THE AUSTRALIAN ENTOMOLOGIST

ABN#: 15 875 103 670

The Australian Entomologist is a non-profit journal published in four parts annually

by the Entomological Society of Queensland and is devoted to entomology of the

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Cover: A basking male of Rhyothemis princeps princeps Kirby 1894 (Odonata:

Libellulidae) at Eubenangee Swamp, North Queensland. This subspecies occurs in

tropical Queensland from Rockhampton north to Iron Range and Weipa and is also

known from southern New Guinea. It is very common at most standing waters In the

Queensland Wet Tropics where it occurs together with its congeners Rhyothemis

graphiptera Rambur, 1842 and the rarer Rhyothemis resplendens Selys, 1878.

Another race with much more extensive hyaline areas occurs in north and east New

Guinea. Pen and ink drawing by Dr Albert Orr whose illustrated books on

dragonflies and butterflies have won awards in Australia and overseas. He lived at

Bramston Beach 1999-2004 and nearby Eubenangee Swamp was a favourite hunting

ground.

Australian Entomologist, 2014, 41 (1): 1-6 ]

THE DISTRIBUTION OF EUPLOEA DARCHIA NIVEATA (BUTLER,

1875) (LEPIDOPTERA: NYMPHALIDAE: DANAINAE) IN

TORRES STRAIT, QUEENSLAND, AUSTRALIA

TREVOR A. LAMBKIN

School of Biological Sciences, University of Queensland, St Lucia, Qld 4072

(Email: trevor.lambkin@ugqconnect.edu.au)

Abstract

A review of the known distribution of Euploea darchia niveata (Butler) showed that the taxon

predominately occurs on mainland Queensland but also encompasses Thursday (Waiben) Island

in the very south of Torres Strait. Based on this review, the record of a single female collected on

Darnley (Erub) Island, Torres Strait in April 1910 by Hermann Elgner seems unusual. Despite

Elgner’s generally high standard in the veracity of his specimen labelling, this particular record

is considered to be erroneous. Based on a reconstruction of Elgner’s 1910 travels through Torres

Strait by using data from specimen labels, a probable explanation for this record is that the

specimen was collected on Thursday Island around 29 March 1910, prior to his departure for

Darnley Island. On reaching Darnley on 3 April 1910 the specimen was then likely mistakenly

labelled as originating from Darnley Island. The suggested narrower range of this taxon

highlights the Australian endemic status of E. d. niveata.

Introduction

The crow butterfly Euploea darchia (W.S. Macleay, 1826) is known from

Timor, Banda, Aru and the Kai Islands and northern Australia (Ackery and

Vane-Wright 1984, Morishita 1985, Braby 2000). Two endemic subspecies

occur within Australian limits. Euploea d. darchia is known from coastal

Northern Territory including Gove Peninsula (Braby 2000 and pers. comm.)

and from the Kimberley area of Western Australia (Braby 2000). Euploea d.

niveata (Butler, 1875) is recorded predominately from coastal areas of

northern Queensland (Braby 2000), where it is known from Thursday

(Waiben) Island in Torres Strait south through Cape York Peninsula,

including its west coast bordering the Gulf of Carpentaria to Weipa

(McCubbin 1971) and Kowanyama (Hopkinson 201 1), then south along the

east coast to Mission Beach (Braby 2000) and Tully. South of the Wet

Tropics, E. d. niveata has been recorded sporadically (possibly as vagrants)

from Murray Upper (recorded here), Mackay, St Lawrence, Yeppoon,

Caloundra, Brisbane and Burleigh Heads (Common and Waterhouse 1981,

Braby 2000, Moss 2010, A.G. Orr pers. comm.).

Euploea d. niveata frequents a variety of wet shady habitats, including

rainforest, monsoonal vine forest and wet coastal swamps (Barrett and Burns

1951, McCubbin 1971, Braby 2000). Although generally uncommon and

erratic in its distribution, it is observed more frequently in the very north of

its range, for example at Bamaga and Lockerbie at the tip of Cape York

Peninsula and on Thursday Island.

While E. d. niveata is known essentially from the Queensland mainland, with

records from the south of Torres Strait (Waterhouse and Lyell 1914, Lambkin

2 Australian Entomologist, 2014, 41 (1)

and Knight 1983, De Baar 1988, T.A. Lambkin unpublished data), there is a

longstanding and puzzling isolated record, documented in Waterhouse and

Lyell (1914), of the butterfly’s occurrence on Darnley (Erub) Island, Torres

Strait, a substantial distance across water (approximately 200 km) from

Thursday Island and Cape York. In this paper I discuss a likely hypothesis

that would account for this record, which in all probability is incorrect, and

outline the species’ ostensibly restricted distribution in Torres Strait.

Abbreviations of collectors’ names or collection repositories are: AIK — A.I.

Knight; AM — Australian Museum, Sydney; HE — H. Elgner; MDB — M. De

Baar; QDAFFC — Queensland Department of Agriculture, Fisheries and

Forestry collection, Brisbane; TAL — T.A. Lambkin; TLIKC — Joint

collection of T.A. Lambkin and A.I. Knight, Brisbane.

Material examined

Specimen under review

QUEENSLAND: 1 9, Darnley Island, 13.iv.1910 [HE] (AM) (recorded as E. hyems

niveata [Butler] in Waterhouse and Lyell 1914, p. 23) (Fig. 1).

Additional material examined

QUEENSLAND: 3 99, Green Hill, Thursday Island, 2.v.1989, 12-15.iv.1992,

3.11.2004, AIK (TLIKC); 2 dd, 1 9, same data except 27-29.iii.1987, MDB

(TLIKC); 1 d, 1 9, same data except 9.iii.2001 (8), 12.1.1994 (9), TAL (TLIKC);

1 d, Lockerbie Scrub, 14 km NE of Bamaga, 23-31.vii.1983, TAL (TLIKC); 1 3,

2 22, same data except, 21-24.vi.1992, AIK (TLIKC); 2 dd, 17-21 km NE of

Bamaga, 3-8.v.1994, AIK (TLIKC); 2 $3, 20 km E of Bamaga, 27.vii.1983, AIK

(QDAFFC); 1 9, Cape Tribulation, 10.v.1979, TAL (TLIKC); 1 d, Kuranda,

21.iii.1975, MDB (TLIKC); 1 3, Cairns, 22.v.1982, TAL (QDAFFC); 1 d, Flying

Fish Point, via Innisfail, 4.v.1978, TAL (TLIKC); 1 8, Coquette Point, via Innisfail,

20.iv.1987, TAL (TLIKC); 1 3, same data except, 22.iv.1987, AIK (QDAFFC); 1 9,

El Arish, 19.iii.1975, MDB (TLIKC); 1 9, Tully, 22.iv.1982, AIK (QDAFFC); 1 d,

Murray Upper, 15.1.1989, MDB (TLIKC); 1 9, Burleigh Heads, 9.iv.1977, TAL

(TLIKC).

Discussion

Euploea darchia niveata (Butler, 1875) was described (as a subspecies of

Calliploea darchia) from material (in the original description) from

‘Queensland’ and ‘Australia’ (Edwards ef al. 2001). In a later publication,

Butler (1878) recorded a male from Cape York and a female from Fitzroy

Island, Queensland as types but Edwards et al. (2001) considered that these

locations could have been from subsequent accessions to the material used in

the original description. In any case, Waterhouse and Lyell (1914) listed

several localities for the butterfly, stretching from Cape York south to

Mackay on the mainland, with additional records from Thursday and Darnley

Islands in Torres Strait. Thursday Island has been a popular destination for

butterfly collectors since the late 1800s-early 1900s (Mathew 1885,

Australian Entomologist, 2014, 41 (1) 3

Waterhouse and Lyell 1914) and more recently in the 1980s and 1990s, with

E. d. niveata being frequently observed (Lambkin and Knight 1983, De Baar

1988, T.A. Lambkin unpublished data).

Fig. 1. Elgner's specimen of Euploea darchia niveata labelled ‘Darnley Island,

13.iv.1910" [forewing length 38 mm]; photograph courtesy of the Australian Museum.

Of particular interest is the Darnley Island record of E. d. niveata by

Waterhouse and Lyell (1914). This specimen (labelled ‘Darnley Island,

13.iv.1910’ and housed in the Australian Museum: Fig. 1), is one of many

Torres Strait butterflies collected by Hermann Elgner, a German collector

who made significant collections of Torres Strait butterflies and dragonflies

predominately in the first decade of the 20^ Century (Moulds 1977, Dunn

2007) (material housed in the Australian Museum and Museum Victoria).

Darnley Island is volcanic in origin (Lambkin 2013) and located in the far

eastern sector of Torres Strait. Since the rediscovery of the islands of Torres

Strait as a focus for butterfly collecting in the late 1970s and 1980s (Monteith

1978), Darnley Island has been surveyed for butterflies by several collectors,

commencing in the 1980s (e.g. Johnson 1983, De Baar 1988), through to the

first decade of this century (T.A. Lambkin unpublished data). While several

species of Euploea Fabricius have been collected over this period on Darnley

Island, E. d. niveata remains the only Euploea taxon not to have been

collected or observed there since Elgner’s 1910 isolated ‘collection’. Apart

from this remote record, the butterfly is known in Torres Strait only from

4 Australian Entomologist, 2014, 41 (1)

Thursday Island in the very south of the region. Trophis (= Malaisia)

scandens (Lour.) Hook. & Arn. (Moraceae), the recorded host plant of E. d.

niveata (Manski 1939), has a patchy distribution in Torres Strait and is

known to occur on Darnley Island (Lambkin and Knight 2005).

Hermann Elgner was based for some of his time in the Torres Strait region at

Cape York (Moulds 1977) and undertook collecting forays to several Torres

Strait islands (Moulds 1977, Dunn 2007). Of particular interest are his

movements through several of the Torres Strait islands in 1910. What can be

best determined, according to Moulds (1977) and Dunn (2007), is that

Elgner’s last reported collecting day on Thursday Island was 29 March 1910,

followed by his (only recorded) visit to Hammond (Keriri) Island on 31

March and then to Moa (Banks) Island en route to Darnley Island. He arrived

at Darnley Island on the 3 April for an extended period and returned to Cape

York in either June or July that same year (Moulds 1977, Dunn 2007).

Dunn (2007) intensively reviewed the movements of H. Elgner, basing his

study on roughly 1400 collected specimens. Dunn (2007) considered the

accuracy of Elgner’s labelling to be ‘exemplary’ at approximately 99%.

While the overall veracity of Elgner’s locality records cannot be disputed

some errors have been detected (Meyer et al. 2004, Dunn 2007). The case of

the E. d. niveata specimen discussed here seems to be possibly one of the rare

Elgner errors in labelling. Although the possibility of a vagrant from the

south-west (i.e. Cape York or Thursday Island) cannot be entirely discounted,

the specimen is of fresh appearance and unlikely to have travelled such a long

distance. It seems more likely that it was collected by Elgner on Thursday

Island on or prior to 29 March 1910 and was not labelled until after he

arrived at Darnley Island on 3 April 1910, thus being mistakenly labelled as

originating from Darnley Island in April of that year.

The distribution of E. d. niveata appears to be almost entirely confined to

mainland Queensland. How far west it extends along the coastal vine scrubs

of the Gulf of Carpentaria, possibly into the Gove Peninsula of the Northern

Territory, is currently unknown, as this region is rarely visited by

lepidopterists. Despite a flurry of collecting, undertaken primarily during the

1980s and 1990s and then, to a lesser degree, since 2000 on several Torres

Strait islands, including the southern group of islands that satellite Thursday

[Waiben] Island (specifically Horn [Narupai], Hammond and Prince of Wales

[Muralag]), Thursday Island remains the only location in Torres Strait where

E. d. niveata has been observed or collected. However, it is likely that the

butterfly might yet be found on those islands that satellite Thursday Island,

since they support potential breeding habitat comprising patches of deciduous

monsoon forest (T.A. Lambkin unpublished data).

Finally, taking into account the temporal movements of Hermann Elgner

through Torres Strait in March and April 1910, and with the weight of

Australian Entomologist, 2014, 41 (1) 5

collecting data indicating that E. d. niveata appears to be endemic to tropical

coastal Queensland and Thursday Island in the extreme south of Torres Strait,

it seems unlikely that the butterfly’s distribution encompasses: Darnley Island

as Elgner’s label indicates.

Acknowledgements

Appreciation is given to J.S. Bartlett (Brisbane) for provision of the image of

the H. Elgner specimen of E. d. niveata and providing access to material held

in the Queensland Department of Agriculture, Fisheries and Forestry

Collection, Brisbane. Thanks also to S. Ginn of the Australian Museum,

Sydney for organising approval for publication of the image. M.F. Braby and

A.G. Orr kindly provided their personal communications. This work partially

fulfills the requirements for a Master of Philosophy degree undertaken by the

author at the University of Queensland, Brisbane.

References

ACKERY, P.R. and VANE-WRIGHT, R.I. 1984. Milkweed butterflies. British Museum (Natural

History), London; ix + 425 pp.

BARRETT, C. and BURNS, A.N. 1951. Butterflies of Australia and New Guinea. N.H. Seward

Pty Ltd, Melbourne; 195 pp.

BRABY, M.F. 2000. Butterflies of Australia: their identification, biology and distribution.

CSIRO publishing, Collingwood; xx + 976 pp.

BUTLER, A.G. 1878. On the butterflies in the collection of the British Museum hitherto referred

to the genus Euploea of Fabricius. Journal of the Linnaean Society of London, Zoology 14: 290-

303.

COMMON, LF.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Angus and

Robertson, Sydney; xiv + 682 pp.

DE BAAR, M. 1988. Insects collected during a trip to Torres Strait 27 March to 10 April, 1987.

News Bulletin of the Entomological Society of Queensland 15: 107-117.

DUNN, K.L. 2007. The geographical movements of Hermann Elgner (d. 1913): a lepidopterist

based in Torres Strait, Australia. Calodema 10: 19-34.

EDWARDS, E.D., NEWLAND, J. and REGAN, L. 2001. Lepidoptera: Hesperioidea,

Papilionoidea. In: Wells, A. and Houston, W.W.K. (eds), Zoological catalogue of Australia. Vol.

31.6. CSIRO Publishing, Collingwood; x + 616 pp.

HOPKINSON, M. 2011. New distribution records for several butterfly species, including

Deudorix democles (Miskin) (Lepidoptera: Lycaenidae), from western Cape York Peninsula.

Australian Entomologist 38(2): 91-95.

JOHNSON, S.J. 1983. A brief look at the butterflies of Darnley Island. News Bulletin of the

Entomological Society of Queensland 8: 117-119.

LAMBKIN, T.A. 2013. A revision of the Euploea batesii C. & R. Felder, 1865 ‘complex’

(Lepidoptera: Nymphalidae: Danainae) in mainland Papua New Guinea and Australia, including

its biology and biogeography. Australian Entomologist 40(4): 187-218.

LAMBKIN, T.A. and KNIGHT, A.I. 1983. Butterflies recorded from the Torres Strait and

northern Cape York Peninsula regions in July 1983. News Bulletin of the Entomological Society

of Queensland 11: 98-103.

6 Australian Entomologist, 2014, 41 (1)

LAMBKIN, T.A. and KNIGHT, A.I. 2005. New Australian butterfly records (Lepidoptera) from

Saibai and Dauan Islands, Torres Strait, Queensland. Australian Entomologist 32(2): 49-54.

MANSKI, M.J. 1939. The Euploeas (Rhopalocera) of the Cairns district. North Queensland

Naturalist 8: 3-4.

MATHEW, G.F. 1885. An afternoon among the butterflies of Thursday Island. Proceedings of

the Linnaean Society of New South Wales 10: 259-266.

McCUBBIN, C. 1971. Australian butterflies. Thomas Nelson, Melbourne; xxxi + 206 pp.

MEYER, C.E., BROWN, S.S. and WEIR, R.P. 2004. The first record of Euploea modesta lugens

Butler (Lepidoptera: Nymphalidae: Danainae) from Australia. Australian Entomologist 31(4):

177-180. Í

MONTEITH, G.B. 1978. An insect collection from the islands of Torres Strait. News Bulletin of

the Entomological Society of Queensland 5: 139-140.

MOULDS, M.S. 1977. Bibliography of the Australian butterflies (Lepidoptera: Hesperioidea

and Papilionoidea) 1773-1973. Australian Entomological Press, Greenwich NSW; 239 pp.

MORISHITA, K. 1985. Danaidae. In: Butterflies of the South East Asian Islands. Vol II. Plapac

Co, Ltd, Tokyo; 623 pp.

MOSS, J.T. 2010. The intriguing white-margined crow butterfly Euploea darchia niveata

(Butler, 1875). Metamorphosis Australia 56: 4-9.

WATERHOUSE, G.A. and LYELL, G. 1914. The butterflies of Australia. A monograph of the

Australian Rhoplaocera. Angus and Robertson, Sydney; vi +239 pp.

Australian Entomologist, 2014, 41 (1): 7-14 7

PHENOLOGY OF THE AUSTRALIAN SOLITARY BEE SPECIES

LEIOPROCTUS PLUMOSUS (SMITH) (HYMENOPTERA:

COLLETIDAE)

MICHAEL BATLEY' and BARRY BRANDLEY?

! Australian Museum, 6 C ollege Street, Sydney, NSW 2010

(Email: michael.batley(à) gmail.com)

75A Jason Place, Ulladulla, NSW 2539

Abstract

A nesting aggregation of the bee species Leioproctus (Leioproctus) plumosus (Smith, 1853) in a

suburban garden was observed for a period of seven years. Between one and three generations

per year were observed and estimates obtained for the development time and lifespan of adults.

The emerging bees were accompanied, at this site, by the parasitic ichneumonid wasp Labium

pettitorium (Erichson, 1842).

Introduction

Surprisingly little is known about the life cycles of Australian bees in the

family Colletidae, which contains 53% of named Australian species (AFD `

2013). Several species in the family Apidae (Cardale 1968a, b, Steen and

Schwarz 2000, Schwarz et al. 2007 and references therein, Halcroft et al.

