Corresponding author: Sarah C. Crews (
Academic editor: Dmitry Logunov
We relimit and revise the family
Spiders of the family
Revisionary and descriptive work for
While it has long been known that species of
Although the Australian fauna is restricted to a single species,
The purpose of this paper is twofold. First, we focus on the genera of the family
Taxa were described and illustrated primarily from specimens stored in 75% ethyl alcohol. In some cases, specimens were not available for direct examination, and we based our conclusions on published descriptions. Female copulatory organs were dissected and cleared using pancreatin (
anterior eye row
anterior lateral eyes
anterior median eyes
posterior eye row
posterior lateral eyes
posterior median eyes
femur
metatarsus
patella
tibia
tarsus
apical
dorsal
prolateral
retrolateral
ventral
median apophysis
retrolateral tibial apophysis
conductor
Australian Museum, Sydney, NSW, Australia (G. Milledge, H. Smith)
California Academy of Sciences, San Francisco, CA, USA (C. Griswold, D. Ubick)
Muséum d’Histoire Naturelle, Geneva, Switzerland (P. Schwendinger)
Queensland Museum, Brisbane, Qld, Australia (R. Raven, O. Seeman)
Royal Museum for Central Africa, Tervuren, Belgium (D. DeBakker, R. Jocqué)
South Australian Museum, Adelaide, SA, Australia (D. Hirst)
University Museum of Zoology, Cambridge University, United Kingdom (M. Lowe)
Western Australian Museum, Welshpool, WA, Australia (J. Waldock)
Museum für Naturkunde, Berlin, Germany (J. Dunlop)
Zoologisches Institut und Zoologisches Museum, Hamburg, Germany (H. Dastych)
Zoological Museum of the Moscow University, Moscow, Russia (K. G. Mikhailov)
Zoological Survey of India, Kolkata, India
The
Females (those of
1 | Two pairs of ventral spines on each Ti I and II, and on Mt I and II (known from Madagascar and Reunion Island) |
|
– | Spination otherwise | 2 |
2(1) | Three pairs of ventral spines on Ti I and II, and two pairs of spines on Mt I and II | 3 |
– | Spination otherwise | 4 |
3(2) | With tarsal scopulae (found in Africa, Asia, southern Europe and the New World) |
|
– | Without tarsal scopulae (found in India and Nepal) | |
4(2) | Found in Africa or Madagascar | 5 |
– | Found elsewhere | 6 |
5(4) | Spination pattern on ventral Ti and Mt I and II 4–3 and found in Madagascar |
|
– | Spination otherwise, or if 4–3, not found in Madagascar, but Africa |
|
6(4) | Tibia I and II with 4 ventral spines, Mt I and II with 3 ventral spines (found in India) | |
– | Other spination pattern | 7 |
7(6) | With 7 pair of ventral spines on Ti I and II, and 5 pairs on Mt I and II (found in Taiwan) | |
– | With a different number of ventral spines on Ti and Mt I and II | 8 |
8(7) | Epigynum not divided into lateral lobes, with very small, simple spermathecae ( |
|
– | Epigynum either divided into lateral lobes and/or with complex, coiled spermathecae | 9 |
9(8) | Epigynum with posterodorsal fold covering part of the extremely coiled spermathecae ( |
|
– | Epigynum without posterodorsal fold covering spermathecae ( |
Males (those of
1 | Two pairs of ventral spines on Ti I and II, and on Mt I and II (found in Madagascar and Reunion Island) |
|
– | Spination otherwise | 2 |
2(1) | Palpus without median apophysis ( |
|
– | Palpus with median apophysis ( |
3 |
3(2) | Spination pattern on ventral Ti and Mt I and II 3–2 | 4 |
– | Spination pattern on ventral Ti and Mt I and II otherwise | 5 |
4(3) | Tarsal scopulae absent (found in India and Nepal) | |
– | Tarsal scopulae present (found in Africa, Asia, southern Europe and the New World) |
|
5(3) | Conductor T-shaped with basally rounded projection ( |
|
– | Conductor otherwise ( |
6 |
6(5) | Chelicerae project forward with long fangs (found in Southeast Asia) |
|
– | Chelicerae and fangs otherwise | 7 |
7(6) | Median apophysis large, complex, and strongly sclerotized, often twisted ( |
|
– | Median apophysis much smaller, simple and tapered, either with one or two branches, lightly sclerotized and never twisted ( |
All members of this genus can be distinguished from other genera by the ventral spination on the tibiae and metatarsi of legs I and II, where there are 3 pairs of spines on the tibiae, and 2 pairs of spines on the metatarsi.
