New species of Agathodesmus Silvestri, 1910 from Australia (Diplopoda, Polydesmida, Haplodesmidae)

Abstract The genus Agathodesmus Silvestri, 1910 is speciose and widespread in high-rainfall parts of eastern Australia. In addition to the type species Agathodesmus steeli Silvestri, 1910 and Agathodesmus johnsi Mesibov, 2009 from New South Wales and Agathodesmus bucculentus (Jeekel, 1986) from Queensland, the following 18 new species are recognised: Agathodesmus adelphus sp. n., Agathodesmus aenigmaticus sp. n., Agathodesmus agnus sp. n., Agathodesmus anici sp. n., Agathodesmus gayundah sp. n., Agathodesmus hahnensis sp. n., Agathodesmus kerensis sp. n., Agathodesmus kirrama sp. n., Agathodesmus millaa sp. n., Agathodesmus parapholeus sp. n., Agathodesmus quintanus sp. n., Agathodesmus sagma sp. n., Agathodesmus summus sp. n. and Agathodesmus yuccabinensis sp. n. from Queensland; Agathodesmus carorum sp. n. from New South Wales and Victoria; Agathodesmus bonang sp. n. and Agathodesmus morwellensis sp. n. from Victoria; and Agathodesmus chandleri sp. n. from South Australia.


Introduction
Agathodesmus Silvestri, 1910 was established for a single female of A. steeli Silvestri, 1910, a small polydesmidan millipede from New South Wales, Australia. In a recent paper (Mesibov 2009) I redescribed A. steeli from freshly collected males and females, and added a second New South Wales species to the genus. The two species share with Atopogonus baccatus Carl, 1926 (New Caledonia) and A. bucculentus Jeekel, 1986 (Queensland, Australia) a distinctive gonopod conformation: there is no cannula or prostatic groove, and the telopodite is sharply bent mid-length at a 'knee' topped with a thin tab. In Mesibov (2009) I made Atopogonus Carl, 1926 a junior synonym of Agathodesmus.
In this paper I describe 18 new species of Agathodesmus and add a few observations on A. bucculentus. I collected specimens of four of the new species during field trips from 2006 to 2011. The remaining 14 species were found in the Australian National Insect Collection and the Queensland Museum, mainly among arthropods extracted from rainforest litter by the Berlese method.
The late C.A.W. Jeekel wrote of A. bucculentus: "With the discovery of a species of Atopogonus in what seems to be a perfectly natural habitat in Queensland it becomes likely that the genus is in essence a continental Australian taxon" (Jeekel 1986, p. 46). Agathodesmus is now known to be widespread and speciose in parts of eastern Australia with a mean annual rainfall greater than 1000 mm (Fig. 1). Agathodesmus is particularly diverse in the Wet Tropics of far north Queensland, which is home to 12 of the 21 known Australian species. The wide gaps in the genus distribution map suggest to me that more species remain to be discovered, especially in the wetter mountain forests of New South Wales, southeast Queensland and Victoria.

Methods
'Male' and 'female' in the text refer to adult individuals. All specimens are stored in 75-80% ethanol in their respective repositories.
Gonopods were cleared in 80% lactic acid and temporarily mounted in 60% lactic acid for optical microscopy, while other body parts were temporarily mounted in a 1:1 glycerol:water mixture. Preliminary gonopod drawings were traced from photomicrographs taken at 160× through a binocular microscope. Measurements were made with a Nikon SMZ800 binocular dissecting microscope using an eyepiece scale, and are reported below to the nearest 0.5 mm. SEM images were acquired digitally using a FEI Quanta 600; some specimens were examined after air-drying and sputter-coating with platinum, while others were air-dried, examined and returned to alcohol. Images and drawings were prepared for publication using GIMP 2.8. Maps were generated using ArcView GIS 3.2.