2013) and one halictine species (Kukuk ef al. 2005 and references therein),

have received significant study but observations of colletids have been

restricted to indirect inferences concerning the number of generations per

year.

Observation of adult activity at times separated by more than 8 weeks has

frequently been taken as an indication of two or more generations per annum

(e.g. Houston 1971, 1975, 1987), based on the expectation that solitary bees

have an active lifespan of only a few weeks (Michener 2007). Exceptions,

like the queens of highly social species or species that overwinter as adults,

are uncommon, although the colletid species Amphylaeus morosus is reported

to live for up to a year (Spessa et al. 2000). The observations need to be made

at the same or similar locations as the activity periods for univoltine species

at different places may be widely out of phase (Houston 1991).

On this basis, six Australasian Leioproctus species are believed to be

univoltine and six bivoltine or multivoltine (Rayment 1935, 1950, Donovan

1980, Houston 1990, Houston and Maynard 2012, Maynard 2013), but no

detailed phenology of an Australian Leioproctus species has been reported.

Leioproctus plumosus (Smith) (Fig. l) is widely distributed throughout

Australia but predominantly in coastal regions in the southern half of the

continent (Fig. 2). While Museum collection data (Fig. 3; ALA 2013)

indicate that L. plumosus is active over an extended period, it is unclear

whether this is an indication of multivoltinism or just the result of combining

data from a range of environments.

8 Australian Entomologist, 2014, 41 (1)

A nesting aggregation of L. plumosus in a domestic garden in Ulladulla, a

coastal site ca 170 km south of Sydney, NSW, provided an opportunity to

examine the emergence of adult bees over a period of several years.

Fig. 1. Leioproctus plumosus: (a) female (b) male. Scale bars = 5 mm.

Methods

The activity of adult bees was monitored by visual observation of males

flying near the ground and above vegetation in the vicinity of the nesting site

and females returning to nesting tunnels in the ground. Observations were

made over eight seasons from October 2005 to April 2013. No attempt was

made to quantify the number of individuals. One male and three female bees,

plus ten female ichneumonid wasps, were collected for identification. These

specimens have been lodged in the Australian Museum.

Leioproctus plumosus specimens were identified using the male genitalia and

hidden sterna (Maynard 2013). Labium pettitorium (Erichson, 1842)

(redescribed by Turner and Waterston 1920) was identified by the coarse

punctures on the scutum, the strong projections from the propodeum, overall

colour and the number of flagellar segments. Labium inflexum (Morley,

1914), which was transferred to the genus Labium by Townes et al. 1961, is

one of the few yellow Labium species and was identified by the colour

pattern, including infuscation of the wing tips (Morley 1914). Records from

the Australian Bureau of Meteorology automatic weather station number

69138 (Station name: Ulladulla AWS), were obtained from the Bureau’s

internet site (BOM 2013). The station is located 550 m from the nest site.

Results

Each year the bees emerged in late October or early November and

disappeared before the end of December (except in 2012). In six of the eight

years, a second emergence was seen between the middle of January and the

middle of February, although there was some variation in the exact starting

date. In three of the years, a third emergence was seen in autumn (Table 1,

Fig. 4), although on both occasions the number of bees seen was small.

Australian Entomologist, 2014, 41 (1) 9

il | | | l

0 T i T =) 1

Jul Aug Sep Oct Nov Dec Jan ` Feb Mar Apr May Jun

month

Figs 2-3. (2) Location and (3) month collected for Leioproctus plumosus specimens

recorded in Atlas of Living Australia.

10 Australian Entomologist, 2014, 41 (1)

Table 1. Emergence dates and length of active periods for Leioproctus plumosus.

year emerged emerged period | emerged | period

(days) (days)

Ems [umo * | nm [ s | omer r

be | wor «| —

Eme | roa | M | mm | | aan | x |

ms [so | o | are | | -

Dosao | soa | & | S | 9 [2^ |

ems [mw |»]|

uum

rur

€:

£8

30 |

S | |

E 25 | |

Bom 2

? 15 |

å | |

300 e à -nnne má i man = i

250

E 200 |

3. |

ius b

NT. all KAn Mi M VN Au I M | M Mi iu | Ml

Fig. 4. Weather parameters and nesting activity of Leioproctus plumosus: (a) weekly

average maximum temperature; (b) weekly rainfall at Ulladulla AWS; (c) observed

periods of activity at nesting site. Vertical scale indicates presence or absence of bees.

year

Australian Entomologist, 2014, 41 (1)

The mean length of the spring and summer activity periods was 45 days (s.d.

11 days), while the intervals between activity periods were estimated as

differences between either the mid-points of activity or the dates of first

emergence. The former gave an average of 75 days (s.d. 8 days), while first

emergences were separated by an average of 83 days (s.d. 9 days).

Ichneumonid wasps, identified as Labium pettitorium (Erichson) (Fig. 5a),

were frequently observed patrolling the nesting site. The wasps were first

noticed in all years, but their number relative to that of the bees was

noticeably high in 2010/11 and noticeably small in 2012/13. A different wasp

species, Labium inflexum (Morley) (Fig. 5b), was found in another L.

plumosus colony in a garden in the Sydney suburb of Lilyfield in October

2000, demonstrating that L. plumosus may be parasitised by more than one

Labium species.

Fig. 5. Labium species: (a) L. pettitorium female; (b) L. inflexum female. Scale bars =

5 mm.

Discussion

The observations demonstrate that Leioproctus plumosus is unambiguously

multivoltine and individuals have short lifetimes like many other solitary

species (Michener 2007). Consequently, the extended period of activity

shown in Fig. 3 can be explained by the blurring of gaps between generations

by small variations in emergence times at different sites.

No attempt was made to estimate the lifespan of the bees by marking

individuals, but the length of the active periods provides a rough estimate.

Given that most female hymenopterans probably mate only once and

consequently males emerge slightly earlier than females (Alcock et al. 1978),

we estimate that individual bees of this species live for between 30 and 50

days.

Adults emerging in the second period of activity are probably offspring of

bees that emerged in the first period rather than of bees active a year earlier.

Consistent with this interpretation are the observations that (i) summer and

autumn hatchings occurred even when there was only a spring hatching in the

12 Australian Entomologist, 2014, 41 (1)

previous year and (ii) no autumn emergence was seen in the absence of a

summer generation in the same year (Fig. 4).

The time intervals between activity periods ranged from 65 to 90 days, which

is comparable with the development times reported for other species. The

minimum time for adult development of the ground-nesting species Amegilla

pulchra was estimated to be 42-65 days (Cardale 1968a), similar to the 55

days found for Austroplebeia australis (Halcroft et al. 2013). Although A.

australis is eusocial, its larvae develop within sealed cells. For other species,

only the time between pupation and eclosion of the adults has been observed.

For Lithurgus atratiformis the length of this pupal stadium was 22 days

(Houston 1971), for Amegilla paracalva 22 days (Houston 1991) and for

Stenotritus greavesi 51 days (Houston and Thorp 1984). Hence the intervals

between periods of activity exhibited by L. plumosus are not shorter than

those expected between generations.

The occasional failure of the summer and/or autumn generations does not

seem to be correlated with either rainfall or maximum temperature (Fig. 4).

In fact, the bees were particularly numerous in the spring of 2012 and

summer of 2013 despite particularly heavy rain shortly before emergence.

Three weeks before the spring emergence 233 mm of rain fell in 24 hours and

one week before the summer emergence there was a fall of 105 mm.

No obvious correlation was observed between bee emergence and flowers in

bloom at the time. Leioproctus plumosus is known to visit a range of flowers

from at least four plant families (Maynard 2013) and the area within a radius

of 400 m around the nesting site consisted entirely of suburban gardens in

which exotic species were predominant. Furthermore, the bees are active over

a period longer than the flowering time of most plants and must therefore

exploit several flower species. Hence it is unlikely that the emergence times

have become synchronised with the blooming of any particular flower.

A more likely explanation for the variations is that the number of bees is

affected by parasitism, as has been suggested for other species (Dolphin

1979). Although the number of L. pettitorium wasps was not measured, their

number was noticeably high in 2010/11, when the summer and autumn

generations failed to appear and noticeably low in 2012/13, when bee

numbers were greater than average.

Conclusion

Relatively little is known about the life history of Australian bees, yet simple

observations of a nesting site over an extended period of time could provide

useful information not available from accumulated museum records.

Collections usually contain a small number of specimens from each site and

the data are, therefore, averaged over a range of sites and times, leading to a

blurring of information. Citizen scientists could make valuable contributions

(Danielson et al. 2003) to the understanding of our bee fauna.

Australian Entomologist, 2014, 41 (1) 13

References

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Australian Entomologist, 2014, 41 (1); 15-28 15

MODIFICATION OF THE ALL PROTOCOL TO CHARACTERISE

THE OVERALL ANT ASSEMBLAGE IN TEMPERATE EUCALYPT

FOREST

DAVID BRAY

260 Gilloglys Rd, Bulga Forest, NSW 2429

(Email: davidmbray@yahoo.com.au)

Abstract

A standardised sampling procedure for ants, the ALL (Ants of the Leaf Litter) Protocol, was

modified for use in temperate eucalypt forest. Terrestrial samples from leaf litter and pitfalls (the

basic ALL Protocol) were supplemented with arboreal samples from tree traps. Four 200 m

transects were sampled using the modified protocol within a 0.5 ha site on a forested property

located near the mid-north coast of New South Wales. Site vegetation was mixed-age eucalypt

forest, which had been undisturbed for more than 30 years. Tree traps captured a total of 54

species and 19 of these were absent from terrestrial samples. The addition of tree traps to the

basic ALL Protocol: i) increased the number of species detected per species occurrence by an

average of 20% per transect, with almost 2/3 of the increase due to species of Camponotus,

Polyrhachis and Myrmecia; ii) increased the proportion of common species collected in

transects, with less variation in species numbers between transects; and iii) provided a more

comprehensive characterisation of this ant assemblage. Three groups of ants were present in the

assemblage: 1) widespread species, active both in litter and on trees; 2) small litter-dwelling

terrestrial species; and 3) larger species which were captured primarily on trees but were mostly

ground-nesting. The genera of ants detected at the site were largely shared with similar east coast

sites. Voucher specimens were deposited at the Australian Museum and images of the ants are

available by email from the author.

Introduction

Australian ants have been extensively used as bioindicators to monitor

environmental changes (Majer et al. 2004) and also as a target group in

biodiversity surveys (Stanisic ef al. 2005, Burwell and Nakamura 2011,

Callan et al. 2011). However, thorough assessments of ant biodiversity are

time-consuming and often involve specialist entomologists (Andersen ef al.

2002). In an effort to make invertebrate assessments easier to conduct, rapid

Survey procedures have been developed using simplified methods for

sampling, sorting and identification of specimens (Oliver and Beattie 1996).

Andersen et al. (2002) simplified the assessment procedure by sorting only

larger ant species from 12 selected genera and thereby reduced the survey

effort while achieving similar conclusions to those of a more intensive

survey. A rapid sampling procedure, the Ants of the Leaf Litter (ALL)

Protocol, has been proposed for assessing ant assemblages of the forest floor

from samples collected by pitfall trapping and extraction of ants from leaf

litter using mini-Winkler sacks. This standardised sampling methodology

enables direct comparisons between sites and between studies (Agosti and

Alonso 2000). The ALL Protocol was developed for tropical forests, where

distinct assemblages of arboreal and litter ants occur, but there is little

evidence for this distinction in temperate forests (Gotelli et al. 2011). In

Australian eucalypt forests few ant species are known to nest and forage

primarily in trees, but ground-nesting species may forage in trees (Andersen

16 Australian Entomologist, 2014, 41 (1)

and Yen 1992) and some of these species appear to do so preferentially when |

leaf litter is well developed on the ground (Andersen 1995). Ants which

forage largely in trees and strictly arboreal species are both likely to be

under-represented in terrestrial samples collected with the basic ALL |

Protocol. Several ant surveys with a focus on the overall ant assemblage in —

tropical habitats have included tree traps to sample the arboreal stratum |

(Andersen et al. 2006, Andersen et al. 2007).

The present study examined the use of baited tree traps to supplement ground —

samples collected with the basic ALL Protocol in a temperate eucalypt forest |

and thereby provide a less biased description of the overall ant assemblage. A |

relatively undisturbed site with a well developed leaf litter layer was

surveyed using multiple transects and the modified ALL Protocol. Results |

from individual transects were inspected for changes in species richness and

species composition due to the addition of tree-trap samples.

Methods

Study site

The study site was located on a 400 ha property on the eastern fall of the

Great Dividing Range, near the NSW mid-north coast (31.513°S, 152.246°E,

500 m altitude, mean annual rainfall 1150 mm). The property was almost

entirely forested and surrounded by similarly forested lands. The topography

consisted of ridges with moderate slopes. Disturbances over the past century

included cattle grazing, timber harvesting and frequent fires. Prior sampling

had indicated the presence of a relatively rich ant fauna on a sheltered slope

with north-west aspect which had been free from these disturbances for more

than 30 years and this was selected for the study area. The study site was

located mid-slope and limited to 0.5 ha to avoid changes in soil moisture and

ground vegetation on the upper slope and lower slope.

Site vegetation consisted of dry eucalypt forest, with an open mixed-age

canopy dominated by Grey Gum (Eucalyptus propinqua), Grey Box (E.

moluccana) and Forest Red Gum (E. tereticornis). The open understorey was

mostly Black Oak (Allocasuarina littoralis) and the shrub layer was sparse.

The closed ground cover consisted of grasses (Themeda, Entolasia), Matrush

(Lomandra), sedges (Gahnia) and a well-developed leaf litter layer of 2-5 cm

depth. The soil was finely textured, compact and stony, with patches of

surface stones, gravel, logs, branches and debris.

Sampling

Four linear 200 m transects (A-D) were sampled with the modified ALL

Protocol at the study site during January and February, 2011. Successive

transects were displaced at least 10 m to avoid overlap. Sampling stations

were located at 10 m intervals along each transect. Terrestrial samples from

litter and a pitfall, together with an arboreal sample from a tree trap, were

collected at each station (a total of 20 samples per method per transect).

Australian Entomologist, 2014, 41 (1) 17

. Each leaf litter sample was collected from an area of 1 m°, sieved to remove

coarse material and placed in 5 mm mesh plastic containers which were

suspended in mini-Winkler sacks for 48 hours (Bestelmeyer et al. 2000) at

temperatures of 17-38°C. Emergent ants were collected in 80% methylated

spirits. Pitfalls and tree traps were constructed from plastic cups, 65 mm

diameter and 90 mm depth, partly filled with 80% methylated spirits as

collecting fluid and operated for 48 hours. Pitfalls were dug into the ground

and shaded with plastic containers. Tree traps were pinned to a tree trunk at

1.5 m height and baited around the rim with honey (Fig. 1). Observations of

ant nests were recorded as: 1) in the ground or in wood on the ground; or 2)

above ground in dead wood of standing trees.

Fig. 1. Tree trap attached to the trunk of a canopy eucalypt by up

1 | holstery pins,

partially filled with 80% methylated spirits and baited around the rim with honey.

Ants were identified to genus using keys by Shattuck (1999) and CSIRO

(2012), then separated into morphospecies (referred to as species hereafter)

and identified to described species or species-group where suitable keys were

available. Voucher specimens were deposited at the Australian Museum and

images of all morphospecies detected are available by email from the author.

18 Australian Entomologist, 2014, 41 (1)

Data analysis

Performance of the modified ALL Protocol was qualitatively assessed from

the increase in the number of species collected and the change in species

composition in each transect relative to data from the terrestrial samples of

the basic ALL Protocol.

At each sampling station within transects, data from litter and pitfall samples |

were pooled for the basic ALL Protocol and this was pooled with the data

from the tree traps for the modified ALL Protocol (i.e. 20 pooled-method

samples per transect). Incidence data were used as it has been advocated as a |

more appropriate unit of ant biodiversity than the abundance of individual |

ants (Ellison et al. 2007, Gotelli et al. 2011). Incidence (occurrence) was

recorded as the presence or absence of a species in each sample of pooled-

methods data or in each sample collected by individual methods.

The numbers of species and species occurrences were recorded and the

software EstimateS 8.2 (Colwell 2009) was used to compute species

accumulation curves for transect data. Species detected were plotted against

species occurrences rather than samples, to reduce bias due to differences in

ant numbers (Gotelli et al. 2011). The number of species collected in each

transect was compared at a value of species occurrences corresponding to the

least total for any transect. At this value, the number of species in each

transect was estimated, where necessary, by interpolation between points in

the species accumulation data.

Changes in species composition were identified by inspection of the

incidence data collected by the basic and modified ALL Protocols for each

transect.

Results

Species richness

The numbers of species occurrences, genera and species collected were

greatest in litter samples and least in pitfalls (Table 1). Transects varied in the

numbers of species occurrences and species detected, with greater variation

between transects collected with the basic ALL Protocol than with the

modified ALL Protocol (Fig. 2).

Data from tree traps supplemented basic ALL Protocol data by an average of

84 species occurrences, three genera and 14 species per transect (Table 1).

The number of species collected in each transect ranged from 47 to 60 for the

basic ALL Protocol and 61 to 70 for the modified ALL Protocol (Fig. 2),

representing differences between transects of up to 28% and 15%

respectively.

The modified ALL Protocol accumulated more species per species

occurrence than the basic ALL Protocol. Transect B with basic ALL Protocol

yielded 309 species occurrences, the least number for any transect and the

Australian Entomologist, 2014, 41 (1) 19

data for the remaining transects were standardised to this value. At 309

species occurrences, transects with the basic ALL Protocol averaged 51

species and this increased by 20% to 61 species with the addition of data

from the tree traps used in the modified ALL Protocol (Table 1).

Table 1. Species occurrences at the site. Totals for each sampling method are the

combined data for four transects. Average values per transect (based on the four

transects) compare the basic (Bas.) and modified (Mod.) ALL Protocol using: a) raw

data; and b) data standardised to 309 species occurrences.

Species Genera Species

occurrences

Totals

Litter 1216 39 79

Pitfalls 265 24 39

Tree traps 485 25 54

Average/transect

a) raw data

Bas. ALL Protocol 331 31 52

Mod. ALL Protocol 415 34 66

b) standardised to 309 sp. occ.

