: 85, Qin & Halliday, 1996b: 1844, figs 28-35.Distribution: Australia (South Australia, Western Australia); EuropePenthaleus sp.Penthaleus sp., McMillan, 1969: 393, 399; 402; Spain & Luxton, 1971: 191

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Distribution: New Zealand (Hurt Valley).

Penthaleus sp.Penthaleus sp.; Luxton, 1982b: 302Distribution: New Zealand (Waikato)

Penthaleus sp.Penthaleus sp.; Luxton, 1983b: 140Distribution: New Zealand (Waikato).

PENTHALODIDAE ThorPenthalodes sp.

Penthalodes sp., McMillan, 1969: 399, 402; Spain & Luxton, 1971: 192.Distribution: New Zealand (Hurt Valley)

Note: species of Penthalodes are not supposed to occur in Southern Hemisphere (Strandtmann1979), and this record needs to be confirmed.

Stereotydeus areolatus WomersleyStereotydeus areolatus Womersley, 1935b: 79; Thor & Willmann, 1941: 70; Strandtmann,1979: 49; Qin, 1994: 1308, figs 1-6.Distribution: Australia (South Australia).

Stereotydeus australicus ThorStereotydeus australicus Thor, 1934b: 65-67; Thor & Willmann, 1941: 69; Strandtmann,1979: 49; Qin, 1994: 1310, figs 7-13.Distribution: Australia (South Australia, Victoria, Western Australia).

Stereotydeus brevipalpus QinStereotydeus brevipalpus Qin, 1994: 1313, figs 14-20.Distribution: Australia (South Australia).Stereotydeus nudisetatus StrandtmannStereotydeus nudisetatus Strandtmann, 1964: 152, fig 4.Distribution: New Zealand (Campbell Island).

Stereotydeus occidentalis WomersleyStereotydeus occidentale Womersley, 1935b: 80; Strandtmann, 1979: 49.Stereotydeus occidentalis Womersley, Thor & Willmann, 1941: 71; Qin, 1994: 1315, figs 21-26.Distribution: Australia (South Australia).

Stereotydeus pseudopulcher QinStereotydeus pseudopulcher Qin, 1994: 1318, figs 27-32.Distribution: Australia (Tasmania).

Stereotydeus pulcher StrandtmannStereotydeus pulcher Strandtmann, 1964: 154, fig. 5.Distribution: New Zealand (Campbell Island)

Stereotydeus tasmanicus QinStereotydeus tasmanicus Qin, 1994: 1320, figs 33-38.Distribution: Australia (Tasmania).Stereotydeus undulatus StrandtmannStereotydeus undulatus Strandtmann, 1964: 150, figs 2 & 3.Distribution: New Zealand (Campbell Island).

Stereotydeus sp.Stereotydeus sp., Wood, 1964: 40; Spain & Luxton, 1971: 192.Distribution: New Zealand.

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Stereotydeus sp.Tectopenthalodes sp., Spain, 1967: 86; Spain & Luxton, 1971: 192.Distribution: New Zealand (Ruahine Mountants: Mt Wharite).

RHAGIDIIDAE Oudemans

Coccorhagidia clavifrons (Canestrini)Norneria clavifrons Canestrini, 1886: 710.Rhagidia clavifrons: Thor, 1934a: 318.Coccorhagidia clavifrons: Thor & Willman, 1941: 128; Strandtmann & Prasse, 1977: 13,figs 61-65; Martin, 1983: 16.Distribution: New Zealand (Nelson); South Africa; North America; Europe.

Coccorhagidia sp.Coccorhagidia sp., Luxton, 1982b: 302Distribution: New Zealand (Waikato).Coccorhagidia sp.Coccorhagidia sp., Luxton, 1983b: 140Distribution: New Zealand (Waikato).Poecilophysis macquariensis (Womersley & Strandtmann)Rhagidia macquariensis Womersley & Strandtmann, 1963: 465, figs 41—47.Poecilophysis macquariensis (Womersley & Strandtmann), Zacharda, 1980: 606, fig. 47.Distribution: Australia (Macquarie Island).

Rhagidia campbellensis ZachardaRhagidia campbellensis Zacharda, 1980: 563, figs 26 B, D.Distribution: New Zealand (Auckland Island).

Rhagidia mildredae StrandtmannRhagidia mildredae Strandtmann, 1964: 157, figs 8 & 9; Zacharda, 1980: 561, figs 26 A, C.Distribution: New Zealand (Auckland Island, Campbell Island).