Despite the amount of recent work that has been done on the family, there is still some difficulty in determining boundaries of genera, in particular with the genus
Additionally, molecular phylogenetic work (
Total length 4–20.
Currently there are 124 species of
The Chinese species have been reviewed by
Male and female syntypes: Japan (ZMB 2679, 2692, 3501-52, not examined).
This species has been found in China, Japan, Korea and Taiwan. In China, the species has been found in Sichuan (Chengdu, Xiushan), Henan (Xinyang), Jiangsu (Suzhou), and Zhejiang (
In molecular phylogenetic analyses (
In China, it has been found on cedar (
Female holotype: Leshan Buddha Temple, Sichuan Province, China [
Only known from the type locality (
Known only from Sichuan Province in China (
It is clear from the drawings provided by
Total length 6.90.
A single species,
Holotype female: San Jose,
The specific epithet comes from the Buhid Mangyan word
This species can be differentiated from all others by the very simple internal copulatory organs (
Copulatory organs of
The type locality only (
Part of Asia showing the known distribution of the
Total length 4.00–17.40.
Currently there are 64 species of
Total length 5.30–6.90.
Currently there are three described extant species of
Total length 4.50.
Known only from the type locality (
The genus contains a single species,
Holotype male: canopy fogging the tree
The specific epithet comes from the Latin word
This species can be separated from all other
This species was collected from canopy fogging the tree
The type locality only (
Papua New Guinea showing the distribution of
Known only from Madagascar and the island of Réunion.
There are currently six described species of
Total length 3.90–10.30.
Species of
India showing the distribution of
In addition to transferring
(males of
1 | Males | 2 |
– | Females | 17 |
2(1) | Ventral spines on tibiae I and II unpaired | 3 |
– | Ventral spines on tibiae I and II paired | 4 |
3(2) | Base of embolus between 3 and 5 o’clock, base of median apophysis subquadrangular, with single branch curving slightly distally in ventral view ( |
|
– | Base of embolus between 4 and 6 o’clock, base of median apophysis ovoid, with single branch pointed retrolaterally in ventral view ( |
|
4(2) | Cheliceral promargin with 4 teeth | 5 |
– | Cheliceral promargin with 3 teeth | 6 |
5(4) | Cymbium pointed at tip, conductor terminates at 1 o’clock, base of embolus large and rounded, extending almost to base of cymbium ( |
|
– | Cymbium rounded at tip, conductor terminates at nearly 3 o’clock, base of embolus angular ( |
|
6(4) | Tibial apophyses with 3 processes | 7 |
– | Tibial apophyses with 2 processes | 9 |
7(6) | Conductor crescent-shaped, narrowing very abruptly in the middle, forming a long, narrow, scythe-shaped terminus ( |
|
– | Conductor shaped otherwise or not narrowing as abruptly ( |
8 |
8(7) | Conductor angular, terminus directed proximally, strongly sclerotized, MA directed retrolaterally ( |
|
– | Conductor with squarish projection medially, terminus directed retrolaterally to proximally, MA directed distally ( |
|
9(6) | MA very small, and attached to base of embolus ( |
10 |
– | MA larger, and attached elsewhere | 11 |
10(9) | Pointed angular projection directed proximally coming off base of embolus at 6 o’clock position ( |
|
– | No angular projection on base of embolus ( |
|
11(9) | MA with two branches ( |
12 |
– | MA with one branch ( |
14 |
12(11) | Base of embolus very large, covering part of MA, quadrangular projection at the tip of the conductor ( |
|
– | Base of embolus not covering part of MA, conductor pointed at tip | 13 |
13(12) | Tip of conductor slightly undulate ( |
|
– | Tip of conductor curved regularly, directed retrolaterally, portion of conductor behind MA ( |
|
14(12) | Dorsal tibial apophysis tapered, slender, and pointed, MA of irregular shape, conductor with pointed terminal projection ( |
|
– | Dorsal tibial apophysis quadrangular, truncate in lateral view, MA with a distinct hook | 15 |
15(14) | Embolus thick, directed distally, bisecting the cymbium, not curving or hook shaped ( |
|
– | Embolus with a thick base, but becoming very long and slender, curving around the edge of the cymbium | 16 |
