Australian Entomologist, 2015, 42 (4): 161-186 161

THE CONTRIBUTION OF STEPHEN JAMES JOHNSON TO THE

STUDY OF AUSTRALIAN BUTTERFLIES

PETER S. VALENTINE

College of Marine and Environmental Sciences, James Cook University, Townsville, Old 4811

(Email: peter.valentine@jcu.edu.au)

Abstract

A summary is given of the life of Dr Steve Johnson (13 April 1950 - 16 December 2014) and his

contribution to the knowledge of Australian butterflies. It includes analyses of his publications

and his private collection, now lodged in the Museum of Tropical Queensland, Townsville, and

photographs of some aspects of his work. He collected mainly in Cape York Peninsula and

Torres Strait, made many new species records for Australia and discovered many new life

histories.

Introduction

Stephen James Johnson (Fig. 1) was born on 13th April 1950 at Wynnum,

third child to Stella and Arnold, younger brother to Bruce and Helene and

soon to be older brother to Ian, Estelle and Elizabeth (Fig. 2). His childhood

years were spent at Hemmant, then a semi-rural suburb of Brisbane. He

attended Wynnum State Central High School until grade 4 and Church of

England Grammar until matriculation in 1968. He was a keen sportsman and

achieved full colours as a member of the undefeated first XV (Rugby Union)

in 1967 and 1968 and a member of the first 8 (Rowing) in 1967 and 1968.

He failed to matriculate in 1967 (due to his sporting activities), repeated

Senior and started at the University of Queensland in 1969, where he studied

Veterinary Science, graduating in 1973.

Steve's professional life included private veterinary practice at Kingaroy

before joining the Queensland Department of Primary Industries as a

veterinary officer. He was stationed at Warwick Meatworks for five years

before moving to Townsville in 1979 as a Veterinary Entomologist at the

Oonoonba Veterinary Laboratory, which he subsequently headed until his

early retirement in 2004.

His premature death in December 2014 brought to a close a productive

contribution to the science and knowledge of Australian butterflies from a

generous and erudite person.

Publications of S.J. Johnson and citation in this paper

All Steve Johnson's publications on Australian butterflies are listed and

numbered in a Bibliographic Appendix to this paper. These papers are cited

by boldface number in the following text. Other publications referred to in

this paper are listed in the References and are cited in the text by author and

date. Steve also made significant contributions to veterinary science, in

particular on arboviruses. These publications, 30 in number, are also listed in

the appendix but are not referred to in the text.

162 Australian Entomologist, 2015, 42 (4)

Fig. 1. Stephen James Johnson (13 April 1950 - 16 December 2014).

Australian Entomologist, 2015, 42 (4) 163

Fig. 2. The Johnson family in the early 1980s. Steve's parents Arnold and Stella

seated centre. Their three sons (L to R) Steve, Bruce and Ian at rear, behind daughters

Helene (L) and Elizabeth. Children at front are Steve's two children Sarah and

Matthew with Helene's daughter Pip in centre. Arnold encouraged and supported

Steve and lan in their butterfly interests from a young age and sometimes

accompanied them on remote field trips.

164 Australian Entomologist, 2015, 42 (4)

Early life in Brisbane

Like many active boys in the era before television, Steve wandered the fields,

paddocks and swamps of greater Brisbane, collecting birds' eggs with brother

Bruce, catching snakes, swimming in Bulimba Creek, diving off Hemmant

Railway bridge and shooting sparrows for the Council bounty for pocket

money. His constant companion was mischief. Steve was able to channel his

boundless energy into rowing and football at school. As a robust second

rower, rugby suited his less than genteel manner and, as his Dad described it,

he went at it like a bull at a gate. Sport proved detrimental to his academic

outcome in 1967 but, like most very bright people, he was able to turn this

around in 1968 and matriculate. Steve's life changed in an even more positive

way in 1967 when he met Bronwyn Clarke. They were inseparable and

married in 1972. She became his loving wife, mother of his children (Sarah

and Matthew) and unswerving supporter and confidant. For 38 years she was

Neddy to his Jim (following Steve's love of the Goons).

While butterflies had featured briefly as a child, it was the necessity of his

brother Ian to prepare a butterfly collection for a high school zoology class in

1969 that started a passion in Steve that lasted the rest of his life. He became

an obsessive collector and observer of butterflies, very soon developing a

focus on learning their habits and their life histories. While studying for his

Veterinary degree, he would frequently visit the University of Queensland

Insect Collection (housed at that time in the Agricultural Sciences Building,

adjacent to the Veterinary Sciences Building). According to Geoff Monteith,

the then Curator, Steve would arrive at lunch time (or during lecture breaks)

to peruse the butterfly collection and discuss all things butterfly, occasionally

accompanied by his father Arnold, known to most as Gator — one of the

nicknames applied by Steve. It was a feature of his behaviour that almost

everyone got a nickname, like it or not. Even then Geoff was very aware of

Steve's unforgettable personality and exuberant approach to butterflies. Was

he aware that his own nickname was Fangs?

Steve's time at Warwick saw a major expansion of his collecting interests

and the development of woodworking skills that enabled him to make his

own drawers and cabinets. For the rest of his life he prepared his own

material, constructed his own drawers and cabinets and refined the design. He

also designed and built collecting boxes and other equipment that

accompanied him on field trips everywhere. Over time, his woodworking

skills and tools expanded significantly and it was part of his love of

butterflies that he was able to house his specimens in high quality storage

facilities that he had built himself.

Moving to the tropics

Shifting to Townsville in 1979 was a significant milestone in Steve and

Bronwyn's life and created many new opportunities for not only their

respective professional careers, but also for Steve's growing passion for

Australian Entomologist, 2015, 42 (4) 165

tropical butterfly discovery. He had already completed a significant butterfly

collecting trip to Iron Range and the Mcllwraith Range of Cape York

Peninsula in 1976 and another to Iron Range in 1978 (with his father,

Murdoch De Baar and Darryl Washbourne), when they camped in Cook's

Hut (Fig. 3), the famous butterfly hunters’ dwelling and legendary abandoned

home of former prospector and road maintenance man Reg Cook. Now Steve

was in an even better location to expand his knowledge of Australian tropical

butterflies. It was also how I came to know him.

Fig. 3. Cook's Hut at Iron Range, now demolished: (a) its exterior in 1980 (Photo by

Alan Walford-Huggins); (b) from left, Arnold ‘Gator’ Johnson, Steve Johnson and

Murdoch De Baar pinning specimens in Cook's Hut, 1978 (Photo by Darryl

Washbourne).

Our paths had crossed even before we met. In 1976, Steve and his father had

a grand adventure, their first trip to Cape York Peninsula and Iron Range. In

that same year I also went to Cape York Peninsula and the Mcllwraith Range.

We both ended up at some point catching butterflies at Peach Creek, but that

was something we only discovered later.

We first came together through the pages of The Australian Entomologist

(then known as the Australian Entomological Magazine), where I read a 1979

paper by Sands, De Baar and Johnson (32) on the discovery of Hypochrysops

cleon Grose-Smith, 1900 in Australia and he read a brief note by myself

(Valentine 1979) on some northern Queensland butterflies. Over the next

thirty years there were many original articles published by Steve and I,

sharing the senior authorship: we jointly authored 30 peer-reviewed papers

on Australian butterflies. On the strength of that first piece, Steve turned up at

my office at James Cook University on the 21st February 1980. I was from a

166 Australian Entomologist, 2015, 42 (4)

farming background in Western Australia and had entered academia with

appointment as lecturer in Geography at JCU in 1975. I had developed a

passion for nature as a boy and first discovered butterflies when I started

research on tropical rainforests in northern Queensland. I set out to use them

as cues to the environment but soon came to enjoy them in their own right.

Steve and I immediately established a rapport and a partnership that persisted

until his death. My diary notes: “Steve Johnson - ex Warwick, now with DPI,

Oonoonba - came out to Uni. Very keen collector with 10 years experience.

Plan some trips together.’ Steve Johnson became my close friend - my best

mate - for 35 years, half my life. We undertook many trips together of

thousands of kilometres by 4WD vehicle, by aircraft and by helicopter over

many parts of Australia.

Steve was a genuine pioneer in the study of Australian tropical butterflies and

was the most accomplished observer of butterflies in the field that I knew.

Much of our knowledge of Australian tropical butterflies is due to Steve's

observations right across the continent. He was not just an expert but was

always generous with his knowledge; dozens of people benefitted from his

personal assistance. In preparing this account I have received numerous

comments from others about how kind and helpful Steve was to them. He

shared his knowledge willingly and he gave strong personal support to

everyone he met. Every request was responded to quickly and generously.

Steve was a very positive person himself and he shared that with others,

finding a way to acknowledge good work by them.

Steve was prolific in writing up his observations and was an excellent author.

He later was a first-rate editor and referee for publications.

For Steve and I the relationship transcended butterflies. Our families were

drawn in to numerous picnics and camping expeditions that, for some reason,

always took us to excellent butterfly places. Our children played together

under the watchful eyes of their mothers, Bronwyn and Valerie, while their

fathers were off stalking butterflies. Some places were close to home around

Townsville but others required the two families to take extensive trips - to

Iron Range and to Rocky River and to other places on Cape York Peninsula.

Steve's children Matt and Sarah and our daughters Polly, Leonie and Kate

were good friends and all loved the farm environment of Oonoonba

Veterinary Laboratory where the Johnson family lived.

While based in Townsville, Steve not only completed his BSc in Entomology

at the University of Queensland (including a stint full time back in Brisbane),

graduating in 1984, but he commenced and completed a PhD in Veterinary

Entomology at James Cook University, graduating in 1989. His PhD thesis,

Studies of Stephanofiliariasis in Queensland, was based on abattoir, field and

laboratory studies and is a major contribution in the field. Since completion it

has been downloaded from the James Cook University online library a total

of 553 times (as at June 2015) and is still being accessed frequently. As an

Australian Entomologist, 2015, 42 (4) 167

indication of its global significance, downloads include those by researchers

in the USA (100), Australia (73), Indonesia (56), India (50), Germany (45),

China (21) and Bangladesh (21). In keeping with his collaborative approach,

Steve Johnson's professional publications in the field of veterinary

entomology include a total of 30 refereed papers with 42 different co-authors

(excluding his butterfly papers; see Appendix). The main focus of Steve's

professional work was in blue tongue viruses and the risks to the Australian

sheep and beef industries. He was a very significant participant in the sentinel

herds established as part of national biosecurity arrangements, involving

regular monitoring of small herds across the tropics in order to detect the

possible arrival of arboviruses. Several major projects he managed required

extensive field work across the northern tropics, from Western Australia to

Queensland.

The focus of Steve Johnson's butterfly work

While there had been significant work on butterflies in tropical Australia

from the very early years of European settlement, there had been few resident

collectors since Frederick Parkhurst Dodd resided in Kuranda (Monteith

1991). Steve brought a substantial scientific knowledge base that was very

effective in revitalizing the biological studies of Australian tropical

butterflies. In the early years his focus was exploratory: field studies in

locations where little was known about the butterfly fauna. Cape York

Peninsula continued to be a major drawcard, with almost every trip producing

new material and new observations. But, in addition, Steve explored local

environments in the Townsville and Cairns region and planned specific

searches in regions wider afield in tropical Australia.

As his knowledge and familiarity with Queensland butterflies advanced,

Steve began to look further afield, both within the tropics (in Western

Australia and the Northern Territory) and also in the southern States. But his

primary focus remained the tropical species and their habitats. A typical trip

to Iron Range during the 1980s might involve three weeks camped in the

rainforest, with tarps for weather protection and everything required for

collection. Cook’s Hut, often used by visiting biologists to Iron Range and

now demolished, was used occasionally but generally not and all camping

needs had to be brought in with the vehicle, including fuel. In those days it

was a three-day effort to get there from Townsville, with very rough roads

beginning at Mt Carbine, where the bitumen ended. Winching was often

required to cross flooded creeks and boggy areas. Major effort was needed at

the large Wenlock and Pascoe Rivers on the final stage of the trip and

crossing the several fords on the Claudie River was rarely easy. Each trip

required shovels, chainsaws and winches just to get there and in the earliest

years fuel was barged in to Lockhart River.

Being the person he was, Steve was often drawn in to provide free veterinary

services for the local community at Lockhart River during his field trips. That

168 Australian Entomologist, 2015, 42 (4)

also involved neutering the many unmanaged dogs in the township at the

request of the council and local people.

Field methodology

Collecting specimens at Iron Range was usually focused along the bush

tracks and mining roads, including the tops of Lamond and Phillip Hills.

Particularly long net handles had been made by Steve to try and access the

higher canopy areas - these poles were carried on the vehicle roof tops

secured by stockings, collected for that purpose from female friends and

relatives. With Steve's typical wit these poles had to have a name and

became the gorgonzolas - with reference to how ‘high’ they could reach.

Even with these long extensions many butterflies were out of reach, a

continuing frustration for decades. Another focus was to use hilltops or adult

leks to try and collect specimens. At Iron Range, Phillip and Lamond Hills,

plus other high points and ridges, worked very well for many species. In the

case of the many Philiris Röber species (Lycaenidae), it was noted that adults

aggregated on certain very tall trees and could be collected using the long

pole extensions, sometimes aided by standing on top of the vehicle roof-rack.

During the three week camp, butterfly collecting occurred along the roads,

which were walked every day multiple times. At that time there were many

flowers that attracted adults and enabled easy collecting.

One approach that Steve adopted was to search for juvenile stages. Any

female was a good target to watch and wait. Typical female oviposition

behaviour, fluttering around plants for example, would elicit a reprieve from

the net and some careful observation. Rewards were not common but did

allow a number of new life histories to be obtained. That nearly always

required multiple steps of identifying the plant, acquiring a specimen for the

purpose-built shade houses back in Townsville and then, on later trips,

searching plants for juvenile stages of the butterfly to take back for captive

rearing. Occasionally, if small plants were evident in the bush they might be

dug up and taken back with larvae or eggs intact. Or, if the plant was already

well known and occurred locally in the Townsville region, the juvenile forms

might be taken back with enough bagged material to last the journey south.

For example, larvae on mistletoe sometimes may be readily transferred from

one species to another. Sometimes larval stages could be found by simply

observing evidence of feeding marks on the plants or, in the case of

Hesperiidae, looking deliberately for larval shelters. On other occasions,

adult females would be collected and bagged with any known or suspected

larval food plant to try and induce oviposition.

Adult specimens, once collected, were always set in the field. Steve normally

set each specimen on the day it was caught and avoided placing it in a paper

triangle for later setting. To this end, he designed and built large setting boxes

with several foam boards that would take many specimens set in the field

(Fig. 4). These were generally given high priority in the packing of the

Australian Entomologist, 2015, 42 (4) 169

vehicle. In more recent years, when 12-volt refrigerators were used,

specimens were sometimes kept fresh or even frozen for the journey home.

Fig. 4. Steve Johnson setting butterflies on Dauan Island in 2004. The polystyrene

foam boards fitted into a specially made rugged box that was essential kit on every

field trip. The silky oak pin boxes (foreground) were made to carry setting equipment

securely.