Bas. ALL Protocol 309 51

Mod. ALL Protocol 309 61

Species

30 -

20 -

10 -

0 100 200 300 400

Species occurrences

Fig. 2. Species accumulation curves (based on species occurrences) for each of the

four transects (A-D) using data from the basic ALL Protocol (Bas.) and modified

ALL Protocol (Mod.).

20 Australian Entomologist, 2014, 41 (1)

Composition of the ant assemblage

Ant species detected at the site are listed in Table 2. The species composition

of the assemblage in terrestrial samples (litter and pitfalls) differed from that

in arboreal samples (tree traps). A total of 54 species was collected in tree

traps (Table 1). Terrestrial samples collected 46 species not captured by tree

traps; another 35 species were present in both strata and 19 species were

captured only in tree traps (Table 2). Ten subfamilies of ants were present in

terrestrial samples, while tree traps contained ants from only five subfamilies

(Table 2). Differences in species composition between ants collected with the

basic and modified ALL Protocols were most conspicuous in the three

subfamilies Formicinae, Myrmicinae and Myrmeciinae. The modification to

the ALL Protocol added totals of 11, four and three species, respectively, to

these subfamilies (Table 2). Twelve of the 19 species unique to tree traps

were from the genera Camponotus, Polyrhachis and Myrmecia (Table 2). The

modification to the ALL Protocol added an average of 8.5 species per

transect from these three genera, representing 61% of the overall increase in

species numbers.

The subfamilies Myrmicinae and Formicinae accounted for 68% of species

collected from transects and 71% of species occurrences. The Myrmicinae

were comprised largely of small species active in leaf litter, with few of these

species active on tree trunks. The Formicinae consisted of a mixture of small

primarily active in the ground litter and large mobile

species which were

n the ground.

species which were active on tree trunks but rarely captured o

Genera which were common in samples from one stratum, but not the other,

included Pheidole, Stigmacros, Hypoponera, Solenopsis and Lordomyrma in

terrestrial samples and Camponotus, Polyrhachis, Myrmecia and

Leptomyrmex on tree trunks. Genera that were common both on the ground

and on tree trunks were Anonychomyrma, Nylanderia, Meranoplus and

Crematogaster. The genus Rhytidoponera included two similar species with

differing habitat preferences: Rhytidoponera victoriae (André) was common

only on the ground, while Rhytidoponera metallica (Smith) was common

both on the ground and on tree trunks.

Forty-four of the collected species were considered common as they occurred

at more than 10% of sampling stations (i.e. incidence greater than 8 in the

Mod column in Table 2). Of these 44 common species, at least 42 (95%)

were collected along each of the four transects with the modified ALL

Protocol, while each transect with the basic ALL Protocol collected at least

36 (82%).

Ant nests were observed on or near the site for 23 of the 54 species active on

tree trunks and nests of 18 of these species were in the ground or in wood

lying on the ground, while those of five species were arboreal in dead

standing trees of Eucalyptus, Acacia and Allocasuarina (Table 2).

Australian Entomologist, 2014, 41 (1) 21

Table 2. Total number of species occurrences (incidence) in 80 samples each from: L,

litter; P, pitfalls; T, tree traps; Bas, basic ALL Protocol; and Mod, modified ALL

Protocol. Nest sites (N) for species active on trees are denoted as: G, in the ground or

in wood on the ground; or A, above ground in standing dead wood.

Species

Amblyoponinae

Amblyopone sp. 1 3 0 0 3 3

Prionopelta robynmae Shattuck 1 0 0 1 1

Stigmatomma sp. 1 1 0 0 1 1

Cerapachyinae

Cerapachys turneri Forel 5 0 0 5 5

Cerapachys larvatus (Wheeler) 4 0 0 4 4

Sphinctomyrmex sp. 1 1 0 0 1 1

Dolichoderinae

Anonychomyrma sp. 1 50 33 33 61 72 G

Anonychomyrma sp. 2 9. 2 1 6 6

Iridomyrmex splendens Forel 7 5 18 10 21 G

Iridomyrmex mayri Forel 3 4 6 6 12 G

Leptomyrmex nigriventris (Guérin) 0 2 18 2 19 A

Ochetellus sp.1 (glaber group) 1 0 0 1 1

Ochetellus sp.2 (glaber group) 1 0 3 1 4 G

Tapinoma sp.1 35 0 2 35 36

Tapinoma sp.2 0 0 1 0 1

Ectatomminae

Rhytidoponera metallica (Smith) I ul 25 25 39

Rhytidoponera victoriae (André) 56 36 3 63 63 G

Formicinae

Acropyga myops Forel 1 0 0 1 1

Camponotus aeneopilosus Mayr 3 2 24 4 28 G

Camponotus elegans Forel 1 0 46 1 46 G

Camponotus sp.6 (intrepidus group) 0 0 26 0 26 G

Camponotus sp.7 (? humilior) 0 0 21 0 21 G

Camponotus sp. 9 (near elegans) 0 0 3 0 3

Camponotus sp.12 (sponsorum group) 0 0 1 0 1

Camponotus macrocephalus Erichson 0 0 1 0 1

Melophorus sp. 1 1 0 0 1 1

Melophorus sp. 2 2 0 1 2 3 G

Notoncus capitatus Forel 2 A d 33 37

Nylanderia sp. 1 0 0 5 0 5 A

Nylanderia sp. 3 58 3

Paraparatrechina sp. 2 (minutula group) 38 3

Paraparatrechina sp. 4 (minutula group) 1

22 Australian Entomologist, 2014, 41 (1)

Polyrhachis sp. 2 M 1 0 1 G

Polyrhachis phryne Forel 2 0 26 2 27

Polyrhachis sidnica Mayr 0 0 5 0 5

Polyrhachis sp. 11 1 0 3 1 4 A

Polyrhachis sp. 15 0 0 1 0 1

Polyrhachis sp. 16 (0 (0 3 0 3

Prolasius sp. 1 14 1 5 15 19

Prolasius sp. 2 AW 3 2 54 54

Prolasius sp. 3 44 6 7 44 46

Prolasius sp. 4 2 1 0 3 3

Prolasius sp. 5 2 0 0 2 2

Pseudonotoncus hirsutus Clark 0 0 1 0 1

Stigmacros sp. 1 19 0 0 19 19

Stigmacros sp. 2 12 0 7 12 17 G

Stigmacros sp. 4 24 0 0 24 24

Stigmacros sp. 5 38 1 0 38 38

Stigmacros sp. 6 1 0 1 1 2

Stigmacros sp. 8 18 1 0 18 18

Heteroponerinae

Heteroponera sp. 1 (imbellis group) 2) $5 0 33 33

Myrmeciinae

Myrmecia nigrocincta Smith 1 ] 12 2 14 G

Myrmecia brevinoda Forel 0 (0 3 0 3 G

Myrmecia sp.3 (gulosa group) YY — (9 1 0 1

Myrmecia fulvipes Roger 1 0 0 1 1

Myrmecia sp.9 (mandibularis group) 0 0 1 0 1

Myrmicinae

Carebara sp. 1 22 EN] 0 23 23

Colobostruma alinodis Forel 1 1 0 2 2

Colobostruma lacuna Shattuck 1 0 0

Crematogaster sp. 1 a. m E

Crematogaster sp. 2 vb i 2

Crematogaster sp. 6 3 0 3

Epopostruma wardi Shattuck 0 0 1

Lordomyrma sp.1 47 0 0

Mayriella spinosior Wheeler 5 3 0

Mayriella sp. 2 (near abstinens) 2 0 4

Mayriella abstinens Forel 1 0 0

Meranoplus sp. 1 43 26 17

Mesostruma browni Taylor 10 0 0

Monomorium rubriceps Mayr 0 0 1

Australian Entomologist, 2014, 41 (1) 23

Species L Bas Mod N

Monomorium tambourinensis Forel 40 1 0 40 40

Monomorium sydneyense Forel 3 0 4 3 7

Monomorium fieldi Forel 7 5 4 11 14 A

Monomorium sp. 7 (? sydneyense) 1 1 0 2 2

Monomorium leae Forel 1 0 0 1 1

Orectognathus phyllobates Brown 1 0 0 1 1

Orectognathus antennatus Smith 0 0 1 0 1 G

Orectognathus rostratus Lowery 7 0 0 7 7

Orectognathus sp. 5 (? clarki) 2 0 5 2 7

Pheidole sp. 1 55 35 4 65 66

Pheidole sp. 2 63 14 0 66 66

Pheidole sp. 4 1 0 0 1 1

Pheidole sp. 6 2 0 0 2 2

Pheidole sp. 7 0 1 2 1 3

Podomyrma sp. 2 0 0 1 0 1

Solenopsis sp. 1 53 12 0 55 55

Solenopsis sp. 2 2 Dp 0 4

Solenopsis sp. 3 2p) di 0 23 23

Strumigenys perplexa Smith 33 0 0 33 33

Strumigenys sp. 2 11 0 0 11 11

Tetramorium confusum Bolton 6 1 4 7 11

Ponerinae

Hypoponera sp. 1 7 S 0 77 77

Hypoponera sp. 2 3 0 0 3 3

Hypoponera sp. 3 1 0 0 1 1

Leptogenys sp. 1 2 1 0 3 3

Pachycondyla sp. 1 12 oj 0 13 13

Pachycondyla sp. 2 l 0 0 1 1

Ponera leae Forel 5 0 0 5 5

Proceratiinae

Discothyrea sp. 1

Discothyrea sp. 2

Discussion

Evaluation of the modified ALL Protocol

In this study the ant assemblage in a temperate eucalypt forest was sampled

with the terrestrial sampling methods of the basic ALL Protocol

supplemented by arboreal samples collected with baited pitfall traps on tree

trunks. The modification to the ALL Protocol yielded a modest increase in

the number of species detected, a clearly identifiable change to the species

24 Australian Entomologist, 2014, 41 (1)

composition of the ants collected in each transect, and less variable estimates

of species richness and species composition.

The addition of tree traps increased the number of species collected in each

transect and some of this increase can be attributed to the extra species

occurrences generated by the tree traps. However, when the data were

standardised to species occurrences the modified ALL Protocol collected an

average of 20% more species per transect, indicating that it accumulated

species more efficiently than the standard ALL Protocol despite the increase

in sampling effort. The additional field time required for the modified ALL

Protocol was minimal as the tree traps were installed and operated

concurrently with the ground pitfalls.

The composition of the ant assemblage collected with the standard ALL

Protocol was partly altered by the addition of tree traps. Substantial changes

were confined to the subfamilies Formicinae and Myrmeciinae, in which the

number of species collected increased by 50% and 250% respectively. The

additional species were mostly from Camponotus, Polyrhachis and

Myrmecia, genera which were infrequently present in the unbaited pitfall and

litter samples collected with the standard ALL Protocol. These results appear

to support the observations of Andersen (1995) that subordinate

Camponotini, such as Camponotus and Polyrhachis, avoid well developed

litter by foraging in trees. However, these ants may have been present and

active on the ground, but not readily collected by unbaited pitfalls in the well

developed litter cover at the site. Ants are more likely to be captured in

pitfalls when the surrounding ground cover has a relatively open structure

(Melbourne 1999). Dense litter cover reduces pitfall capture rates

(Bestelmeyer et al. 2000) and Andrew et al. (2000) found that Camponotus

was more common on the ground at burnt rather than unburnt forest sites.

The use of bait may also influence the capture rate for pitfalls as Romero and

Jaffe (1989) captured more ant species in savanna habitats when pitfalls were

baited with meat, although Wang et al. (2001) reported that unbaited pitfalls

were more effective in temperate oak forest. Greenslade and Greenslade

(1971) collected more Camponotus ants when pitfalls were baited with syrup.

Thus, the probability of detecting any Camponotus species that is active on

the ground may improve if honey bait is added to the ground pitfalls used in

the basic ALL Protocol. Other baits, such as meat or fish, have potential to

collect additional species as Kaspari and Yanoviak (2001) found that canopy

ants in a tropical forest preferred meat baits to sugar baits.

Although individual transects collected with the modified ALL Protocol did

not detect all species collected at the site they captured a greater proportion of

the common species than transects with the basic ALL Protocol. The number

of species per transect differed less between transects collected with the

modified ALL Protocol than between those collected with the basic ALL

Australian Entomologist, 2014, 41 (1) 25

Protocol. Together, these results indicated that the modified ALL Protocol

provided less variable estimates of species richness and species composition

than the basic ALL Protocol.

For studies which aim to maximise the species inventory of an overall ant

assemblage, the improved sampling efficiency of the modified ALL Protocol

should outweigh the slight increase in field time required to implement the

tree traps.

The ant assemblage

Features of the ant assemblage at the study site were the large number of

species active on tree trunks and the proportion of these which were under-

represented in terrestrial samples. Previous studies indicated limited and

patchy arboreal ant activity in southern Australian eucalypt forests (Majer

1990), although 37 species were recorded as active in eucalypt canopies in

New South Wales (Majer et al. 2000) and 44 species were detected in the

canopy of mallee eucalypts in northwestern Victoria (Andersen and Yen

1992). In the present study, 54 species were present in tree trap samples and a

third of these were absent from the ground samples collected with the basic

ALL Protocol.

The overall assemblage was comprised of three groups of species: 1) those

widespread and common in both strata; 2) those primarily in terrestrial

samples; and 3) those primarily in arboreal samples. Species in the

widespread group included ants in a range of sizes from the subfamilies

Dolichoderinae, Formicinae, Ectatomminae and Myrmicinae and they were

from functional groups described by Andersen (1995) as comprising species

which are usually abundant and unspecialised. The terrestrial group was the

most diverse and consisted mainly of small cryptic species of Formicinae,

Myrmicinae and Ponerinae, which are typical of habitats with well developed

leaf litter (Andersen 1986, 1995, Hoffmann and Andersen 2003). The

arboreal group was characterised by larger species of Formicinae and

Myrmeciinae, which have been reported to be predominantly ground nesting

(Andersen and Yen 1992). Within this group only five of the 23 nests

observed during the present study were located above ground. Although

ground nests of Camponotus were found up to 10 m distant from canopy

trees, these ants were uncommon or rare in ground samples but more

commonly collected on tree trunks. The arboreal group represented a

substantial proportion of the species present and the combination of terrestrial

and arboreal samples provided a more comprehensive inventory of the ant

assemblage at this eucalypt forest site.

The genera found at the site were similar to those found elsewhere in eastern

Australian eucalypt forests, with the majority also occurring 40 km to the east

at Bulls Ground State Forest (York 2000, Andrew ef al. 2000) and at forest

26 Australian Entomologist, 2014, 41 (1) |

sites near Brisbane (Stanisic et al. 2005). However, shared genera can mask |

differences in species composition and to facilitate comparisons of species.

composition, Callan et al. (2011) retained reference specimens and provided

online images of all ant species detected in their study at Barrow Island.

Ecological studies typically employ small plots and generate data which may

not be immediately applicable at larger scales (Andersen 1997). However, the

use of standardised sampling, as in the modified ALL Protocol, enables

baseline data to accumulate from successive studies and provides the

potential for comparative analysis at both local and regional scales. The

results of this study support the use of multiple sampling methods for

biodiversity assessment in order to offset the bias of individual methods and

to improve detection rates for species which utilise more than one habitat.

Acknowledgements

I would like to thank Derek Smith (Australian Museum) for assistance with

ant identification and the referees for their valuable advice during preparation

of the manuscript.

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Australian Entomologist, 2014, 41 (1); 29-44 29

TWO NEW SPECIES OF ABANTIADES HERRICH-SCHAFFER

(LEPIDOPTERA: HEPIALIDAE) FROM WESTERN AUSTRALIA

M.D. MOORE! and E.D. EDWARDS?

!South Australian Museum, Adelaide, SA 5000

^CSIRO Ecosystems Sciences, GPO Box 1700, Canberra, ACT 2601

Abstract

Abantiades lineacurva sp. n. and Abantiades argentangulum sp. n. are described, illustrated and

compared with other Australian species of Hepialidae.

Introduction

The genus Abantiades Herrich-Schaffer is endemic to southern Australia,

with the northernmost occurrence listed by Tindale (1932) as Rockhampton,

Queensland. The genus was last revised by Tindale (1932), who included 14

Species, six described as new. Tindale’s revision formed the basis of the

works by Nielsen (1996), Nielsen et al. (2000), Edwards (2007) and Kallies

and Douglas (2008). Since 1932, nothing of significance has been published

on this genus and no new species have been described.

The genesis of this paper was a reorganisation of the South Australian

Museum hepialid collection, which contains approximately 500 specimens of

Abantiades, including all the described species. Curation revealed specimens

of two unnamed species from Western Australia. Subsequent examination of

the hepialid collection in the Australian National Insect Collection revealed

two more unnamed Western Australian species.

In this paper we describe two new species from these collections.

Terminology used for the genitalia follows Dugdale (1994); that of the wing

venation follows Kristensen (1998).

Abbreviations: ANIC — Australian National Insect Collection, Canberra;

CSIRO — Commonwealth Scientific and Industrial Research Organisation;

BMNH - The Natural History Museum, London, UK; SAMA — South

Australian Museum, Adelaide; WAM — Western Australian Museum, Perth.

Systematics

Tindale (1932) described Abantiades thus: ‘Antennae unipectinate often

broad and lamellate in males, reduced in females. Labial palpi three-

segmented, first and second segments approximately equal, apical one short,

about as long as wide and subspherical, maxillary pair reduced, forming ill-

articulated protuberances at base of labium. Forewings R1 separating from Rs

before the branching of R5; R4 and R5 forked. Hindwings with R4 and R5

branching before the radio-median cross-vein. The type species is A.

hyalinatus Herrich-Scháffer, 1853, designated by Kirby in 1892.

30 Australian Entomologist, 2014, 41 (1) ,

Abantiades lineacurva sp. n.

(Figs 1-7).