Rhagidia sp.Rhagidia sp., Wood, 1964: 40.Distribution: New Zealand

Rhagidia sp.Rhagidia sp., Strandtmann, 1964: 160Distribution: New Zealand (Campbell Island).

Rhagidia sp.Rhagidia sp., McMillan, 1969: 393, 399, 402.Distribution: New Zealand (Hurt Valley).

Rhagidia sp.Rhagidia sp., Luxton, 1982a: 562Distribution: New Zealand (Waikato).Rhagidia sp.Rhagidia sp., Luxton, 1982b: 302Distribution: New Zealand (Waikato).

Rhagidia sp.Rhagidia sp., Luxton, 1983a: 97Distribution: New Zealand (Waikato).

Rhagidia sp.Rhagidia sp., Luxton, 1983b: 140

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Distribution: New Zealand (Waikato).

Rhagidia sp.Rhagidia sp., Martin, 1983: 16.Distribution: New Zealand (Nelson).

PROVISIONAL KEY TO THE ADULTS OF EUPODOID SPECIES OFAUSTRALIA, NEW ZEALAND AND THEIR SUBANTARCTIC ISLANDSThis key is provisional because I have seen no specimens ofEupodes minutus, Poecilophysismacquariensis or Rhagidia campbellensis, and key features for these species are derivedfrom published descriptions and illustrations. Only the 30 named species are included.Anatomical nomenclature follows that of Baker (1990, 1995), Qin (1994, 1996) and Qin &Halliday (1997) and important features are illustrated in Figures 1-11. Features illustrated inpreviously published works are also referred to in the key where appropriate, e.g. fig. 2 ofStrandtmann(1981).1. Dorsal hysterosoma with 7 or 8 pairs of setae (Fig. 1); coxal field II with only 1 pair

of setae (Fig. 2); aggenital setae arranged in less than 3 rows (Fig. 2); genital setaein 1 or 2 rows (Fig. 2) 11Dorsal hysterosoma with 9 or more pairs of setae (sometimes numerous, see Fig 7);coxal field II with 3 or more pairs of setae (Fig. 8); aggenital setae arranged in 4 ormore rows (Fig. 8); genital setae in 3 or more irregularly rows (Fig. 7) 2

2(1) Palp tibia with 6 or 7 setae (Fig. 11); gnathosomatic base with 15-34 setae (Fig. 10);naso-prodorsal process present (Fig. 7) ERIORHYNCHIDAE ... 3

- Palp tibia with 3 setae; gnathosomatic base without setae; naso-prodorsal processabsent PENTHALEIDAE (in part) ... 7

3 (2) Leg tibiae with only smooth setae; female with 6 pairs of eugenital setae (fig. 34,Qin & Halliday, 1997); sbc2 branched (fig. 37, Qin & Halliday 1997); naso-prodorsal process ornamented Eriorhynchus ramosus

- Leg tibiae with some ciliate setae (e.g. fig. 7, Qin & Halliday 1997); female with 7pairs of eugenital setae (e.g. fig. 41, Qin & Halliday 1997); sbc2 not branched; naso-prodorsal process unornamented 4

4 (3) Three rhagidial organs present on each of tarsi I and II (figs 21, 23, Qin & Halliday1997) Eriorhynchus hadesTwo rhagidial organs present on each of tarsi I and II (e.g. figs 6, 8, Qin & Halliday1997) 5

5 (4) All setae on leg tarsi ciliate (figs 43, 44, Qin & Halliday 1997)Eriorhynchus walteri

Most dorsal setae and some lateral setae on leg tarsi smooth 66 (5) Tarsi III and IV each with 2 rows of ventral setae (fig. 11, Qin & Halliday 1997)

Eriorhynchus australicusTarsi III and IV each with 3 or more irregular rows of ventral setae (fig. 57, Qin &Halliday 1997) Eriorhynchus womersleyi

7 (2) Coxal field I with 3 setae (fig. 2, Strandtmann 1981); prodorsum with 3 pairs ofsetae plus a pair of trichobothria (fig. 1, Strandtmann 1981)

Linopenthaloides novazealandicusCoxal field I with more than 3 setae (Fig. 7; fig. 2, Qin & Halliday 1996b);prodorsum with unpaired (always more than 8) setae plus a pair of trichobothria(Fig. 7; figs 1, 13, 21, 28, Qin & Halliday 1996b) 8

