taxonID	type	description	language	source
03EE7A4DC902FD0DFF5EFF09FDAEFE85.taxon	materials_examined	Type genus: Tumerozetes Hammer, 1966	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC902FD0AFF5EFE57FB90FF26.taxon	description	Definition. Relatively small, sub-globose non-poronotic brachypyline oribatid mites with the hysterosoma moreor-less as broad as long; with modified plate-like extensions of humeral, notogastral and / or prodorsal region forming convex coverings of the prodorsal surface. Subcapitulum secondarily anarthric with genal tectum divided by medial infrabuccal cleft; labiogenal articulation absent; bases of setae a and h positioned on tectum rather than covered by it. Rutellum bifurcate; chelicerae modified, with digitus fixus bearing lobe-like scale apically; digitus mobilis with comb-like fringe. Prodorsum with well-developed lamellae with cusps. Humeral processes well-developed; either relatively short and projecting anteriomedially or massive, covering lateral prodorsum between dorsosejugal furrow and bothridia. Tutorium and pedotectum I present; pedotectum II present or absent. Well-developed discidium present. Notogaster with 9 - 11 pairs of marginal setae. With well-developed circumpedal carinae. Epimeral plates discrete, separated in mid-line. Genital setae 4 - 5 pairs; aggenital setae 1 pair, anal setae 2 pairs, adanal setae 2 pairs; monodactylous.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC902FD0AFF5EFE57FB90FF26.taxon	discussion	Remarks. The redefinition of the family is required to accommodate Sacculella gen. nov., defined below. Hitherto, the definition of Tumerozetidae was based on that for Tumerozetes by Hammer (1966), as modified by Luxton (1985). Hammer did not define the family, stating only “ Tumerozetes does not belong to any family known so far but forms its own family, the Tumerozetidae. ” Family definitions were given by Piffl (1972) and Woas (2002). Piffl (1972) defined the family as Polypterozetoidea with medial notogastral sculpture extending over the dorsosejugal boundary, without scalps, with 10 pairs of notogastral and 6 or 5 pairs of genital setae, and only 2 pairs of adanal setae. Woas (2002) defined the Tumerozetidae as follows: “ Prodorsum with swollen, plate-like, interlamellar projections dorsal of well-developed lamellae; digitus fixus of chelicera separated from main cheliceral body by distinct step, with hook-like tooth ks (shared with Polypterozetidae). Cuticle with extremely well-developed cerotegument (shared with Polypterozetes, Gymnodamaeoidea and most of the ancient Eupheredermata) covering larger parts of prodorsum; rostral region bill-shaped, distinctly incurved, ending bluntly and turned down sharply at its distal end; tutoria developed; central portion of notogaster with two longitudinal dorsal carinae; humeral region of notogaster with shoulder pieces (reminiscent of Polypterozetes); notogaster encircled by a line of setae (resembling immatures of Polypterozetes), in some species the setae bifurcate; in ventral aspect, acetabular region with smaller triangular processes in place of pedotecta II (shared with Polypterozetidae); epimeral region with four pairs of distinct epimera (shared with Polypterozetidae); genital opening flanked by distinct aggenital ridges (reminiscent of Carabodes); preanal organ tubular-claviform, almost reaching rear border of genital opening (shared with Megeremaeidae); anal opening wider posteriorly than anteriorly and at most as long as genital opening; ventral plate with carina circumpedalis, dorsal branch qp in front of dorsosejugal line of notogaster; bases of rutella fused with mentum, Y-shaped tips distinctly separated from mentum (shared with Polypterozetidae); genu of leg I abruptly calyx-like, broadened distally. ” There are some issues with this definition. Woas (2002) confused the well-developed interlamellar plates (bearing the interlamellar setae) with the lamellae, though they are clearly different and separate when viewed in lateral orientation (Figure 1 c). The description of the chelicera with hook-like tooth ks, as in Polypterozetes cherubin (Figure 4 j) does not accord with the morphology of the chelicera of Tumerozetes roughleyi sp. nov. (Figure 2 d), the only species in the Tumerozetidae for which the chelicera has been described. The cerotegumental covering is unique in that it consists of ovoid-rectangular lumps of a dark, possibly waxy, secretion (Hammer 1966, p. 81 stated “ thick solid strings of wax are attached to the lamellar plates, to the lateral sides of the hysterosoma, etc. ”), but the cerotegument is not necessarily extensive and in T. roughleyi sp. nov. it tends