Types. Holotype á, WESTERN AUSTRALIA: Kojonup, 18.iv.1960, Wallace, in

ANIC. Paratypes: 21 63, 6 99, WESTERN AUSTRALIA: 1 3, 1 9, Kellerberrin,

14.v.1939, Tindale; 1 3, Kojonup, 6.iv.1960, Beresford, in SAMA; 1 3, Kojonup,

11.iv.1961, Priest; 1 J, Kojonup, 11.iv.1962, Peters; 1 3, 1 9, Nedlands, 22.iv.1964,

21.iv.1964, Wallace; 15 $3, 2 99, Kojonup, 10.iv.1963, 13.iv.1963, 17.iv.1963,

21.iv.1963, 22.iv.1963, 23.iv.1963, 14.iv.1964, 16.iv.1964, 17.iv.1964, 19.iv.1964,

21.iv.1964, 4.v.1964, 12.iv.1965, 19.iv.1966, 8.v.1967, Rogers; 19, Bakers Hill, |

9.v.1968, I. Southey, in ANIC; 1 d, Jacup, 1.v.1914, W.B. Alexander, 33?46'S

119°17°E, in WAM.

Additional material examined. WESTERN AUSTRALIA: 1 9, Katanning, 1879; 1 3,

Hale, 8.vii.1872, Bailey; 2 $$, Jacup, v. 1914; 1 ĝ, Jacup, v.1919; 1 3, Carlingup, v.

1919, in WAM.

Diagnosis. Male and female: antennae paler at tip with broad flattened rami;

forewing dark brown with a prominent ‘T’ shaped silver-white marking

which does not reach apex, without intricate scroll-like pattern; hind wing

dark brown, only slightly paler towards base.

Description. Male (Fig. 1). Head: proboscis absent; labial palpi three

segmented, apical one short sub-spherical, narrow, directed forwards;

antennae red brown, paler towards tip, short, about a third length of costa,

unipectinate, rami in middle about 2.0-2.5 x shaft, sub-elliptical in shape,

covered in very short fine cilia (Fig. 7), leading edge flattened; frons and

vertex with dark brown hair scales. Thorax: densely covered in dark brown

hairs, legs dark brown. Abdomen: densely covered in dark brown hairs.

Forewing: length 30-34 mm, costa straight, gently rounded towards tip,

termen and dorsum evenly and continuously rounded; upperside dark brown

with silver markings and without intricate scroll-like pattern, a fine silver

subterminal disjointed line from Rs2 to below CuAl, a parallel silver

disjointed line subapically from Rs1 to postmedially below CuA1, from the

middle of this line there extends a long longitudinal angularly undulating line

to the costal vein near base, a short disjointed silver line, sometimes obsolete,

runs from Rs1 inwards and not reaching Rs3, cilia dark brown. Hindwing:

costa slightly arched, apex rounded, termen and dorsum evenly rounded, dark

brown, slightly paler basally, cilia dark brown; underside of both wings dark

brown with muted silver pattern showing through forewing.

Genitalia. (Figs 2-4). Pseudotegumen heavily sclerotised, wedge shaped,

ventral apex projected rearwards in a strong curve, tip obtuse, anterior vertex

*beak-like', anterior margin somewhat serrate, posterior margin less. Each

side of pseudotegumen curves outwards then narrows as it rises towards

apex, lateral edge curves outwards at apex producing a mild horned effect in

most specimens. Twin processes short, set well back. Valva short, paddle

Australian Entomologist, 2014, 41 (1) 31

shaped, dorsal edges more heavily sclerotised. Trulleum sclerotised, roughly

trapezoid in shape with anterior apices lobed. Juxta; weakly bilobed, weakly

sclerotised. Saccus broadly ‘V’ shaped, posterior edge heavily sclerotised

‘with two acute points at its midline. Sternite 8; posterior margin with sizeable

concave notch.

Fig. 1. Adult male of Abantiades lineacurva sp. n.: upperside

pst _- Ventral

apex

Figs 2-4. Male genitalia of Abantiades lineacurva sp. n.: (2) lateral, (3) ventral, (4)

ventrolateral. j = juxta; pst = pseudotegumen; s8 = sternite 8; sa = saccus; tpr = twin

processes; tr = trulleum; va = valva.

32 Australian Entomologist, 2014, 41 a)

Fig. 6. Female genitalia of Abantiades lineacurva Sp. n.: ventral view. dp — dorsal

piate; mp = median piece; sp = side piece. Ductus bursae and corpus bursae not

shown.

Australian Entomologist, 2014, 41 (1) 33

| Female (Fig. 5). Head: proboscis and palpi as male; antennae as male but

shorter, about one-sixth length of costa, unipectinate, rami mid antenna about

| 1.5 x shaft, strongly flattened and subelliptical, covered in very fine short

cilia (Fig. 7). Thorax as in male. Abdomen densely covered in dark brown

| hairs. Forewing: length 50-70 mm, colour and pattern similar to male but

more elongate. Hindwing similar to male but more elongate.

Genitalia (Fig. 6). Anogenital field wider than high. Dorsal plate lobes

triangular, setose, lightly sclerotised. Median piece distinctly bi-cuminate,

| Setose, heavily sclerotised. Side pieces sub-elliptical.

JMLO we

ag bo (9c Og ©9

Fig. 7. Antennal segments of male (left) and female (right) of Abantiades lineacurva

Sp. n.: (a) terminal ramus, (b) mid rami-lateral view, (c) mid ramus.

Variation. The extent and detail of the silver markings on the forewings vary

as in other species of Abantiades.

Etymology. lineacurva (Latin) — curved line referring to the longitudinal,

angularly undulating silver line that is the most conspicuous forewing mark.

Distribution. Most specimens were collected at the CSIRO light trap at

Kojonup. It is also known from the Perth area, Bakers Hill in the eastern

Darling Range, Kellerberrin in the wheat belt east of Perth, and Jacup and

Carlingup in the southern mallee (Fig. 8).

Comments. Abantiades lineacurva is the only Western Australian species that

has grey-brown fore and hind wings and a prominent, single white line on the

forewing. Abantiades albofasciatus Swinhoe, 1892 also has a single major

white stripe on its forewing but it is wider, angled rather than curved, and

reaches the apex. Abantiades albofasciatus also has intricate scroll-like

markings on the upperside of the forewing and a hind wing that is white.

Abantiades aurilegulus Tindale, 1932 is golden-brown rather than dark

brown in colour and has two prominent lines on its forewing.

34 Australian Entomologist, 2014, 41 (1)

Kellerberrin

* Bakers hill

Hale

Nedlands

Carlingup

Kojunup

Fig. 8. Capture sites of Abantiades lineacurva sp. n.

. Abantiades argentangulum sp. n.

(Figs 9-16)

Types. Holotype 6, WESTERN AUSTRALIA: Yanchep National Park, 5 miles north

of Yanchep, 12.iv.1968, I. Common and M. Upton, in ANIC. Paratypes: 7 53,2 99,

WESTERN AUSTRALIA: 1 c, Kojonup, 8 iv 1964 A. L. Rogers, in SAMA; 2 33,

1 9, Konjunup? 2-11.iv.1962, A.L. Peters; 3 34, 21 miles west of Kojonup,

28.11.1968, I. Common and M. Upton; 1 d, Pithara, 25.iv.1968; 1 ĝ, Peak Charles

NP, 16.iv.2007, A. Zwick and C. Cocking; 1 9, Coolgardie Goldfield Woodlands NP,

17.iv.2007, A. Zwick and C. Cocking, in ANIC; 1 ĝ, 1 9, Dryandra State Forest,

12.8 km SE of Caballin, 5.iv.1984, R.P. McMillan, in WAM.

Additional material examined. WESTERN AUSTRALIA: 4 cd, Lake Magenta

Reserve, 17 m E. Pingrup, 4.iv.1971, D. Kitchener and L. Smith; 1 $, Lake Magenta

Reserve, iv.1971, L. Smith; 2 $$, Dryandra State Forest, 12.8 km S.E. of Caballing,

5.iv.1984, R.P. McMillan; 1 $, 20 m N.E. of Corrigan, 15.iv.1963, A.C. Morton; 1 3,

Gooseberry Hill, v.1970, E. Mullins; 1 c, Valema Farms, Corrigan, iii.2001—2002

Insect Survey, M. Golding and I. Pitman; 1 ĝ, Gosnells, 21.iv.1939, W. Stevens; 1 á,

Fitzgerald River National Park, 9-11.iv1982, T. Houston; 1 6, Dryandra State Forest,

iv.1972, A. Burbridge; 1 $, Glen Forest, 16.iv.1974, I. Lyon; 1 6, Carlingup,

3.vii.1914, W.B. Alexander; 1 4, West Perth, 1.iv.1937, A. Holder; 1 ĝ, Shackleton,

Australian Entomologist, 2014, 41 (1) 35

31.v.1922, L. Morrison; 1 4, no data, M. Powell Collection; 1 9, Cheritons Find,

11.iv.1971, B. Evans, all in WAM.

Diagnosis. Male: antennae with broad flattened spade shaped rami; forewing

dark grey-brown with two prominent, silver-white, triangle shaped markings;

hind wing mid brown, more yellowish towards base. Female: antennal rami

absent; forewing grey with scroll pattern.

- Fig. 9. Adult male of Abantiades argentangulum sp. n.: upperside.

Description. Male (Fig. 9). Head: proboscis absent; labial palpi three

segmented, apical segment short, subspherical, directed anteriorly; antennae,

65-73 segments, dark chestnut-brown, short, about a third length of costa,

unipectinate, rami mid filament 3 x shaft, strongly flattened, at base sub-

elliptical, mid antenna spade shaped, at termen near triangular, densely

covered in short fine cilia (Fig. 16); frons, covered in long grey hairs. Thorax,

covered in long brown hairs; legs, covered in stiff hairs; forelegs, femur inner

yellow, outer black, edge white; tibia, brown and yellow; tarsi, grey tipped

white; mid legs, all grey tipped white; hind legs yellow. Abdomen covered in

long dark, grey-brown hairs.

Forewing: length 52-60 mm, costa straight gently rounded towards tip, tip

acute, termen and dorsum continuously gently curved giving wing a narrow

lanceolate shape, upperside, pale brown, darker at base; two large silver-

white markings, one thin and lanceolate from near tip to below CuAI

approximately parallel to termen, second triangular with vertices at the origin

of M3, join CuA1 with M3 and 0.4 distance along Rs4, both edged dark

36 Australian Entomologist, 2014, 41 (1)

brown, between, a sparse line of small circles edged mid brown; slight

scrolling near dorsum, fine mid brown lines fill other spaces; close to termen

a thin off white line runs between Rs4 and Al; underneath, light brown

tending darker at the costa, coating of yellowish hairs to the basal part,

Hindwing: costa straight, curving towards tip, tip obtuse, termen and dorsum

evenly rounded; upperside light brown grading to yellowish buff at base;

underside light brown, yellowish hairs at base.

Genitalia (Figs 10-12). Pseudotegumen; heavily sclerotised, ventral apex

acute, triangular, with slight posterior projection, tip obtuse, anterior apex

heavily hooked, anterior margin somewhat concave, posterior sinuously

curved, thickened, bumpy. Twin process; short, strongly pointed. Valva;

‘arms’ short, broad at base, rugose, curved and sclerotised along ventra]

surface. Trulleum; large, lightly sclerotised, angled posteriorally, anterior

apices lobed. Juxta; weakly sclerotised, large, posterior roundly bilobed,

anterior lateral acute lobes. Saccus; broadly ‘V° shaped, curved anterior apex,

posterior with two sclerotised points along midline. Sternite 8; posterior

margin gentle convex curve.

Ventral

apex

—tpr

Figs 10-12. Male genitalia of Abantiades argentangulum sp. n.: (10) lateral, (11)

ventral, (12) ventro-lateral. j = juxta; pst = pseudotegumen; s8 = sternite 8; sa =

saccus; tpr = twin processes; tr = trulleum; va = valva.

Female (Fig. 13). Head: proboscis and palpi as male, antennae dark chestnut-

brown, short, one fifth costa, non pectinate, rami reduced to bumps that abut,

producing an ongoing series of undulations along the length of the filament,

well covered with short white setae over the entire surface, some longer setae

on underside of filament (Fig. 16). Thorax covered in long brown hairs; legs

mid to dark brown. Abdomen as male. Forewing: 77-90 mm, similar to male

except the silver-white markings are replaced with subdued, non-contrasted,

pale grey-brown areas, the spaces between filled with scroll-like markings.

Hindwing: upperside similar to male but paler; underside light brown.

| Australian Entomologist, 2014, 41 (1) 37

Fig. 13. Adult female of Abantiades argentangulum sp. n.: upperside.

Genitalia (Figs 14-15). Anogenital field wider than high. Dorsal plate; deeply

cleft, lobes distinct, pear-shaped, setose, lightly sclerotised. Median plate,

shape distinctly mound-like, setose, heavily sclerotised. Side plates roughly

triangular. Intergenital lobe wide trapezoidal. The bursae copulatrix occupies

the entire length of the female abdomen. Ductus bursae and corpus bursae of

about equal length, diverticulum at intersection.

Etymology. argentangulum (Latin) — referring to the angular silver marking

prominent on the forewing of the male.

Distribution. Widely distributed over the southwestern part of WA (Fig. 17).

Comments. Abantiades argentangulum is easily separated from all other

Western Australian Abantiades species: males by the striking silver-white

triangles on the forewings, females with rami absent from antennae. This

species has been collected extensively in the past and has for many years

been referred to as the ‘WA magnificus’. It has only now been studied in

sufficient depth to establish its specific status.

Biology

We know nothing significant of the biology of either species, although

rainfall data for the capture sites and times of the above mentioned specimens

indicate that emergence of this species is linked to autumn and winter frontal

rainfall events, similar to that known for other hepialid species.

38 Australian Entomologist, 2014, 41 (1)

igl

CER db

‘E —— div

IS T

Figs 14-15. Female genitalia of A. argentangulum sp. n.: (14) anogenital area; Ventral

view, (15) bursae copulatrix. dp = dorsal plate; igl = intergenital lobe; sp = side piece;

mp = median piece; db = ductus bursae; cb = corpus bursae; div = diverticulum.

Australian Entomologist, 2014, 41 (1) 39

()c (bo

(a) g?

Fig. 16. Antennal segments of male (left) and female (right) of Abantiades

argentangulum sp. n.: (a) terminal ramus, (b) mid rami-lateral view, (c) mid ramus.

PN lC EN

Pithara

Coolgardie

E .

Yanchep NE 5 Cheritons

Perth Shackleton S

Gen Fo t "C

en Forres "orrigi

Goosebu Hill IMS

Gosnells ^

Cablling Peak Charles

A NP

Carlingup

‘Pingrup

Kojunup

Fig. 17. Capture sites of Abantiades argentangulum sp. n.

Discussion

Abantiades lineacurva is similar to another Western Australian species,

Bordaia moesta Tindale, 1932 in size, shape, colour and markings but differs

in its appearance, antenna and genitalia. When describing Bordaia Tindale

(based on the type species B. pica Tindale), Tindale (1932) noted that it has

an appearance like Oxycanus Walker but venation like Abantiades, where R4

40 Australian Entomologist, 2014, 41 (1)

and R5 on both wings were stalked, whereas in Oxycanus R5 arises from the

stem of Ry+3+4. Unlike Abantiades, which has unipectinate antennae, Bordaia

has bipectinate antennae.

Bordaia moesta was known for many years only from the holotype in SAMA

but recently a further male has been discovered in the WAM collection. Both

specimens have yellow bipectinate antennae, whereas A. /ineacurva has deep

reddish brown, unipectinate antennae.

Although the overall structure and form of the genitalia of A. lineacurva and

B. moesta are similar, there are differences in the structures of various parts

(Figs 2-4 and 18-20). In A. lineacurva the genitalia are more heavily

sclerotised, the valva are broader and thicker, the trulleum smooth in outline

with diverging anterior projections, the posterior margin of the juxta is

distinctly linear, the ventral projection of the pseudotegumen is curved,

acutely angular and backward pointing, and the twin processes much longer.

The notch in S8 also differs. The anterior projections on the trulleum of B.

moesta have an interesting spear-pointed shape.

pst Ventral

apex

pst

Figs 18-20. Male genitalia of Bordaia moesta: (18) lateral, (19) ventral, (20) ventro-

lateral: j = juxta; pst = pseudotegumen; s8 = sternite 8; sa = saccus; tpr = twin

processes; tpr = trulleum; va = valva.

The close structural similarity of the male genitalia of Abantiades and

Bordaia do, however, hint at a closer similarity than the antennal differences

would indicate. A careful study of the generic limits of Bordaia and

Abantiades is needed to clarify the position of these genera and this is being

undertaken by Thomas Simonsen (BMNH) (pers. comm).

Abantiades argentangulum most resembles the eastern Australian A.

magnificus (Lucas, 1898). The similarities in the two species indicate a close

relationship. Males of both species have large, silver-white triangles on their

forewings but in 4. argentangulum the silver-white markings are relatively

larger with a smaller area of dark brown scales surrounding them. Hind wings

Australian Entomologist, 2014, 41 (1) 41

on A. argentangulum are yellowish and distinctly lighter than the forewing,

whereas in A. magnificus the hind wings are grey-brown and darker than the

forewings. Antennae: in A. argentangulum the rami are larger, 3.0 x the

filament width and, although not bilateral, are distinctly spade shaped,

whereas in A. magnificus the plates are smaller, 1.5 x the filament width and

orbicular to lanceolate in shape. Females of the two species can only be

separated using genital structures.

Comparison of the genitalia shows specific differences. In males (Figs 10-12

and 21-23), the slope of the posterior margin of the pseudotegumen in A.

argentangulum is curved but distinctly stepped and steeper in decline in A.

magnificus, the trulleum in 4. argentangulum is longer with anterior

projections present but pentagonal shaped and lacking projections in A.

magnificus; the juxta in A. argentangulum has apical projections, is

proportionally larger and has the posterior margin lobed but in A. magnificus

is without apices, is proportionally smaller and lacking lobes on the posterior

margin; valves with ‘arms’ relatively shorter in 4. argentangulum.

Ventral

pst—

Figs 21-23. Male genitalia of Abantiades magnificus (21) lateral, (22) ventral, (23)

ventro-lateral: j = juxta; pst = pseudotegumen; sa = saccus; tpr = twin processes; tr —

trulleum; va = valva.