During the field trips at Iron Range, a list of species collected and observed

was made. It was not unusual to record over 120 species during a three-week

field trip and there was an attempt to improve the list each visit. I think the

maximum recorded was 128 species in three weeks. Another dimension to his

Iron Range work was the adaptation of canopy traps, following a design from

Mike Barnett who had used them in Uganda. These were suspended under the

canopy, baited with rotting fruit and other delights, and proved very effective

for Charaxes latona Butler, 1865 and a number of other nymphalids (Fig 5).

Iron Range has strong challenges for collecting and two species illustrate this

well. The hesperiid butterfly Rachelia extrusa C. & R. Felder, 1867 was

initially collected in Australia on the slopes of Mt Tozer by Geoff Monteith

(Atkins 1975) and very few specimens were known until Steve collected

males from a tree canopy near the second crossing of Gordon's Creek. For

the next two decades all specimens collected were from this exact same site

and all were males. It was clear that this was a lek for males. Females were

discovered later at other rainforest-edge sites, including specimens reared by

Steve from the Mcllwraith Range (45). The lycaenid butterfly Hypochrysops

cleon Grose-Smith, 1900 was first found by Steve at Iron Range on a treetop

170 Australian Entomologist, 2015, 42 (4)

from a ridge just south of the old track up Phillip Hill (32). Since that time all

other specimens of this species have come from that same treetop, except for

the only two females known, collected by David Lane and myself in 1984,

near the first crossing of Gordon's Creek (unpubl. obs.). There are still no

other females known but males continue to lek at the original tree. Specimens

of many species from Iron Range (Fig. 6), in collections across Australia,

have been made possible by Steve's discoveries.

Fig. 5. Charaxes latona (Nymphalidae: Charaxinae): (a) specimen perched in the

canopy at Iron Range, photographed from Steve's cherry picker; (b) Steve with one of

the fruit-baited traps hauled to the canopy to catch Charaxes and other nymphalids.

Using historical records and environmental data

Steve was a great advocate of learning from earlier workers and would

frequently consult the literature or talk to other butterfly people to gain an

insight on how to learn more. A good example was the work on the life

history of the Moth Butterfly, Liphyra brassolis Westwood, 1864

(Lycaenidae). The published observations and F.P. Dodd’s speculations

(Dodd 1902) were studied carefully and collection data from others then was

used to refine the search. It was apparent that G.A. Waterhouse had found the

species on Great Palm Island (Waterhouse 1932) and consulting with an

Ingham fruit grower (Herb Bosworth) led to the conclusion that a careful

search of old citrus orchards on Palm Island might be worthwhile. Having

arranged for permission, the expedition proceeded with great success and

resulted in the discovery of many larvae and pupae. Because direct evidence

of butterfly larval carnivory was being sought, Steve set up a glass aquarium

so that the feeding behaviour could be observed and photographed. This

enabled the first photo of actual carnivory to be taken and published (16).

Australian Entomologist, 2015, 42 (4) 171

Fig. 6. Steve Johnson and Peter Valentine at Iron Range, one of dozens of trips made

over forty years (Photo by Andrew Rankin).

172 Australian Entomologist, 2015, 42 (4)

Another example of field planning was an attempt to predict maximum

butterfly activity in the wet-dry tropics, especially in central Cape York

Peninsula. There were good grounds to expect very high early wet season

activity amongst butterflies that specialised in avoiding the dry conditions

and Steve set up a target to visit Mt White, a well known hilltop location near

Coen. The timing, it was reasoned, needed to be not long after the first major

rainfall - presumed to break the diapause of adults or pupae. Such rain might

also initiate fresh growth on potential larval food plants. Coordinating with a

local cattle station owner, the situation was monitored and, in January 1988,

the trip to Mt White was timed for 10 days after a major rain event. While

conditions were appalling for field work (very hot and very steamy), the

outcome was astonishing, with both a huge abundance of adult butterflies

(17) and two new life histories recorded as a result - Libythea geoffroy

nicevillei Olliff, 1891 (18) and Graphium aristeus (Stoll, [1780]) (36).

The Torres Strait islands

A significant frontier for Steve was to investigate the butterfly fauna of the

Torres Strait islands. This era of his work began in 1983, when Steve made a

brief trip to Thursday and Darnley Islands (4). Steve's quarantine work took

him to Torres Strait occasionally and he developed connections that

facilitated later butterfly trips. He was able to gain an occasional berth with

customs and quarantine missions up the east coast and within the Torres

Strait. While collecting samples for arbovirus work, he was also able to

inspect the butterfly fauna. This part of his butterfly activity was further built

up after his physician brother Ian undertook some medical work on Thursday

Island in the 1980s and provided a knowledge base from which Steve could

later explore the more remote islands, including Dauan in 1989. Visits to

Murray and Darnley Islands (in 1993 and 1994 respectively) and Moa Island

in 1993 (38) added to the Torres Strait exploration. Dauan became a favourite

destination and was very prolific. Initially, the trips to Dauan involved a

flight to Saibai and accommodation at the school and then persuading a local

inhabitant to provide transport across to Dauan. After purchasing a drum of

outboard petrol, a walk along the beach usually revealed an aluminium dingy,

with the owner willing to provide the crossing. On one trip it was a pair of

young boys who seemed to delight in ensuring the passengers (Steve and I)

suffered the maximum impacts from wave thumps. Later, a ferry service was

initiated and access became a little more reliable, but not much. On our 2004

trip the ferry had drifted and washed up on rocks the night before we were

due to return to Saibai. After a fair delay the boat was retrieved and brought

around for loading. The other passenger was the Dauan Council chair, Mrs

Margaret Mau, which is probably why the vessel was made ready. By the

time we reached Saibai we were up to our shins in sea water and sinking fast,

just making the jetty in time with setting boxes held high.

Australian Entomologist, 2015, 42 (4) 173

An increase in species collected at Dauan occurred when Steve (and Ian)

arranged helicopter support to get to the top of Mt Cornwallis, the island's

central 275 m peak surrounded by vine thicket and suspected of being a good

hilltop for collecting. It was here that Steve first collected the Map Butterfly,

Cyrestis achates nedymnus C. & R. Felder, 1865, having seen it previously

on one of the beaches in the same year it was originally collected (Lambkin

and Knight 2005). In his last year of life, while badly impacted by the cancer,

Steve went up to Dauan and camped overnight on the peak of Mt Cornwallis

with his brother Ian. He always loved his time on this island.

Tropical Western Australia

Partly connected with his professional quarantine and biosecurity work on

monitoring the tropical coastline for invading arboviruses, Steve was invited

on a cruise through the offshore islands along the Kimberley coast in April

199]. Support included a helicopter to access some of the islands and Steve

was able to sample many of the island butterfly populations (5).

This also stimulated further his interest in exploration of this part of Australia

and, to this end, Steve arranged for a fixed-wing aircraft to fly in to the

Mitchell Plateau in the 1995 wet season. As space was at a premium, priority

was given to butterfly collecting material, the inevitable setting boxes

included. A bush pilot, John Collins, took us in to a remote airstrip and hired

us his old 4WD vehicle, which he kept near the airstrip, so we could get

around. We were the only people on the plateau at this time (the road was

still closed). Because the plane was very small we had very little luggage.

After the critical butterfly equipment we had room for a sleeping bag each, a

gas stove and bottle, a billy and two plates. Setting butterflies was always a

high priority, so Steve built a collapsible table that could fit into the plane so

we had a work surface. The only food we could fit in was dehydrated - add

water and boil type: the water came from a nearby creek. We would consume

these packet meals having to imagine the bit of the instruction that said *add

steak’ or ‘add chicken’. Steve referred to his designer table as the ‘have table,

will travel’ and it later made many more trips with us. The trip was a great

success and was followed by further time at Kalumburu before a cyclone

drove us out (20).

Discovering cancer

Cancer became part of the Johnson family's life when his wife Bronwyn was

diagnosed and treated for breast cancer in the mid 1990s. In 1998 Steve had

an off-road caravan built and he began to use that with Bron to spend time in

remote parts of Queensland, including Iron Range, looking for butterflies. In

2001 we took long service leave and, with our wives, spent three months in

northwestern Western Australia, down the west coast and through the

southwest. This was a combination butterfly trip and family enjoyment but,

for a lot of the time, butterflies won. It was Steve's way of getting more time

with Bronwyn and we completed useful butterfly observations also (24).

174 Australian Entomologist, 2015, 42 (4)

In April 2004, Steve and I returned from a trip to Dauan Island in Torres

Strait only to discover that Steve had been diagnosed with terminal cancer

and was initially given only a few weeks to live. This devastating news

caused an immediate shift to Brisbane, to be closer to family. Sadly,

expectations to the contrary, it was Bronwyn who passed away first, a very

sad loss. Subsequently, despite declining health, Steve was keen to get out

into butterfly locations in other parts of Australia that he had not visited. He

went on many trips, including Kangaroo Island (October 2004), Alice

Springs (February 2005, with Grant Miller), Lockerbie Scrub (April 2005)

and Tasmania (January 2006). During these ventures Steve was often very

sick from his cancer; it was tough on him but he was determined to

experience some of the local species in the wild. We also took several trips to

Western Australia up until 2013 (two trips in that year for the Julimar form of

Neolucia agricola occidens Waterhouse & Lyell, 1914 in March and Ogyris

subterrestris petrina Field, 1999 in October). His brother Ian also spent a lot

of time with Steve in his last few years, especially on trips to Torres Strait

chasing new species and life histories.

The cherry picker innovation

No account of Steve's butterfly life can ignore the cherry picker innovation

(Fig. 7). Over many years of our visits to Iron Range, on the long drive back

to Townsville there would eventually be a reference to all the butterflies we

could see up in the canopy of the rainforest but which we could not catch,

despite our extension poles. Steve would eventually bemoan ‘If only we had

a cherry picker'. Following his diagnosis of cancer he concluded it was now

or never and decided to have a cherry picker made to suit his purpose. Based

on an Isuzu rigid truck, the tower reached to 20 metres with room for two

people in the bucket. He drove it up from Brisbane to northern Queensland in

June 2006 to try it out. We went to Cooktown and Shipton's Flat to test its

suitability and gain some experience. In July that year, Steve drove it to Iron

Range for its first field outing. It served well for many more years. We would

drive it up at the end of the wet season and it would stay at Iron Range

through the dry season, parked at the National Park headquarters between

visits. Several trips to Iron Range were made each year, usually with one or

more collectors joining Steve. These included Peter Wilson, Peter Samson,

Kerrod Beattie, Grant Miller, Darryl Washbourne, David Lane, Bill Graham

and others. I went with Steve on many occasions.

He also used the cherry picker in southern Queensland, on one occasion

supporting an Entomological Society of Queensland field trip to Mt Glorious

in April 2007. He gave rides to the canopy to a few people and helped a PhD

student collect scale insects in the canopy (Figs 7a-b).

These trips with the cherry picker were much enjoyed by Steve and all who

went with him. They did result in some excellent outcomes for his butterfly

work. For example, after seeing the large high-flying nymphalid Apaturina

Australian Entomologist, 2015, 42 (4) 175

erminea (Cramer, 1779) on many occasions, the cherry picker enabled

specimens to be collected as they rested at height on tree trunks (unpubl.

obs.). Description of the complete life history of Charaxes latona Butler,

1865 (Fig. 5a) (47) was enabled after eggs were observed being laid on

leaves in the canopy and subsequently collected. Work on other lycaenids

that laid eggs high in the forest was also facilitated, including Hypochrysops

hippuris Hewitson, 1874 (22), Hypochrysops elgneri (Waterhouse & Lyell,

1909) (30) and Philiris ziska (Grose-Smith, 1898) (29). In December 2008, a

journalist and photographer joined Steve and I at Iron Range. They wanted a

story on butterflies but in the end they loved the story of Steve. This story

and photos appeared in the 2009 April/June edition of Australian Geographic

(Van Tiggelin 2009). The journalist, John Van Tiggelin, had many positive

things to say about Steve in the article. When I spoke with him at the time of

Steve's funeral he described the Iron Range experience that year as his

favourite assignment and Steve as his favourite character.

, di z l adj c

Fig. 7. Steve Johnson's cherry picker truck: (a) in use on an Entomological Society of

Queensland field trip to Mt Glorious in April 2007; (b) Steve (on right) with student

Ben Nomark searching for scale insects; (c) Steve (on right) with Peter Valentine high

in the canopy at Iron Range in 2008. (Photos a and b by Noel Starick, c by Andrew

Rankin).

Publications and influence

Steve was a prolific author and contributed a total of 47 peer-reviewed papers

on Australian butterflies (see Bibliographic Appendix). His collaborative and

supportive approach also meant that he published with many of Australia's

most knowledgeable butterfly scholars. His co-authors number 15 and, while

176 Australian Entomologist, 2015, 42 (4)

the bulk of his publications were shared with me (30 papers), he also shared

authorship more than once with four other authors and was a co-author with

10 other researchers. The publications include range extensions and habitat

notes through many life history descriptions to several descriptions of new

taxa (Table 1). Much of the focus was on Queensland but papers also covered

Western Australia (both the south-west and the north) and the Northern

Territory. The majority of Stephen Johnson's butterfly publications were in

Australian Entomologist, which he also supported as a reviewer.

Significant work on the life histories of Australian butterflies covered many

species but there were other contributions of lesser extent covering many

other species. These were often published in papers covering biogeography or

regional records of species.

Table 1. Butterfly species for which Steve Johnson made a significant published life

history contribution. The relevant publications are shown in brackets after each

species.

Family / Species

HESPERIIDAE RIODINIDAE

Allora doleschallii (C. Felder, 1860) (7) Praetaxila segecia (Hewitson, 1861) (31)

Rachelia extrusa (C & R. Felder, 1867)

(45) LYCAENIDAE

Trapezites atkinsi Williams, Williams & ` Liphyra brassolis Westwood, 1864 (16)

Hay, 1998 (44)

Trapezites taori Atkins, 1997 (23) Hypochrysops elgneri (Waterhouse &

Lyell, 1909) (30)

Neohesperilla senta (Miskin, 1891) (25) Hypochysops polycletus (Linnaeus, 1758)

(14)

Hesperilla sarnia Atkins, 1978 (15) Hypochrysops miskini (Waterhouse, 1903)

(35)

Telicota brachydesma Lower, 1908 (43) Hypochrysops hippuris (Hewitson, 1874)

(22)

PAPILIONIDAE

Graphium aristeus (Stoll, [1780]) (36) Philiris diana Waterhouse & Lyell, 1914

(40)

NYMPHALIDAE Philiris ziska (Grose-Smith, 1898) (29)

Taenaris artemis (Vollenhoven, 1860)

(10) Deudorix epirus (C. Felder, 1860) (3)

Melanitis constantia (Cramer, 1777)

(13) Nesolycaena medicea Braby, 1996 (21)

Orsotriana medus (Fabricius, 1775) (13)

Charaxes latona Butler, 1865 (47) Catopyrops ancyra (C. Felder, 1860) (39)

Libythea geoffroy Godart, 1824 (18)

Australian Entomologist, 2015, 42 (4) 177

Steve Johnson was also the first to record several taxa from Australia,

including. Paraduba metriodes (Bethune-Baker, 1911) (Lycaenidae) (11),

Charaxes latona Butler, 1865 (Nymphalidae) (6), Melanitis constantia

(Cramer, 1777) (Nymphalidae) (13), Euploea netscheri Snellen, 1889

(Nymphalidae) (20), Philiris azula Wind & Clench, 1947 (Lycaenidae) (9),

Hypochrysops cleon Grose-Smith, 1900 (Lycaenidae) (32), Prosotas gracilis

(Róber, 1886) (Lycaenidae) (20), Nacaduba calauria (C. Felder, 1861)

(Lycaenidae) (20) and Cethosia cydippe damasippe C. & R. Felder, 1867

(Nymphalidae) (27). In addition, Steve rediscovered Taenaris artemis

queenslandica Rothschild, 1916 (Nymphalidae) at the Lockerbie scrub, Cape

York Peninsula (10) and also helped rediscover Tagiades nestus (C. Felder,

1860) on Dauan Island in Torres Strait (46). He contributed to our knowledge

of many other species, including Delias lara (Boisduval, 1836) (Pieridae)

(28).