The Appendix tabulates specimen data for all museum lots noted in the text. Locality details are given in all cases with latitude and longitude based on the WGS84 datum. Latitude/longitude data for Queensland Museum specimen localities are mainly taken from the Queensland Museum insect collection database, but some Mt Bellenden Ker data have been corrected following Mesibov (2012b). My estimate of the uncertainty for each locality is the radius of a circle around the stated position, in metres or kilometres.
Remarks. The diagnosis above slightly amends the one given in Mesibov (2009). Agathodesmus as a genus is easily recognised by the distinctive structure of the gonopod telopodite (Fig. 2). The proximal portion (pp) is typically straight and usually more or less cylindrical, but with the medial and posterior surfaces flattened. The pp arises from the distomedial corner of the small, oblong gonocoxa, where its base is partly contained in a small concavity. The telopodite base may extend basally as a short, rounded projection (be) to overlap the apex of the gonocoxa in ventral or posterior view. The apex of the pp extends distally as a thin tab (at), and on or just below the tab on the posterior surface there are three long, closely adjacent, apical setae (as) in a row; scattered smaller setae may be present on the posterior surface of the pp. Arising just basal to the as on the posterodistal surface of the pp is the distal portion (dp) of the telopodite. Jeekel (1986) used the word 'complicated' three times in his description of the dp ('acropodite') in A. bucculentus, and while the details of its structure can be very hard to put into words, some generalities are clear and are applicable to all known Australian Agathodesmus species. The dp is always directed posterobasally or laterobasally, giving the telopodite as a whole the appearance of a leg tightly bent at the knee. The main branch (mab) of the dp is flattened into a lamella and is usually divided into lobes. The lamella is typically curved so that the surface seen in posterior view is slightly convex, and the distal margin of the mab and portions of its inner, concave surface may be thickened or folded. A smaller, medial branch (meb) of the dp arises near the base of the dp on its medial side and usually curves laterally so that its tip is hidden behind the mab in ventral view. Portions of the meb are sometimes densely covered with fine, hair-like structures.
In this paper I provide posterior or posterolateral gonopod views of all species described here. These views are convenient when examining males with tightly flexed telopodites, and with careful attention to shape and position, a posterior or posterolateral view is diagnostic. However, readers should be aware that important diagnostic details of the telopodites may be hidden in these views, and that in the SEM views published here, thin portions of the mab may have been distorted by drying. Agathodesmus spp.
are unusual among Australian Polydesmida in their 'hidden' gonopod complexity, and manipulation of the telopodite to partially unbend it and separate the meb is often necessary for a positive identification.
When an Agathodesmus sp. gonopod is cleared with 80% lactic acid, the telopodite sometimes extends so that the angle between the pp and dp is greater than 90°. It is easy to imagine that the telopodite extends in this way during mating, and that the apical tab (at) serves as a check on the rotation of the telopodite 'knee'. A tab in a similar location is present anterior to the 'joint' in the pseudo-articulated gonopod telopodite of species in the Australian genus Ginglymodesmus (Dalodesmidae) (Mesibov 2005). However, I have as yet no evidence to support the idea that the telopodite extends in living males. No Agathodesmus have been collected and preserved in copula, and in all of the 430+ Agathodesmus males I have examined the uncleared gonopod telopodites are flexed.
Variation in most non-gonopod characters across the genus is minor and the redescription of the type species A. steeli in Mesibov (2009) applies to most details in the new species. The 'diagnostic descriptions' given below include only those characters known to vary significantly among Australian Agathodesmus spp. Generalised gonopods of Agathodesmus spp., posterior view. as = apical setae, at = apical tab, be = basal extension of telopodite, dp = distal portion of telopodite, mab = main branch of distal portion of telopodite, meb = median branch of distal portion of telopodite, pp = proximal portion of telopodite.