In females (Figs 14-15 and 24-25), differences can be found in the

morphology of the dorsal plate (lobes in A. magnificus are more evenly

rounded than in A. argentangulum) and of the bursae copulatrix (A.

argentangulum has a diverticulum, A. magnificus does not).

Abantiades argentangulum is widespread in southwestern Australia, living in

a wide variety of open woodland types. Abantiades magnificus is from

eastern Australia, ranging from the New England area of NSW along the

great divide to Moe and the Grampians. Neither species has been recorded in

South Australia. On the basis of distinct morphological differences in overall

appearance (in males), antennal shape (in males), genitalia structures, their

geographical separation and their differences in habitat preference, we

believe that 4. argentangulum and A. magnificus are separate species.

42 Australian Entomologist, 2014, 41 (1)

F- re E

UN cb

| M. 25

N uj

Figs 24-25. Female genitalia of Abantiades magnificus. (24) anogenital area, (25)

bursae copulatrix. dp = dorsal plate; sp = side plate; mp = medial plate; igl =

intergenital lobe; db = ductus bursae; cb = corpus bursae.

Australian Entomologist, 2014, 41 (1) 43

Figs 26-30. Ventrolateral views of pseudotegumen; (26) Abantiades lineacurva, (27)

Bordaia moesta, (28) A. argentangulum, (29) A. aurilegulus, (30) A. hydrographus.

Remarks

When working with older specimens of Australian hepialids, care is required

when describing colour differences. The appearance of specimens when first

caught may be brighter and more colourful, with some colours fading or

disappearing as the specimen ages. Irrespective of the age of the specimen,

differences outlined above hold true.

In the genus Abantiades there seems to be two forewing shapes exhibited by

the various species: longer, more lanceolate forewings such as in Abantiades

argentangulum and a broader forewing as exhibited by A. lineacurva.

Identifying Abantiades species without dissection

Sometimes it is convenient not to dissect the male genitalia, in which case

parts of the pseudotegumen may be seen by brushing away some scales

around the genital area (Figs 26-30). Tindale (1935) used this technique in

studying Oxycanus. It has the disadvantage that important characters may be

hidden; the degree of extrusion of the male genitalia in different specimens

will differ and so will be more or less visible.

Acknowledgements

We thank Peter Hudson for suggesting the project, his ongoing support,

patience and forbearance in all things and for critically reading the

manuscript, Chris Watts for his support, detailed reading and critical analysis

of our writing, You Ning Su for his instruction on the use of the montaging

camera and help and support with the imaging and computer graphics and

Howard Hamon for his construction of the final images and distribution

maps, Thanks must also be given to the staff of SAMA, WAM and ANIC for

access to specimens and their equipment.

44 Australian Entomologist, 2014, 41 (1)

References

DUGDALE, J.S. 1994. Hepialidae (Insecta: Lepidoptera.), Fauna of New Zealand 30: 1-164.

EDWARDS, E.D. 2007. Australian Faunal Directory. Hepialidae. URL: http//www.environment.

gov.au/biodiversity/abrs/onlineresources/fauna/afd/taxa/HEPIALIDAE

KALLIES, A. and DOUGLAS, F. 2008. Checklist of the Victorian ghost moths (Lepidoptera,

Ditrysia, Hepialidae). Victorian Entomologist 38(4): 50-56.

KRISTENSEN, N.P. 1998. The Homoneurous Glossata. In: Kristensen, N.P. (ed.), Lepidoptera,

moths and butterflies Vol. 1, evolution, systematics and biogeography. Handbook of Zoology 4

Arthropods; Insects Part 35. Walter de Gruyter, New York.

NEILSEN, E.S. 1996. Hepialidae s./at. Pp 24-26, in: Neilsen, E.S. Edwards, E.D. and Rangi,

T.V. (eds), Checklist of the Lepidoptera of Australia. Monographs of Australian Lepidoptera

Vol. 4. CSIRO Publishing, Collingwood.

NEILSEN, E.S., ROBINSON, G.S. and WAGNER, D.L. 2000. Ghost moths of the World: a

global inventory and bibliography of the Exoporia (Mnesarchaeoidea and Hepialoidea)

(Lepidoptera). Journal of Natural History 34: 823-878.

TINDALE, N.B. 1932. Revision of the Australian ghost moths (Lepidoptera, Homoneura,

Family Hepialidae) Part 1. Records of the South Australian Museum YV(4): 510 -536.

TINDALE, N.B. 1935. Revision of the Australian ghost moths (Lepidoptera, Homoneura,

Family Hepialidae) Part 3. Records of the South Australian Museum V(3): 280-331.

| Australian Entomologist, 2014, 41 (1): 45-54 45

AN ANNOTATED KEY TO THE RIOXA COMPLEX OF GENERA

(DIPTERA: TEPHRITIDAE: ACANTHONEVRINI)

DAVID L. HANCOCK

8/3 McPherson Close, Edge Hill, Cairns, Qld 4870

! Abstract

The Rioxa complex of Indo-Australian fruit flies is reviewed and keyed. The 14 recognised

species are referred to 6 genera: Cribrorioxa Hering (1 sp.) Ectopomyia Hardy (2 spp),

Hexacinia Hendel (3 spp), Hexamela Zia (1 sp.), Rioxa Walker (6 spp) and Sophiroides Hendel

(1 sp.) Hexacinia punctifera (Walker, 1861), H. multipunctata Malloch, 1939 and H.

flavipunctata Hering, 1940 are placed as new synonyms of H. stellipennis (Walker, 1860), while

Rioxa quinquemaculata Bezzi, 1913 and R. vittata Zia, 1963 are placed as new synonyms of R.

parvipunctata de Meijere, 1911, stat. rev. and R. sexmaculata (van der Wulp, 1880) is placed as

a new synonym of R. discalis (Walker, 1861). Acinia marginemaculata Macquart, 1851 is

excluded from Hexacinia. A record of Hexacinia stellipennis from Sumatra is regarded as an

error, while those of Rioxa lanceolata Walker from Sri Lanka and China are regarded as

misidentifications of R. parvipunctata. Known larval hosts are fallen logs.

Introduction

This is the third in a series of papers reviewing and keying Indo-Australian

and East Asian fruit flies referred to the Acanthonevra group in tribe

Acanthonevrini (sensu Korneyev 1999) of subfamily Phytalmiinae. It deals

with the Rioxa complex, which occurs from Sri Lanka and India eastwards to

the Philippines and Papua New Guinea. Previous papers dealt respectively

with the Acanthonevra and Sophira complexes (Hancock 2011, 2012).

Korneyev (1999) placed Rioxa Walker, Ectopomyia Hardy and Hexacinia

Hendel in an apparently monophyletic clade within his Acanthonevra

subgroup in the Acanthonevra group of genera. Hexamela Zia was regarded

as a close ally of Hexacinia by Zia (1963) and Wang (1998). Cribrorioxa

Hering was included in this complex by Hancock (2005) and Sophiroides

Hendel also appears to belong here (Hancock 2012).

All available biological information is recorded under the [incorrect] name

‘Rioxa sexmaculata’. Hardy (1986) collected specimens of [R. parvipunctata

de Meijere] ‘in mating flight around buttress of a large forest tree in botanical

garden' in Bogor, Java and suspected that they might breed in rotting wood.

This was confirmed by Kovac et al. (2010), who recorded oviposition in

holes made by bark beetles or other insects in recently fallen trees in northern

Thailand. Permkam (1995) collected individual specimens of [R. discalis

(Walker)] (Fig. 1) attracted to cut shoots of the bamboos Thyrostachys oliveri

and Bambusa arundinacea on three separate occasions in southern Thailand,

suggesting that a casual relationship with bamboo also exists.

An annotated key to the 6 genera and 14 species recognised in this study is

provided below, with several other taxa previously included in Hexacinia or

Rioxa here synonymised or excluded (see Discussion). Illustrations of most

species may be found in Hardy (1973, 1974, 1986) and Wang (1998).

46 Australian Entomologist, 2014, 41 (1)

Fig. 1. Rioxa discalis (Walker): male from West Malaysia.

Key to genera and species

* — New distribution records based on material in The Natural History

Museum, London (BMNH).

1 Wing pattern largely brown, yellowish basally almost to apex of cell c

and with a faint brown transverse band medially, with a transverse

posterior yellow indentation from wing margin across cells cu; and dm

into cell br and with a narrow yellow transverse band from cell r; near

apex of stigma into cell dm between R-M and DM-Cu crossveins that is

broadly produced as a 2-pronged marking into subapical parts of cells rp,3

and r4; a shallow hyaline indentation in cell m along wing margin;

scutum yellow with 4 dark longitudinal vittae, the dorsocentral pair not

extending onto scutellum; face concave in profile; presutural setae absent;

secondary scutellar setae weak and hair-like [Sri Lanka; Rioxa magnifica

Senior-White, 1921 is a synonym; illustrated by Hardy 1968] .............

Loro bo COSE FH eiii Sophiroides flammosus Hendel, 1914

‘) Australian Entomologist, 2014, 41 (1) 47

— Not as above; wing pattern blackish brown with hyaline or subhyaline

Spotsjand indentations lessees eee PRAE 2

2 Scutum pale medially with a pair of dark dorsocentral vittae extending

over sides of scutellum; pleura normally with dark vittae but not distinctly

spotted; wing with vein Sc distinctly arcuate and cell m with a narrow

marginal band or single small hyaline indentation; head with upper frontal

setae often not close to lower setae and not reclinate; anepisternum

without an additional dark seta near ventral margin centrally

— Scutum largely yellow with or without dark spots or dark medially with

pale submedial vittae extending most of its length; pleura yellow or with

brown to black spots; wing with vein Sc not distinctly arcuate and cell m

with 1 large or 2 small hyaline indentations; head with upper frontal setae

very close to lower setae and reclinate; anepisternum with or without an

additional dark seta near ventral margin centrally

3 Wing with numerous subhyaline discal spots but no distinct hyaline

marginal indentations, at most small hyaline spots towards apex of cell r;

and over apices of veins Ro; and R45 and a narrow marginal band in

cells m and cu;; cell c apically blunt; head setae mostly yellow [eastern

Indonesia (Sumba)] ..................... Cribrorioxa perforata Hering, 1952

— Wing without numerous discal spots but with distinct hyaline indentations

in cells r243, r4+5 and m, that in cell r4,5 oval and almost filling apex; cell c

apically acute; almost all head setae brown to black [Ptilonina Enderlein,

I9lisasynonym| eee Walker, 1856 ... 4

4 Wing veins R; and R33 approximate at apex, with vein R}, less than 1/4

distance between veins R; and Ra,5; cell r45 with a large hyaline basal

spot that is more than half width of cell; cell dm often with a small

subapical hyaline spot; either cell br with a broad hyaline streak or cell

cu; with an isolated anteromedial spot -a.a 5

— Wing vein Rə more than 1/3rd distance between veins R, and R445; cell

dm entirely dark, cell br with at most a faint narrow streak and cell r4,5

normally with at most a tiny basal spot (large in most megispilota) ...... 6

5 Wing cell br with a broad and elongate hyaline streak extending almost to

-M crossvein [Malaysia (West, Sarawak), Singapore and Indonesia

(Sumatra, Nias, Java); R. nox Rondani, 1875 is a Synonym; records from

Sri Lanka and China are errors] ................. R. lanceolata Walker, 1856

— Wing cell br without an elongate hyaline streak; cell cu; with an isolated

anteromedial spot alongside vein Cu, [East Malaysia (Sarawak) and

Indonesia (Kalimantan: Babidjoelan*)] ........... R. erebus Rondani, 1875

6 Wing without a distinct hyaline indentation in cell c; cell r; with hyaline

Spot near base absent and indentation at apex of cell sc absent or small

and not crossing cell; cell br sometimes with a narrow and indistinct pale

i=)

Australian Entomologist, 2014, 41 (1)

streak and base of cell r4.5 sometimes with a small spot [India, Sri Lanka,

Burma, China (Yunnan), Thailand, Philippines (Palawan, Balabac),

Malaysia (West, Sarawak) and Indonesia (Sumatra, Java, Kalimantan:

Babidjoelan*); R. quinquemaculata Bezzi, 1913, syn. n., R. infirma

Hering, 1941, R. vittata Zia, 1963, syn. n. and ‘R. sexmaculata‘ of Hardy

(l1086)rarezregardedzasisynonyms]Eeeee een tees

dotbeseopbooo oeneocinqon codon R. parvipunctata de Meijere, 1911, stat. rev.

Wing with distinct hyaline spots or indentations in cells c-sc and in cell rj

near base and at apex of cell sc, the latter crossing cell ..................... 7

Wing cell cu; with at most an elongate, non-quadrate hyaline indentation

at apex of vein A,+Cu, [Burma, China (Yunnan), Thailand, Malaysia

(West, Sarawak, Sabah*), Singapore* and Indonesia (Sumatra, Java);

Ptilona sexmaculata van der Wulp, 1880, syn. n. and R. sumatrana

Enderlein, 1911 are synonyms; Fig. 1] .......... R. discalis (Walker, 1861)

Wing cell cu; with a large quadrate hyaline indentation at apex of vein

At Gwithaticrossesitheicelll e eT Es 8

Wing with a large hyaline indentation in cell rj, below apex of vein R43

and with (females and some males) or without a large hyaline spot at base

of cell r4} near R-M crossvein [southern Philippines (Tawi Tawi,

Mindanao); illustrated by Hardy (1970)] ...... R. megispilota Hardy, 1970

Wing with at most a small hyaline indentation in cell rj, below apex of

vein R?.5 and without a large hyaline spot at base of cell r4,; [Philippines

(including Mindanao and Tawi Tawi but not Palawan or Balabac);

illustrated by Hering (1941)] ......................... R. lucifer Hering, 1941

Wing base largely hyaline; apex of cell r4+s entirely dark; cell rı with a

large quadrate indentation crossing cell; 2 large discal spots either side of

R-M crossvein; single elongate hyaline indentations in cells m and cui,

the latter crossing cell; anepisternum without an additional seta; arista

long plumose [China (Yunnan)] ............ Hexamela bipunctata Zia, 1963

Wing base largely dark; apex of cell r4,; largely hyaline or subhyaline,

discal spots normally small and subhyaline; cells m and cu; usually with 2

and 3 small marginal hyaline indentations respectively (combined and

large in Ectopomyia males); anepisternum often with an additional dark

seta near ventral margin centrally; arista short plumose .................- 10

Almost all head setae brown to black; abdomen largely dark but without

distinct dark spots; scutum mostly dark (including medially) with pale

submedial vittae and a distinct yellow patch between prescutellar setae;

scutellum yellow with a pair of submedial dark vittae; wing with apical

hyaline indentations in cells r}, and r4,5 large and broadly ovate or

subquadrate and with few discal spots; anepisternum with additional seta

weakiOnabDSelitveessstrisnsscesesteasetectee sss Ectopomyia Hardy, 1973 ... 11

| Australian Entomologist, 2014, 41 (1) 49

Head setae mostly yellow; abdomen yellow with distinct brown to black

spots; scutum and scutellum yellow with or without small dark spots;

wing with apical hyaline indentation in cells r;,4 small and rounded and

that in cell rj,5 subquadrate or narrowly hyaline or subhyaline and with

numerous hyaline or subhyaline discal spots; anepisternum with

additional seta prominent ..................... Hexacinia Hendel, 1914 ... 12

Wing veins Rz and R4+s diverging apically, with the apex of vein R;45

about equidistant between veins R4+5 and M at wing margin; cell r» with

a hyaline central spot near line of R-M crossvein; anepisternum with

additional seta absent; sexes dimorphic in wing pattern and male with a

large ventral appendage on front femur [China (Yunnan) and Laos] .......

OCHOODOOPOCOBOCODUOOEDCU OOOO OÖ red E. baculigera Hardy, 1973

Wing veins Rọ and R4,5 almost parallel, with the apex of vein R45

distinctly closer to vein R45 than to vein M at wing margin; cell r23

without a hyaline central spot near line of R-M crossvein; anepisternum

with additional seta present; male unknown [West Malaysia; illustrated by

(:hua2009] E E oe EN ene E. hancocki Chua, 2009

Wing cell r4+5 with a broad subquadrate hyaline spot at apex not filling

cell; antennae brown to black on apical half [India, Sri Lanka, Burma,

China (Yunnan), Thailand, Vietnam, West Malaysia, Indonesia (Sumatra)

and Brunei (Chua 2002); presence in Philippines (Luzon) requires

confirmation: the type locality, Manila, is possibly incorrect; H. stellata

(Macquart, 1851) (a homonym) and H. nigroantennata Hering, 1956 are

regarded as synonyms] ....................... H. radiosa (Rondani, 1868)

Wing cell r4.5 narrowly subhyaline at apex; antennae entirely yellow ... 13

Wing cell r; with 2 hyaline to subhyaline indentations from costa, both

crossing cell; discal spots large, consisting mostly of transverse, elongate

spots equal or nearly equal to width of cell; scutum and pleura almost

entirely pale or with only a few faint brown spots [Philippines (including

Palawan)] $9.2 er EE RE ETE H. pellucens Hardy, 1970

Wing cell r; with 2 or 3 hyaline to subhyaline indentations from costa, the

medial spot often vestigial or absent; discal spots small, generally

rounded and occupying only a fraction of width of cell; scutum and pleura

usually with numerous dark brown to black spots [Philippines (including

Tawi Tawi but not Palawan), Brunei (Chua 2002), East Malaysia (Sabah),

eastern Indonesia (Sulawesi, Maluku, West Papua) and Papua New

Guinea (including Bismarck Archipelago: Hancock and Drew 2003 and

Bougainville: Hardy 1986); H. punctifera (Walker, 1861), syn. n.,

H. stigmatoptera Hendel, 1928, H. multipunctata Malloch, 1939, syn. n.,

H. flavipunctata Hering, 1940, syn. n. and H. celebensis Hering, 1941 are

regarded as synonyms; a record from Sumatra (Hardy 1986) appears to be

QNLCITOL] exerceri Donna kxcccedbn nent H. stellipennis (Walker, 1860)

50 Australian Entomologist, 2014, 41 (1)

Discussion

Hexacinia

Acinia marginemaculata Macquart, 1851, described from ‘Asia’ (Macquart

1851), was placed in Hexacinia by Foote (1984) and Norrbom et al. (1999),

possibly following a suggestion by Bezzi (1913), but its narrow wing and

pattern of markings, particularly the numerous (5-6) marginal spots in cell m,

suggest it does not belong there. It is possibly a species of Paracanthella

Hendel (subfamily Tephritinae) but examination of the type is needed to

determine its identity and relationships.