16(15) | MA large, directed laterally, with a small hook distally, tapering toward hook, widening to a flat, truncate tip. Space between conductor and MA. Embolus curving, but not curving around the edge of the cymbium ( |
|
– | MA directed distally, with a distal hook, rounded at the tip. Conductor with long distal processes leaving no space between MA and conductor. Embolus curving around edge of cymbium ( |
|
17(1) | Tibiae I and II each with 6 pairs of ventral spines | 18 |
– | Tibiae I and II each with 5 pairs of ventral spines | 21 |
18(17) | Sperm ducts highly coiled, with 4–5 coils, no obvious oval or round large spermathecae ( |
|
– | Sperm ducts with less than 4 coils, large oval to round spermathecae | 19 |
19(18) | Lateral lobes form diamond shape around median septum, epigynal pockets absent ( |
|
– | Lateral lobes not forming diamond shape around median septum, epigynal pockets present | 20 |
20(19) | Internal ducts coiled ( |
|
– | Internal ducts not coiled ( |
|
21(17) | Cheliceral promargin with 4 teeth | |
– | Cheliceral promargin with 3 teeth | 22 |
22(21) | Spermathecae not large and round, but small and ovoid to elongated ( |
23 |
– | Spermathecae large and round ( |
27 |
23(22) | Epigynal pockets present ( |
|
– | Epigynal pockets absent | |
2424(23) | Lacking a clearly defined median field, lateral lobes indistinct ( |
|
– | Median field and lateral lobes more distinct | 25 |
25(24) | Median field large and keyhole-shaped ( |
|
– | Median field otherwise | 26 |
26(25) | Internal ducts long and gently curving, with less than 5 coils ( |
|
– | Internal ducts tightly coiled with more than 5 coils ( |
|
27(22) | Internal ducts not coiled ( |
|
– | Internal ducts coiled at least once | 28 |
28(27) | Lateral lobes forming diamond shape around median septum ( |
|
– | Lateral lobes and median septum shaped otherwise | 29 |
29(28) | Lateral lobes widely separated ( |
30 |
– | Lateral lobes close together or fused ( |
33 |
30(29) | Median septum quadrangular, no sclerotization at copulatory openings ( |
|
– | Median septum shaped otherwise, copulatory openings sclerotized | 31 |
31(30) | Spermathecae huge, nearly touching near the midline, median septum with some wrinkling ( |
|
– | Spermathecae well-separated, median septum smooth | 32 |
32(31) | Sides of median septum parallel, median septum quadrangular ( |
|
– | Sides of median septum coming together near the epigastric furrow, median septum subtriangular ( |
|
33(29) | Median septum and lateral lobes forming a keyhole shape ( |
|
– | Median septum and lateral lobes otherwise | 34 |
34(33) | Median septum and lateral lobes fused ( |
|
– | Boundaries of median septum and lateral lobes distinct | 35 |
35(34) | Copulatory openings without proximal bilobal sclerotization ( |
|
– | Copulatory openings with proximal bilobal sclerotization ( |
36 |
36(35) | Copulatory openings located medially ( |
|
– | Copulatory openings located in the upper 1/3 of the epigynal plate ( |
|
– | Copulatory openings located in the upper 1/3 of the epigynal plate ( |
The following synopsis of
Holotype immature (ZMH, not examined): Bowen [
The male coiled, the small MA that is attached to the base of the cymbium (
Copulatory organs of
The holotype from Bowen, north-eastern Queensland, is an immature (
This species has been collected from trees fogged with pyrethrum in vine scrub, and has been seen under the bark of eucalypts (R. Atkinson, pers. comm.).
This specieshas been collected from Northeast Queensland to the southern Cape York Peninsula (
Northeast Queensland Australia (inset) showing the distribution of
Holotype female (QM S52315): under bark in rainforest near Amos Bay [
The specific epithet comes from the indigenous word for Cooktown, the type locality, in the Guugu Yimithirr language.
Males can be distinguished from other species by a triangular projection directed basally coming off of the base of the embolus (
This species has been collected under bark in rainforest, and under bark of
Northeast Queensland (
Holotype female (QM S61052): Upper Lankelly Creek [
This species is named for the collector, G.B. Monteith, in honor of his amazing collecting prowess.
Females of
Copulatory organs of
No data.