Although Steve was not focused on taxonomic work but rather encouraged

and supported others in their endeavours, he did take the initiative in

describing several taxa. These were Hesperilla malindeva dagoomba Johnson

& Valentine, 1994 (Hesperiidae) (19), Hesperilla crypsargyra binna Johnson

& Wilson, 2005 (Hesperiidae) (26), Jalmenus notocrucifer Johnson, Hay &

Bollam, 1992 (Lycaenidae) (8) and Jalmenus inous bronwynae Johnson &

Valentine, 2007 (Lycaenidae) (24). The subspecific epithet dagoomba was

chosen by Steve as an Aboriginal place name for Magnetic Island and his

choice of bronwynae was to honour his wife.

Many of the species on which Steve Johnson made a significant contribution

are shown in Figs 8 and 9.

The Johnson Collection at the Museum of Tropical Queensland

Steve Johnson put together, over many years, an outstanding collection of

Australian butterflies. It 1s the product of over forty years of collecting,

although most of the material was added after Steve moved to Townsville.

Given the initial diagnosis of terminal cancer in 2004, with perhaps a few

weeks to live, Steve arranged for the collection to go to the Museum of

Tropical Queensland. The Johnson Collection, housed in drawers and

cabinets built by Steve over the years, is one of the most comprehensive

collections of Australian butterflies (Table 2, Fig. 10). Consisting of over

12,000 specimens, it covers all families and is missing only two genera and a

handful of species (Table 2). Note that Steve did not include the five species

only known from Christmas Island and the one species only known from

Norfolk Island as part of the Australian Biogeographic Region. Of the

missing species, Steve had seen two in the wild (Prosotas gracilis and

Tagiades nestus) and had made several trips to Burrell's Trig in the Northern

Territory in search of another, Acrodipsas decima Miller & Lane, 2004. The

vast majority of the specimens in the collection were netted by Steve in the

wild or reared by him from material he collected in the wild.

178 Australian Entomologist, 2015, 42 (4)

Fig. 8. Australian butterfly species strongly associated with Steve Johnson: (a)

Taenaris artemis, (b) Apaturina erminea, (c) Charaxes latona; (d) Melanitis

constantia; (e) Liphyra brassolis (9); (f) Graphium aristeus (3), (g-h) Libythea

geoffroy (LO & 9). All of these species are from tropical Queensland, mostly Cape

York Peninsula.

Australian Entomologist, 2015, 42 (4) 179

Fig. 9. Australian butterfly species about which Steve Johnson contributed significant

new knowledge: (a-b) Hypochrysops elgneri (LO & 9); (c-d) H. hippuris (3 & 9);

(e-f) H polycletus (8 & 9); (g-h) H cleon (3 & 9); (i-j) Philiris diana papuana

(£ & 9); (k-I) Nesolycaena medicea (3 & 9); (m-n) Nacaduba calauria (À & 9);

(o) Prosotos gracilis (9); (p-q) Catopyrops ancyra (£ & 9); (r) Tagiades nestus (É);

(s-t) Allora doleschallii ($ & 9), (u) Telicota brachydesma (3); (v-w) Rachelia

extrusa (3 & 9); (x) Trapezites taori (9); (y) T. atkinsi (8). All but one of these are

from the Queensland tropics, the last from southwestern Australia.

180 Australian Entomologist, 2015, 42 (4)

Table 2. Details of the Johnson Collection at the Museum of Tropical Queensland.

These totals do not include specimens Steve collected during 2015, which added a

few more individuals from Dauan Island and Weipa but no new species.

Family Specimens Species Species (and Australian species

number) missing

Reared

Hesperiidae 3750 122 86 (1366) Tagiades nestus

Papilionidae 303 18 16 (176) None

Pieridae 867 36 2] (348) Appias celestina

Nymphalidae 1867 83 46 (580) Euploea modesta,

Junonia erigone,

Lexias aeropa,

Taenaris catops

Libytheidae 26 1 1 (3) None

Lycaenidae 5499 151 113 (2643) Acrodipsas decima,

Nothodanis schaffera,

Prosotas gracilis

Totals 12,312 411 283 (5116) 9 species missing

Fig. 10. (a) Part of the Johnson Collection cabinets and drawers built by Steve

Johnson; now in the Museum of Tropical Queensland; (b) Steve Johnson and Peter

Valentine working on the Johnson Collection, Museum of Tropical Queensland,

Townsville, 2009. (Photos by (a) Niel Bruce, MTQ; (b) Andrew Rankin).

Australian Entomologist, 2015, 42 (4) 181

Conclusion

Stephen Johnson is very well remembered by many in the Australian

butterfly world. He personally supported all he met and shared his excellent

knowledge with anyone who had an interest. He was generous in every way

and an enthusiast for all things butterfly. A nice example was the ‘birthday

gift' to Grant Miller of a pair of Pithecops dionisius (Boisduval, 1832). Grant

had been trying to find these for so long he had taken to calling them

“Mythecops'. Steve was similarly generous with specimens to others and

many collections include excellent material with a label indicating S.J.

Johnson as the collector. His premature death leaves a gap in our butterfly

science, especially in northern Australia. My own life has hundreds of

wonderful memories from our intertwined butterfly songlines.

Acknowledgements

In preparing this account I have benefited very much from the support of

Steve's immediate family, especially his brother Ian and his son Matthew. Ian

prepared a moving eulogy to Steve at his funeral and that provided me with

much of the framework about Steve's early life. Ian was also able to fill in

details about some of the more recent collecting trips to Dauan as well as his

recollections of early trips to Iron Range. Many other butterfly people were

pleased to share their recollections about Steve and I thank them all, in

particular Geoff Monteith, Grant Miller, Peter Wilson, Don Sands, Kerrod

Beattie, Trevor Lambkin, Darryl Washbourne, Ted Edwards, Steve Brown,

Cliff Meyer, Max Moulds, David Lane, Mike Barnett, Bill Graham and John

Young. Geoff Monteith has given much editorial assistance in preparing the

text and illustrations.

References

ATKINS, A. 1975. The first record of Rachelia extrusa (C. & R. Felder) (Lepidoptera:

Hesperiidae: Trapezitinae) in Australia. Journal of the Australian Entomological Society 14:

237-241.

DODD, F.P. 1902. Contribution to the life-history of Liphyra brassolis, Westw. Entomologist

35: 153-156.

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

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

MONTEITH, G.B. 1991. The Butterfly Man of Kuranda - Frederick Parkhurst Dodd.

Queensland Museum, Brisbane; 32 pp.

VALENTINE, P.S. 1979. Notes on the biology and distribution of some north Queensland

lycaenid butterflies. Australian Entomological Magazine 6: 55-56.

VAN TIGGELIN, J. 2009. The butterfly effect. Australian Geographic 94: 80-89.

182 Australian Entomologist, 2015, 42 (4)

BIBLIOGRAPHIC APPENDIX

I: LEPIDOPTERA PUBLICATIONS OF STEPHEN JAMES JOHNSON

Listed alphabetically by author and numbered sequentially. These

publications are cited in the text by number.

1. ATKINS, A.F., EDWARDS, E.D., BRABY, M.F., JOHNSON, S.J. and VALENTINE, P.S.

2003. The butterflies of White Mountains National Park, Northern Queensland, and adjoining

localities. Pp 7-10, in White Mountains Scientific Study Report, Geography Monograph Series

No 9, Royal Geographical Society of Queensland, Brisbane.

2. DE BAAR, M. and JOHNSON, S J. 1980. Rhopalocera notes and hosts from Iron Range trip,

June-July 1978. News Bulletin of the Entomological Society of Queensland 8: 45.

3. DE BAAR, M. and JOHNSON, S.J. 1996. Notes on the foodplant of Deudorix epirus agimar

Fruhstorfer (Lepidoptera: Lycaenidae). Australian Entomologist 23: 36.

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

Entomological Society of Queensland 11: 117-119.

5. JOHNSON, S.J. 1993. Butterfly records of interest from northern Australia. Australian

Entomologist 20: 75-76.

6. JOHNSON, SJ. and DE BAAR, M. 1993. First record of Charaxes latona Butler

(Lepidoptera: Nymphalidae) from Australia. Australian Entomological Magazine 6: 94-96.

7. JOHNSON, S.J. and DOHERTY, W.M. 1991. The life history and distribution of A/lora

doleschallii doleschallii (Felder) (Lepidoptera: Hesperiidae) from northern Queensland.

Australian Entomological Magazine 18: 111-112.

8. JOHNSON, SJ., HAY, R.W. and BOLLAM, H.H. 1992. Jalmenus notocrucifer sp. n.

(Lepidoptera: Lycaenidae) from south Western Australia. Australian Entomological Magazine

19: 69-74.

9. JOHNSON, S.J. and JOHNSON, LR. 1984. First record of Philiris azula Wind and Clench

(Lepidoptera: Lycaenidae) from Australia. Australian Entomological Magazine 10: 89-90.

10. JOHNSON, S.J. and JOHNSON, IR. 1991. Notes on the life history of Taenaris artemis

(Snellen von Vollenhoven) (Lepidoptera: Nymphalidae) in Australia and the rediscovery of 7. a.

queenslandica Rothschild. Australian Entomological Magazine 18: 85-86.

11. JOHNSON, SJ. and JOHNSON, LR. 2013. The first record of Paraduba metriodes

(Bethune-Baker) (Lepidoptera: Lycaenidae) within Australian limits. Australian Entomologist

40: 173-174.

12. JOHNSON, S.J., JOHNSON, LR. and VALENTINE, P.S. 1994. New and interesting

butterfly records (Lepidoptera) from Torres Strait Islands. Australian Entomologist 21: 121-124.

13. JOHNSON, S.J., JOHNSON, I.R. and VALENTINE, P.S. 1995. Notes on the early stages of

Orsotriaena medus moira Waterhouse & Lyell and Melanitis constantia Cramer (Lepidoptera:

Nymphalidae: Satyrinae) from Torres Strait, Australia. Australian Entomologist 22: 65-68.

14. JOHNSON, S.J. and SAMSON, P.R. 1999, The life history of Hypochrysops polycletus

rovena Druce (Lepidoptera: Lycaenidae). Australian Entomologist 25: 121-123.

15. JOHNSON, S.J. and VALENTINE, P.S. 1983. Notes on the biology and morphology of

Hesperilla sarnia Atkins (Lepidoptera: Hesperiidae). Australian Entomological Magazine 10: 6-

Australian Entomologist, 2015, 42 (4) 183

16. JOHNSON, S.J. and VALENTINE, P.S. 1986. Observations on Liphyra brassolis Westwood

(Lepidoptera: Lycaenidae) in north Queensland. Australian Entomological Magazine 13: 22-26.

17. JOHNSON, SJ. and VALENTINE, P.S. 1988. Butterflies collected on Mount White in

January 1988. News Bulletin of the Entomological Society of Queensland 16: 12-13.

18. JOHNSON, S.J. and VALENTINE, P.S. 1989. The life history of Libythea geoffroy nicevillei

OIliff (Lepidoptera: Libytheidae). Australian Entomological Magazine 16: 59-62.

19. JOHNSON, SJ. and VALENTINE, P.S. 1994. An insular subspecies of Hesperilla

malindeva Lower (Lepidoptera: Hesperiidae) from northern Queensland. Australian

Entomologist 21: 33-36.

20. JOHNSON, S.J. and VALENTINE, P.S. 1997. Further observations and records for

butterflies (Lepidoptera) in northern Australia. Australian Entomologist 24: 155-158.

21. JOHNSON, S.J. and VALENTINE, P.S. 2001. The life history of Nesolycaena medicea

Braby (Lepidoptera: Lycaenidae). Australian Entomologist 27: 109-112.

22. JOHNSON, S.J. and VALENTINE, P.S. 2001. Notes on the life history of Hypochrysops

hippuris nebulosis Sands (Lepidoptera: Lycaenidae). Australian Entomologist 28: 13-16.

23. JOHNSON, S.J. and VALENTINE, P.S. 2004. Notes on the biology and distribution of

Trapezites taori Atkins (Lepidoptera: Hesperiidae). Australian Entomologist 31: 13-19.

24. JOHNSON, S.J. and VALENTINE, P.S. 2007. A new subspecies of Jalmenus inous

Hewitson (Lepidoptera: Lycaenidae) from Shark Bay, Western Australia. Australian

Entomologist 34: 77-83.

25. JOHNSON, S.J., VALENTINE, P.S. and LANE, D.A. 1994. Notes on life histories and

biology of the species of Neohesperilla Waterhouse and Lyell (Lepidoptera: Hesperiidae).

Australian Entomologist 21: 55-59.

26. JOHNSON, S.J. and WILSON, P.R. 2005. A new subspecies of Hesperilla crypsargyra

(Meyrick) (Lepidoptera: Hesperiidae) from southern Queensland and a new status for Hesperilla

hopsoni Waterhouse. Australian Entomologist 32: 163-171.

27. LAMBKIN, T.A. and JOHNSON, S.J. 2015. Cethosia cydippe damasippe C. & R. Felder,

1867 (Lepidoptera: Nymphalidae: Heliconiinae) in Torres Strait, Queensland, Australia.

Australian Entomologist 42: 67-72.

28. MILLER, C.G. and JOHNSON, S.J. 2015. Further records of Delias lara (Boisduval, 1836)

(Lepidoptera: Pieridae) from Australia. Australian Entomologist 42: 35-37.

29. SAMSON, P.R. and JOHNSON, S.J. 2009. The life history and adult morphology of Philiris

ziska titeus D’ Abrera (Lepidoptera: Lycaenidae). Australian Entomologist 36: 113-118.

30. SAMSON, P.R., JOHNSON, S.J. and VALENTINE, P.S. 1997. The life history of

Hypochrysops elgneri barnardi Waterhouse (Lepidoptera: Lycaenidae). Australian Entomologist

24: 159-163.

31. SAMSON, P.R., JOHNSON, S.J. and WILSON, P.R. 1999. The life history of Praetaxila

segecia punctaria (Fruhstorfer) (Lepidoptera: Lycaenidae: Riodinidae). Australian Entomologist

26: 57-63.

32. SANDS, D.P.A., DE BAAR, M. and JOHNSON, S.J. 1979. First record of Hypochrysops

cleon Grose-Smith (Lepidoptera, Lycaenidae) from Australia. Australian Entomological

Magazine 6: 23-24.

33. VALENTINE, P.S. and JOHNSON, S.J. 1982. New records of Lycaenidae and Hesperiidae

(Lepidoptera) from northern Queensland. Australian Entomological Magazine 9: 1-3.

184 Australian Entomologist, 2015, 42 (4)

34. VALENTINE, P.S. and JOHNSON, S.J. 1988. Some new larval food plants for north

Queensland Lycaenidae (Lepidoptera). Australian Entomological Magazine 14: 89-91.