Hexacinia palpata Hendel, 1915, from China, Taiwan and far eastern Russia,

was retained in that genus by Hardy (1973, 1974) but was included in genus

Hexaptilona Hering by Zia (1963) and Wang (1998). Together with the

closely related H. hexacinioides (Hering, 1938) from Burma, it is currently

placed in subfamily Blepharoneurinae (Norrbom and Condon 1999).

Specimens referable to both H. stellipennis (Walker) and H. punctifera

(Walker) occur in Sulawesi (Hardy 1959, DLH pers. obs. of BMNH

specimens), Borneo (Hardy 1986, Chua 2002) and New Britain (Hancock and

Drew 2003) and the two taxa do not appear to be separable. The size of the

medial spot in cell rı is variable and sometimes vestigial or absent (Hancock

and Drew 2003), with a small spot present in the holotype of H. celebensis

(Hering 1941). The number and intensity of the dark pleural spots also appear

to be variable (Hardy 1974). Accordingly, I am treating H. punctifera

(Walker, 1861), H. multipunctata Malloch, 1939 and H. flavipunctata Hering,

1940 as new synonyms of H. stellipennis, together with the previously

synonymised H. stigmatoptera Hendel, 1928 and H. celebensis Hering, 1941.

Whereas the male epandrium and surstyli appear to be relatively longer and

more slender in H. pellucens Hardy [and even more so in H. radiosa

(Rondani)] than in H. stellipennis and H. punctifera, there appear to be no

discernible differences between the latter two taxa (Hardy 1974, 1986).

Hardy's (1986) record of H. stellipennis from Sumatra, based on specimens

allegedly in BMNH, appears to be an error, since no Sumatran specimens of

the genus were found there during a visit in 2012 (DLH pers. obs.).

Rioxa

There has been considerable confusion in the literature concerning the

nomenclature and identity of Rioxa sexmaculata (van der Wulp, 1880).

Originally described from Sumatra (van der Wulp 1880), it was regarded as a

senior synonym of R. sumatrana Enderlein, 1911, also described from

Sumatra (Enderlein 1911), by both de Meijere (1914) and Hardy (1974), the

latter at least basing his synonymy on examination of the types. Later,

however, Hardy (1986) again separated the two taxa, with R. sumatrana

subsequently placed as a junior synonym of R. discalis (Walker, 1861) by

Hancock (1998). Hardy (1973, 1986) regarded R. quinquemaculata Bezzi,

Australian Entomologist, 2014, 41 (1) 51

1913, as a junior synonym of R. sexmaculata and appears to have used the

former taxon, described from Tenasserim, Burma (Bezzi 1913) as the basis

for his 1986 definition of R. sexmaculata, incorrectly listing the ‘Type 9’ of

the latter as in the Zoological Museum, University of Copenhagen. However,

he had previously (Hardy 1969) correctly recorded the ‘Lectotype @’ as in

the Zoological Museum, University of Amsterdam, so his 1986 concept of R.

sexmaculata (unlike that in Hardy 1974) appears not to be based upon the

types. Indeed, the type series of R. sexmaculata, as illustrated by van der

Wulp (1881), is clearly synonymous with R. discalis and R. sumatrana. The

apical extension to wing cell bcu is shown incorrectly elongate in van der

Wulp’s figures 10 and 11 but correctly in his figure 7; the incorrect state was

used by Enderlein (1911) to define his new genus Ptilonina and separate R.

sexmaculata from R. sumatrana.

Based on Hardy’s (1986) concept of R. sexmaculata, Hancock and Drew

(1995) synonymised R. parvipunctata de Meijere, 1911 with it. However, the

latter species, originally described from Java (de Meijere 1911) as a variety

[subspecies] of R. sexmaculata, is recognised here as distinct, with R.

quinquemaculata Bezzi, 1913, R. infirma Hering, 1941 and R. vittata Zia,

1963 placed as synonyms, two newly so. Hardy (1973) had previously

recognised the synonymy of R. infirma.

Records of R. parvipunctata (as ‘R. sexmaculata’) and R. discalis (as ‘R.

sumatrana’) from West Papua, Indonesia and Malaita, Solomon Islands

respectively (Hardy 1986) were regarded as errors by Hancock and Drew

(2003), the former being the result of a misinterpreted specimen label that

actually refers to Soekaboemi, Java. Records of R. lanceolata from Sri Lanka

and Yunnan, China are also errors, with both based on misidentifications of

`R. parvipunctata. Specimens recorded from Sri Lanka (Hendel 1928) lack the

hyaline markings either side of the R-M crossvein, while that from Yunnan

(Wang 1998) has a faint pale streak in cell br and a small basal spot in cell

745. Similar variation in wing markings was noted from Sri Lanka by Hering

(1956, as ‘R. infirma"), while Hancock and Drew (1995) also recorded a

Malaysian specimen (as *R. sexmaculata ’) with a small basal spot in cell r445.

Two additional species included in Rioxa by Norrbom et al. (1999), viz.

Trypeta manto Osten Sacken and Rioxa vinnula Hardy, were transferred to

Freyomyia Hardy, in the Acanthonevra complex, by Hancock (2011).

Biogeography

The Rioxa complex is almost entirely restricted to South and Southeast Asia,

extending as far west as India and Sri Lanka but only as far north as Yunnan

Province in southern China. Although well represented in the Philippines,

only two species are otherwise known east of Borneo (Cribrorioxa perforata

and Hexacinia stellipennis), with only H. stellipennis reaching the island of

New Guinea and the Bismarck Archipelago.

52 Australian Entomologist, 2014, 41 (1)

Three monotypic genera are known, with limited and peripheral distributions: |

Sophiroides in Sri Lanka, Cribrorioxa on Sumba in the Lesser Sunda Islands |

and Hexamela in Yunnan, China. All have relatively broad wings with an

almost or entirely dark apex to cell r4+s and an often weak or narrow hyaline

marginal indentation in cell m. Cribrorioxa shares with Rioxa the distinct

dark vittae over the scutum and scutellum and an arcuate vein Sc, while

Hexamela shares with Hexacinia the lack of scutal vittae and the mostly

yellow head setae with the upper frontals reclinate. The affinities of |

Sophiroides are uncertain but it is possibly an ancestral relict.

Hexacinia, Ectopomyia and Rioxa are centred in SE Asia. Some species of

Hexacinia and Rioxa are widespread, with two (H. radiosa and R.

parvipunctata) recorded from India and Sri Lanka to at least Borneo and one

(H. stellipennis) known from the Philippines and Borneo to Papua New

Guinea. Rioxa discalis is known from Burma and Yunnan to Borneo and

Java, while Ectopomyia has a more restricted distribution, with the two

species (E. baculigera and E. hancocki) known from Yunnan-Laos and West

Malaysia respectively. Rioxa lanceolata is known from West Malaysia to

Borneo and Java and R. erebus is known only from Borneo. Three species

(H. pellucens, R. lucifer and R. megispilota) appear to be endemic to the

Philippines. All three genera have a distinctly slender epandrium, surstyli and

proctiger and a hyaline or subhyaline apex to cell r4+s, best developed in

Rioxa, Ectopomyia and Hexacinia radiosa.

The spotted pleura, reclinate upper frontal setae and presence of 2 hyaline

marginal indentations in cell m and (usually) 3 in cell cu; in Ectopomyia and

Hexacinia, plus the presence of a weak anepisternal seta near the ventral

margin centrally in E. hancocki and the subquadrate shape of the hyaline

apical spot in cell r5 in H. radiosa, suggest a close (and possibly sister-

group) relationship. Rioxa differs from the other two genera in scutal and

scutellar markings, in having the upper frontal setae incurved and the vanes

of the phallapodeme fused into a Y-shaped structure, and from all the other

genera in vein Sc reaching the costa at a distinctly acute angle.

Biology

The log-breeding biology of Rioxa parvipunctata is very similar to that seen

in the Dacopsis and Acanthonevra complexes (Permkam and Hancock 1995,

Hancock 2013) and the relationship between them and the Sophira complex,

which uses living bamboo (Hancock 2012), requires further investigation.

Acknowledgements

I thank Allen Norrbom (USDA, Washington DC) for help in assessing the

identity of Acinia marginemaculata, Kim Goodger (The Natural History

Museum, London) for access to material in her care and Susan Phillips for

preparing Figure 1.

Australian Entomologist, 2014, 41 (1) 53

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BEZZI, M. 1913. Indian trypaneids (fruit-flies) in the collection of the Indian Museum, Calcutta.

Memoirs of the Indian Museum 3: 53-175, pls 8-10.

CHUA, T.H. 2002. New records of Trypetinae from Brunei Darussalam (Diptera: Tephritidae).

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CHUA, T.H. 2009. Ectopomyia hancocki, a new species of acanthonevrine fly (Diptera:

Tephritidae: Phytalmiinae) from Peninsular Malaysia. Raffles Bulletin of Zoology 57(1): 25-27.

de MEIJERE, J.C.H. 1911. Studien über südostasiatische Dipteren. VI. Tijdschrift voor

Entomologie 54: 258-432.

de MEIJERE, J.C.H. 1914. Studien über südostasiatische Dipteren. IX. Tijdschrift voor

Entomologie 57: 137-276.

ENDERLEIN, G. 1911. Trypetiden-Studien. Zoologische Jahrbucher. Abteilung für Systematik,

Oekologie und Geographie der Tierre 31: 407-460.

FOOTE, R.H. 1984. Family Tephritidae (Trypetidae). Pp 66-149, in: Soós, A. and Papp, L. (eds),

Catalogue of Palaearctic Diptera. Volume 9, Micropezidae — Agromyzidae. Akademiai Kiado,

Budapest & Elsevier Science Publishers, Amsterdam; 460 pp.

HANCOCK, D.L. 1998. Notes on some fruit flies (Diptera: Tephritidae) described by Francis

Walker. Australian Entomologist 25(3): 67-68.

HANCOCK, D.L. 2005. A note on three unusual species of Phytalmiinae (Diptera: Tephritidae)

from Papua New Guinea. Australian Entomologist 32(2): 65-66.

HANCOCK, D.L. 2011. An annotated key to the species of Acanthonevra Macquart and allied

genera (Diptera: Tephritidae: Acanthonevrini). Australian Entomologist 38(3): 109-128.

HANCOCK, D.L. 2012. Bamboo-stem flies: an annotated key to the species of the Sophira

complex of genera (Diptera: Tephritidae: Acanthonevrini). Australian Entomologist 39(1): 5-32.

HANCOCK, D.L. 2013. Themara maculipennis (Westwood) and Themara hirtipes Rondani

(Diptera: Tephritidae: Acanthonevrini): a case of confused synonymies. Australian Entomologist

40(2): 93-98.

HANCOCK, D.L. and DREW, R.A.I. 1995. New genera, species and synonyms of Asian

Trypetinae (Diptera: Tephritidae). Malaysian Journal of Science 16A: 45-59,

HANCOCK, D.L. and DREW, R.A.I. 2003. New species and records of Phytalmiinae (Diptera:

Tephritidae) from Australia and the south Pacific. Australian Entomologist 30(2): 65-78.

HARDY, D.E. 1959. The Walker types of fruit flies (Tephritidae-Diptera) in the British

Museum collection. Bulletin of the British Museum (Natural History), Entomology 8(5): 159-

242, pls 11-16.

HARDY, D.E. 1968. The fruit fly types in the Naturhistorisches Museum, Wien (Tephritidae-

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ptera) of Thailand and bordering countries.

Pacific Insects Monograph 31: 1-353, pls 1-8.

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Monograph 32: 1-266, pls 1-6.

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HENDEL, F. 1928. Neue oder weniger bekannte Bohrfliegen (Trypetidae) meist aus dem

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HERING, E.M. 1941. Neue Dacinae und Trypetinae des Zoologischen Museums der Universitat

Berlin. Siruna Seva 3: 1-25.

HERING, E.M. 1956. Trypetidae (Dipt.) von Ceylon. Verhandlungen der Naturforschenden

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KORNEYEV, V.A. 1999. Phylogenetic relationships among higher groups of Tephritidae. Pp

73-113, in: Aluja, M. and Norrbom, A.L. (eds), Fruit flies (Tephritidae): phylogeny and

evolution of behavior. CRC Press, Boca Raton; xviii + 944 pp.

KOVAC, D., DOHM, P. and FREIDBERG, A. 2010. Field observations on the mating behaviour

of the Oriental Rioxa sexmaculata (van der Wulp) (Diptera: Tephritidae) and a review of the

reproductive behaviour patterns in Acanthonevrini. Biosystematica 4(1): 5-14.

MACQUART, P.J.M. 1851. Diptéres exotiques nouveaux et peu connus. Suite du 4e

supplement. Mémoires de la Société National des Sciences, de l'Agriculture et des Arts de Lille

1850: 134-282, pls 15-28.

NORRBOM, A.L. and CONDON, M.A. 1999. Phylogeny of the subfamily Blepharoneurinae. Pp

135-155, in: Aluja, M. and Norrbom, A.L. (eds), Fruit flies (Tephritidae): phylogeny and

evolution of behavior. CRC Press, Boca Raton; xviii + 944 pp.

NORRBOM, A.L., CARROLL, L.E., THOMPSON, F.C., WHITE, I.M. and FREIDBERG, A.

1999. Systematic database of names. Pp 65-251, in: Thompson, F.C. (ed.), Fruit fly expert

identification system and systematic information database. Myia 9: ix + 524 pp.

PERMKAM, S. 1995. Bamboo shoot fruit flies in southern Thailand. Songklanakarin Journal of

Science and Technology 17(3): 229-238.

PERMKAM, S. and HANCOCK, D.L. 1995. Australian Trypetinae (Diptera: Tephritidae).

Invertebrate Taxonomy 9: 1047-1209.

van der WULP, F.M. 1880. Eenige Diptera van Nederlandsch Indie. Tijdschrift voor

Entomologie (1879-1880) 23: 155-194, pls 10-11. i

van der WULP, F.M. 1881. Midden-Sumatra. Reizen en onderzoekingen der Sumatra-Expeditie,

uitgerust door het Aardrijkskundig Genootschap, 1877-1879, beschreven door de leden der

expeditie, onder toezicht van Prof. P.J. Veth. Vol. 4, Natuurlijke Historie, Part 1(2), Fauna,

Chapter 9, Diptera. E.J. Brill, Leiden; 60 pp, 3 pls. [Ptilona sexmaculata: Pp 51-52, pl. 3, figs 7-

11].

WANG, X.-J. 1998. The fruit flies (Diptera: Tephritidae) of the East Asia Region. Acta

Zootaxonomica Sinica 21(Supplement): viii + 338 pp + 268 figs + 41 pls.

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| Australian Entomologist, 2014, 41 (1): 55-56 55

A NOTE ON THE STATUS OF A SUPPOSED ‘TYPE’ OF THEMARA

ENDERLEINI HERING (DIPTERA: TEPHRITIDAE)

DAVID L. HANCOCK

8/3 McPherson Close, Edge Hill, Cairns, Qld 4870

Abstract

The ‘Type 4” of Themara enderleini Hering recorded by Hardy in 1974 and 1986 is found not to

belong to the original series. The true Type 3 and Type 9 are located in Warsaw.

Discussion

In discussing the status of Themara enderleini Hering, 1938, Hancock (2013)

noted there was some uncertainty regarding the location of much of Hering’s

(1938) type series, particularly his ‘Type ^ and ‘Type 9’. Images of T.

enderleini were provided by Hancock (2013); for comparison, the related 7.

hirtipes Rondani, with well developed ‘eye-stalks’, is shown in Figure 1.

Clyde Wild, Griffith University).

56 Australian Entomologist, 2014, 41 (1)

Hardy (1974) recorded the ‘Type 4” as deposited in the Zoological Museum,

Berlin (ZMHU), this later corrected (Hardy 1986) to the Senckenberg

Deutsches Entomologisches Institut (DEI). However, examinations of the

ZMHU and DEI collections (T. Dikow and A. L. Norrbom pers. comm.)

confirmed that there are no valid syntypes of T. enderleini in either of those

collections, although there is a specimen in DEI with the following label data:

“Sumatra / R. Weber Collection / Themara enderleini 3 Hering det. M.

Hering 1939 (handwritten except for ‘det. M. Hering 193"). Evidently it was

seen by Hering only after his description was published and therefore has no

status as a type.

Hering’s (1938) ‘Type (^ and ‘Type 9’ from Soekeranda, Sumatra, are

indeed in the Polish Academy of Sciences Museum and Institute of Zoology,

Warsaw (PAN) and bear his determination labels (T. Huflejt pers. comm.).

These were both designated paralectotypes by Hancock (2013) who, in

designating a male from Liangagas as Lectotype, considered all specimens

used by Hering (1938) in his description of 7. enderleini to be part of his type

series [i.e. syntypes], with none expressly excluded by him (ICZN 1999:

Article 72.4.1) and the designation ‘8-, Q- Type’ used, rather than the strictly

defined term ‘types’ alone (ICZN 1999: Article 72.4.6).

Acknowledgements

I thank Allen Norrbom and Torsten Dikow for discussions and information

on the DEI specimen, Tomasz Huflejt for information on those in PAN and

Clyde Wild (Griffith University) for the photograph of T. hirtipes.

References

HANCOCK, D.L. 2013. Themara maculipennis (Westwood) and Themara hirtipes Rondani

(Diptera: Tephritidae: Acanthonevrini): a case of confused synonymies. Australian Entomologist

40(2): 93-98.

HARDY, D.E. 1974. The fruit flies of the Philippines (Diptera: Tephritidae). Pacific Insects

Monograph 32: 1-266, 6 pls.