to be mostly confined to the anterior part of the rostrum. The small triangular process referred to ‘ in place of pedotecta II’, and found in Polypterozetes, is the patronium, positioned between pedotecta I and II. This structure is clearly illustrated for Tumerozetes bifurcatus Hammer, 1966 by Norton and Behan-Pelletier (2009, Fig. 15.51 F therein), is present in T. roughleyi sp. nov. (Figures 1 b, 1 c) and probably all other Tumerozetes spp. described by Hammer (1966), though it was not illustrated or mentioned in her descriptions. Finally, the preanal organ is claviform and reaching the posterior margin of the genital plate in T. bifurcatus (Hammer 1966, Figure 110 a therein) but in T. roughleyi sp. nov. it is broad, T-shaped and does not extend as far as the genital plate (Figure 2 g).	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD0AFF5EFE92FC27FE7F.taxon	materials_examined	Type species: Tumerozetes bifurcatus Hammer 1966, p. 81: Luxton 1985, p. 55.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD0AFF5EFE18FDB4FADE.taxon	description	Definition. Tumerozetidae with broad, convex interlamellar plates bearing setae in, in addition to lamellae. With well-developed anteriomedian projection of notogastral plate covering central part of prodorsum and connected with paired longitudinal ridges on notogaster.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD0AFF5EFE18FDB4FADE.taxon	discussion	Remarks. Hammer (1966, p. 81) defined Tumerozetes as follows: propodosoma and hysterosoma fused. Propodosoma swollen. Lateral edges of lamellae with two vertical and medially concave plates fused anteriorly in the midline and bearing setae in. Setae le on apices of long cusps, reaching apex of rostrum. Bothridia pointing laterally; bothridial setae hyaline with finger-like branches on end of long stalk. Pedoctectum I well-developed. A ‘ chitinous figure’ situated at the transition between prodorsum and notogaster, consisting of a short, narrow anterior part and two long, narrow longitudinal ridges on the notogaster. With humeral projections. With 10 pairs of notogastral setae; seven along lateral margins, three on posterior border. With six pairs of genital, one pair of aggenital, two pairs of anal, and two pairs of adanal setae. Legs monodactylous. With thick strings of cerotegument. A qualification of Hammer’s (1966) definition is that the interlamellar plates (the two ‘ vertical and medially concave plates’ she refers to) are not necessarily fused anteriorly and may simply be adjacent or slightly overlapping (Figures 2 a and 2 b). Also, a large, heart-shaped anteriomedian projection of the notogastral plate is present in T. roughleyi sp. nov. (Figure 1 a) and T. bifurcatus (as illustrated by Norton and Behan-Pelletier 2009, Figure 15.51 E therein, labelled ‘ anteriomedial projection’). However, its configuration in the four other Tumerozetes spp. is less clear. This structure was referred to by Hammer (1966, p. 83) as the anterior part of the ‘ chitinous figure at the transition between the propodosoma and the hysterosoma’ (the ‘ chitinous figure’ also includes the paired notogastral ridges). The anteriomedian projection was not mentioned in the family definition by Woas (2002). In T. circularis Hammer 1966 and T. parallelus Hammer 1966 it appears to be a rectangular structure with lateral, hornlike extensions; in T. indistinctus Hammer 1966 it is short and ovoid and in T. pumilis Hammer 1966 it is indistinct, seemingly covered in cerotegument.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD09FF5EFA85FDCFFA8A.taxon	description	(Figures 1, 2) Dimensions. Holotype female length 309 μm, breadth 205 μm. Paratype males: mean length (n = 4) 256 μm (range 239 – 283 μm); mean breadth 165 μm (range 154 – 183 μm). Paratype females: mean length (n = 10) 307 μm (range 296 – 321 μm); mean breadth 200 μm (range 194 – 211 μm). Ratio of prodorsum to total length: 0.47 (holotype). Adult. Prodorsum: rostrum very broad, rounded, with margin covered by blocks of cerotegument (Figure 2 c). Rostral setae (ro) short, smooth; barely visible in dorsal view. Lamellae (la) long, thin, narrow; lamellar setae (le) well-developed, barbed, on parallel lamellar cusps extending anteriorly as far as apex of rostrum (Figure 1 a). Lamella (la) connecting anteriorly with tutorium (tu) in anterior view (Figure 2 a). Prodorsum inflated, with broad, lateral ridge-like interlamellar plates (ilp) extending from base of lamellae almost to apex of rostrum, bearing smooth, curved interlamellar setae (in). Setae in projecting vertically from interlamellar plates in anterior view (Figure 2 a). Interlamellar plates sub-rectangular, slightly