In 12 of the 21 known Australian species, males and females have 19 body rings, in five species 20 rings, and in one species males have 19 rings and females 20; three species known only from males have 19, 19 and 20 rings. The smaller species generally have 19 rings and the larger 20 rings, but the correlation of ring number and body size is loose, and two of the largest Australian species have 19 rings (A. kirrama sp. n. and A. yuccabinensis sp. n.).   (Jeekel, 1986), short bipartite seta with slightly flared tip (male ex ANIC 64-000332) B A. quintanus sp. n., very long bipartite setae with slightly flared tips (male paratype ex QM S96066). Scale bars: A = 0.02 mm, B = 0.2 mm.
There is also no apparent relationship between body size and the development (or absence) of the 'pseudo-paranota' formed by lateral metatergal tubercles on posterior rings (Fig. 3).
Although the metatergal setae in most Australian Agathodesmus spp. are short with slightly flared tips (Fig. 4A), three of the newly described species have very long setae (Fig. 4B) and appear 'hairy' at low magnification.
While examining the new Agathodesmus species I was able to identify what appear to be spiracular openings, something I failed to do earlier (Mesibov 2009). The openings are minute (Fig. 5) and on diplosegments are both located close to the anterior leg base.
Finally, included here as Fig. 10D   Diagnostic description. Male and female with head + 20 rings. Colour in alcohol very faintly reddish. Male/female ca 8.0/8.5 mm long; ring 12 maximum diameter ca 0.6/0.7 mm, maximum width ca 0.85/0.9 mm. Metatergal tubercles in 8-10 irregular transverse rows, mostly without setae; metatergal setae short with slightly flared tips; lateralmost two rows of tubercles not enlarged, together forming narrow pseudo-paranotum with 6 marginal tubercles. Male leg 6 coxa with small, rounded, mediodistal projection. Telopodite ( Fig. 6A) with pp straight; at in longitudinal plane, short and rounded-triangular; dp directed laterobasally at base; mab concave medially, deeply divided with smaller medial lobe bent posteriorly, tapering and with sharp subterminal tooth on lateral surface; larger, lateral mab lobe widening to thickened, emarginate apex; meb curving strongly behind mab, divided at ca one-third length into 2 thin processes.
Distribution. Known only from rainforest high on Mt Bartle Frere in tropical north Queensland (Fig. 13A). Possibly co-occurs on Mt Bartle Frere with A. quintanus sp. n. and A. summus sp. n., although both these species are found at higher elevations on the mountain. Name. Latin adelphus, 'brother', a punning reference to the 'Frere' in the name of the type locality; adjective.
Distribution. Known from rainforest at two localities ca 450 km apart in north Queensland (Figs 12, 13A; see Remarks).
Remarks. The very wide disjunction between the two known localities is puzzling, and I suspect that the Bellenden Ker locality on the specimen label is incorrect. I have not noticed any differences in gonopod or non-gonopod features between the two populations. Diagnostic description. Male and female with head + 19 rings. Colour in alcohol pale yellow. Male/female ca 8.0/8.0 mm long; ring 12 maximum diameter ca 0.65/0.8 mm, maximum width ca 0.7/1.1 mm. Metatergal tubercles in ca 12 irregular transverse rows, mostly without setae; metatergal setae short with slightly flared tips; lateralmost tubercles not enlarged, not forming pseudo-paranotum. Male legs 6 coxa with rounded, mediodistal projection. Telopodite ( Fig. 6C) with pp straight; at in transverse plane, short, triangular and curving posteriorly; dp directed posterobasally at base; mab deeply divided into large posterior and small posterolateral lobes; posterior lobe of mab distally with small folds and with medial edge produced near mab base as short, pointed lobe; posterolateral lobe of mab folded, the posteriormost fold with apex tooth-like and pointing anteromedially; meb divided at base into 2 needlelike processes curving behind mab.
Distribution. Rainforest in the Lamb Range and adjacent hills inland from Gordonvale, tropical north Queensland (Fig. 13A). Co-occurs with A. quintanus sp. n. on North Bell Peak. There is also a doubtful record from Cammoo Caves in central Queensland ( Fig. 1; see Remarks).