35. VALENTINE, P.S. and JOHNSON, S.J. 1989. Polyphagy in larvae of Hypochrysops miskini

miskini (Waterhouse) (Lepidoptera: Lycaenidae). Australian Entomological Magazine 16: 1-3.

36. VALENTINE, P.S. and JOHNSON, S.J. 1989. Observations on the life history of Graphium

aristeus parmatus (Gray) (Lepidoptera: Papilionidae). Australian Entomological Magazine 16:

17-20.

37. VALENTINE, P.S. and JOHNSON, S.J. 1992. Late dry season butterflies on Cape York

Peninsula. Victorian Entomologist 22: 87-91.

38. VALENTINE, P.S. and JOHNSON, S.J. 1993. The butterflies of Moa Island, Torres Strait.

Victorian Entomologist 23: 116-121.

39. VALENTINE, P.S. and JOHNSON, S.J. 1997. Life history notes on the genus Catopyrops

Toxopeus (Lepidoptera: Lycaenidae) from northern Australia. Victorian Entomologist 27: 101.

40. VALENTINE, P.S. and JOHNSON, S.J. 1997. Ecological observations and notes on the life

history of Philiris diana papuana Wind & Clench (Lepidoptera: Lycaenidae). Australian

Entomologist 24: 35-36.

41. VALENTINE, P.S. and JOHNSON, S.J. 2000. Butterflies of southwestern Queensland with

new life history notes. Victorian Entomologist 30: 59-62.

42. VALENTINE, P.S. and JOHNSON, S.J. 2001. Observations of the butterflies at the

Carnarvon Range, Queensland. Victorian Entomologist 31: 2-3.

43. VALENTINE, P.S. and JOHNSON, S.J. 2001. The life history of Telicota brachydesma

Lower (Lepidoptera: Hesperiidae). Australian Entomologist 27: 103-108.

44. VALENTINE, P.S. and JOHNSON, S.J. 2003. Notes on the distribution and conservation

status of Trapezites atkinsi Williams, Williams & Hay (Lepidoptera: Hesperiidae). Australian

Entomologist 30: 87-91.

45. VALENTINE, P.S. and JOHNSON, S.J. 2004. The life history and distribution of Rachelia

extrusa (C. & R. Felder) (Lepidoptera: Hesperiidae: Trapezitinae) in Australia. Australian

Entomologist 31: 29-36.

46. VALENTINE, P.S. and JOHNSON, S.J. 2005. The rediscovery of Tagiades nestus (C.

Felder) (Lepidoptera: Hesperiidae: Pyrginae) in Australia. Australian Entomologist 32: 155-156.

47. VALENTINE, P.S. and JOHNSON, S.J. 2009. The complete life history of Charaxes latona

Butler (Lepidoptera: Nymphalidae) from Cape York Peninsula, Queensland, Australia.

Australian Entomologist 36: 63-66.

2: VETERINARY PUBLICATIONS OF STEPHEN JAMES JOHNSON

Listed alphabetically by author. These publications are not cited in the text.

BELLIS, G.A., GIBSON, DS POLKINGHORNE, I.G., JOHNSON, S.J. and FLANAGAN, M.

1993. Infection of Culicoides brevitarsis and C. wadai (Diptera: Ceratopogonidae) with 4

Australian bluetongue virus serotypes. In Arbovirus research in Australia: Proceedings Sixth

Symposium, December 7-11, 1993, Brisbane, Australia.

BELLIS, G.A., GIBSON, D.S., POLKINGHORNE, I.G., JOHNSON, S.J. and FLANAGAN, M.

1994. Infection of Culicoides brevitarsis and C. wadai (Diptera: Ceratopogonidae) with four

Australian serotypes of bluetongue virus. Journal of Medical Entomology 31(3): 382-387.

Australian Entomologist, 2015, 42 (4) 185

BUDD, T., CLINTON, P., DELL, A., DUCE, I.R., JOHNSON, S.J., QUICKE, D.L., TAYLOR,

G.W., USHERWOOD, P.N. and USOH, G. 1988. Isolation and characterisation of glutamate

receptor antagonists from venoms of orb-web spiders. Brain Research 448(1): 30-39.

DOHERTY, W.M., BISHOP, A.L., MELVILLE, L.F., JOHNSON, SJ., BELLIS, G.A. and

HUNT, N.T. 2004. Protection of cattle from Culicoides spp. in Australia by shelter and chemical

treatments. Veterinaria Italiana 40(3): 320-323.

DOHERTY, W.M., GIBSON, D.S., JOHNSON, S.J., BELLIS, G.A. and DYCE, A.L. 1993.

Culicoides survey of northern Australia, 1990-1992. In Arbovirus research in Australia:

Proceedings Sixth Symposium, December 7-11, 1993, Brisbane, Australia.

DOHERTY, W.M., JOHNSON, SJ. and REID, A.E. 2001. Suppression of Culicoides

brevitarsis (Kieffer) (Diptera: Ceratopogonidae) on cattle in Queensland with deltamethrin and

cypermethrin. General and Applied Entomology 30: 45-47.

FLANAGAN, M., DASHORST, ME WARD, M.P., MORRIS, C.M. and JOHNSON, S.J..

1993. The current bluetongue situation in Queensland. In Arbovirus research in Australia:

Proceedings Sixth Symposium, December 7-11, 1993, Brisbane, Australia.

FLANAGAN, M. and JOHNSON, S.J. 1995. The effects of vaccination of Merino ewes with an

attenuated Australian bluetongue virus serotype 23 at different stages of gestation. Australian

Veterinary Journal T2(12): 455-457.

FLANAGAN, M., JOHNSON, S.J., HOFFMANN, D., POLKINGHORNE, I.G., REID, D.J. and

SHEPHERD, M.A. 1993. Clinical pathology of Australian bluetongue virus serotype-16

infection in Merino sheep. Australian Veterinary Journal 70(3): 101-104.

JOHNSON, S.J. 1984. The effects on pigs and chickens of feeding grain contaminated with

seeds of Corchorus olitorius. In Proceedings of the Second Australia-United States Symposium

on Poisonous Plants, St. Lucia, Brisbane, 14-18th May, 1984. Brisbane, The Symposium.

JOHNSON, S.J. 1987. Stephanofilariasis - a review. Helminthological Abstracts, Series A

(Animal and Human Helminthology) 56(8): 287-299.

JOHNSON, S.J. 1991. Vector competence of Culicoides sp. for bluetongue serotypes and

pathogenicity studies in sheep (DAO31P/DAO4). Final project report to Australian Wool

Corporation, Queensland Department of Primary Industries, Townsville.

JOHNSON, S.J. 1992. Lack of anthelmintic effect of Calliandra calothyrsus in sheep. Reply.

Australian Veterinary Journal 69(3): 70.

JOHNSON, S.J. 1994. Evaluation and development of vaccination strategies for recombinant

bluetongue vaccine produced in a Baculovirus expression system (DAQ6S). Final project report

to AWRAP, Queensland Department of Primary Industries, Townsville.

JOHNSON, S.J., ARTHUR, R.J. and SHEPHERD, R.K. 1986. The distribution and prevalence

of stephanofilariasis in cattle in Queensland. Australian Veterinary Journal 63(4): 121-124.

JOHNSON, S.J., HOFFMANN, D., FLANAGAN, M., POLKINGHORNE, I.G. and BELLIS,

G.A. 1989. Recent experience with bluetongue in Queensland. In Arbovirus research in

Australia. Proceedings Fifth Symposium, August 28-September 1, 1989, Brisbane, Australia.

JOHNSON, S.J., HOFFMANN, D., FLANAGAN, M., POLKINGHORNE, I.G. and BELLIS,

G.A. 1992. Clinico-pathology of Australian bluetongue virus serotypes for sheep. In B/uetongue,

African horse sickness, and related orbiviruses: Proceedings of the Second International

Symposium. 1992.

JOHNSON, S.J., PARKER, R.J., NORTON, J.H., JAQUES, P.A. and GRIMSHAW, A.A. 1981.

Stephanofilariasis in cattle. Australian Veterinary Journal 57(9): 411-413.

186 Australian Entomologist, 2015, 42 (4)

JOHNSON, S.J., POLKINGHORNE, I.G., FLANAGAN, M. and TOWNSEND, W.L. 1992. The

Australian experience: results of a bluetongue vaccination program. In B/uetongue, African horse

sickness, and related orbiviruses: Proceedings of the Second International Symposium. 1992.

JOHNSON, S.J. and ROY, P. 1996. Bluetongue recombinant vaccines. In B/uetongue disease in

Southeast Asia and the Pacific: Proceedings of the First Southeast Asia and Pacific Regional

Bluetongue Symposium, Greenlake Hotel, Kunming, P.R. China, 22-24 August 1995.

JOHNSON, S.J. and TOLEMAN, M.A. 1982. The toxicity of jute (Corchorus olitorius) seed to

pigs. Australian Veterinary Journal 58(6): 264-265.

JOHNSON, S.J. and TOLEMAN, M.A. 1984. Apparent lack of toxicity of jute (Corchorus

olitorius) seed for poultry. Australian Veterinary Journal 61(4): 124.

JOHNSON, S.J. and TOLEMAN, M.A. 1988. Prevalence of stephanofilariasis in young Bos

indicus cattle in northern Australia. Veterinary Parasitology 29(4): 333-339.

KIRKLAND, P.D., ELLIS, T., MELVILLE, L.F. and JOHNSON, S.J. 1996. Australian National

Arbovirus Monitoring Program - a model for studying bluetongue epidemiology in China. In

Bluetongue disease in Southeast Asia and the Pacific: Proceedings of the First Southeast Asia

and Pacific Regional Bluetongue Symposium, Greenlake Hotel, Kunming, P.R. China, 22-24

August 1995.

PRITCHARD, LI DANIELS, P.W., MELVILLE, LE KIRKLAND, P.D., JOHNSON, S.J.,

LUNT, R. and EATON, B.T. 2004. Genetic diversity of bluetongue viruses in Australasia.

Veterinaria Italiana 40(4): 438-445.

PRITCHARD, L.I., DANIELS, P.W., MELVILLE, L.F., KIRKLAND, P.D., JOHNSON, SJ.

and LUNT, R.A. 2001. Molecular monitoring of exotic and reassortant bluetongue viruses in

Australia. In Arbovirus Research in Australia.

TAYLOR, L.F., BLACK, PF. PITT, DI MACKENZIE, AR. JOHNSON, SJ. and

RODWELL, B.J. 2006. A seroepidemiological study of bovine pestivirus in Queensland beef

and dairy herds conducted in 1994/95. Australian Veterinary Journal 84(5): 163-168.

WARD, M.P., CARPENTER, T.E. and JOHNSON, S.J. 1996. Spatial analysis of seroconversion

of sentinel cattle to bluetongue viruses in Queensland. Australian Veterinary Journal 74(2): 128-

131.

WARD, M.P., DOHERTY, W.M. and JOHNSON, S.J. 1997. Association between risk of

seroconversion of sentinel cattle to bluetongue viruses and Culicoides species (Diptera:

Ceratopogonidae) in Queensland, Australia. Preventive Veterinary Medicine 32(3-A): 267-274.

WARD, M.P., FLANAGAN, M., CARPENTER, T.E, HIRD, D.W., THURMOND, M.C.,

JOHNSON, S.J. and DASHORST, M.E. 1995. Infection of cattle with bluetongue viruses in

Queensland, Australia: results of a sentinel herd study, 1990-1992. Veterinary Microbiology

45(1): 35-44.

Australian Entomologist, 2015, 42 (4): 187-188 187

VARIABILITY IN THE ADULT COLOUR FORMS OF

THECLINESTHES ALBOCINCTA (WATERHOUSE, 1903)

(LEPIDOPTERA: LYCAENIDAE: POLYOMMATINAE)

FROM COASTAL SOUTH AUSTRALIA

A.M.P. STOLARSKT and C.E. MEYER?

'PO Box 423, Tailem Bend, SA 5260 (Email: roseberrynursery@bigpond.com)

?29 Silky Oak Avenue, Moggill, Old 4070 (Email: cnameyer@bigpond.com)

Abstract

The brown form of Theclinesthes albocincta (Waterhouse, 1903), previously known only from

‘inland form’ populations occurring in arid or semi-arid regions of Australia, is recorded from

coastal South Australia for the first time.

Introduction

Theclinesthes albocincta (Waterhouse, 1903) is endemic to Australia with a

very wide but disjunct distribution (Common and Waterhouse 1981, Braby

2000). Braby (2000) recognised three forms of the butterfly, viz: the ‘eastern

form’ recorded from Peak Downs near Emerald and 13 km west of Nebo

(C.G. Miller pers. comm.) in central eastern Queensland; the ‘inland form’

recorded from the arid and semi-arid areas of Western Australia, Northern

Territory, northwestern Victoria and central southwestern Queensland; and

the ‘southern coastal form’ restricted to the coastal areas of South Australia.

All forms show seasonal variation but specimens of the southern coastal form

and the eastern form can usually be separated from specimens of the inland

form by having more extensive pale greyish blue scales on the upperside in

both males and females (Braby 2000, Grund 2010). Grund (2010) noted that

the blue form is only found in coastal areas and that a blue-brown hybrid

population has been recorded from the Gawler Ranges, South Australia. To

our knowledge, the brown form has not been recorded previously from

southern coastal populations.

Results and discussion

On 21 February 2015, AMPS collected several eggs and larvae from Adriana

quadripartita (Labill.) Muell. Arg. (Euphorbiaceae) growing in sandy areas

near Meningie, South Australia. These were successfully reared by CEM in

Brisbane, with 15 adults emerging between 22-31 March 2015. Of the 15

adults that emerged, nine (1 £, 8 99) were the greyish blue form (Fig. 1)

typical of southern coastal populations, four (1 8, 3 29) were a brown form

(Figs 2-3) with no greyish blue scales near the basal regions typical of inland

populations, and the remaining two (2 9 ?) showed intermediate colouration

(Figs 4-5). Specimens of the brown form of the butterfly have also been

reared or collected by the authors from Robe and from 6 km north of

Malinong, in coastal South Australia.

188 Australian Entomologist, 2015, 42 (4)

Figs 1-5. Adult colour forms of Theclinesthes albocincta from Meningie, coastal

South Australia: (1) typical blue form female; (2) brown form male; (3) brown form

female; (4) intermediate female with minimal greyish blue basal scales; and (5)

intermediate female with more greyish blue scales.

References

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

CSIRO Publishing, Melbourne; xxvii + 976 pp.

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised Edition.

Angus and Robertson, Sydney; xiv + 682 pp.

GRUND, R. 2010. South Australian butterflies data sheet, Theclinesthes albocincta. [Accessed

24 July 2015]. <http://www.chariot.net.au/~rbg/albocincta_ds.htm>

Australian Entomologist, 2015, 42 (4): 189-196 189

POLYMORPHIC FORM PALLESCENS FEMALES OF HYPOLIMNAS

BOLINA (LINNAEUS) (LEPIDOPTERA: NYMPHALDAE) FROM

TONGA

R.B. LACHLAN

Entomology Department, Australian Museum, 6 College St, Sydney, NSW 2010

Abstract

Thirty polymorphic females of Hypolimnas bolina (Linnaeus, 1758) were collected from the

Tongan islands of Tongatapu and ‘Eua during May 1985 and April 2010. The abnormally large

proportion of pale f. pallescens specimens, particularly from ‘Eua, is discussed and 14 of these

are illustrated.