longer than broad, fused posteriomedially (Figure 2 b). Apices of interlamellar plates not fused anteriorly in dorsal orientation. Heart-shaped anteriomedian projection of notogastral plate (ap) with fine alveolate ornamentation and median ridges covering central part of prodorsum, positioned below and not fused with interlamellar plates, extending from anterior of in, converging toward dorsosejugal furrow. Anteriomedian projection of notogastral plate strongly convex in lateral view. Bothridia prominent, projecting laterally, with well-developed rim and internal circular ridges (Figure 2 h). Bothridial seta long (65 μm; holotype), setiform, slightly inflated apically and with fine, long spinules. Short, smooth exobothridial seta (ex) present. Notogaster: margins of relatively small humeral processes (h) convex laterally, curving ventrally and with median protuberance; humeral processes not extending as far as bothridia (Figure 1 a). Seta c 2 positioned immediately posterior of humeral process on small, rounded projection. Dorsosejugal furrow (ds) straight, transverse, with paired, curved, longitudinal median notogastral ridges (mnr) forming ovoid shape, extending posteriorly as far as level of lyrifissure iad, fused anteriorly, connecting at dorsosejugal furrow with anteriomedian projection of notogastral plate. Notogaster smooth, broader (215 μm; holotype) than long, with 9 pairs of marginal setae (c 1 plus la, lp, h and p series); la, lp and h 3 with fine spines basally. Lyrifissurae ia present. Ventral aspect: subcapitulum rounded apically, with tectum derived from genae, bearing a median infrabuccal cleft (ibc) on its ventral surface (Figure 7 e); setae a and m with bases not hidden by tectum in ventral orientation; setae h positioned posteriomedially on subcapitulum. Labiogenal articulation absent. Rutellum Y-shaped (Figure 2 f). Chelicerae short, with digitus fixus reduced to flat, smooth, lobe-like structure; digitus mobilis curved, projecting beyond digitus fixus, bearing a comb-like fringe of several thin, straight teeth (Figure 2 d). Palp setal formula 0 - 2 - 1 - 3 - 9; setae on palpfemur, genu and seta d on tibia with sparingly spinose ornamentation, others smooth (Figure 2 e). Eupathidia not on prominent tubercles; eupathidium acm not fused with solendidum ω which is positioned posteriorly on palptarsus. Epimeral plates discrete, all broadly separated in midline, sub-rectangular (I-II) or sub-triangular (III-IV); epimeral setation 3 - 1 - 3 - 3 (Figure 1 b). Pedotectum I (pd I) and II (pd II) and patronium (pa) strongly developed; humeral processes appearing lobe-like in ventral orientation; discidium (di) with prominent, sharp point; welldeveloped circumpedal carina (cc) present (Figure 1 b). Ventral plate sub-circular, broader than long. Genital and anal plates each surrounded by ring of sclerotized cuticle; genital plates 46 µm long (holotype), with 5 pairs of setae; 1 pair of aggenital setae, 2 pairs of adanal setae and 2 pairs of anal setae; lyrifissure iad in para-anal position (Figure 2 g). Anal plates lozenge-shaped, 55 µm long (holotype); only slightly separated from genital plates. Preanal organ (po) broad, flat T-shaped. Lateral aspect: anterior prodorsum strongly concave. Tutorium (tu) pointed apically, similar shape as lamella, but without cusp; pedoctectum I (pd I) with sharp, well-developed cusp; pedotectum II (pd II) well-developed with blunt apex; discidium (di) prominent, pointed. Interlamellar plates convex, strongly inflated; seta in pointed vertically. Anteriomedian projection of notogastral plate convex, strongly inflated. Notogastral setae positioned on faint horizontal lath of more heavily sclerotized cuticle that connects anteriorly with humeral process. Legs: monodactylous; claw lacking dorsal tooth.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD09FF5EFA85FDCFFA8A.taxon	materials_examined	Type designation, material examined and locality data. Holotype, wet moss, Cumberland Creek Valley, Marysville-Woods Point Road, nr. Cambarville, Yarra Ranges National Park, Victoria, 37 º 33 ’ S 145 º 53 ’ E, coll. R. E. Roughley, 12. iii. 1996. Paratypes: four females, one male, ANIC 297, Moss and litter, rainforest with Nothofagus cunninghami, Cumberland Creek Valley, near picnic area [Cumberland Memorial Reserve], Yarra Ranges National Park, Victoria. 