8 (7) Rhagidial organs present on tarsi I, II and III (figs 5, 7, 9, Qin & Halliday 1996b);palp long and slender, with all tibial setae ciliate (figs 11, Qin & Halliday 1996b);fixed cheliceral digit 2-tined Chromotydaeus quartus

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Fig. 1-6 (reproduced from Qin, 1994): Morphological features of eupodoids used in the key.Stereotydeus areolatus Womersley: 1. dorsum, 2 venter; 3. chelicera; 4. hypostome and palp, ventralview; 5. leg I, dorsal view; 6. leg II, dorsal view. Abbreviations: ag = aggenital setae, FD = fixed digit,g = genital setae, MD = movable digit, or = adoral seta, ps= pseudanal setae, ro = rhagidial organ, sbc= subcapitular seta. Open circles on legs = ventral setae.

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\jr:.M , & . • • • • • ' • • •

MDvFD

chaor

tarsus

7 1/'gnathosomal base

I

10

Fig. 7-11 (reproduced from Qin & Halliday, 1997): Morphological features of eupodoids used in thekey. Enorhynchus austrahcus (Womersley): 7. dorsum; 8. venter; 9. chelicera; 10. subcapitulum,ventral view; 11. palp. Abbreviations: cha = cheliceral seta, FD = fixed digit, MD = movable digit, or= adoral seta, sbc = subcapitular seta.

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Rhagidial organs present on tarsi I and II only; palp short and stout, with all tibialsetae smooth or only 1 with cilia (e.g. fig. 19, Qin & Halliday 1996b); fixedcheliceral digit 3-tined (fig. 20, Qin & Halliday 1996b) 9

9 (8) Chelicerae with movable digit short and tapered, not sickle-like (fig. 34, Qin &Halliday 1996b); palp tarsus truncate apically, and tibia with 1 ciliate seta (fig. 35,Qin & Halliday 1996b); dorsodistal papilla absent from genu of legs I and II; with2 pairs of pseudanal setae (figs 28, Qin & Halliday 1996b) Penthaleus minorChelicerae with movable digit long and slender, sickle-like (figs 20, 26, Qin &Halliday 1996b); palp tarsus tapered apically, tibia without ciliate setae (figs 19,27,Qin & Halliday 1996b); papillae present on genu of legs I and II (figs 15,17, Qin &Halliday 1996b); with 3 pairs of pseudanal setae (figs 13, 21, Qin & Halliday1996b) 10

10 (9) Dorsal idiosomal setae sparse (21 pairs), arranged roughly in 6 longitudinal rows,long (generally more than 30 (xm) (fig. 21, Qin & Halliday 1996b); rhagidial organsrecumbent, 3 present on tarsus II (fig. 23, Qin & Halliday 1996b); tarsi II and IIIeach with a single solenidion present Penthaleus major

- Dorsal idiosomal setae numerous (at least 50 pairs), distributed irregularly, short(generally less than 30 (im) (fig. 13 Qin & Halliday 1996b); rhagidial organs erector semi-erect, 2 present on tarsus II (fig. 17, Qin & Halliday 1996b); tarsi II and IIIwithout a solenidion Penthaleus falcatus

11(1) Body sclerotized; cuticle with polygonal or granular ornamentations; dorsum with adistinctive V- or Y-shaped furrow, or with longitudinal parallel furrows onopisthonotal shield (Fig. 1); epirostrum (a sclerotised anterior projection extendingover gnathosoma) present (Fig. 1) PENTHALODIDAE ... 12

- Body unsclerotized or weakly sclerotized; cuticle not ornamented, dorsum withoutV- or Y-shaped or longitudinal parallel furrows; epirostrum absent 20

12 (11) All femora divided (Figs 5, 6) 13- All femora undivided (fig. 5b, Strandtmann 1964) 1713 (12) Palp tarsus longer than palp tibia (Fig. 4) 14

Palp tarsus shorter than palp tibia (e.g. fig. 11, Qin 1994) 1514 (13) Aggenital setae arranged in 4 pairs (Fig. 2) Stereotydeus areolatus- Aggenital setae arranged in 5 pairs (fig. 4b, Strandtmann 1964)

Stereotydeus nudisetatus15 (13) Cuticle ornamented with granules (fig. 7, Qin 1994) Stereotydeus australiacus- Cuticle ornamented with polygonal areas (fig. 33, Qin 1994) 1616(15) Epirostrum stout and short, middle and lateral lobes all dotted; 3 rhagidial organs

not arranged in a longitudinal row on tarsus I (fig. 3e, Strandtmann 1964)Stereotydeus undulatus