Introduction

Hypolimnas bolina pallescens (Butler, 1874) is the subspecies found on the

Pacific Ocean islands of Tokelau, Fiji, Tonga, Samoa (previously Western

Samoa) and American Samoa (Tennent 2006). The name pallescens was

originally used by Butler to describe a female Hypolimnas bolina (Linnaeus,

1758) from the Solomon Islands, as Diadema pallescens Butler. For the sake

of clarity, the present author acknowledges that all specimens collected in

Tonga presently belong to the subspecies H b. pallescens but is referring

only to the pale form of this subspecies, also called pallescens, in this paper.

The form name pallescens, along with numerous other names cited by

Poulton (1924), Clark and Sheppard (1975), Lachlan (2014) and a number of

other authors, have been used to describe some of the many different forms

of the extremely polymorphic H. bolina females found on many of the Pacific

islands and in other parts of its known range.

The present author visited the Kingdom of Tonga in May 1985 and April

2010. While conducting a survey of the butterflies of the island of 'Eua,

approximately 18 km southeast of the main island of Tongatapu, during April

2010, it quickly became apparent that there was an unusually high percentage

of very pale H. b. pallescens f. pallescens females to be found. The author

had never seen such numbers of this unique female form on any Pacific

island up to that time or more recently (Lachlan 2014).

Not a lot has been written about H. bolina from Tonga over the years. Four

forms of H. bolina were listed as occurring there, including the pale forms,

by Clark and Sheppard (1975) in their Localities Table under the name

Friendly Islands, an old name for Tonga. Miller and Miller (1993) described

H. b. pallescens as a common species and noted that ‘Tongan females are

highly variable’ and ‘display a bewildering range of forms’.

Surveys

In May 1985 the author collected several species of butterflies on the main

island of Tongatapu, including three H. b. pallescens females. In 2010 an

intensive survey of the nearby island of ‘Eua was conducted from 10 to 18

190 Australian Entomologist, 2015, 42 (4)

April. It was primarily hot and sunny each day and a variety of butterfly

species were encountered daily. Several H. b. pallescens females were seen

and collected on each day of the survey along the forest roads and trails. As

noted by Lachlan (2014), many of the very pale female pallescens appeared

to fly quite rapidly and did not seem to alight as often as the other, darker

forms. Therefore many additional specimens of this form, in particular,

evaded capture.

Hypolimnas bolina pallescens (Butler, 1874)

(Figs 1-14)

Material examined. First survey: 9 SJ, 3 99, TONGA : 3 km east of Nuku'alofa,

Tongatapu Is. 15-16.v.1985, R.B. Lachlan. (All in RBL collection).

Second survey: 13 Sd, 27 PP, TONGA : ‘Eua Island, 21?23'35.3"S, 174956742.5”W

and 21?25'43"S, 174*56'45"W, 10-18.iv.2010, R.B. Lachlan. (All in RBL collection).

The different female forms collected during both surveys and their numbers

are recorded in Table 1.

Specimen sizes. All measurements are set wingspans. The largest recorded

was 75 mm (Fig. 3); the smallest recorded was 57 mm (Fig. 14). The average

was 67.6 mm, (n — 30). The smallest specimen recorded is abnormally small

for this species. The next smallest specimen was 62 mm. Thirteen specimens

ranged between 66-68 mm. Eight specimens ranged between 70-75 mm.

Table 1. Combined results for Hypolimnas bolina pallescens females from both

surveys.

Form Numbers collected / %

euploeoides 0/0

euploeoides-naresi 0/0

naresi 9/30

euploeoides-nerina 3/10

nerina 2/6.6

euploeoides-pallescens 1/3.3

pallescens 15/50

Discussion

Clark and Sheppard (1975) used the varietal names for the four main forms

given by Poulton (1924) to group the four basic phenotypes of this extremely

polymorphic species. They are: form euploeoides, a very dark, relatively

plain form; form naresi, often male-like in appearance; form nerina, having a

bright orange-brown variable patch two-thirds of the way along the inner

border of the forewing; and form pallescens, with the base of the fore and

hindwings tending to be dark brown (variable), the rest of the wing areas

replaced with orange-brown (can be very pale and variable) and with white

patches (variable in size) visible on the fore and/or hindwings.

Australian Entomologist, 2015, 42 (4) 191

Figs 1-3. Females of Hypolimnas bolina from Tonga: (1) f. euploeoides-pallescens;

(2-3) f. pallescens.

Australian Entomologist, 2015, 42 (4)

192

Figs 4-6. Females of Hypolimnas bolina from Tonga: f. pallescens.

193

Australian Entomologist, 2015, 42 (4)

Figs 7-9. Females of Hypolimnas bolina from Tonga: f. pallescens.

194 Australian Entomologist, 2015, 42 (4)

Figs 10-12. Females of Hypolimnas bolina from Tonga: f. pallescens.

Australian Entomologist, 2015, 42 (4) 195

Figs 13-14. Females of Hypolimnas bolina from Tonga: f. pallescens.

Clark and Sheppard (1975) also added three intermediate forms: euploeoides-

naresi, euploeoides-nerina and euploeoides-pallescens but, for reasons

unknown, they left out the intermediate form euploeoides-naresi from their

very comprehensive Table | ‘Localities of the various forms of H. bolina’. In

that Table they recorded the forms naresi, nerina, euploeoides-pallescens and

pallescens from Friendly Is (Tonga).

The present author recorded three morphs of the form euploeoides-nerina

(not recorded by Clark and Sheppard 1975) and, surprisingly, fifteen morphs

of the form pallescens. This high percentage of form pallescens is significant

and very unusual given that this form is not encountered very often, even in

other Pacific countries where it has been recorded such as New Caledonia,

Vanuatu, Fiji, Samoa, Tokelau and the Cook Islands. It is rarely encountered

anywhere else. Tennent (2009) indicated an increasing number of female

196 Australian Entomologist, 2015, 42 (4)

forms in the southern islands of Vanuatu but the 25 H. bolina females he

illustrated in colour did not show any individuals as pale as those collected in

Tonga. Tennent (2006) was more specific when he stated that ‘there is a

considerable increase of [form] pallescens’ in the southern Vanuatu islands

of Tanna, Futuna and Aneityum.

It is interesting to note that Miller and Miller (1993) examined a collection of

26 female H b. pallescens taken by D. Thomas in August-September 1986,

mostly from Vava'u, north of Tongatapu, and described seventeen of the

specimens as 'similar to, or even paler than, the one figured by d'Abrera

(1977: 222) as typical of pallescens.’ This represented 65% of the entire

collection. The present author's recent survey also supports the fact that

higher numbers of form pallescens, including very pale morphs, seem to be

commonly encountered in Tonga, perhaps more than anywhere else in the

Pacific region.

Acknowledgements

I am very grateful to Russel Cox (Entomology Department, Australian

Museum, Sydney) for all his help in the production of the digital images. I

also wish to thank John Tennent (Natural History Museum, London) for

providing references and encouragement.

References

CLARK, C.[A.] and SHEPPARD, P.M. 1975. The genetics of the mimetic butterfly Hypolimnas

bolina (L.). Philosophical Transactions of the Royal Society of London, Series B 272: 229-265.

D'ABRERA, B. 1977. Butterflies of the Australian Region. 2nd edition. Lansdowne Press,

Melbourne; 415 pp.

LACHLAN, R.B. 2014. Extreme polymorphism in Hypolimnas bolina (Linnaeus) (Lepidoptera:

Nymphalidae) females from a single locality on the southwestern Coral Coast of Viti Levu, Fiji.

Australian Entomologist A1(4): 233-263.

MILLER, J.Y. and MILLER, L.D. 1993. The butterflies of the Tonga Islands and Niue, Cook

Islands, with the descriptions of two new subspecies. Bishop Museum Occasional Papers 34: 1-

24.

POULTON, E.B. 1924. Mimicry in the butterflies of Fiji considered in relation to the Euploeine

and Danaine invasions of Polynesia and to the female forms of Hypolimnas bolina L in the

Pacific (with an appendix on the numerical aspect of reciprocal mimicry (diaposematic

resemblance) by H.H. Turner). 7ransactions of the Entomological Society of London 1923: 564-

691, 23 pls.

TENNENT, W.J. 2006. A checklist of the butterflies of Melanesia, Micronesia and Polynesia

and some adjacent areas. Zootaxa 1178: 209 pp.

TENNENT, W.J. 2009. A field guide to the butterflies of Vanuatu: Ol buttaflae blong Vanuatu.

Storm Entomological Publications. Ardley, Oxfordshire; 192 pp.

Australian Entomologist, 2015, 42 (4): 197-218 197

REVIEW OF VARIATION IN ACRODIPSAS CUPREA (SANDS, 1965)

AND A. AURATA SANDS, 1997 (LEPIDOPTERA: LYCAENIDAE),

WITH DESCRIPTIONS OF A NEW SUBSPECIES OF A. CUPREA

AND A NEW SPECIES OF ACRODIPSAS SANDS FROM INLAND

SOUTHERN QUEENSLAND

D.P.A. SANDS! and M.C. SANDS?

'CSIRO Ecosciences Precinct, GPO Box 2583, Brisbane, Qld 4001

267 Haven Road, Upper Brookfield, Old 4069

Abstract

Variation in Acrodipsas cuprea (Sands, 1965) and A. aurata Sands, 1997, both part of the cuprea

species-complex, is reviewed. A new subspecies, A. cuprea variabilis subsp. n., from northern

New South Wales and southeastern Queensland, is described and compared with nominotypical

A. cuprea cuprea found from subcoastal New South Wales to Victoria. Acrodipsas violacea sp.

n., a closely related species from southern inland Queensland, is described, figured and assigned

to the myrmecophila species-group on the basis of morphology of the femora of both sexes.

Introduction

The genus Acrodipsas Sands, 1980 is a group of morphologically similar

Lycaenidae assigned to the Tribe Luciini (Edwards 1996). All species are

restricted to the Australian mainland. Little is known of their biology but

evidence suggests that the larvae are predatory on immature stages of ants

(Common and Waterhouse 1972, Samson 1989). Adult females emerge from

the pupa with almost fully mature eggs (Sands 1979), unusual for Australian

Lycaenidae. Two species-groups were recognised by Miller and Edwards

(1978) and Sands (1979): the myrmecophila species-group, in which the adult

legs are unmodified, and the i/lidgei species-group, in which the fore and

hind femora are swollen and the mid tibia is greatly reduced. Geographical

variation in Acrodipsas cuprea (Sands) a member of the myrmecophila

species-group, was noted when the species was first described (Sands 1965).

Many species are difficult to distinguish because of the rarity of the females

and because of variation in the colour of adults, in this case of A. cuprea

(Common and Waterhouse 1972, Sands 1979, 1997, Braby 2000, Eastwood

and Hughes 2003a, Orr and Kitching 2010). Similar species, including A.

cuprea and A. aurata Sands, are referred here to a ‘cuprea species-complex’.

Variation in males of A. cuprea includes distinct, obscure or absent coppery

scaling of basal areas on the upperside of the forewings, or presence or

absence of blue scales on the upperside of both wings (Braby 2000, Orr and

Kitching 2010). Braby (2010) and Field (2013) referred to these males as

copper, brown or blue ‘forms’ of A. cuprea. Variation in the extent of blue

scaling on the upperside of females of A. cuprea and A. aurata is also

considerable and in A. aurata the presence and extent of blue scaling may be

seasonal (Sands 1997). Eastwood and Hughes (2003a) studied the DNA of

mounted specimens from various populations and referred to the different

forms as morphotypes. They also examined the phylogenetic relationships of

198 Australian Entomologist, 2015, 42 (4)

Acrodipsas spp., based on the study on DNA samples from dried museum

specimens, and grouped the differing populations according to their

molecular similarity (Eastwood and Hughes 2003b).

In this paper we discuss variation in the cuprea species-complex, including A.

aurata, and revise the taxonomic identities of the previously recognised

forms of A. cuprea. We describe the variable subspecies A. cuprea variabilis

subsp. n. from northern New South Wales, coastal southeastern Queensland

and the main Dividing Range from the New South Wales border to the

Blackdown Tableland, Qld. We also describe and figure the new species

Acrodipsas violacea sp. n. from inland southern Queensland and compare it

with A. cuprea variabilis.

Abbreviations: AMS — Australian Museum, Sydney; ANIC — Australian

National Insect Collection, CSIRO, Canberra; MCS — collection of Michael

Sands, Brisbane; NMV — National Museum of Victoria; QM — Queensland

Museum, Brisbane; RM - collection of Russell Mayo, Pomona, Queensland.

Acrodipsas cuprea cuprea (Sands)

(Figs 1-8)

Pseudodipsas cuprea Sands, 1965: 69-71. Holotype £, New South Wales: Toronto, in

AMS.

Acrodipsas cuprea (Sands): Sands 1979: 251-265.

Acrodipsas cuprea (Sands) copper form: Braby 2000 (Plate 43: 6a), Field 2013,

Eastwood and Hughes 2003a.

Diagnosis. The uppersides of males of the southern subspecies, A. cuprea

cuprea, are distinguished from other Acrodipsas species, including A.

brisbanensis (Miskin) and A. cuprea variabilis subsp. n., by the well defined

subtriangular coppery base to the forewings and the position of the

postmedian bands on the underside of the forewings. The forewings are more

strongly bowed than those of A. c. variabilis. On the underside of the

forewings of both sexes, the postmedian bands are aligned or only slightly

displaced at M; and CuA» in A. c. cuprea and A. c. variabilis, a useful

characteristic for separating the southern subspecies A. c. cuprea from A.

brisbanensis, in which there is strong displacement. Above, the upperside of

both wings of females of A. c. cuprea are grey-black or black, with variable

basal and subbasal areas of dark blue usually reaching the cells but not

extending to the tornus of the hindwing. Females of A. c. cuprea are very

similar to females of A. c. variabilis but the forewings of A. c. variabilis are

usually broader and the areas of blue on the upperside are more extensive.

Variation. Males of A. c. cuprea can be distinguished from those of 4. c.

variabilis by the subtriangular coppery basal area of the forewings. This

coppery area is well defined in A. c. cuprea but may vary in brightness and,

in specimens from north of Newcastle, NSW, the area may be less well

defined than in specimens from southeastern NSW and Victoria.

Australian Entomologist, 2015, 42 (4) 199

Figs 1-8. Acrodipsas cuprea cuprea (Sands) (1, 3) Ĉĉ uppersides; (5, 7) Jd

undersides; (2, 4, 6) 9 9 uppersides; (8) 9 underside. Localities: (1) Stanwell Park,

NSW; (2) Moe, Victoria; (3, 7) Newcastle, NSW; (4, 6, 8) Port Stephens, NSW; (5)

Narara, NSW.