37 ° 34 ’ S 145 ° 52 ’ E, 920 m., coll. R. W. Taylor and R. J. Bartell, 4. xi. 1970. Paratypes: five females, moss (Dicranoloma billiardieri), Nothofagus cunninghami forest, The Beeches, Lady Talbot Drive near Warburton, Yarra Ranges National Park, Victoria, 37 ° 29 ’ S, 145 ° 50 ’ E, 800 m., coll. G. Perdomo, May, 2009. Paratypes: 22 males, 49 females, moss on Sassafras (Atherosperma moschatum), cool temperate rainforest, 1077 m., Errinundra Saddle, Errinundra National Park, Victoria, 37 ° 19 ’ S 148 ° 51 ’ E, 1030 m., coll. M. J. Colloff, 29. ix. 2009. Paratype: male, moss, Nothofagus cunninghami rainforest, Melba Gully, Otway National Park, Victoria, coll. M. J. Colloff, 20. vii. 2011. Paratypes: 12 males, 24 females, moss, Nothofagus cunninghami rainforest, rainforest walk, Maits Rest, Otway National Park, Victoria, coll. M. J. Colloff, 19. vii. 2011.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD09FF5EFA85FDCFFA8A.taxon	etymology	Etymology. This species is named in honour and memory of the collector of the holotype specimen, Robert E. Roughley.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD09FF5EFA85FDCFFA8A.taxon	diagnosis	Diagnosis. Tumerozetes roughleyi sp. nov. can be distinguished from all other species in the genus by the following combination of characters: (1) the paired, curved, longitudinal median notogastral ridges forming an ovoid structure and extending posteriorly as far as level of lyrifissure iad: (2) setae la, lp and h 3 with fine spines basally; (3) with heart-shaped anteriomedian projection of the notogastral plate; (4) the presence of a patronium; (5) the dorsosejugal furrow coinciding with the junction between the longitudinal notogastral ridges and the anteriomedian projection of the notogastral plate and positioned markedly posterior of the level of the bothridia.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC905FD09FF5EFA85FDCFFA8A.taxon	discussion	Remarks. Tumerozetes roughleyi sp. nov. is morphologically most similar to T. pumilis in the shape of the median notogastral ridges and to T. bifurcatus in the shape of the anteriomedian projection of the notogastral plate. However, in T. pumilis each notogastral ridge is considerably narrower and they converge posteriorly at a position level with setae h 3. Tumerozetes bifurcatus Hammer, 1966 and T. circularis Hammer, 1966 both have short, parallel longitudinal dorsal ridges set some distance apart and extending posteriorly only as far as a position level with setae la. Also, T. bifurcatus has bifurcate notogastral setae. Tumerozetes indistinctus Hammer, 1966 and T. parallelus Hammer, 1966 both have very long, parallel longitudinal dorsal ridges, almost contiguous and extending posteriorly as far as a position level with setae h 3. Other than dimensions, there were no obvious external morphological differences between males and females.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC906FD06FF5EFA70FE83FD5A.taxon	description	(Figures 3, 4)	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC906FD06FF5EFA70FE83FD5A.taxon	type_taxon	Type species: Sacculella yarra sp. nov.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC906FD06FF5EFA70FE83FD5A.taxon	description	Definition. Adults. Relatively small, sub-globose, non-poronotic brachypyline oribatid mites. Rostrum covered by broad lamellae diverging posteriolaterally; with lamellar setae positioned ventrally on lamellae. Subcapitulum secondarily anarthric; apex of rutellum Y-shaped. Digitus fixus of chelicera a lobe-like structure; digitus mobilis comb-like. Rostrum truncated, anterior margin vertical in lateral view (Figure 3 a). Bothridium well-developed, with peripheral and internal sclerotized thickening; bothridial setae elongated, setiform. Notogaster broader than long, smooth, without scalps; with 11 pairs of short, marginal notogastral setae; large, inflated, sub-trapezoid humeral processes. Humeral processes with V-shaped dorsal keels and extending as far as posterior margin of bothridia. Dorsosejugal furrow straight, transverse. Epimeral plates well-defined, sub-rectangular, separate in medial line. Pedotectum I and II, patronium and discidium well-developed. Circumpedal carina present. Genital setae 4 pairs, aggenital setae 1 pair, anal setae 2 pairs, adanal setae 2 pairs; anal plates lozenge-shaped; pre-anal organ T-shaped. Legs monodactylous, claw lacking dorsal tooth.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC906FD06FF5EFA70FE83FD5A.taxon	diagnosis	Diagnosis. Adults of this genus are unique among the non-poronotic Brachypylina and the Polypterozetoidea in having the following combination of character states: subcapitulum secondarily anarthric; rostrum covered by broad lamellae; rostrum truncated, vertical in lateral aspect; lamellar seta on ventral surface of lamella; with well-developed pointed discidium and circumpedal carina; notogaster broader than long, smooth, with 11 pairs of marginal notogastral setae; humeral processes large, inflated, sub-trapezoid, bearing setae c 2; 4 pairs of genital setae and 2 pairs of adanal setae.