Name. Latin agnus, 'lamb', for the type locality, the Lamb Range; adjective.
Remarks. Like A. agnus sp. n., Asphalidesmus magnus Mesibov, 2011 andProsopodesmus crater Mesibov, 2012 were found in ANIC berlesate 535 from Cammoo Caves (Mesibov 2011(Mesibov , 2012a. For all three species, all other specimens are from localities on or near the Lamb Range, ca 800 km to the north of Cammoo Caves. The locality labelling for ANIC berlesate 535 appears to be incorrect (Mesibov 2012a). In March 2013 I searched briefly for millipedes in rainforest in the Cammoo Caves area but found no specimens of Agathodesmus, Asphalidesmus or Prosopodesmus.
Other material. None. Diagnostic description. Male with head + 20 rings. Colour in alcohol pale white. Male ca 7.5 mm long; ring 12 maximum diameter ca 0.6 mm, maximum width ca 0.75 mm. Metatergal tubercles in 7-8 irregular transverse rows, mainly without setae; metatergal setae long, pointed; 4 lateralmost tubercles not enlarged, forming very narrow pseudo-paranotum. Male leg 6 coxa with prominent mediodistal projection. Telopodite ( Fig. 6D) with pp straight; at in transverse plane, short, narrowly triangular and with tip curving posteriorly; dp directed posterobasally and laterally at base; mab shallowly divided into narrower anterior and wider posterior lobes; meb curving behind mab and divided at ca 1/3 length into short, needle-like medial and broader lateral processes, the latter following the curve of the posterior mab lobe and nearly as long.
Distribution. Known only from the type locality on the Cape York Peninsula in far north Queensland (Fig. 1).
Name. In honour of ANIC, the Australian National Insect Collection, whose collection of berlesates has yielded many new species of Australian millipedes.
Other material. None. Diagnostic description. Male and female with head + 19 rings. Colour in alcohol pale white. Male/female ca 3.5/4.0 mm long; ring 12 maximum diameter ca 0.3/0.4 mm, maximum width ca 0.4/0.5 mm. Metatergal tubercles in ca 4-5 irregular transverse rows, mostly without setae; metatergal setae short with slightly flared tips; 3 lateralmost tubercles enlarged, forming narrow pseudo-paranotum. Male leg 6 without coxal projection. Telopodite (Fig. 7A) with pp straight; at in transverse plane, very short, rounded-triangular and bent posteriorly; dp directed posterobasally and slightly laterally at base; mab deeply and widely divided into 2 subequal lobes with bluntly pointed apices; meb not divided, bent first posteriorly, then laterobasally and only very slightly curved, apex behind medial edge of medial lobe of mab.
Distribution. Known from wet eucalypt forest in East Gippsland, Victoria, over a linear extent of ca 20 km (Fig. 11).
Name. For the Bonang Highway, a narrow and winding road through the eastern Victorian mountains, type locality of this species; noun in apposition.
Remarks. Like A. carorum sp. n., A. bonang sp. n. is white in colour when alive, and contrasted well with the wet leaf litter and rotting wood in which I found it. Diagnostic description. Male and female with head + 19 rings. Colour in alcohol pale white. Male/female ca 4.0/4.5 mm long; ring 12 maximum diameter ca 0.4/0.5 mm, maximum width ca 0.5/0.6 mm. Metatergal tubercles in ca 6 irregular transverse rows, mostly without setae; metatergal setae short; 3 lateralmost tubercles enlarged, forming narrow pseudo-paranotum. Male leg 6 without coxal projection. Telopodite (Fig. 7B) with pp slightly curved posteriorly, widening distally; at in transverse plane, long, tapering to blunt point and curving posteromedially; dp directed posterobasally at base; mab directed posterolaterally at ca 90° to pp axis, then curving basally, the distal margin emarginate in its medial portion; meb not divided, bent sharply and directed posterolaterally near base, apex behind medial edge of mab.