200 Australian Entomologist, 2015, 42 (4)

Males from between Boambee and Port Macquarie, NSW, have the basal

coppery suffusions less clearly defined, whereas in males from Stanwell Park

and Pambula, NSW, the coppery areas are clearly developed. The blue areas

on the forewings of A. c. cuprea females are somewhat variable and may be

restricted to the subbasal region of both wings or extend into the cell. The

male genitalia of A. c. cuprea differ only slightly from those of A. c.

variabilis, the valvae of the latter being slightly more slender apically and the

apical fold slightly narrower than that of A. c. cuprea, but not as narrow as in

A. violacea.

Distribution. Victoria: Western Gippsland and Warragul to the NSW-Victoria

border (Field 2013); New South Wales: southeastern coastal and subcoastal

ranges from Pambula to Boambee south of Coffs Harbour, westwards from

the Hunter Valley to Denman and the main Dividing Range south of Ebor.

Biology. Immature stages are associated with the ant genus Crematogaster

Lund and larvae are probably predatory on the immature stages of these ants

(Braby 2000, Field 2013). Eggs of A. c. cuprea (figured by Field 2013) are

similar to those of A. myrmecophila (Waterhouse & Lyell) (Sands 1979) and

are said to be carried by ants into their nests (Field 2013). At Newport, NSW,

Crematogaster sp. ants were seen tending eggs (Sands 1997) before carrying

newly-hatched larvae into a hollow branch of a mature Acacia implexa

Benth, already occupied by larvae of Hypochrysops delicia delicia

Hewitson. In Victoria, 17 specimens of A. c. cuprea (initially thought to be A.

brisbanensis) were reared from nests of Crematogaster sp. in senescing

hollow branches of an Acacia baileyana (Field 1978 and L. Gooding pers.

comm. ).

Acrodipsas cuprea variabilis subsp. n.

(Figs 9-18, 46-48)

Acrodipsas cuprea (Sands) 'brown and blue forms': Braby 2000, Field 2013,

Eastwood and Hughes 2003a.

Types. Holotype 3, labelled ‘QUEENSLAND, Mt Ngungun, 253 m, Glasshouse Mtns

Nat Pk, 26953759”S, 152°56’06E, 11 March 2005, M.C. Sands’, “Reg. No. T232286’,

in QM. Paratypes: QUEENSLAND: 1 9, Mt Tibberoowuccum, 232 m, 26953759”S,

152°56’06”E, 23 March 2003, M.C. Sands, Reg. No. T232287, in QM; 2 oo, Mt

Tibberoowuccum, 232 m, 26°55’S, 152º55"58”E, 26 January 2003, M.C. Sands, Reg.

Nos. T232167 & T232168, in QM; 1 3, Mt Tibberoowuccum, 232 m, 26°55’S,

152555'58"E, 23 February 2003, M.C. Sands; 1 9, same data except 2 March 2003, in

AMS; 1 3, 1 9, Mt Tibberoowuccum, 232 m, 26°55’S, 152°55’58”E, 15 February

2003, M.C. Sands, in NMV; 1 &, Mt Maroon, 882 m, 28°12’2”S, 152*43'52"E, 15

December 2002, M.C. Sands; 1 &, 1 9, same data except 22 December 2002; 1 9,

same data except 16 March 2003; 2 8, Bunya Mountains, 735 m, 4.8 km Nth

Burtons Well, 26947758?S, 151°34’11”E, 9 February 2006, M.C. Sands; 1 &, Coochin

Hills (west peak), 240 m, 26?47'58"S, 151?34' 11" E, 03 September 2006, M.C. Sands;

1 3, Mt Tibberoowuccum, 232 m, 26°55’S, 152°55’58”E, 26 January 2003, M.C.

Sands; 1 4, same data except 3 May 2003; 1 4, same data except 1 April 2006;

Australian Entomologist, 2015, 42 (4) 201

Figs 9-16. Acrodipsas cuprea variabilis subsp. n.: (9, 11, 13, 15) 64 uppersides; (10,

12) PP uppersides; (14, 16) 99 undersides. Localities: (9-10, 12-14, 16) Glasshouse

Mountains, Old; (11, 15) Toowoomba, Old.

202 Australian Entomologist, 2015, 42 (4)

Figs 17-18. Acrodipsas cuprea variabilis subsp. n.: Ac undersides, Glasshouse

Mountains, Qld.

1 9, same data except 20 September 2003; 1 &, Mt Tibrogargan, 264 m, 26955737”S,

152°56’60”E, 3 January 2003, M.C. Sands; 1 3, North Stradbroke Island, Tripod

Lookout, 27°26’24’S, 153°27 52 E, 20 September 2006, M.C. Sands; 1 £, Flinders

Peak, 679 m, 27948747”S, 152°45’36”E, 1 December 2002, M.C. Sands; 1 6, Rocky

Knob via Toowoomba, 580 m, 27?36'50"S, 152?02'14"E, 29 December 2004, M.C.

Sands; 1 £, same data except 28 May 2009; 1 9, same data except 10 March 2005, all

in MCS; 26 SJ, 9 99, Toowoomba, J Macqueen, dated 29-10-64 (13), 27-10-65

(13), 15 Nov 65 (13), 15-12-65 (18), 18-12-65 (14), 21-12-65 (288), 14-1-66 (18),

9-2-66 (18), 22-2-66 (29 9), 2-3-66 (14), 12-10-66 (14), 13-10-66 (19), 27-10-66

(19), 28-10-66 (28), 10 Nov 66 (13), 10-11-66 (18), 15-11-66 (13), 17-11-66

(13), 6 Feb 67 (1d), 4-3-67 (14), 31-1-67 (29 9), 20-2-67 (19), 25-2-67 (19),

10 Oct 68 (233), 11-12-68 (18), 9 March 71 (388), 21-3-71 (18, 19); 1 9,

Toowoomba, 2 Oct 66, D. Sands; 1 9, Mt Tinbeerwah, Noosa, 6 Sept. 82, R.

Eastwood, all in ANIC; 1 3, Mt Cooroora, Pomona, 10 March, 2015, R Mayo; 1 d,

Mt Cooroora, Pomona, 21 September, 2003, R Mayo; 1 £, Mt Tinbeerwah, Tewantin,

10 September, 2003, R Mayo; 1 9, Mt Tinbeerwah, Tewantin, 10 April 2003, R

Mayo; 1 9, Mt Tinbeerwah, Tewantin, 24 August 2004, R Mayo; 1 l, Blackdown

Tableland, 17 September 2006, R Mayo; all in RM. NEW SOUTH WALES: 1 d,

Grafton, 10 Nov 1944; 1 &, Slaughterhouse Pinnacle, 29°44’S, 150°18’E, 28 xii 1994,

H. (Mike) Groth, in ANIC.

Description. Male (Figs 9, 11, 13, 15, 17, 18). Antenna length (of holotype)

7.3 mm; shaft black with segmental bands white; club long, dorsally and

ventrally dark brown-black, apex tipped with orange; head dorsally black,

ventrally overlain with long white hairs scales; frons grey-black with narrow

median white line and white line narrowly above eyes; eyes grey-black; palpi

dorsally dark grey-brown, ventrally with patches of white scales, apex tipped

white; thorax and abdomen dorsally grey-black, edged with long brown

scales extending over wing bases, ventrally overlain with white hair scales;

legs brown with fore and hind femora not swollen, tarsi brown, femur and

tibia with long white hair scales, apical fore-tibial spines (modified setae)

moderately developed, thickened basally, longer than spurs on tarsi. Forewing

length (of holotype) 14.2 mm, costa and inner margin almost straight, apex

Australian Entomologist, 2015, 42 (4) 203

strongly acute, termen weakly bowed M;-CuA;, tornus obtuse; upperside

dark brown, medium grey-brown or coppery brown, veins often dark brown,

termen darker brown, submedian and postmedian areas occasionally with

indistinct coppery areas reaching inner margin, often with some basal blue

scales, cilia light brown. Hindwing costa almost straight, termen bowed.

between M; and CuA;, squared CuA; to 1A+2A at tornus, inner margin

straight; upperside grey-brown, darker towards base, often with some basal

blue scales (rarely on coastal specimens), veins and termen darker brown,

transverse apical cell bar on vein M, to Ms, dark brown; termen narrowly

dark grey-brown; an oval black spot at termen between CuA, and CuUA», two

smaller spots between CuA» and 1A+2A, a narrow blue subterminal line M3

to CuA, to tornus, extending broader CuA, to tornus separating black spots

and dark termen; cilia light brown, black and broadly tufted at CuA;, CuA»

and at tornus, inner marginal fold light grey. Forewing underside, pale

brown-grey with pale bands rarely indistinct, edged with dark brown then

white, three transverse brown bands in cell, one narrow at base, broader and

dark at mid cell, broadest between bases of Mi M» and M3; termen dark

brown with subterminal subcrescentic line closer to tornus than termen,

separated at veins R, to 1A+2A, a subparallel postmedian band R; to Ms,

closer to termen at tornus, a submedian spot between CuA, and CuA; and

band below cell beneath CuA». Hindwing underside, grey-brown with brown

bands edged darker brown then white; termen dark brown, narrow at apex,

broader at tornus, with subterminal crenulated pale brown line, parallel to

termen except closer Rs to Mu a median band between Rs and Mj, a

postmedian band M; to tornus, displaced between veins with sections M, to

M; and CuA, to CuA; closer to termen, section between 1A+2A and inner

margin closer to 3A; a prominent subterminal black spot between CuA, and

CuA»; two black spots, sometimes merged, between CuA» and 1A+2A, both

edged anteriorly and prominently with orange, spots separated from dark

termen by narrow white line CuA; to tornus.

Male genitalia (Figs 46-48). Unmounted: vinculum + tegumen subcircular,

slightly longer than wide in posterior view. Slide mounted: margins of socius

strongly bowed, sociuncus lobe broad, uncus bifurcate, branches well

separated at base; brachium long, slender, apical section ca twice as long as

basal section; valva with base broadly oval, concave near midpoint, apical

section subequal in length to basal section, apically narrower than at median

fold, apex subsquared, tip pointed, outwardly directed, fold slender, weakly

slerotised; juxta U-shaped with relatively long apical arms; aedeagus basally

broad, prezonal sheath rounded anteroventrally, zone narrow, postzonal

sheath ca two-thirds length of prezonal sheath, curved ventrally near

midpoint, slender, smooth, apically tapered to a point.

Female (Figs 10, 12, 14, 16). Antenna length (of allotype) 6.5 mm; head,

palpi, thorax, abdomen and legs similar to male, palpi with second segment

longer than male, abdomen considerably larger than male abdomen, ventral

204 Australian Entomologist, 2015, 42 (4)

surface of thorax, anterior surface of abdomen, femur and tibia with long

white hair scales. Forewing apex obtuse, termen strongly bowed, tornus

obtuse, uppersides of both wings brown-black with basal and submedian

areas of medium to dark blue; forewing basal 1/2-2/3 blue, mostly filling cell,

extending to mid inner margin or slightly beyond posteriorly; apical cell vein

at bases of M, and M- dark brown-black; cilia white. Hindwing costa bowed,

grey-brown, termen straight between M; and CuA,, produced and squared

between CuA, to CuA;, slightly concave between CuA» and 1A+2A; above

blue area from base to cell postmedian region, extending from cell to beyond

inner margin, with transverse black cell bar between bases of Mı and Mb,

occasionally a series of subterminal crenulated blue markings from M, to

1A+2A, a large subtriangular black spot between CuA; and CuA», a narrow

black spot or band between CuA» and 1A+2A, a narrow blue subterminal line

M; to 1A+2A edging black spots termen at tornus; cilia white, black and

broadly tufted at CuA;, CuA» and tornus, basal half of inner marginal fold

grey-brown, grading to dark brown towards tornus. Forewing underside, light

brown with brown bands or spots narrowly edged with dark brown, then

white; cell with transverse bands dark, narrow at base and at mid cell, a spot

between subbases of CuA; and CuA», a subbasal band between CuA, and

1A+2A, a broad postmedian band, slightly displaced at Mi, closer to termen

at tornus than apex, a narrow subterminal band, dark brown, somewhat

crenulated, slightly closer to termen at tornus than apex; area between

subterminal band and termen slightly darker brown, darker and narrower

towards termen. Hindwing underside with basal and submedian narrow bands

or dark spots, two at base of cell; mid and postmedian bands wider, sections

M; to Ms at cell apex and CuA; to CuA» closer to termen, section between

CuA, and 1A+2A angled anteriorly towards base; two black spots separated

at tornus, edged orange anteriorly, one a larger, oval spot between CuA, and

CuA», the second a smaller spot at tornus overlying 1A+2A; spots at termen

edged narrowly blue CuA; to tornus; a dark postmedian band, angled and

closer to base at inner margin; inner half of marginal fold light grey; cilia

light brown-cream, black and broadly tufted at CuA;, CuA»and tornus.

Female genitalia. Slide mounted: apophyses anteriores slender, apically

tapered, moderate in length; papillae anales broadly rounded, finely setose;

ostium bursae small, U-shaped, weakly sclerotised, surrounded by broad U-

shaped anterior wall of ostium; ductus bursae not sclerotised, membranous,

broad, long, gradually expanded to merge with anteriorly rounded,

membranous corpus bursae.

Comments. Males of A. c. variabilis are readily distinguishable from the

nominotypical subspecies by absence or obscurity of the subtriangular

coppery basal area on the upperside of the forewings. The ground colour of

males of A. c. variabilis is usually much darker than that of A. violacea, but

may vary from dark brown to coppery brown with occasional blue basal

scales on both wings, mostly on specimens from the main Dividing Range.

Australian Entomologist, 2015, 42 (4) 205

The ground colour is similar to that of the upperside of A. brisbanensis

(Miskin) but in A. c. variabilis the termen of males, although variable in

shape, is more strongly bowed than in either A. brisbanensis or A. c. cuprea.

The ground colour of females and areas of blue of A. c. variabilis are very

similar to the nominotypical subspecies, but in A. c. variabilis the termen of

the forewing in females is more strongly bowed than in either A. c. cuprea or

A. violacea.

Male and female genitalia are similar to those of nominotypical A. c. cuprea

(see Sands 1997). The apices of the valvae of males of A. c. variabilis appear

to be slightly longer than in A. c. cuprea but not as slender as in A. violacea.

Variation. The forewings of small males of A. c. variabilis are often more

strongly bowed than in larger specimens, accompanied by extraordinary

variation in wingspans (Sd 16-24 mm, 9 9 17-28 mm), usually in specimens

from the Glasshouse Mountains, where small males are unusually abundant.

The upperside of males of A. c. variabilis from Stradbroke Island 1s darker

grey-brown than in specimens from the mainland and is not known to have

areas of blue, whereas specimens from the Glasshouse Mountains and other

mainland coastal areas occasionally have areas of blue on both wings. Blue

basal scales are uncommonly seen on coastal specimens of males of A. c.

variabilis. The ground colour of males from Toowoomba is sometimes light

brown with basal coppery suffusions; this variation 1s also seen in specimens

from other parts of the main Divide and from Blackdown Tableland. The

upperside of males from the main Divide from Amiens and between

Toowoomba and Blackdown Tableland (8%, n = 61) sometimes has blue

scales, which rarely fill the cell of the forewing and do not extend to the

postmedian region of the hindwing.

In males of A. c. variabilis the apical fore-tibial spines are more strongly

developed than in A. violacea, but appear to vary according to the size of the

specimen, with longer and basally broader spines on larger specimens.

The ground colour of females of A. c. variabilis is darker than that of A.

violacea and the extent of blue on the uppersides of both wings is less

extensive than the violet-blue areas of females of A. violacea.