HARDY, D.E. 1986. Fruit flies of the subtribe Acanthonevrina of Indonesia, New Guinea, and

the Bismarck and Solomon Islands (Diptera: Tephritidae: Trypetinae: Acanthonevrini). Pacific

Insects Monograph 42: 1-191.

HERING, [E].M. 1938. Neue palaearktische und exotische Bohrfliegen.

Entomologische Zeitschrift 1938: 397-417.

ICZN (International Commission on Zoological Nomenclature). 1999. International code of

zoological nomenclature. 4th edition. International Trust for Zoological Nomenclature, London;

306 pp.

Deutsche

Australian Entomologist, 2014, 41 (1): 57-76 57

PSYCHOPSOID NEUROPTERA (PSYCHOPSIDAE,

OSMYLOPSYCOPIDAE) FROM THE QUEENSLAND TRIASSIC

K.J. LAMBKIN

18 Carey Street, Bardon, Qld 4065 (Email: megapsychops@bigblue.net.au)

Abstract

Six species of psychopsoid neuropterans are recorded from the southeastern Queensland Late

Triassic Blackstone Formation at Denmark Hill and Dinmore, the Late Triassic Mount Crosby

Formation at Mount Crosby, and the early Middle Triassic Gayndah Formation at Gayndah.

Triassopsychops superbus Tillyard (previously known from Denmark Hill, newly recorded from

Dinmore) has a suite of apomorphies confirming its earlier placement in Psychopsidae.

Additional material of Osmylopsychops spillerae Tillyard (type species of the family

Osmylopsychopidae) (previously known from Denmark Hill and Mount Crosby, newly recorded

from Dinmore) confirms the presence of a recurrent humeral vein and demonstrates its high

degree of intra-specific variation in size and venation. Petropsychops superbus Riek (previously

known from Denmark Hill, newly recorded from Mount Crosby), with M and Rs uniquely

structured basally, is retained in Osmylopsychopidae, although its strongly pectinate M is similar

to some species of the Mesozoic psychopsoid family Kalligrammatidae. Archepsychops

triassicus Tillyard (Denmark Hill), possibly conspecific with O. spillerae, is known only from its

fragmentary holotype and is retained for convenience in Osmylopsychopidae. Protopsychopsis

venosa Tillyard (Denmark Hill) is known only from its holotype, which is too fragmentary to be

considered any more than Neuroptera incertae sedis. The sixth species, Gayndahpsychops

carsburgi gen. et sp. n., from the early Middle Triassic of Gayndah, is a small osmylopsychopid

with CuA extensively pectinate.

Introduction

In the fossil insect literature, the term ‘psychopsoid’ is used for those fossil

lacewing species with a suite of wing characters broadly similar to those of

the extant Psychopsidae, viz. wing broad, generally no more than twice as

long as wide, somewhat triangular, with apical margin truncate; costal space

broad, especially in the forewing, with recurrent humeral vein; Sc, R; and Rs

strongly aligned; venation dense, Rs with numerous, usually closely parallel,

pectinate branches. The psychopsoid clade is generally considered to

comprise four families: the extant and fossil Psychopsidae, and the

extinct Mesozoic families Osmylopsychopidae, Brongniartiellidae and

Kalligrammatidae (Makarkin et al. 2013, and see discussion in Yang et al.

2012). The Mesozoic Aetheogrammatidae (Ren and Engel 2008) has also

been included, but the placement of this aberrant family remains

problematical.

In Australia, fossil psychopsoids have previously been recorded from two

Triassic localities in southeastern Queensland. Tillyard (1917, 1919, 1922,

1923) described Protopsychopsis venosa Tillyard, Archepsychops triassica

Tillyard (now A. triassicus, see ICZN Article 30.1.4.3), Triassopsychops

superba Tillyard (now T. superbus, ICZN Article 30.1.4.3), and

Osmylopsychops spillerae Tillyard from the Blackstone Formation at

Denmark Hill. Riek (1955) recorded A. triassicus and O. spillerae from the

Mount Crosby Formation at Mount Crosby and, in 1956, described

58 Australian Entomologist, 2014, 41 (1)

Petropsychops superba Riek (now P. superbus, ICZN Article 30.1.4.3) from

Denmark Hill. Both Tillyard and Riek considered 7. superbus as the oldest

true psychopsid and Riek (1955) placed the other four species in his newly

established family Osmylopsychopidae, unaware that Martynova had

established the same family in 1949 (but as Osmylopsychopsidae, see

Makarkin and Archibald 2005).

In recent years, local fossil enthusiast Allan Carsburg has collected many

valuable new specimens of fossil psychopsoids from Mount Crosby and from

a different exposure of the Blackstone Formation at Dinmore, also in

southeastern Queensland. There is now a second specimen of T. superbus

(from Dinmore) and a nice series of O. spillerae from both Mount Crosby

and Dinmore. Additionally, a new psychopsoid species has been collected by

the author from the Middle Triassic Gayndah Formation near Gayndah, again

in southeastern Queensland. The availability of this new material has

prompted the present review of the Queensland Triassic psychopsoid fauna.

The purpose of the present work, therefore, is to re-examine and, for the first

time, provide accurate line drawings of the holotypes of P. venosa, A.

triassicus, T. superbus, and P. superbus, provide new information on the

venation of O. spillerae further to Lambkin (1992), describe the new species

from Gayndah and discuss the family relationships of the species.

Specimens have been identified as fore- or hind wings by the relative width

of the costal space (much wider in forewings) and the convexity or concavity

of CuA (convex in forewings, concave in hind wings). Conservative

venational nomenclature is used and the view expressed by Makarkin et al.

(2009) that the radius and media are not fused basally in Neuroptera is

adopted. The division of the media into MA and MP is only used where the

stem of M has a clear primary dichotomous basal fork with each branch

forming a reasonably distinct vein system. All figures are inked line drawings

prepared using a camera lucida attachment on a Motic stereomicroscope. To

facilitate comparison, all specimens are figured as right wings. Abbreviations

for collections are as follows: ACC — Allan Carsburg Collection, Brisbane (to

be deposited in QM); GSQ - Geological Survey of Queensland (all

specimens now transferred to the QM); QM — Queensland Museum; UQ —

University of Queensland (all specimens now transferred to the QM).

The fossil localities

The Denmark Hill fossil lacewings recorded by Tillyard (1917, 1919, 1922,

1923) and Riek (1956) were collected in grey shales of the Late Triassic

(Carnian) Blackstone Formation, the uppermost member of the Ipswich Coal

Measures (Purdy and Cranfield 2013). The exposure, which is no longer

accessible, was in a small quarry in what is now the Denmark Hill

Conservation Park (-27.622? 152.756?) in the city of Ipswich. The site was

documented in detail by Dunstan (1923).

| Australian Entomologist, 2014, 41 (1) 59

The Dinmore locality (-27.606° 152.827°), also in the Blackstone Formation,

is a small commercial clay pit in Dinmore, a suburb of Ipswich,

approximately seven kilometres east of Denmark Hill. The site has-been well

| documented by Rozefelds and Sobbe (1987). The lithology and composition

| of the insect fauna are more or less identical to those of Denmark Hill.

The Mount Crosby insects occur in green shales of the Mount Crosby

Formation and have been collected at five separate exposures designated as

Fossil Insect Localities A, B, C, D and E (details in Allen 1961), in the

vicinity of Mount Crosby, north of Ipswich. The specimens discussed herein

were collected at Localities B (-27.552? 152.782?) and C (-27.550?

152.769°). The Mount Crosby Formation, also dated as Carnian, is the basal

sedimentary member of the Ipswich Coal Measures (Purdy and Cranfield

2013). The fossil insects are thus slightly older than those of the Blackstone

Formation at Denmark Hill and Dinmore.

The Gayndah species described herein was collected in grey shales of the

early (Anisian) Middle Triassic Gayndah Formation (Purdy 2013), in a road

cutting (-25.615? 151.640?) approximately three kilometres ENE of the town

of Gayndah. A species of the lacewing genus Lithosmylidia Riek has also

been recorded from the same site (Lambkin 1988).

Systematics

Family Psychopsidae Handlirsch

Triassopsychops superbus Tillyard

(Figs 1, 3)

Triassopsychops superba Tillyard, 1922: 467-469, text-fig. 89, plate 52.

Material examined. Holotype GSQ 284a, Denmark Hill, incomplete forewing, part

only, length (along Sc) 21 mm, width 21 mm; ACC I.100, Dinmore, Queensland, right

forewing fragment, part only, length 14.5 mm, width 15 mm.

Notes. The only previous illustrations of T. superbus have been photographs

by Tillyard (1922, plate 52) and Jell (2004, p. 84), and a line drawing by

Tillyard (1922, text-fig. 89). While Tillyard's photograph is adequate, his line

drawing is one of his ‘restorations’, with much surmise and many

inaccuracies and, for some reason, was printed upside down. Jell’s new

photograph is not all that clear and, apparently based on Tillyard’s

restoration, is also printed upside down.

Due to the state of preservation the identification of the veins in the M and

Cu fields is problematical. This is especially true of the holotype where the

wing in this area is broken and over-folded. The bases of both specimens are

crushed, with the veins broken and/or artificially approximated. The holotype

is flattened and has some veins showing as moulds and some as castes. It has

therefore not been possible to determine if it is a left or right wing or make

any confident determinations of vein convexities or concavities.

Australian Entomologist, 2014, 41 (1 )

rn,

p Ui

zg, Wi WL.

A LLL 6 a

TEN

EE EE

ac Se

PTA

ey ey E Mo rH AN

SS ee ore SS l=

MSS SSS:

b SESS

Figs 1-2. Psychopsidae. (1) Triassopsychops superbus, holotype GSQ 284a, Denmark

Hill. (2) Psychopsis illidgei, 9, Tamborine Mountain, Queensland, in QM, right

forewing. n nygma; rs-m basal crossvein between Rs and M.

Australian Entomologist, 2014, 41 (1) 61

Fig. 3. Triassopsychops superbus, ACC 1.100, Dinmore.

The venational interpretation herein, therefore, is based on as close as

possible examination of the wing bases, the vein convexities or concavities in

ACC I.100, and comparison of the relative length and extent of the M and Cu

fields with those in the most similar living psychopsid, Psychopsis illidgei

Froggatt (Fig. 2).

The drawings of the holotype (Fig. 1) and the new Dinmore specimen (Fig. 3)

illustrate the following pertinent information about the wing and the venation

of T. superbus:

1. The angle of the basal subcostal veinlets suggests the presence of a

recurrent humeral vein (Fig. 1).

2. No trichosors were detected on the preserved wing margins.

3. The costal space does not taper and is almost as wide apically as it is

basally (Fig. 1).

4. The crossveins of the costal space are random basally (Figs 1, 3),

although there is a suggestion of an alignment apically in the holotype

(Fig. 1).

5. Sc, R, and Rs appear to form a true ‘vena triplica’: strongly aligned,

apparently almost parallel and not noticeably convergent apically

(although the holotype is not clearly preserved in this area) and braced by

crossveins for the entire length (Figs 1, 3).

62 Australian Entomologist, 2014, 41 (1)

6. Two nygmata are present in the central radial field (Fig. 1).

7. Most branches of Rs are dichotomously forked within the discal area of

the wing (Fig. 1).

8. The radial and medial fields have numerous irregularly spaced crossveins,

although there is alignment into gradate series in apical half of sectors in

the holotype (Figs 1, 3).

9. M occupies a small area of the central portion of the wing and is

dichotomously branched into MA and MP extreme basally, before the

origin of Rs (Figs 1, 3).

10. The stem of MA curves anteriad basally and either just touches (Fig. 1),

or is fused for a short distance with (Fig. 3), the basal branch of Rs.

11.MA has three long dichotomous branches, simple for most of their

lengths (Figs 1, 3).

12.MP is simple to beyond half its length, then dichotomously branched

(Figs 1, 3).

13. The space between MP and CuA is braced by numerous crossveins (Fig.

3).

14. The base of CuA in ACC 1.100 is convex, although its longitudinal

section is neutral. CuA is simple to about half its length, then

dichotomously branched.

15. CuP is concave in ACC L100 and is extensively pectinately branched

(Figs 1, 3).

16. 1A is extensively pectinately branched (Figs 1, 3).

I consider that the presence of a uniformly broad costal space, an apparent

true ‘vena triplica’, the outer radial and medial cross-veins at least partly

aligned into gradate series and the richly pectinate CuP and 1A, are all

apomorphies of Psychopsidae and provide good evidence that T. superbus is

indeed the oldest true psychopsid. In this regard, it is worth noting the

similarities in these characters to those in the only living species of similar

size, P. illidgei (Fig. 2). Triassopsychops superbus, however, has a suite of

plesiomorphies not found in extant psychopsids, viz. the much more

randomly placed crossveins, the presence of nygmata in the radial field, the

extensive dichotomous branching of the Rs branches within the discal area,

the retention of MP as a separate longitudinal vein (only the weakly

sclerotised stem remains in P. illidgei - see Fig. 2) and the dichotomous

branching of CuA. The peculiar up-curving of the stem of MA to touch or

fuse for a short distance with the basal branch of Rs is considered an

autapomorphy for the genus. In living psychopsids, e.g. P. illidgei (Fig. 2),

Psychopsis mimica Newman, Psychopsis elegans (Guérin), Psychopsis

Australian Entomologist, 2014, 41 (1) 63

barnardi Tillyard, and Psychopsis insolens McLachlan (specimens in QM

collection; see also New 1988, Oswald 1993) and in the Baltic amber

Propsychopsis lapicidae MacLeod (MacLeod 1970, fig. 8), the stem of MA

= does not curve towards the Rs but is joined to its base by an upright or

forwardly inclined crossvein (Fig. 2). Triassopsychops superbus must have

been an impressively large lacewing, with a total forewing length, based on

the size of the holotype, of around 30 mm.

Peng et al. (2011) listed 35 Mesozoic and Tertiary fossil species that have

been ascribed to Psychopsidae and, with additions by Jepson et al. (2012) and

Makarkin et al. (2012), the number now stands at 41. Many of these are

fragmentary and/or poorly illustrated, so their relationships thus remain

uncertain. Based on the available illustrations, many have similarities with

Osmylopsychops Tillyard, the type genus of the family Osmylopsychopidae,

and may be more accurately placed in that family. Peng ef al. (2011)

suggested a division of the fossil psychopsids into two groups. The first

group, which includes all extant genera as well as five from the Cretaceous

and Eocene, has the branches of Rs in the discal field simple, crossveins of

the costal space arranged in one or two gradate series and those of the radial

field in two to four gradate series. The second group, which includes the

remaining Mesozoic species in their list, including Triassopsychops, has the

branches of Rs in the discal field dichotomously branched and the crossveins

of the costal space and the radial field random or at most forming many short,

irregular gradate series.

Family Osmylopsychopidae Martynova

Osmylopsychops spillerae Tillyard

(Figs 4-8)

Osmylopsychops spillerae Tillyard, 1923: 496-497, text-fig. 93, plate 43.

Material examined. QM F57529, Dinmore, incomplete right forewing, part and

counter-part, length 8 mm, width 7 mm; ACC 1.101, Dinmore, incomplete right

forewing, part only, length 14 mm, width 11.5 mm; QM F57530, Mount Crosby,

Locality B, right forewing fragment, part only, collected by K.J. Lambkin, length 10.5

mm, width 7.5 mm; ACC L102A, B, Mount Crosby, Locality B, incomplete left

forewing, part and counter-part, length 12 mm, width 8.5 mm; ACC I.19A, B, Mount

Crosby, Locality C, incomplete left forewing, part and counter-part, length 14 mm,

width 6.5 mm.

Notes. Osmylopsychops spillerae is the most frequently collected and best

documented lacewing from the Queensland Triassic. In 1992, I reconstructed

an almost complete forewing based on several well-preserved specimens

from Mount Crosby (Lambkin 1992). There are now five additional

forewings, three from Mount Crosby and two from the Dinmore locality of

the Blackstone Formation (Figs 4-8), which tell us more about the wing and

venation of this species.

Australian Entomologist, 2014, 41 (1)

SES ao

3A CuA

// d 6

Bie <

SS e

Figs 4-6. Osmylopsychops spillerae: (4) QM F57529, Dinmore; (5) QM F57530,

Mount Crosby; (6) ACC I.19A, B, Mount Crosby. n nygma; nd dilation of veins

suggesting presence of nygma; RAV recurrent humeral vein; sc-r; crossvein between

Sc and Ry.

Australian Entomologist, 2014, 41 (1) 65

WILL

CuP CuA

: WY Vila

Figs 7-8. Osmylopsychops spillerae: (7) ACC 1.102A, B, Mount Crosby; (8) ACC

1.101, Dinmore. n nygma; nd dilation of veins suggesting presence of nygma.

66 Australian Entomologist, 2014, 41 (1)

In all specimens CuA is noticeably convex, which identifies them as

forewings. The vein system interpreted as MA is identified as such on the

grounds that it has a common basal stem with the system identified as MP, as

demonstrated in the clearly preserved specimen illustrated in Lambkin (1992,

fig. 6) and also strongly suggested in new specimens QM F57529 and ACC

I.19A, B (Figs 4, 6).

QM F57529 (Fig. 4) confirms the presence of a recurrent humeral vein at the

base of the costal space, as conjectured by Tillyard (1923, text-fig. 93). This

is the first specimen to preserve this area of the wing. The recurrent humeral

vein is generally present in both fossil and extant psychopsoids (Makarkin et

al. 2013) and is clearly present in the only other well enough preserved

osmylopsychopid, an undescribed species from the Middle to Late (Ladinian-

Carnian) Triassic Madygen Formation of Kyrgyzstan (Shcherbakov 2008,

fig. 6; V.N. Makarkin pers. comm.).