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC906FD06FF5EFA70FE83FD5A.taxon	discussion	Remarks. Sacculella gen. nov. is included in the Tumerozetidae on the basis of a set of shared character states with Tumerozetes: the secondarily anarthric subcapitulum, with a tectum derived from the genae and with a median infrabuccal cleft and the bases of setae a and h not hidden by the tectum in ventral orientation; the digitus fixus of the chelicera terminating in a lobe-like structure and the digitus mobilis with a comb-like margin; the presence of a circumpedal carina; two pairs of adanal setae; the sub-globose hysterosoma, more-or-less as broad as long; with 9 pairs of marginal notogastral setae.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC906FD06FF5EFA70FE83FD5A.taxon	etymology	Etymology. The generic prefix saccul - is derived from the Latin sacculus (a small bag or sac) and refers to the inflated humeral regions of the type species. The suffix - ella is the Latin diminutive, referring to the small size of the species.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC909FD05FF5EFD00FB77FDE2.taxon	description	(Figures 3, 4) Dimensions. Holotype male length 302 μm, breadth 208 μm. Paratypes: male: length 300 μm, breadth 217; females (n = 3) 327 μm (range 323 – 330 μm); mean breadth 244 μm (range 236 – 252 μm). Ratio of prodorsum to total length: 0.58 (holotype). Adult. Prodorsum: rostrum broad in dorsal view, almost covered by broad, apically rounded lamellae, extending from apex of rostrum to bothridia and diverging posteriolaterally and with a faint, short ventral translamella (Figure 3 a). With short, smooth rostral setae (ro), only visible in lateral view (Figure 3 c). Lamellar setae (le) smooth, emerging from ventral surface of lamellae, only visible in ventral view (Figure 3 b). Interlamellar seta (in) welldeveloped, positioned anteriomedially of bothridium. Bothridium well-developed, cylindrical, with sclerotized rim and internal circular thickening. Bothridial setae elongated, setiform, with faint spinose ornamentation; extending posteriorly as far as bases of setae c 1. Lateral region of prodorsum posterior of bothridium strongly concave. Notogaster: Margins of humeral processes (h) convex laterally, then slightly concave anteriorly, extending as far as bothridia, medially with thin cuticle with polygonal alveolate microsculpture and dorsal V-shaped keels, forming sacculate structures covering lateral parts of prodorsum (Figure 3 a). Notogaster smooth, U-shaped, broader (216 μm; holotype) than long; with 11 pairs of marginal setae (c 1, c 2 plus l, h and p series); l, h and p series ornamented with fine spines basally and borne on faint lateral region of more heavily sclerotized cuticle. Setae c 1 and c 2 smooth, well developed, positioned laterally and adjacent on humeral process. Dorsosejugal furrow transverse, heavily sclerotized, overhanging prodorsum. Lyrifissurae ia present. Ventral aspect: apex of subcapitulum in form of broad, inverted V, with tectum derived from genae, bearing a median infrabuccal cleft (ibc) on its ventral surface (Figure 8 f); short, setiform setae a and m with bases not hidden by tectum in ventral orientation; setae h short, spiniform, positioned anteriolaterally, immediately posterior of setae m. Labiogenal articulation absent. Rutellum Y-shaped (Figure 3 c). Chelicerae short, with digitus fixus reduced to flat, smooth, lobe-like scale, with tooth kk present; digitus mobilis curved, almost as long as digitus fixus, bearing a comb-like fringe of several thin teeth (Figures 4 b, 4 c). Palp setal formula 0 - 2 - 1 - 3 - 8; all setae smooth. Eupathidia not on prominent tubercles; eupathidium acm not fused with solendidum ω which is positioned posteriorly on palptarsus (Figure 4 a). Tutorium with 4 - 5 short spines, sub-tutorium with cup-shaped invagination and acute point (Figure 4 d). Epimeral plates discrete, all broadly separated in midline and from each other, sub-rectangular; epimeral setation 2 - 1 - 2 - 3 (Figure 3 b). Pedotectum I and II relatively weakly developed; patronium present; discidium (di) with prominent, sharp point; well-developed circumpedal carina (cc) present. Ventral plate sub-circular, broader than long. Genital and anal plates each surrounded by ring of sclerotized cuticle; genital plates 36 µm long, with 4 pairs of setae; 1 pair of aggenital setae positioned posteriorly level with preanal organ (po), 2 pairs of adanal setae and 2 pairs of anal setae; lyrifissure iad in para-anal position (Figure 3 b). Anal plates lozenge-shaped, 53 