The type locality only (
Holotype male (WAM T93/1330): northwest tip of Degerando Island, Champagny Islands,
This species is named after Alan Longbottom, collector of the holotype and many other interesting arachnids for the Western Australian Museum.
Males can be separated from other species by having three processes of the RTA as well as a crescent-shaped conductor with a scythe shaped terminus (
Collected from under rocks.
The type locality only (
Northern Australia (inset) showing the distribution of
Holotype male (WAM T55002): Drysdale River Station,
This species is named for the late Keith Longbottom, collector of many interesting arachnids for the Western Australian Museum.
Males can be differentiated from other species by having an RTA with three processes, a crescent shaped conductor with a medial quadrangular lateral projection, and a MA that is directed distally (
No data.
The type locality only (
Holotype male (WAM T93/1333): north of Larryoo, Drysdale River National Park,
This species is named for the type locality, and is to be treated as a noun in apposition.
Males can be differentiated by the presence of three processes on the palpal RTA and the tip of the conductor being heavily sclerotized and directed basally (
Collected from under rocks.
The type locality only (
Holotype male (WAM T55003): 11 km NW of Roe’s Rock (16A), Fitzgerald River National Park,
The specific name comes from the Nyoongar
Males of this species can be separated from all other species except
Copulatory organs of
Though the male and female have not been collected together, it is clear from their morphologies that they are the same species. Additionally, and interestingly, this species is morphologically similar to
Found in pitfall traps, and at night along overland conveyors.
Near the south and west coasts of southwestern Australia (
The southwest coast of Western Australia (inset) showing the distribution of
Holotype female (AM KS19743): Kitty’s Creek [
The specific epithet comes from the indigenous Dharug word for spider. Dharug is the language indigenous to the type locality. The name is to be treated as a noun in apposition.
This species can be separated from
There is some ambiguity as to where exactly this specimen is from as it is rather old. The region of Kitty’s Creek in Sydney has been searched recently, but the area has been developed a great deal since the specimen was originally collected.
No data.
The type locality only (
Eastern Australia (inset) showing the distribution of
Holotype male (QM S50593):Brooyar State Forest,
The specific epithet is in honor of Dr. Robert Raven from the Queensland Museum in recognition of his work on Australian spiders.
Males can be distinguished from all other males by having unpaired setae ventrally on tibiae I and II, and from
Copulatory organs of
Thus far, this is the most widespread species of
Found under bark and under rocks, and on trees at night (
Widespread in eastern Australia, from the Tinderry Range in the south of New South Wales to the Bundaberg Forest in Queensland in the north (
Holotype female (WAM T97214): site 5, Badgeradda Range, Muggon Station,
The specific epithet comes from the type locality. The name is to be treated as a noun in apposition.
This species can be separated from all other species by genitalic characteristics, including a bilobed sclerotized area covering the copulatory openings located in the posterior third of the epigynal plate (
Immatures are matched with this species as they were collected from the same locality as the holotype.
Collected under rocks (
This species is only known from the Muggon Station area of Western Australia (
The northwest of Western Australia (inset) showing the distribution of
Holotype male (WAM T55000): John Wayne Country, Barrow Island, Western Australia, Australia,
This species is named for Andrew Burbidge in recognition of his conservation activities in Australia.
Males of this species can be differentiated from other species by their large, transparent, laterally directed MA (
Copulatory organs of
Collected from on/under rocks at night, or in pitfall traps around rocks.
Known only from Barrow Island, Western Australia (
Holotype male (WAM T55001): Bush Bay, site BB3,
The specific name comes from the Yinggarda word
Males can be differentiated from other species by having a palpal conductor with a long, fleshy terminal process (
No data.
The type locality only (
Holotype female (WAM T64748): quadrat 19, Lorna Glen Station,
The specific epithet is in honor of Julianne Waldock for all of her work on Australian terrestrial invertebrates.
Females of this species can be differentiated from others by a well-defined keyhole-shaped median field, and a single genital opening that internally is divided into two copulatory ducts leading to medium-sized oval spermathecae (
Copulatory organs of
No data.