Distribution. New South Wales: Grafton (Ramornie Trig) and northwards on

the main Dividing Range to the NSW Border and west to Slaughterhouse

Pinnacle; Queensland: coastal hills and southern ranges, including North

Stradbroke Island, Mount Maroon, Flinders Peak, Glasshouse Mountains

(including Mts Tibberoowuccum, Tibrogargan and Ngungun), Wild Horse

Mountain, Coolum, Mt Tinbeerwah, Mt Cooroora, Pomona, Rainbow Beach,

Main Dividing Range including Amiens, Mt Ferguson, Toowoomba,

Kroombit Tops, Mt Moffatt, Carnarvon National Park (Monteith and Yeates

1986, 1988) and the Blackdown Tableland.

206 Australian Entomologist, 2015, 42 (4)

Biology. Unknown. Freshly emerged males have been observed resting on

stumps (ca 1 m) infested with an ant, Crematogaster sp. (R. Mayo, pers.

comm.). Females emerge with ovaries mature.

Acrodipsas violacea sp. n.

(Figs 19, 20-35, 43-45, 49-51)

Acrodipsas cuprea (Sands) ‘brown and blue forms’: Braby 2000, Eastwood and

Hughes 2003a.

Types. Holotype 3, labelled ‘QUEENSLAND, Commodore Peak via Millmerran, 609

m, 27956715”7S, 151?13' 17" E, 17 September 2004, M.C. Sands’, ‘Reg. No. T232282’,

in OM. Paratypes: QUEENSLAND: 1 &, Commodore Peak, via Millmerran, 609 m,

27°56 15”S, 151913'17”E, 27 September 2004, M.C. Sands, genitalia slide DPAS 77,

Reg. No. T232284; 13. same data except without slide and 26 April 2005, Reg. No.

T232283; 1 9, same data except 25 October 2004, MC & DPA Sands, genitalia slide

DPAS78, Reg. No. T232285; 2 $$, Dunmore via Dalby, SW of Cecil Plains, 9 Dec.

1984, M. De Baar, 224 M, Reg. Nos. T232169' & T232173; 2 Ĉĉ, Dunmore via

Dalby, 11 Dec. 1987, M. De Baar, Reg. Nos. T232170 & T232171; 1 &, same data

except 22 Dec. 1981, A. cuprea det D.P. Sands, Reg. No. T232172, all in QM; 1 d,

Commodore Peak, via Millmerran, 609 m, 27?56'15"S, 151?13'17"E, 17 September

2004, M.C. Sands; 1 6, 1 9, same data except 19 September 2005; 1 £, same data

except 31 October 2009; 1 3 same data except 24 September 2005; 2 $4, Dunmore

via Dalby, 9 Dec. 1984, M. De Baar; 1 9, Millmerran, 28 Mar. 1941, J. Macqueen; all

in ANIC; 14, Dunmore Forest, 402 m, 16 km S.W. Cecil Plains, 27937730”S,

151?01'09"E, 31 October 2009, M.C. Sands; 2 Ĉĉ, Commodore Peak, via

Millmerran, 609 m, 27?56'15"S, 151?13'17"E, 17 September 2004, M.C. Sands;

2 33, same data except 19 September 2005; 6 SS), same data except 24 September

2005; 6 do, same data except 27 September 2004; 1 8 same data except 28

September 2004; 1 (4 same data except 4 October 2008; 1 £, same data except 23

October 2004; 1 &, same data except 24 January 2006; 2 OO. same data except 26

April 2005; 2 3, same data except 9 October 2004, M.C. & D.P.A. Sands, all in

MCS.

Descriptions. The descriptions of adults and adult variation are based on

specimens either unaffected by grease, or after grease has been removed by

immersion of mounted specimens in ethyl acetate.

Male (Figs 20, 22, 24, 26, 28, 30, 32, 34). Antenna length (of holotype) 6.5

mm; shaft black with segmental bands white; club long, dorsally and

ventrally brown-black, apex faintly tipped with orange; head dorsally black,

ventrally overlain with long white hairs scales; frons black with narrow

median white line and white line narrowly above eyes; eyes grey-black; palpi

dorsally dark grey-brown, ventrally with patches of white scales, apex tipped

white; thorax and abdomen dorsally grey-black, edged with long pale brown

scales extending over wing bases, dorsally grey-black, ventrally overlain with

white hair scales; legs (Figs 19, 51) brown with fore and hind femora not

swollen, tarsi lighter brown, apical fore-tibial spines weakly developed, not

thickened basally, not longer than spines on tarsi, femur and tibia with long

Australian Entomologist, 2015, 42 (4) 207

Length = 122.67 um

- GAIA,

Length = 127.15 um

Fig. 19. Acrodipsas violacea sp. n.: male fore-tarsus and tibia and length of spines.

white hair scales. Forewing length (of holotype) 13.7 mm, costa and inner

margin almost straight, apex acute, termen weakly bowed M;-CuA;, tornus

obtuse; forewing upperside, basal half pale coppery brown extending to inner

margin, grading to brown at subtermen, cell with variable scattered or dense

blue-violet scales, extending below cell to base of CuA»; subcostal vein and

postcellular veins brown; termen narrowly dark grey-brown, broader at

tornus; cilia cream. Hindwing costa weakly bowed, termen straight, angled at

Mi, straight between M; and CuA,, squared CuA; to 1A+2A at tornus, inner

margin straight; upperside pale coppery brown, with variable areas of blue-

violet in cell and most broadly in postmedian region M, to CuA;, a transverse

apical cell bar dark grey, darkest between M; and Mb, veins brown; termen

narrowly brown-black, an oval black spot at termen between CuA, and CuA;,

two small spots between CuA» and 1A+2A, a narrow blue subterminal line

separating black spots and termen, CuA; to tornus; cilia cream, black and

broadly tufted at CuA;, CuA» and tornus, inner marginal fold grey-brown.

Forewing underside, pale brown-grey with pale bars or bands, edged darker

brown then white, often indistinct except subterminal dark line, three

transverse dark brown bars in cell, one narrow and darkest at base, broader

and dark at mid cell, broadest and pale between base of Mj, M> and Ms;

termen dark brown with subterminal brown line, closer to tornus than apex,

not edged with white and interrupted at veins R; to 1A+2A, a subparallel

postmedian band R; to M3, pale or indistinct towards inner margin or absent

between M3 and CuA», two submedian spots, one between CuA; and CuA;

and one below cell beneath CuA».

Australian Entomologist, 2015, 42 (4)

208

Figs 20-27. Acrodipsas violacea sp. n. uppersides: (20, 22, 24, 26) Ĉc; (21, 23, 25, 27)

99. Localities: (20-25) Commodore Mountain, Qld; (26) Dunmore via Cecil Plains,

Qld; (27) Millmerran, Qld.

Australian Entomologist, 2015, 42 (4) 209

Figs 28-35. Acrodipsas violacea sp. n. undersides: (28, 30, 32, 34) SS; (29, 31, 33, 35)

OO. Localities: (28-29, 31-34) Commodore Range, Qld; (30) Dunmore via Cecil

Plains, Qld; (35) Millmerran, Qld.

210 Australian Entomologist, 2015, 42 (4)

Hindwing pale brown with brown bands edged darker brown then white,

often indistinct or absent from basal half of wing; termen dark brown with

subterminal brown crenulated line, parallel to termen, except closer between

Rs and M;, a subparallel postmedian band costa to inner margin, sometimes

indistinct or absent, displaced at veins with sections M; to M3 and CuA, to

CuA, closer to termen, section between 1A+2A and inner margin closer to

3A; a prominent subterminal black spot between CuA; and CuA;; a black

band or two merging spots CuA» and 1A+2A, edged anteriorly and faintly

with cream-orange; black spots and subtornal black band separated from dark

termen by narrow bluish white line from CuA; to tornus.

Male genitalia (Figs 43-45, 49). Unmounted: vinculum + tegumen

subcircular, slightly longer than wide in posterior view, saccus expanded.

Slide mounted: margins of socius strongly bowed, sociuncus lobe broad,

uncus bifurcate, branches tapered, well separated at base, curved apically;

brachium long, slender, apical section ca twice as long as basal section,

apical point outwardly directed; valva with base broadly oval, concave before

midpoint, apical section longer than basal section, tapered, narrowing at

midpoint, subapical fold semicircular, weakly slerotised 2/3 distance from

base, apex squared, tip outwardly directed; juxta with arms relatively long,

apically curved inwardly, outwardly turned at base; aedeagus basally broad,

prezonal sheath ca two-thirds length of prezonal sheath, postzonal sheath

concave near midpoint (viewed laterally), curved ventrally, apically tapered

to a strongly slerotised apex.

Female (Figs 21, 23, 25, 27, 29, 31, 33, 35). Antenna length (of allotype) 6.4

mm; colour of head, palpi, thorax, abdomen and legs similar to male, palpi

with second segment longer than male, abdomen relatively large; ventral

surface of thorax, anterior surface of abdomen, femur and tibia with long

white hair scales. Forewing length (of allotype) 13.4 mm, apex obtuse,

termen bowed, tornus obtuse; upperside broadly violet-blue from base to

postmedian region and inner margin, sometimes extending between

subtermen and termen, a prominent brown-black bar at apex of cell between

M; and M»; subtermen broadly brown from costa to tonus, broader at apex

than tornus, termen narrowly dark brown, cilia cream-brown. Hindwing costa

bowed, mostly at apex, termen straight between M; and CuA;, produced and

squared between CuA; to CuA,, slightly concave between CuA» and 1A+2A;

upperside broadly violet-blue from base to postmedian region, broadest at

1A+2A, apex of cell with prominent brown-black bar between bases of M,

and M», subtermen broadly brown costa to tornus, a series of subterminal

crenulated violet-blue markings enclosing dark brown from Rs to 1A+2A at

subtornus; termen narrowly brown-black, a large subtriangular black spot

between CuA, and CuA», a black band between CuA» and 1A+2A, a narrow

blue subterminal line M; to 1A+2A edging black spots at tornus; cilia cream-

brown, black and broadly tufted at CuA;, CuA», broadest at tornus, inner

marginal fold pale grey-brown, grading to darker brown towards tornus.

Australian Entomologist, 2015, 42 (4) 211

Forewing underside, pale grey-brown with brown bands or spots narrowly

edged with dark brown, then white; cell bands darkest and merged at base,

narrow and dark at mid cell, paler at cell apex, a spot between subbases of M,

and CuA, and CuA, and CuA;, postmedian band broad, closer to termen at

tornus than apex, section between R4 and R4.5 and M; to CuA» displaced

towards base, subterminal band narrow, somewhat crenulated, slightly closer

to termen at tornus than apex, area between subterminal band and termen

brown with ground colour darker and narrower towards termen; hindwing

underside basal and submedian bands narrower than postmedian band, cell

basal, mid and apical bands progressively wider, postmedian band broad,

sections M; to M; at cell apex and CuA; to CuA;» closer to termen, section

between CuA» and 1A+2A angled anteriorly towards base, two prominent

subterminal black spots separated at tornus, edged orange anteriorly, one

larger oval spot between CuA, and CuA», a second smaller spot at tornus

overlying 14724, spots edged narrowly blue, a dark postmedian band.

Female genitalia (Fig. 50). Slide mounted: apophyses anteriores slender,

apically tapered, moderate in length; papillae anales broadly oval, finely

setose; ostium bursae small, moderately sclerotised, shallowly U-shaped with

out-turned arms, surrounded by U-shaped anterior wall of ostium, wider than

long; ductus bursae not sclerotised, membranous, broad, gradually expanded

to merge with small, anteriorly rounded, membranous corpus bursae.

Comments. The ground colour on the upperside of males of A. violacea is

coppery brown, rather than the brown or dark grey-brown usual for A. cuprea

variabilis. Violet-blue basal areas are mostly present in males of A. violacea,

except for some specimens from the Dunmore area, where the upperside is

uniformly pale coppery brown. The forewing termen of A. violacea is less

bowed than that of A. c. variabilis, somewhat resembling the termen of A.

brisbanensis. On the hindwing underside, the orange edging of the black

spots near the tornus of A. violacea is paler than the orange of A. c. variabilis

and in some specimens of A. violacea the orange edging is replaced by

cream. The underside bands in males of A. violacea may be clearly marked,

obscure or absent and the narrow subterminal brown band, interrupted by

veins, may be the only postmedian marking on the forewing. In male A.

violacea when bands are present on the underside, the white edging tends to

be more clearly marked than in the bands of A. c. variabilis. On the legs of

males, the apical fore-tibial spines are less developed when compared with A.

cuprea variabilis and, although not longer than spines on the tarsi, appear to

vary according to the size of the specimen, with more robust tibial spines

occurring on larger specimens. The male genitalia are similar to those of A.

cuprea and A. aurata (Sands 1997), but the apices of valvae of A. violacea

are narrower than those of A. c. variabilis and resemble somewhat the shapes

of valvae of A. mortoni Sands, Miller & Kerr (see Sands et al. 1997, fig. 12).

212 Australian Entomologist, 2015, 42 (4)

In A. violacea females, the violet-blue areas extend to the postmedian region

or beyond and to the inner margin of both wings, whereas in females of A.

cuprea variabilis the blue areas rarely extend beyond the cell. The ground

colour of females of A. violacea from Millmerran and Commodore Peak is

light to medium brown, with distinctive violet-blue areas extending beyond

the cells of the upperside of both wings and reaching the postmedian region

and subtornus, whereas in A. cuprea variabilis the ground colour is grey-

black and the dark blue area is restricted to the basal half of the forewings. To

date, no females of A. violacea are known from Dunmore for comparison

with the females from Millmerran and Commodore Peak, which have

extensive areas of violet-blue on both wings. Based on one specimen

available for dissection, the female genitalia (Fig. 50) could not be

distinguished from those of A. cuprea.

Variation. The upperside of males of A. violacea varies in the shade of the

brown ground colour, the basal coppery areas on the forewings and subtornus

of hindwings, the extent of blue scales and the prominence of cell and

postmedian veins. Blue areas on the upperside of both wings are present on

males of A. violacea from Commodore Peak, but are less extensive or absent

on the few males known from Dunmore. The shade of light brown on the

upperside varies and the underside of males is very variable, with median

bands on the underside often indistinct or absent and, except for the

prominent brown subterminal line, interrupted at the veins of both wings.

Orange edging to the black spots near the tornus of A. violacea may be pale

and in some specimens orange is replaced by cream. The violet-blue areas on

the upperside of females of A. violacea are variable in extent but are much

more extensive on both wings than the areas of blue of A. cuprea variabilis.

Distribution. To date, A. violacea has been recorded at Millmerran,

Commodore Range and Dunmore, via Cecil Plains, and observed at Captains

Mountain. Several males from Mount Moffatt may be this species but the

absence of females from this locality prevents confident identification or

designation of these as paratypes of A. violacea. Males of A. cuprea

variabilis have been found at some inland localities where A. violacea and A.

cuprea may prove to be sympatric when more material, particularly females,

becomes available.

Biology. The habitats of this species are Eucalyptus and Allocasuarina

woodlands on isolated inland ridges of southern Queensland, where males

congregate to intercept unmated females that visit hilltops. Males of A.

violacea are rapid in flight, resembling the behaviour of A. brisbanensis

rather than that of A. cuprea variabilis. A female of A. violacea was collected

from blossom by J. Macqueen on his property about 24 km south of

Millmerran and females were observed (M.C. Sands) flying close to a hilltop

around a dead tree (ca 7 m) supporting a colony of ants.