µm long; only slightly separated from genital plates. Preanal organ broad, flat T-shaped. Lateral aspect: Rostrum straight, vertical, almost square; tutorium pointed apically; pedoctectum I and II rounded apically; discidium prominent, pointed (Figures 3 b, 3 c). Lamellae projecting as far as dorsal apex of rostrum, markedly elevated immediately anterior of bothridia. Interlamellar seta (in) projecting vertically. With a small, blunt, triangular lobe (bl) posterior of bothridium. Humeral processes covering prodorsum between bothridium and dorsosejugal furrow, posterior prodorsum strongly invaginated and overhung by anterior notogaster. Notogaster markedly deeper than long. Legs: monodactylous; claw lacking dorsal tooth.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC909FD05FF5EFD00FB77FDE2.taxon	materials_examined	Type designation, material examined and locality data. Holotype male, paratype female, wet moss, Cumberland Creek Valley, Marysville-Woods Point Rd, near Cambarville, Yarra Ranges National Park, Victoria, 37 º 33 ’ S 145 º 53 ’ E, coll. R. E. Roughley, 12. iii. 1996. Paratypes: one female, one male, ANIC 297, moss and litter, rainforest with Nothofagus cunninghami, near picnic area at Cumberland Memorial Reserve, Cumberland Creek Valley, Yarra Ranges National Park, Victoria. 37 ° 34 ’ S 145 ° 52 ’ E, 920 m. Coll. R. W. Taylor and R. J. Bartell, 4. xi. 1970.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC909FD05FF5EFD00FB77FDE2.taxon	etymology	Etymology. This species is named for its type locality in the Yarra Ranges, Victoria.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC909FD05FF5EFD00FB77FDE2.taxon	diagnosis	Diagnosis. Sacculella yarra sp. nov. can be distinguished from all other species in the genus by the following combination of characters: (1) the lamellae broad, converging anteriomedially, with setae le positioned ventrally; (2) the bothridia prominent, internally ridged; bothridial setae elongated, setiform, smooth; (3) the humeral processes large, convex, covering lateral regions of prodorsum; (4) the presence of a patronium; (5) the rounded notogaster, considerably broader than long, with 11 pairs of marginal setae.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC909FD05FF5EFD00FB77FDE2.taxon	discussion	Remarks. The shared character states between Sacculella yarra gen. et sp. nov. and Tumerozetes were outlined above in the remarks section on the description of Sacculella gen. nov. However, Sacculella yarra sp. nov. lacks the interlamellar plates, the paired median longitudinal notogastral ridges and the anteriomedian projection of the notogastral plate of Tumerozetes spp.; it has 4, rather than 5 - 6 pairs of genital setae; broad lamellae with ventral lamellar setae, rather than narrow pointed ones with the lamellar setae apical; the humeral process are in the form of massive, convex plates, rather than small curved projections; the tutorium is serrated, rather than smooth; pedotectum I is weakly developed with a rounded apex, rather than well developed and pointed; the palptarsus has 8 setae, rather than 9 and the epimeral setal formula is a bi-deficient 2 - 1 - 2 - 3, rather than the holotrichous 3 - 1 - 3 - 3. Other than dimensions, there were no obvious external morphological differences between males and females.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90AFD05FF5EFD58FDA4FCD5.taxon	materials_examined	Type genus: Nodocepheus Hammer, 1958	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90AFD05FF5EFBD4FAC7F836.taxon	description	Definition. Polypterozetoid mites with subcapitulum secondarily anarthric (Figure 5 b) or diarthric, though the labiogenal articulation may be incomplete (Figure 7 d); rutellum bifurcate, apex Y-shaped (Figure 4 g); chelicerae modified: either with digitus fixus pointed, lacking teeth and digitus mobilis comb-like (Figure 4 e) or both digits with reduced, blunted teeth, offset from each other (Figure 4 l). With serrated tutorium and cusped sub-tutorium. With short, rounded single-lobed or bilobed humeral processes. With 11 pairs of smooth notogastral setae positioned marginally; c 1 and c 2 marginal, on or near humeral process, sometimes on ventral fold. With five pairs of genital setae. Epimeral setal formula 3 - 1 - 3 - 3. Genital setae 6 pairs; aggenital setae 1 pair, anal setae 2 pairs, adanal setae 3 pairs. Legs monodactylous.