The type locality only (
Holotype female (WAM T97482): Trinity Bore South TBRC078, vicinity of Cardo Camp, Red Hill, Pilbara, Western Australia, Australia,
The specific name comes from the Kurrama word
Females of this species can be differentiated from all others by the indistinct median septum and lateral lobes, and the median field being a long, narrow depression, with round spermathecae, and the sperm ducts nearly touch medially (
Collected from under rocks (
Found in the Pilbara region of Western Australia (
Holotype female (AM KS43756): Kempsey Road, West of Armidale, above Macleay River [
The specific epithet comes from the word for spider,
Females can be differentiated from all other species by having a large quadrangular medium septum (
Found on a rock wall and under bark (
The type locality only (
Holotype male (SAM N199353): Arcoona Creek, near Sambot Waterhole, Gammon Ranges National Park,
The specific name comes from the Adnyamathanha words
Males can be differentiated from all other species by having a portion of the conductor behind the MA, and the conductor being sclerotized terminally. Additionally, the embolus is short and hook shaped, and in the center of the bulb rather than the lateral edge (
Copulatory organs of
Collected at night.
Known from throughout the Gammon and Flinders Ranges in South Australia (
Central Australia (inset) showing the distribution of
Southern central Australia (inset) showing the distribution of
Holotype male (WAM T76590): base of Trig Hill, Old Telegraph Station, Alice Springs, Northern Territory, Australia,
This species is named in honor of Karl Pilkington.
The male has a thick and short embolus that bisects the bulb (
Collected from under rocks.
Only from Alice Springs (
Holotype female (SAM N199350): Mount Lindsay, South Australia, Australia [
The specific epithet comes from the Latin word
Females can be differentiated from other species by the median septum of the copulatory organs tapering posteriorly, giving it a subtriangular appearance (
Copulatory organs of
Collected from under rocks.
The type locality only (
Holotype male (SAM NN10914): 8 km northeast of Mount Woodroffe, South Australia, Australia, in gorge,
The specific epithet refers to the indigenous Pitjantjatjara name for Mount Woodroofe, the type locality. The name is to be treated as a noun in apposition.
Males of this species can be differentiated from others by having a sinuate anterior margin of the conductor, having two branches on the MA, and a hook-shaped embolus that bisects the palpal bulb (
Although the male and female specimens were not collected together (they were collected some 46 km apart), it seems reasonable to assume they are conspecific.
No data
The type locality only (
Holotype male (WAM T54996): Fitzgerald River National Park, northeast slope of West Mount Barren,
This species is named in honor of the second author’s (MSH) daughter, Frances Harvey.
Copulatory organs of
Throughout its range, this species is subject to variation in body size, ventral tibial and metatarsal spination of legs I and II, as well as in the number of promarginal teeth. There are 6 pairs of spines ventrally on tibiae I and II in most specimens, but some specimens have 5 pairs. Typically, there are 4 pairs of spines located ventrally on the metatarsus, however, at least one specimen has a 2–2-1 pattern. This species has either 4 or 5 promarginal teeth. The leg lengths also vary a great deal and have been found to be 3421, 23=41, 3241 and 2341. There is no genitalic variation.
This species has been collected from under bark, on trees, and under rocks. The female guards her egg-sac (
Found along the south coast of Western Australia from Mount Lindesay east to Mount Diamond, and in the islands of the Recherche Archipelago (
Holotype female (WAM T76592): Stirling Range National Park, Toolbrunup,
The specific epithet refers to the type locality, which means ‘drizzle carrier’ in the indigenous Nyoongar language. The name is to be treated as a noun in apposition.
This species can be separated from most others by having 4 teeth on the cheliceral promargin, and from
Collected from under rocks on a scree slope (
The type locality only (
Holotype male (WAM T93/1366): Mount Cooke,
This species is named in honor of the second author’s (MSH) daughter, Ellen Harvey.
Males of this species can be separated from others by the irregular shaped and unsclerotized MA, and the conductor has a pointed projection terminally (
Copulatory organs of
Collected from beneath bark, rocks and other debris.
Along the west coast of southwest Australia (
Holotype female (WAM T65078): Oscar Range, Western Australia, Australia,
The specific epithet is named in honor of the first author’s (SCC) sister, Jennifer Crews.
This species can be differentiated from all others by the small round median septum and the lateral lobes comeing into contact posteriorly. The internal ducts are not coiled and the spermathecae are ovoid and slightly pinched medially (
This large species has been collected from limestone rocks at night.