Distribution. Wet eucalypt forest in far southeastern New South and far eastern Victoria (Fig. 11).
Name. In honour of Catherine and George Car, co-collectors of this species at the holotype and paratype localities; adjective.
Remarks. Like A. bonang sp. n., A. carorum sp. n. is white in colour when alive. Diagnostic description. Males and females with head + 19 rings. Colour in alcohol pale yellow. Male/female ca 5.0/5.5 mm long; ring 12 maximum diameter ca 0.45/0.5 mm, maximum width ca 0.55/0.7 mm. Metatergal tubercles in 6-7 irregular transverse rows, mostly without setae; metatergal setae short with slightly flared tips; 5 lateralmost tubercles enlarged, forming very narrow pseudo-paranotum. Male leg 6 without coxal projection. Telopodite (Fig. 7C) with pp straight; at in transverse plane, short, narrowly triangular with rounded tip curving posteriorly; dp directed posterobasally at base; mab directed laterobasally and curving anterobasally, narrow and not divided into lobes, distal margin thickened and with small emargination; meb curving behind mab, then following anterobasal curve of mab and terminating with it.

Agathodesmus chandleri
Distribution. Wet and dry eucalypt forest in the Adelaide Hills east of Adelaide, South Australia (Fig. 1).
Name. In honour of the American entomologist Don Chandler, who collected the first known specimens of this species while on a field trip to Australia; adjective.
Remarks. At the type locality I found an isolated aggregation of individuals of this species in very wet, friable material inside a rotting eucalypt log (http://www.polydesmida. info/polydesmida/thanks.html). Other specimens were in wet litter close to rotting wood.
Distribution. Known only from rainforest on Hinchinbrook Island, east of Cardwell in tropical north Queensland (Fig. 13A).
Distribution. Rainforest in tropical north Queensland from Mt Finnigan near Cooktown south to the Graham Range, a linear extent of ca 180 km (Fig. 13A) Other material. None. Diagnostic description. Male with head + 19 rings. Colour in alcohol brownish yellow. Male ca 8.0 mm long; ring 12 maximum diameter ca 0.6 mm, maximum width ca 0.65 mm. Metatergal tubercles in ca 10 irregular transverse rows, mainly without setae; metatergal setae short with slightly flared tips; lateralmost row of tubercles not enlarged, not forming pseudo-paranotum. Male leg 6 coxa with small, rounded, mediodistal projection. Telopodite (Fig. 9A) with pp curving anteriorly, then distally at base, slightly flattened anteroposteriorly; at in oblique plane (facing posterolaterally), short, narrowly triangular, arising abruptly from medial side of truncate pp apex, curving slightly posterolaterally; dp directed posterobasally at base; mab divided into 2 distally notched lobes, the posterior lobe curving medially and terminating in a basally directed point; meb broad, not divided, curving behind mab, apex slightly expanded.
Distribution. Known only from rainforest on the summit of Mt Bellenden Ker in tropical north Queensland (Fig. 13A). A. quintanus sp. n. occurs lower down on the same mountain.
Name. For the type locality, Mt Bellenden Ker; adjective. Remarks. Latitude/longitude data corrected following Mesibov (2012b).  Diagnostic description. Male and female with head + 19 rings. Colour in alcohol pale yellow. Male/female ca 10.0/10.5 mm long; ring 12 maximum diameter ca 0.75/0.8 mm, maximum width ca 0.8/0.9 mm. Metatergal tubercles in 12-16 irregular transverse rows, mainly without setae; metatergal setae short with slightly flared tips; lateralmost row of tubercles not enlarged, not forming pseudo-paranotum. Legs 6 and 7 coxae with long mediodistal projections (shorter on leg 7) with rounded tips. Telopodite (Fig. 9B) with pp straight; at in transverse plane, short and narrowly triangular, tip not curving posteriorly (curve in Fig. 9B is artefact of drying); dp directed posterobasally and laterally at base; mab divided into 3 lobes increasing in width and length from anterior to posterior, with anterior lobe curving medially and pointed, middle lobe distally truncate, posterior lobe distally expanded, the distal margin curving cup-like anteriorly and with small notch near medial edge; meb curving behind mab, divided at ca one-quarter length into 2 subequal, closely appressed, needle-like processes.