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90AFD05FF5EFBD4FAC7F836.taxon	discussion	Remarks. The above definition is based on those of Piffl (1972) and Woas (2002). Mahunka (1983) provided a key to the six species of Nodocepheus known at that time, stressing the morphology of the humeral lobe, bothridial seta, tutorium and sub-tutorium. The presence of Trägårdh’s organ is based on the examination by Piffl (1972) of a single specimen of Nodocepheus dentatus Hammer 1958 in which the chelicerae were damaged. The structure is described as short, broad and thus lacks the characteristic elongated form of this structure. I could not identify a Trägårdh’s organ on the chelicerae Nodocepheus luxtoni sp. nov. In Nodocepheus dentatus, N. laterodentatus Piffl, 1972 and N. luxtoni sp. nov., setae c l are shifted laterally to the humeral region, positioned on a fold of the humeral process adjacent to setae c 2 (Piffl 1972, Plate 13 therein; Figure 5 a herein). It is likely that N. barbatus Hammer, 1966 (originally described as N. dentatus var. barbatus) shares this character state, giving a likely notogastral setal complement of 11 pairs (cf. Piffl 1972, p. 292), rather than 9 pairs as illustrated by Hammer (1966, Figure 85 therein). In N. minimus Mahunka, 1985, setae c 1 appear to be absent and c 2 are apparently reduced to their alveoli, giving a complement of 10 pairs (including c 2). In N. cerebralis Mahunka, 1980 and N. hammerae Balogh, 1961 neither of the c series are illustrated, yet in the figures of the humeral regions of N. baloghi Mahunka, 1983, N. cerebralis, N. dentatus and N. hammerae by Mahunka (1983, Figures 47 - 53 therein), c 1 is present on the ventral surface of the median lobe of the humeral process, as illustrated for N. dentatus by Piffl (1972, Plate 13, Figure Nd 3 therein). It seems likely that c 2 is also present in N. cerebralis, N. hammerae, and N. minimus on the anterioventral surface of the rugose lateral lobe of the humeral process, but has been overlooked when viewed dorsally because of its position below the heavily textured surface of the lobe.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90AFD05FF5EFC8DFC80FC0B.taxon	materials_examined	Type species: Nodocepheus dentatus Hammer, 1958, p. 65, fig. 75.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90DFD03FF5EFF41FC7FF823.taxon	description	(Figures 4, 5) Dimensions. Holotype length 204 μm, breadth 117 μm. Paratype male: length 192 μm, breadth 113 μm. Paratype females: mean length (n = 11) 211 μm (range 199 – 221 μm); mean breadth 125 μm (range 113 – 132 μm). Ratio of prodorsum to total length: 0.41 (holotype). Adult. Prodorsum: rostrum bluntly point in dorsal view, with short rostral setae on short tubercles, linked by short, transverse ridge (Figure 4 i). Broad lamellae extending to just posterior of rostrum, with strongly incurved apical cusps at right angles to the rest of the lamellae, bearing short, smooth, setiform lamellar setae (le) and with well-developed translamella (Figure 5 a). Short, smooth interlamellar setae (in) positioned on centrodorsal region. Bothridium well-developed, cylindrical, with broad sclerotized rim and internal circular thickening. Bothridial setae elongated, club-shaped, with strong spinose ornamentation apically. Notogaster: humeral processes (h) well-developed, abutting bothridia, with large, elongated lateral lobe bearing short, setiform seta c 1 and smaller medial lobe bearing seta c 2; lateral lobe extending to point posterior of base of seta la (Figure 5 a). Humeral processes smooth, not ornamented or sculpted. Notogaster smooth, ovoid, longer (216 μm; holotype) than broad; with 11 pairs of smooth, setiform setae positioned marginally (c 1 and c 2 plus l, m and p series); l and h series on short, squat tubercles. Dorsosejugal furrow transverse; with faint, incurved ridges on anteriomedian region of notogaster. Ventral aspect: subcapitulum secondarily anarthric, with tectum derived from genae, bearing median infrabuccal cleft (ibc) on its ventral surface (Figure 5 b); short, setiform setae a and m with bases not hidden by tectum in ventral orientation; setae h short, spiniform, positioned just posteriomedially of setae m. Labiogenal articulation absent. Rutellum Y-shaped (Figure 4 g). Chelicerae with digitus fixus pointed, smooth; digitus mobilis comb-like with sparse, fine teeth (Figure 4 e). Palp setal formula 0 - 2 - 1 - 3 - 8; all setae smooth. Eupathidia not on prominent tubercles; eupathidium acm not fused with solendidum ω which is positioned posteriorly on palptarsus (Figure 4 f). Tutorium well-developed, with a long apical tooth and 4 short teeth; sub-tutorium with blunt cusp (Figure 4 h). Epimeral plates I fused in mid-line, others discrete, broadly separated in midline and longitudinally; plates I and II sub-rectangular, III and IV oval; epimeral setation 3 - 1 - 3 - 2 (Figure 5 b). Pedotectum I and II large, prominent, with pointed anterior and