The other important information that the new material demonstrates is the

degree of intra-specific variation in wing size and venation. Figures 4-8 are

all drawn to the same scale and demonstrate the size range of the species,

from ACC I.19 (Fig. 6) of width 6.5 mm, to the comparable fragment of ACC

1.101 (Fig. 8), almost twice as large at width 11.5 mm. Similar variation also

occurs in extant psychopsids that demonstrate a degree of sexual dimorphism

in size, the males being generally smaller than the females. For instance, the

good series of specimens of Psychopsis insolens in the QM range from

forewing length 11 mm and width 6 mm (d, Toowoomba, Queensland), to

forewing length 18 mm and width 11 mm (9, Killarney, Queensland). The

size variation in the QM specimens of the spectacular Psychopsis illidgei is

equally as stark, ranging from forewing length 23 mm and width 14 mm (d,

Bunya Mountains, Queensland), to forewing length 31 mm and width 20 mm

(9, Tamborine Mountain, Queensland) In many specimens, the size

difference is also reflected in the richness of the venation, particularly the

number of pectinate branches of Rs. In P. insolens these vary from eight in

the Toowoomba d to 13 in the Killarney 9, and in P. illidgei from 19 in the

Bunya Mountains 3 to 28 in the Tamborine Mountain 9.

With regard to the venation, even though the basic pattern is consistent across

all specimens (Figs 4-8; Lambkin 1992, figs 3-8), the venational details can

vary quite considerably. Much of the variation appears to be related to wing

size, with the larger wings generally possessing more vein branches and more

bracing cross-veins. Details of the variation are as follows:

1. The crossveins of the costal space may be apparently absent or almost so

(Figs 4, 8; Lambkin 1992, fig. 3), sparse in number and forming a simple

gradate series (Lambkin 1992, figs 6, 8), or anastomosed with the stems

of the subcostal veinlets to form a network running parallel to Sc (Fig. 6;

Lambkin 1992, fig. 5).

Australian Entomologist, 2014, 41 (1) 67

2. A basal crossvein between Sc and R, just beyond the origin of Rs is

present in the three specimens where this area of the wing is well

preserved (Fig. 6; Lambkin 1992, figs 5, 6). Its apparent absence in QM

F57529 (Fig. 4) is considered to be preservation related. No other

subcostal crossveins were detected in any of the specimens examined.

. Rj-Rs crossveins seem to vary in position and angle (Figs 4, 8; Lambkin

1992, fig 5), and were not detected at all in ACC I.19 (Fig. 6).

. The number of pectinate branches of Rs and the pattern of their

dichotomous branching are not the same in any two specimens (Figs 5-8;

Lambkin 1992, fig. 5), ranging from approximately 24 branches with only

one dichotomously branched in the large ACC I.101 (Fig. 8), to only

approximately 14 branches with three of these dichotomously branched in

the much smaller ACC I.19 (Fig. 6).

. A nygma is consistently present between the second and third, third and

forth, or forth and fifth branches of Rs, either as a detectable spot (Figs 5,

7; Lambkin 1992, fig. 5), or at least evidenced by a slight dilation of the

veins (Figs 6, 8).

. Crossveins of the radial, medial, cubital and anal fields are generally

difficult to detect and are as indicated in Figs 4-8 and the figures in

Lambkin (1992); those that have been tentatively detected vary

considerably in number and position, from very few (Fig. 5; Lambkin

1992, fig. 5), to more numerous and forming irregular gradate series (Figs

7, 8).

. The basic branching pattern and shape of M is quite constant, i.e. primary

forking into MA and MP at the base of the wing, well before the origin of

Rs; MA forked close to its origin and multi-branched, occupying a large

area in the centre of the wing; MP forked dichotomously well beyond the

primary fork of MA, the branches parallel, somewhat sinuous, and simple

for most of their length. The number and pattern of branching of MA,

however, varies considerably, from simply dichotomous with six major

branches (Figs 5, 6), to strongly pectinate with as many as six anteriorly-

directed branches mirroring the pectination Rs (Figs 4, 7). The only

variation in MP is the presence in some specimens of an enclosed cell on

MP3,+4 (Fig. 8; Lambkin 1992, fig 3).

. The pattern of Cu is fairly constant. CuA is simple and straight for much

of its length, but then curved anteriad and forked into eight or nine

branches to form a broad, somewhat triangular area apically (Figs 7, 8;

Lambkin 1992, fig. 5). CuP forks earlier than CuA, about at the level of

the primary fork of MP (Figs 5-8; Lambkin 1992, fig 6) or more basally

(Fig. 4) and the subsequent branching is a little variable (Figs 5-8;

Lambkin 1992, figs 3, 5).

68 Australian Entomologist, 2014, 41 (1)

9. The anal area of the wing is incomplete in most specimens and the

identification of 1A, 2A and 3A in Figs 5-8 is a best estimate based on the

pattern in QM F57529 (Fig. 4) and in the wing base illustrated in

Lambkin (1992, fig. 6). Comparison of the figures again indicates a

consistent pattern of dichotomous branching, but with variation in the

details of the more apical branching of the veins.

In summary, the new specimens recorded herein further augment our

knowledge of the wing and venation of O. spillerae, making it one of the best

documented fossil lacewings. The degree of intra-specific variation is of

particular note as a cautionary guide to the proliferation of species names for

fossil Neuroptera based only on trivial venational differences.

Petropsychops superbus Riek

(Figs 9-10)

Petropsychops superba Riek, 1956: 104-105, fig. 2.

Material examined. Holotype UQ C2135-6, Denmark Hill, incomplete left forewing,

part and counterpart, length 28 mm, width 18 mm; QM F57531, Mount Crosby,

Locality B, left forewing fragment, part and counterpart, collected by K. J. Lambkin,

length 10.5 mm, width 4 mm.

Notes. This species, previously known only from the holotype specimen

collected at Denmark Hill, is now also recorded (QM F57531) from the

slightly older Mount Crosby Formation. The only previous illustrations of P.

superbus have been photographs by Riek (1956, fig. 2), Jell (2004, p. 83) and

Grimaldi and Engel (2005, 9.15). The drawings of the holotype (Fig. 9) and

the new Mount Crosby specimen (Fig. 10) presented herein illustrate the

following pertinent information about the wing and the venation of P.

superbus:

1. The angle of the basal subcostal veinlets suggests the presence of a

recurrent humeral vein (Fig. 9).

2. No trichosors were detected on the preserved wing margins.

3. The costal space tapers to be almost as half as wide apically as it is

basally (Fig. 9).

4. The crossveins of the costal space are few in number and not arranged

into gradate series (Fig. 9).

5. SC, R; and Rs are more or less parallel for their entire lengths, but with

the apex of Sc curved posteriad to fuse with R,. There are several

randomly placed crossveins (Fig. 9).

6. Most of the branches of Rs are dichotomously forked within the discal

area of the wing (Fig. 9).

Australian Entomologist, 2014, 41 (1)

Ne}

Hill

Figs 9-10. Petropsychops superbus: (9) holotype UQ C2135-6, Denmark Hill; (10)

QM F57531, Mount Crosby. n nygma; rs-m basal crossvein between Rs and M.

7. There is a raised spot which may be a nygma between the 2™ and 3"!

branches of Rs (Fig 9). It is, however, not clearly preserved and is not

centred between the veins as nygmata usually are.

8. The proximal radial and medial fields have numerous randomly spaced

crossveins, but none is present beyond the level of the apex of Sc (Fig. 9).

9. The first branch of Rs (Rs;) and the stem of M form a peculiar structure at

the base of the wing, interpreted as follows: Rs; curves posteriad just

beyond the origin of Rs, throws off three or four pectinate branches

parallel with the remaining branches of Rs and those of M, and is then

connected to the stem of M by a short, oblique crossvein (rs-m in Fig. 10)

that appears to be a continuation of Rs;. An alternative interpretation of

the structure is that the system identified as Rs; is indeed a multi-

branched MA formed by the sharp recurving of the stem of MA (rs-m in

70 Australian Entomologist, 2014, 41 (1)

the interpretation above), which then throws off four pectinate branches

and is joined to the base of Rs by a short Rs-MA crossvein.

10. The stem of M is strongly concave and has at least 11 anteriorly-directed

pectinate branches, mostly dichotomously forked, which parallel those of

Rs. M thus occupies a considerable area of the centre of the wing (Fig. 9).

11. The stem of CuA is strongly convex, runs very closely and parallel with

the stem of M and has numerous, mostly dichotomously forked, pectinate

branches that almost precisely mirror those of M (Fig. 9).

12. CuP is deeply dichotomously forked with four or five main branches and

numerous crossveins (Figs 9, 10).

13.1A is deeply dichotomously forked, with approximately nine main

branches and numerous crossveins (Fig. 9).

Petropsychops superbus, as its species name alludes, must have been a

superbly impressive lacewing with a total forewing length, judging by the

size of the holotype, of just over 30 mm. The peculiar structure of Rs; and the

stem of M appears to be unique among Neuroptera and is a strong

autapomorphy for the genus. The relationship of Petropsychops to other

psychopsoids, however, remains problematical. As noted by Riek (1956), the

strong anterior pectination of M is a characteristic of some genera of

Kalligrammatidae, the psychopsoid family considered by Makarkin et al.

(2009) as the sister group of Osmylopsychopidae. Indeed, the similarity of

both M and CuA in P. superbus to those veins in the recently described

kalligrammatid Apochrysogramma rotundum Yang, Makarkin and Ren is

striking (see Yang et al. 2011, fig. 5). P. superbus, however, lacks the very

characteristic remigial ‘eye’ spot and extremely dense crossvenation of

Kalligrammatidae (Ren and Engel 2008). On these grounds, as well as its

similarity to Osmylopsychops, which, as demonstrated herein, also shows a

tendency to the anterior pectination of at least MA (Figs 4, 7), it is considered

that, for the present, Petropsychops is best retained in Osmylopsychopidae.

The difficulty of its family placement again emphasises the issue raised by

many authors (e.g. Makarkin 2010, Peng et al. 2010, Peng et al. 2011) of the

need for a comprehensive analysis of the family classification of the

Mesozoic psychopsoids, which as a result of the many recent discoveries are

now known to be highly diverse and species-rich (see lists in Yang ef af.

2009, and Peng et al. 2011).

Archepsychops triassicus Tillyard

(Fig. 11)

Archepsychops triassica Tillyard, 1919: 205-211, text-fig. 27.

Material examined. Holotype GSQ 137a, Denmark Hill, right forewing fragment, part

only, length 7 mm, width 6.5 mm.

Australian Entomologist, 2014, 41 (1) 71

Notes. Tillyard's text figure is again somewhat of a ‘restoration’ and contains

more veins than the specimen actually possesses. The only other previous

illustration of the specimen is the photograph in Jell (2004, p. 83). The new

drawing (Fig. 11) of A. triassicus demonstrates the fragmentary nature of its

type specimen, which is a cast with both R; and CuA strongly convex thus

indicating a forewing. The vein labelled Cu by Tillyard is actually a deep

groove in the wing and not a vein.

Riek (1955) placed A. triassicus in the Osmylopsychopidae and suggested

that O. spillerae might be its junior synonym. Indeed, the very fragmentary

specimen from Mount Crosby (UQ C679), which Riek tentatively referred to

A. triassicus, is in fact a wing base fragment of O. spillerae showing the anal

veins and multi-branched MA similar to those illustrated in Lambkin (1992,

fig. 2). Although it is quite likely that A. triassicus and O. spillerae are

indeed the same species (compare Fig. 11 with Figs 4-8 and the figures in

Lambkin 1992), due to the fragmentary nature of the type of A. triassicus it

would be unwise to synonymise the very well documented O. spillerae with

it. It also remains possible that A. triassicus is indeed distinct. As indicated in

Fig. 11 it does have at least four crossveins between Sc and Ry, whereas no

specimen of O. spillerae has been found with more than one. The vein

interpreted as CuP is much more deeply forked than in most specimens of O.

spillerae but, as listed above, the level of this forking in O. spillerae is

variable and the level in A. triassicus is the same as that in O. spillerae

specimen QM F57529 (Fig. 4).

Genus Gayndahpsychops gen. n.

Type species Gayndahpsychops carsburgi sp. n.

Diagnosis. Forewing. Relatively small size, width approximately 7.5 mm; M

occupying a large area in centre of wing; MA with two deep dichotomous

forks at level of origin of Rs; MA, fused with the apparent posterior branch

of Rs), this fused vein then fused with the apparent anterior branch of Rs;;

MP simple for about half its length, then dichotomously forked; CuA with

seven long posteriorly-directed pectinate branches which occupy a large area

of the wing; CuP deeply dichotomously forked basal to the origin of Rs, each

branch simple for most of its length.

Etymology. The generic name recognises the locality of the type species, the

Gayndah Formation at Gayndah in southeastern Queensland.

Notes. Gayndahpsychops is placed in Osmylopsychopidae because of its

basic psychopsoid facies, its small size, lack of well defined gradate

crossvein series, lack of extremely dense crossvenation and an eye-spot and,

most importantly, the extensive nature of M, which I consider is

synapomorphic with the type genus of the family, Osmylopsychops. It differs

12 Australian Entomologist, 2014, 41 (1)

from Osmylopsychops in the complex fusions of MA, and Rs; (these veins

are not connected in any of the many specimens of O. spillerae), the much

more apically forked MP (forked just beyond origin of Rs in

Osmylopsychops), the richly pectinate CuA, which occupies a large area of

the wing (dichotomously forked and much less extensive in Osmylopsychops)

and the deep dichotomous forking of CuP with the two branches simple for

most of their length (mostly forked at about half length in Osmylopsychops,

with CuP, deeply dichotomously forked). Beyond the type genus, the generic

composition and limits of Osmylopsychopidae are poorly known. Other

psychopsoid genera that have been ascribed to Osmylopsychopidae, e.g.

Actinophlebia Handlirsch, Parhemerobius Bode (Makarkin and Archibald

2005), are not well enough documented to allow reliable comparison with

Gayndahpsychops. It is of interest to note, however, that the undescribed

osmylopsychopid from the Triassic Madygen Formation illustrated by

Shcherbakov (2008, fig. 6) also seems to have CuA posteriorly pectinate.

Gayndahpsychops carsburgi sp. n.

(Fig. 12)

Type and only specimen. Holotype, QM F57532, Gayndah fossil insect locality,

incomplete right forewing, part and counterpart, collected by K.J. Lambkin, length

11.5 mm, width 7.5 mm.

Description. Forewing. Subcostal veinlets strongly inclined, dichotomously

forked with at least two connecting crossveins; a basal Sc-R, crossvein not

detected (wing not well preserved in this area); R, strongly convex; one R;-

Rs crossvein detected; Rs richly pectinate, the basal two branches apparently

dichotomously forked, the other eight preserved branches simple at least

basally; four randomly placed crossveins detected in Rs field; nygma not

detected, but a small dilation in the 3" and 4" apparent Rs branches suggests

its presence; stem of M not preserved; one basal crossvein between MA; and

MA;; MA, dichotomously forked with at least seven marginal branches; MP

fused with CuA for a short length, with six long apical branches; several

randomly placed crossveins in the area of the forkings of MP; CuA strongly

convex; one basal crossvein between CuA and CuP); at least five of the

pectinate branches of CuA dichotomously forked; anal veins apparently quite

extensive and dichotomously forked (the notional identification of 1A, 2A

and 3A in Fig. 12 based on the pattern in O. spillerae); trichosors not

detected.

Etymology. Named in honour of Mr Allan Carsburg who collected most of

the specimens which made this study possible.

Formation and Age. Gayndah Formation, early (Anisian) Middle Triassic.

Australian Entomologist, 2014, 41 (1) 73

BT ee

Figs 11-13. (11) Archepsychops triassicus, holotype GSQ 137a, Denmark Hill. (12)

Gayndahpsychops carsburgi, holotype QM F57532, Gayndah. (13) Protopsychopsis

venosa, holotype GSQ 160a, Denmark Hill. nd dilation in veins suggesting presence

of nygma.

1 Australian Entomologist, 2014, 41 (1)

Neuroptera incertae sedis

Protopsychopsis venosa Tillyard

(Fig. 13)

Protopsychopsis venosa Tillyard, 1917: 178-180, plate VIII, fig. 3.

Material examined. Holotype GSQ 160a, Denmark Hill, wing fragment, part only,

length 9 mm, width 7 mm.

Notes. Tillyard’s figure is a line drawing and there is also a photograph of the

holotype in Jell (2004, p. 83). As Fig. 13 demonstrates, the species is based

on a specimen preserving only the apex of a wing that is too fragmentary to

warrant further consideration. The only point of note is that the numerous

crossveins illustrated in Tillyard’s figure could not be detected in the

specimen. For the sake of convenience, Riek (1956) placed the species in

Osmylopsychopidae; however, it is simply too fragmentary for even tentative

family placement and is thus designated Neuroptera incertae sedis.

Acknowledgements

I sincerely thank Allan Carsburg for allowing me to examine specimens from

his private collection. I am also very grateful to my daughter Jennifer

Lambkin, who brought my drawings to publication standard and to Kristen

Spring and Pam Wilson of the Queensland Museum, who kindly made

available specimens in their care. I also very much appreciate the comments

of Vladimir Makarkin and Shaun Winterton, which greatly improved the

quality of this paper.

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THE AUSTRALIAN

Entomologist

Volume 41, Part 1, 3 March 2014

CONTENTS

BATLEY, M. AND BRANDLEY, B.

Phenology of the Australian solitary bee species Leioproctus plumosus (Smith)

(Hymenoptera: Colletidae)

BRAY, D.

Modification of the ALL Protocol to characterize the overall ant assemblage in

temperate eucalypt forest

HANCOCK, D.L.

An annotated key to the Rioxa complex of genera (Diptera: Tephritidae: Acanthonevrini)

HANCOCK, D.L.

A note on the status of a supposed ‘type’ of Themara enderleini Hering

(Diptera: Tephritidae)

LAMBKIN, K.J.

Psychopsoid Neuroptera (Psychopsidae, Osmylopsycopidae) from the Queensland Triassic

LAMBKIN, T.A.

The distribution of Euploea darchia niveata (Butler, 1875) (Lepidoptera:

Nymphalidae: Danainae) in Torres Strait, Queensland, Australia

MOORE, M.D. AND EDWARDS, E.D.

Two new species of Abantiades Herrich-Schiaffer (Lepidoptera: Hepialidae)

from Western Australia :

ISSN 1320 6133