Distribution. Known only from rainforest on Mt Hosie and Mt Pershouse, ca 3 km apart in tropical north Queensland (Fig. 13A) Other material. None. Diagnostic description. Male with head + 19 rings. Colour in alcohol pale yellow. Male ca 7.5 mm long; ring 12 maximum diameter ca 0.5 mm, maximum width ca 0.6 mm. Metatergal tubercles in ca 10-12 irregular transverse rows, mainly without setae; metatergal setae short with slightly flared tips; lateralmost row of tubercles not enlarged, not forming pseudo-paranotum. Male leg 6 without coxal projection. Telopodite (Figs 8C, 8D) with pp curving posteriorly, anteroposte- riorly flattened (wider in posterior view than lateral); at in oblique plane (facing posterolaterally), short, narrowly triangular, curving posterolaterally; dp directed posterobasally and laterally at base, a small, rounded, mediolaterally flattened tab arising on posterior surface just above (basal to) mab origin; mab divided into 2 lobes; lateral mab lobe terminating in basomedially curving, finger-like process; medial mab lobe curving medially, divided into broad medial and narrow, distally expanded lateral process with truncate distal margin; meb curving behind mab, divided near base into needle-like, basally directed medial process and long, broad lateral process curving medially and terminating in short, broad hook behind medial lobe of mab.
Distribution. Known only from rainforest at the type locality on the Atherton Tableland in tropical north Queensland (Fig. 13A).

Agathodesmus morwellensis
Distribution. Known from eucalypt forest in the Latrobe River valley in West Gippsland, Victoria, at two sites ca 7 km apart (Fig. 11). Non-male specimens (in NMV) only tentatively assigned to Agathodesmus have been found in the mountains northwest of the type locality.
Name. For the type locality, Morwell National Park; adjective.
Diagnostic description. Male and female with head + 19 rings. Colour in alcohol pale yellow. Male/female ca 8.0/9.0 mm long; ring 12 maximum diameter ca 0.75/0.85 mm, maximum width ca 0.75/0.85 mm. Metatergal tubercles in ca 12 irregular transverse rows, mainly without setae; metatergal setae short with slightly flared tips; lateralmost tubercles not enlarged, not forming pseudo-paranotum. Male leg 6 coxa with prominent mediodistal projection. Telopodite (Fig. 9D) with pp straight; at in transverse plane, short, rounded-triangular, tip not curving posteriorly; dp directed posterobasally and laterally at base; mab deeply divided into 2 lobes; anterior mab lobe subdivided, its posterior portion curving medially; posterior mab lobe with cuplike partial fold at midlength, the distal margin bent into narrow shelf and with small notch near medial edge; meb curving behind mab, divided at base into 2 subequal, closely appressed, needle-like processes.
Distribution. Known from rainforest on the Atherton Tableland and the coastal range just southeast of Cairns in tropical north Queensland (Fig. 13B); the two areas are ca 70 km apart. Co-occurs with A. quintanus sp. n. on the Atherton Tableland. Name.
Greek para, 'near', and pholeos, 'hole'; adjective. The type locality is close to Mt Hypipamee Crater, a vertical volcanic pipe.
Distribution. Rainforest southwest of Babinda in tropical north Queensland, with a known east-west extent of ca 50 km (Fig. 13B). Co-occurs with A. hahnensis sp. n. in the Graham Range and with A. parapholeus sp. n. near Mt Hypipamee Crater. Possibly co-occurs on Mt Bartle Frere with A. adelphus sp. n.