posterior lobes; discidium (di) broad, well-developed, pointed. Perigenital carina present, bearing enantiophysis E 4. Ventral plate ovoid, markedly longer than broad. Genital and anal plates each surrounded by ring of sclerotized cuticle; genital plates 33 µm long (holotype), with 5 pairs of setae; 1 pair of aggenital setae positioned just posterior of genital plates; 3 pairs of adanal setae, 2 pairs of anal setae; lyrifissure iad in para-anal position (Figure 5 b). Anal plates lozenge-shaped, 41 µm long (holotype); separated some distance from genital plates. Preanal organ broad, flat T-shaped. Lateral aspect: Rostrum straight, vertical; tutorium (tu) and sub-tutorium (stu) each with acute point, tu strongly serrated dorsally; pedoctectum I and II pointed rounded apically; discidium leaf-shaped, bluntly pointed (Figure 5 c). Lamellae projecting as far as dorsal apex of rostrum. Exobothridial seta (ex) present. Anterior notogastral margin overhanging prodorsum. Legs: monodactylous; claw with a dorsal tooth.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90DFD03FF5EFF41FC7FF823.taxon	materials_examined	Type designation, material Examined and Locality Data. Holotype female and five paratype females, moss on Sassafras (Atherosperma moschatum), cool temperate rainforest, 1077 m., Errinundra Saddle, Errinundra National Park, Victoria, 37 ° 19 ’ S 148 ° 51 ’ E, 1030 m., coll. M. J. Colloff, 29. ix. 2009. Paratype: female, ANIC 3757, sifted litter and mossy logs, rainforest with Eucalyptus sp., State Forest, 2.2 km north-east of Corinna, Tasmania, 41 ° 39 ’ S 145 ° 06 ’ E., 45 m., coll. T. Weir and C. Lemann, 14. iii. 2008. Paratype: female, TAS- 183 Litter, eucalypt forest, Old Farm Road, Mount Wellington, Tasmania, 42 ° 54 ’ S 147 ° 14 ’ E, coll. P. Greenslade, 20. vi. 1989. Paratypes: four females, moss, Nothofagus cunninghami rainforest, rainforest walk, Maits Rest, Otway National Park, Victoria, coll. M. J. Colloff, 19. vii. 2011.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90DFD03FF5EFF41FC7FF823.taxon	etymology	Etymology. This species is named in honour and memory of my friend and colleague, the late Dr Malcolm Luxton, in recognition of his contribution to our understanding of the taxonomy and biogeography of the oribatid mites of the Australasian region.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90DFD03FF5EFF41FC7FF823.taxon	diagnosis	Diagnosis. Nodocepheus luxtoni sp. nov. can be distinguished from all other species in the genus by the following combination of characters: (1) most of the notogastral setae are on squat tubercles; (2) with the epimeral setal formula 3 - 1 - 3 - 2; (3) the very large, pedotecta I and II with an indented apex, appearing shallowly bifurcate; (4) the prominent, smooth, elongated lateral lobe of the humeral process, extending to a point posterior of base of seta la; (5) the broad, discidium, bluntly pointed; (6) the rostral setae on short tubercles, linked by a short, transverse ridge; (7) the strongly incurved and opposing lamellar cusps (8) the smooth, short setae le positioned sub-apically on the lamellar cusps; (8) the bothridial seta with a long stalk, the head club-shaped and covered in long, well-developed spinules; (9) the tutorium with a long apical tooth and 4 short teeth and the sub-tutorium with a blunt cusp.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
03EE7A4DC90DFD03FF5EFF41FC7FF823.taxon	discussion	Remarks. Mahunka (1983, p. 164) stated that of the six species described by that time, all of them except N cerebralis “ stand very near to each other ” and that, based on an examination of types, they could only be distinguished by the morphology of the humeral processes, tutorium and sub-tutorium, with chaetotaxy and the morphology of the sensillus considered as secondary characters. Nodocepheus luxtoni sp. nov. belongs to a group with N. baloghi and N. minimus that share the following character states: (1) the bothridial seta has a long stalk and a club-shaped head densely ornamented with well-developed spines and (2) the humeral process is large, expanded and extends from the bothridium to a position at least as far posteriorly as the level of the base of setae la. However, N. baloghi and N. minimus both have setae le with spinose ornamentation, rather than smooth, and are positioned apically on broad lamellar cusps, rather than sub-apically on narrow, strongly incurved cusps.	en	Colloff, Matthew J. (2022): First records of Tumerozetidae and Nodocepheidae from Australia, with descriptions of new taxa and a re-assessment of the Polypterozetoidea (Oribatida, Brachypylina). Zootaxa 5194 (1): 33-57, DOI: 10.11646/zootaxa.5194.1.2