Epirostrum extended, middle and lateral lobes not dotted; 3 rhagidial organs arrangedin a longitudinal row on tarsus I (fig. 37, Qin 1994) Stereotydeus tasmanicus

17 (12) Podosomal setae as long as dorsal hysterosomal setae 18- Podosomal setae distinctly shorter than dorsal hysterosomal setae 1918(17) Rhagidial organs of tarsus I parallel to each other and to long axis of tarsus (fig. 31,

Qin 1994); whole surface of middle lobe of epirostrum striate-spiculate as onchelicerae and legs Stereotydeus pseudopulcherBasal rhagidial organ of tarsus I oblique (fig. 5d, Strandtmann 1964); basal portionof surface of middle lobe of epirostrum polygonal, not striate-spiculate

Stereotydeus pulcher19(17) Length of palp tibia plus tarsus less than half length of femorogenu (fig. 17, Qin

1994) Stereotydeus brevipalpus

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- Length of palp tibia plus tarsus more than 80% length of femorogenu (fig. 24, Qin1994) Stereotydeus occidentalis

20 (11) Cheliceral shaft enlarged and swollen, and digits large, forming opposed cheliceralshears (fig. 1, Qin 1996); rhagidial organs oblique on tarsus I (fig. 9a, Strandtmann1964) RHAGIDIIDAE ... 21

- Cheliceral shaft not enlarged and swollen, and digits small (e.g. Figs 3,9); rhagidialorgans not oblique on tarsus I (e.g. Figs 5, 6) 24

21 (20) Trichobothria clavate (fig. 61, Strandtmann & Prasse 1977)Coccorhagidia clavifrons

- Trichobothria not clavate 2222 (21) Cheliceral shaft long, its posterior margin arcuate; palp tarsus less than twice as

long as tibia (fig. 44, Womersley & Strandtmann 1963)Poecilophysis macquariensis

- Cheliceral shaft short, its posterior margin straight; palp tarsus twice as long as tibia(fig. 9e, Strandtmann, 1964) 23

23 (22) Famulus on tarsus I located proximal and lateral to basal rhagidial organ (fig. 26D,Zacharda 1980) Rhagidia campbellensis

- Famulus on tarsus 1 located between 2 basal rhagidial organs (fig. 26C, Zacharda1980) Rhagidia mildredae

24 (20) Sejugal furrow present; naso lobe-like, extended anteriorly from prodorsum; genitalshields each with 6 setae EUPODIDAE ... 25

- Sejugal furrow absent; naso dome-like, confined inside anterior prodorsal margin;genital shields each with 7 or more setae PENTHALEIDAE (in part) ... 27

25 (24) Trichobothria clavate (fig. 42, Strandtmann & Prasse 1977)Claveupodes delicatus

- Trichobothria not clavate (fig. lb, Strandtmann 1964) 2626 (25) Femur IV greatly enlarged (fig. la, Strandtmann 1964) Eupodes longisetatus- Femur IV normal, not enlarged (fig. la, Strandtmann 1967) Eupodes minutus27 (24) Internal vertical setae located in a single insertion socket (fig. 5, Qin & Halliday

1996a); tarsus I with 24 or fewer setae; tarsi II-IV each with 17 setaeHalotydeus bakerae

- Internal vertical setae located in separate insertion sockets; tarsus I with at least 32setae; tarsi II-IV each with at least 22 setae 28

28 (27) Dorsal setae short (usually less than 30 |am), plumose; venter with numerous setaebetween legs II and III, aggenital setae 20 or more pairs (fig. 4, Qin & Halliday1996a) Halotydeus destructor

- Dorsal setae long (usually more than 30 (Am), pilose in proximal half only (fig 16,Qin & Halliday 1996a); venter with only one pair of setae between legs II and III,aggenital setae 6 or 8 pairs 29

29 (28) Aggenital setae in 6 pairs; coxal setal formula 3-1-3-3 (fig 17, Qin & Halliday1996a) Halotydeus spectatusAggenital setae in 8 pairs; coxal setal formula 3-1-3-5 (fig 17, Qin & Halliday1996a) Halotydeus castellus

ACKNOWLEDGMENTSI thank my colleagues Jo Berry, Nick Martin,Wim Wouts, and anonymous referees for comments onthe manuscript. This study was supported by the Foundation for Research, Science and Technologyunder contract C09308.

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