Name. Latin quintanus, 'of the fifth'; adjective. The type locality was recorded as the fifth tower supporting the cableway to the top of Mt Bellenden Ker. This tower has since been renumbered '4' (Mesibov 2012b) Remarks. A. quintanus sp. n. has a particularly complicated gonopod telopodite and I am not certain that I have clearly seen all its details. The species is distinguished by the wide initial curve of the medial meb lobe and the basomedially directed, roundly pointed medial mab lobe.
Specimens from North Bell Peak are larger than those from the type locality and have more prominent dorsal tubercles. Latitude/longitude data for the types have been corrected following Mesibov (2012b Mesibov, 2009 (star), A. morwellensis sp. n. (filled squares) and A. steeli Silvestri, 1910 (crosses Diagnostic description. Male and female with head + 19 rings. Colour in alcohol very pale yellow. Male/female ca 8.0/8.0 mm long; ring 12 maximum diameter ca 0.7/0.8 mm, maximum width ca 0.7/0.85 mm. Metatergal tubercles in 10-12 irregular transverse rows, mainly without setae; metatergal setae short with slightly flared tips; lateralmost tubercles not enlarged, not forming pseudo-paranotum. Male leg 6 coxa with small, rounded, mediodistal projection. Telopodite (Fig. 11A) with pp straight; at in oblique plane (facing posterolaterally), short, rounded-triangular, curving posterolaterally; dp directed posterobasally and laterally at base; mab somewhat expanded distally, divided into 2 lobes with large anterior fold; meb divided at base into 2 needle-like processes, the smaller medial process directed basally, the longer medial process curving behind mab.
Distribution. Wet forest in tropical north Queensland from the Cape tribulation area south to the Atherton Tableland, a north-south extent of ca 120 km (Fig. 13B).
Other material. None.

Diagnostic description.
Male with head + 19 rings, female with head + 20. Colour in alcohol very pale yellow. Male/female ca 9.0/9.5 mm long; ring 12 maximum diameter ca 0.75/0.9 mm, maximum width ca 0.8/1.0 mm. Metatergal tubercles in 10-12 irregular transverse rows, mainly without setae; metatergal setae short with slightly flared tips; lateralmost tubercles not enlarged, not forming pseudo-paranotum. Male leg 6 without coxal projection. Telopodite (Fig. 11B) with pp slightly curving posteriorly; at in oblique plane (facing posterolaterally), short, narrowly triangular, curving posterolaterally; dp directed laterobasally and slightly posteriorly at base; mab expanded distally, divided into 2 distally rounded lobes, the medial lobe with truncate medial projection at ca midlength; meb curving behind mab and nearly as long, divided at ca midlength into shorter, needle-like lateral process and broad medial process terminating in upturned hook.
Distribution. Known only from rainforest at the type locality in tropical north Queensland (Fig. 13B). Possibly co-occurs on Mt Bartle Frere with A. adelphus sp. n. and A. quintanus sp. n.
Name. Latin summus, 'highest'; adjective. This species was found at the top of Queensland's highest mountain, Mt Bartle Frere.
Remarks. The Bartle Frere females were assigned to this species rather than to the co-occurring A. adelphus sp. n. and A. quintanus sp. n. because the females lack the narrow pseudo-paranota found in the other two species.
Distribution. Rainforest in the Cardwell Range (includes Kirrama Range), inland between Tully and Ingham in tropical north Queensland (Fig. 13B). The linear extent of the known range is ca 20 km. Co-occurs with A. kirrama sp. n. on Mt Hosie.
Name. For Yuccabine Creek, the type locality; adjective.
Agathodesmus bucculentus (Jeekel, 1986) http://species-id.net/wiki/Agathodesmus_bucculentus Figs 3A, 3B, 4A Note. This species was described from three specimens collected by Dr and Mrs Jeekel in 1980, under logs in rainforest along the 'Broken Hill' [probably 'Broken River'] track in Eungella National Park (Jeekel 1986). Another 183 specimens from the same general area are in Australian collections (see below), and the A. bucculentus