The type locality only (
Holotype female (WAM T54998): Kakadu National Park, Kapalga, primary site E,
The specific epithet comes from the word for spider,
This species can be distinguished from all others by having an abdominal pattern of light spots on a dark background, and by the copulatory organs, as the lateral lobes come into contact for nearly half of the length of the epigynal plate, and the copulatory ducts are long, laterally positioned, and lead to a mass of winding ducts (
This species has been found on
Known only from the northern region of the Northern Territory (
Here we describe two new species, move three species from
It appears that at least two genera,
It is likely that several more species from this region will be found with further exploration, and it is of note that the male of only one species of
Total length 6.70–9.70.
The genus contains five species:
1 | Tibiae I and II with 4 pairs of ventral spines | 2 |
– | Tibiae I and II with 3 pairs of ventral spines | 3 |
2(1) | Epigynal pockets not reaching sinuous margin covering genital openings ( |
|
– | Epigynal pockets reaching margin covering copulatory openings ( |
|
3(1) | Very large posterodorsal fold covering internal ducts and spermathecae ( |
|
– | Posterodorsal fold absent | 4 |
4(3) | Internal ducts asymmetrical, convoluted, twisting numerous times ( |
|
– | Internal ducts only twisted 3 or so times, internal ducts symmetrical ( |
Holotype female (apparently lodged in ZSI; not examined):Jaonsar, Kumia, Uttarakhand, India [
Males can be separated from other selenopids by the RTA, a single, bifid process, the large T-shaped conductor, and the small, hooked MA (
Copulatory organs of
Collected under bark and rocks, and taken from over 2500 m elevation.
This species is found in northern India and Nepal, near the Himalaya Mountains (
Holotype female (ZSI; not examined):Agumbe Ghat, District Shimoga, Mysore, India, B.K. Tikader, 15.III.1965.
We were unable to examine the type of
Holotype female (CAS 9031589): 2 miles northwest of Punjur, Karnataka, India [
The specific epithet comes from the Kannada word ಮೈಸೂರ = Maisūru referring to Mahishasura, a Hindu asura, for which the city of Mysore, or the region of the type locality, was named. The name is to be treated as a noun in apposition.
This species can be differentiated from all others by a combination of characters including tibiae I and II with three pairs of ventral spines, sperm ducts only coiled a few times, and ducts symmetrical (
No data.
Known only from southern India (
Holotype female (MHNG): Karteri Valley, Tamil Nadu, India [
The species can be separated from other species by the raised epigynal plate and very large posterodorsal fold (
No data.
The type locality only (
Holotype female (CAS 9031584): Bhimashankar, Maharashtra, India,
The specific epithet refers to the Hindu god Shiva, as the type locality is the location of one of the 12 traditional Jyotirlingas of Shiva. The name is to be treated as a noun in apposition.
This species can be differentiated from all others by having 4 pairs of ventral tibial spines on legs I and II, and by the epigynal pockets reaching the sinuous margin where copulatory openings are located (
Copulatory organs of
No data.
The type locality only (
Habitats of
Habitus of representatives of various selenopid genera.
Habitus of representatives of various selenopid genera.
Although we have not examined specimens of this species, the published descriptions (
Total length 6.10.
Taiwan, near Taipei (
A single species,
Kayashima’s collection was thought to have been left in Taiwan when he went to Malaysia (H. Ono pers. comm.), but the material has not been located.
Total length 6.00–7.90.
Four species,
Female holotype (UMZC I.47430): Gunong, Malaysia [
The female of this species can be easily distinguished from all others by the copulatory organs, as the sperm ducts are coiled 16 times, and a posterodorsal fold is present (
No data
The type locality only (
This project was funded by Aquila Resources to whom we are grateful. The project was kindly facilitated by Garth Humphreys from Biota Environmental Sciences, Leederville, Western Australia. We are grateful to the various museum staff for access to the specimens utilized in this study: G. Milledge and H. Smith (Australian Museum, Sydney); C. Griswold and D. Ubick (California Academy of Sciences, San Francisco); P. Schwendinger (Muséum d’Histoire Naturelle, Geneva); R. Raven and O. Seeman (Queensland Museum, Brisbane); D. DeBakker, R. Jocqué (Royal Museum for Central Africa, Tervuren); D. Hirst (South Australian Museum, Adelaide); M. Lowe (University Museum of Zoology, Cambridge University); and J. Waldock and V. Framenau (Western Australian Museum, Welshpool). We also thank Volker Framenau for helpful discussions on Australian