Australian Entomologist, 2015, 42 (4) 213

Larvae of A. violacea are likely to be predatory on the immature stages of

ants and, as in the case with other freshly captured female Acrodipsas spp.,

the expanded abdomens of freshly emerged females contain many fully

developed eggs.

Acrodipsas aurata Sands

(Figs 36-42)

Acrodipsas aurata Sands, 1997: 339-344. Holotype &, Mount Ainslie, Australian

Capital Territory, in ANIC Canberra.

Diagnosis. Sands (1979) noted distinctive features in the apices of valvae of

male genitalia of specimens thought at that time to be a form of Acrodipsas

cuprea from coastal and montane NSW. Subsequently, differences in wing

patterns of both sexes, wingspan and shape of both wings, and male genitalia,

enabled separation of A. aurata from A. cuprea, with recognition of A. aurata

as a distinct species (Sands 1997).

Variation. Some variation has been observed in males of A. aurata, mostly in

the extent of the subtriangular coppery basal area of the forewing upperside,

the prominence of dark postcellular veins and the presence of the coppery

suffusions on the hindwing upperside (Braby 2000). Females from Canberra

and the Southern Highlands of NSW vary considerably, with the ground

colour varying from dark brown to grey-black on the upperside, sometimes

with an indistinct subbasal coppery suffusion, and in the extent of basal blue

scales (Dunn and Dunn 1991, Braby 2000). Females from the Blue

Mountains, NSW, have more extensive blue areas on both wings, from the

base to the cell or beyond, whereas in females from the ACT the blue scales

are restricted to the base, obscure or sometimes absent, as in a coppery brown

female from Mt Ginini, ACT (Sands 1997). In both sexes, the taxonomically

important postmedian subcostal band, between veins Rọ and R45 on the

underside of the forewing, is occasionally indistinct. Females from ACT

exhibit seasonal variation in the extent of blue scales, with those collected in

spring thought to have less blue than those collected in late summer or

autumn (C. Meyer pers. comm. ).

Distribution. Australian Capital Territory (> ca 580 m): including Canberra

and Brindabella Range; New South Wales: Southern Highlands, including

Queanbeyan, Cooma and Blue Mountains; Victoria: Pine Mountain (Field

2013). To date, A. cuprea cuprea and A. aurata have not been collected from

the same hilltops despite their occurrence within several kilometres of each

other in the Southern Highlands of NSW (E. Edwards pers. comm. ).

Biology. An infertile female of A. aurata (originally thought to be A. cuprea),

was induced to oviposit when confined with a colony of ants of the genus

Crematogaster (Atkins 1984, Sands 1997). The larvae of A. aurata are likely

to be predatory on immature stages of ants.

214 Australian Entomologist, 2015, 42 (4)

Figs 36-42. Acrodipsas aurata Sands: (36-39, 41) uppersides, (40, 42) undersides: (36,

38, 40) 33; (37, 39, 41-42) 9 9. Locality: Canberra, ACT.

Australian Entomologist, 2015, 42 (4) 215

Figs 43-51. Acrodipsas spp: (43-45, 48, 50-51) Acrodipsas violacea sp. n., (46, 47, 49) A.

cuprea variabilis subsp. n., (43-49) 2 genitalia: (43) unmounted, lateral view; (44-51)

slide mounted: (44, 46) sociuncus, (45, 47) valvae, (48-49) aedeagus; (50) 9 genitalia;

(51) £ legs. Scale bars = 1 mm.

216 Australian Entomologist, 2015, 42 (4)

As observed in other Acrodipsas species, mounted specimens, particularly

males, are prone to become greasy. Females emerge with mature ovaries, as

evidenced by large numbers of fully developed eggs in abdomens of freshly

captured, unmated specimens. Acrodipsas aurata is associated with eucalypt

woodlands of the ACT, Victoria and the Southern Highlands and Blue

Mountains of NSW, where males congregate on hilltops and freshly emerged

females visit the sites to mate.

Discussion

Our decision to recognise two subspecies of A. cuprea is based on the two taxa

having discrete geographical distributions. Males of A. c. variabilis exist in

several colour forms but the variation is not consistently linked to distribution;

thus there is no justification for recognising other additional forms of A. cuprea

as subspecies. However, this study does not fully resolve the identities of

representatives from all populations considered to belong to subspecies A. c.

variabilis, with insufficient females known from some localities where males

have been collected, preventing a comprehensive overview of variation and

distribution of A. c. variabilis. For example, a series of males from the

Glasshouse Mountains can be divided into two groups, one with smaller

wingspans and a more convex termen when compared with variation in other

localities, but without other detectable morphological differences.

Recognition of A. violacea as a distinct species is based on the upperside

coloration of males and females, the distribution, slight differences in the male

genitalia and, in most male specimens, the relative lengths of spines on the tarsi

and tibiae. While there is a superficial resemblance between the blue males of A.

violacea and some males of A. c. variabilis, the few females of A. violacea

available for study have more extensive areas of blue-violet than the areas of

blue in females of A. c. variabilis and the ground colour of A. violacea is a

considerably lighter brown. Differences between the length of fore-tibial spines

on male forelegs of A. violacea and A. c. variabilis appear to be important, but

the spine lengths can also vary according to size of the adult specimens. More

specimens are required for detailed comparisons of slide-mounted material

before this structural difference between the legs of the two species can be better

understood. The decision to describe A. violacea as a distinct species, rather

than as a subspecies of A. cuprea, took into account similar decisions made

for separating several other closely related species of Lycaenidae. For

example, Eastwood ef al. (2008) elevated Jalmenus eubulus Miskin to

specific rank as a species distinct from J. evagoras (Donovan), based on

differences in the morphology, ecology and genetics of the two taxa. In so

doing, the distributions of J. eubulus and J. evagoras were shown to almost

overlap in some inland areas, while adults of some populations of J. eubulus

were considered to be possibly hybridising.

We have seen a few specimens of male Acrodipsas from Toowoomba,

Barakula State Forest and Cecil Plains that cannot be placed with certainty

Australian Entomologist, 2015, 42 (4) 217

with either A. violacea or A. cuprea variabilis. Other adults (in QM),

collected by the late Murdoch De Baar from the Bunya Mountains, are

clearly affiliated with the cuprea-species complex but cannot reliably be

identified to species. When more material becomes available, particularly

females, the two species A. cuprea variabilis and A. violacea may prove to be

sympatric at some localities and hybridisation between the two cannot be

ruled out.

Eastwood and Hughes (2003a, 2003b) studied the species relationships of the

genus Acrodipsas using a base pair fragment from the 3’-end of COI. They

included a single specimen from Millmerran that did not differ in sequence from

specimens from the main Divide and other inland localities, seemingly

contradicting the decision to separate the new species A. violacea from A.

cuprea. However, the haplotypes they derived for all specimens of A. cuprea

differed only by a single or at most four nucleotides, indicating that this gene

evolves too slowly to examine population structures and species-level

relationships in the cuprea species-complex. To investigate this complex further,

it would be advisable to examine multiple, more rapidly evolving genes,

including nuclear genes. Future DNA studies on Acrodipsas spp. need to be

combined with morphological studies and based on adequate numbers of both

sexes.

Seasonal variation as seen in A. aurata might also occur in other species of

Acrodipsas and, when sufficient specimens of both sexes become available,

such seasonal effects on morphology will need to be considered in future

taxonomic studies of this genus.

Acknowledgements

The authors are grateful to the Queensland Department of Environment and Heritage

Protection for issuing Scientific Purposes permits to collect specimens from protected

areas. We thank Yu Ning Su (CSIRO Canberra), for preparing several photographs,

Cecilia Melano for helping with label data from specimens in the ANIC and Mike

Groth, Russell Mayo and the late Murdoch De Baar for provided specimens for study.

Special thanks to Albert Orr for constructive comments on the manuscript and

Andreas Zwick and Anita Juen for valuable comments on the molecular biology of the

genus Acrodipsas. We also thank Federica Turco, Laura Jones and Susan Sands for

preparing or modifying images for publication, and Anthony Livesley, Ross Field,

Tim New, John Kerr, Ted Edwards, Suzanne Metcalfe and Russell Mayo for loans of

specimens, advice or many helpful comments.

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Australian Entomologist, 2015, 42 (4): 219-252 219

REVIEW OF AUSTRALIAN PHILIRIS RÓBER (LEPIDOPTERA:

LYCAENIDAE), WITH NOTES ON VARIATION AND

DESCRIPTIONS OF TWO NEW SUBSPECIES FROM CAPE YORK

PENINSULA

D.P.A. SANDS

CSIRO Ecosciences Precinct, GPO Box 2583, Brisbane, Old 4001

Abstract

The taxonomic status of the Australian species of Philiris Róber and variation in their adult

morphology are reviewed. P. diana Waterhouse & Lyell is shown to be a species distinct from P.

papuana Wind & Clench, P. /ucina Waterhouse & Lyell is shown to be a species distinct from P.

nitens (Grose-Smith) and P. innotata evinculis Wind & Clench is formally synonymised with P.

innotata (Miskin). Two new subspecies from Cape York Peninsula, P. papuana kerri subsp. n.

and P. azula johnsoni subsp. n., are described, figured and compared with the nominotypical

subspecies from mainland Papua New Guinea. Nine species or subspecies of Philiris are

recognised as occurring in Australia: P. diana, P. nitens and P. lucina, all with Lectotypes newly

designated, plus P. fulgens kurandae Waterhouse, P. papuana kerri subsp. n., P. sappheira

manskiei Ring & Olive, P. ziska titeus d' Abrera, P. azula johnsoni subsp. n. and P. innotata.

Introduction

The genus Philiris Rober is a group of Lycaenidae limited in distribution to

parts of the Australian Region, including tropical and subtropical eastern

Australia, eastern Indonesia and Papua New Guinea, but the genus is not

known to occur in the Solomon Islands. Based mainly on specimens lodged

in the Natural History Museum, London, the genus was revised by Tite

(1963) and referred to the Tribe Luciini by Tite (1963) and Eliot (1973).

Subsequently, d'Abrera (1971) illustrated representatives of these specimens

and included new Australian records of Philiris spp. by Kerr (1967); he also

described a new subspecies, P. ziska titeus d’Abrera, from Cape York

Peninsula. Edwards (1996) provided a revised synopsis of Australian

Lycaenidae and included new records of Philiris spp. by Johnson and

Johnson (1984), while Ring and Olive (1997) recorded an additional species

(P. sappheira Sands) from northern Queensland, previously known only from

mainland New Guinea. In a major work on the butterflies of Papua New

Guinea, Parsons (1998) considered New Guinea to be the major centre of

diversity of the genus Philiris, with 53 species recorded, and referred to

species also known from Australia. More recently, Muller (2002, 2014)

described new Philiris spp. from New Guinea, while Braby (2000) and Orr

and Kitching (2010) published comprehensive accounts of most of the

Australian species, including their life histories.

The male genitalia of Philiris spp. were shown by Tite (1963) to greatly

assist identification of similar species, but some identifications have

continued to be difficult because of variation in adult morphology,

particularly in females (Forbes 1977). This variation has led to unsupportable

descriptions of new taxa from New Guinea (e.g. by Wind and Clench 1947,

Tite 1963) and difficulties in associating females with conspecific males.

Australian Entomologist, 2015, 42 (4)

Figs 1-8. Philiris diana Waterhouse & Lyell: (1-2) 88, (3-8) 99; (1, 3, 5, 7)

uppersides, (2, 4, 6, 8) undersides. Localities: (1-4) Kuranda, Qld; (5-6) Paluma, Qld;

(7-8) Lake Eacham, Qld.

Australian Entomologist, 2015, 42 (4) 221

Here, two new Australian subspecies of Philiris, P. papuana kerri subsp. n.

and P. azula johnsoni subsp. n., are described and changes to the taxonomic

status of P. diana Waterhouse & Lyell, P. nitens (Grose-Smith) and P. lucina

Waterhouse & Lyell are noted, together with lectotype designations for these

three species in order to ensure stability of their names. A lectotype is also

recognised for P. fulgens kurandae Waterhouse. Variation in morphology and

summaries of their biology, when known, are reviewed for each species.

Abbreviations: AM — Australian Museum, Sydney; ANIC — Australian

National Insect Collection, CSIRO, Canberra; BMNH — The Natural History

Museum, London; MCZ - Museum of Comparative Zoology, Harvard

University, Cambridge, USA; QM — Queensland Museum, Brisbane; PNG —

Papua New Guinea; Qld — Queensland; FWL — forewing length.

PHILIRIS RÓBER, 1891

Philiris Róber, 1891: 317; Druce 1897: 14; Druce 1902: 115; Waterhouse 1903a: 650;

Waterhouse 1903b: 188-189; Waterhouse 1913: 699; Waterhouse and Lyell 1914:

76; Waterhouse 1924: xxvii; Waterhouse 1932: 137; Waterhouse 1937: 115; Tite

1963: 222-225; Hemming 1967: 358; Tite 1969: 58; d'Abrera 1971: 368;

McCubbin 1971: 72; Common and Waterhouse 1972: 424; Eliot 1973: 429; Sands

1979: 127; Sands 1981 b: 89; Parsons 1998: 361-362; Edwards 1996: 250.

Parachrysops Bethune-Baker, 1904: 369; Hemming 1967: 338; d'Abrera 1971: 349;

Eliot 1973: 429; Parsons 1998: 361; = Philiris: syn. Müller 2014: 34.

Type species Thecla ilias C. Felder, 1860: 454. Subfamily Theclinae, Tribe Luciini,

Hypochrysops section (Eliot 1973).

Philiris diana Waterhouse & Lyell, 1914

(Figs 1-8, 53)

Philiris diana Waterhouse & Lyell, 1914: 76; Waterhouse 1932: 138; Edwards 1996:

250.

Philiris diana diana Waterhouse & Lyell: Wind and Clench 1947: 6; Tite 1963: 235;

Common 1964: 124; d'Abrera 1971: 370; McCubbin 1971: 172; Common and

Waterhouse 1972: 425; Braby 2000 and 2010; Orr and Kitching 2010: 236.

Types. Lectotype É (here designated), QUEENSLAND: labelled ‘Kuranda Qld, Mch

07 F.P. Dodd’, “G. Lyell collection’, “Type male PHILIRIS DIANA Waterhouse and

Lyell KL21453', in AM. Paralectotypes (here designated): 7 $$, 7 99, with same

locality data as lectotype but with dates and registration numbers as follows: 4 33,

April 1907, KL21456; 2 GG, April 1907, KL21459; 1 &, March 1909, KL21458; 1

9. April 1907, KL21455; 2 99, April 1907, KL21456; 2 99, April 1908, KL21457;

1 9, April 1907, KL21459; 1 9, 16 April 1902, KL21454, with R.E. Turner instead of

F.P. Dodd, all in AM. The Lectotype bears a red label and was listed as the holotype,

in the handwriting of G.A. Waterhouse, in a register of specimens held in AM.

Diagnosis. Both sexes of P. diana were adequately described and illustrated

by Waterhouse and Lyell (1914). The brownish purple upperside of males of

P. diana is distinctively lighter and duller than the purple ground colour of P.

papuana kerri subsp. n., previously considered to be a subspecies of P. diana,

222 Australian Entomologist, 2015, 42 (4)

and the white patch on the forewing of males of P. diana, referred to by

Waterhouse (1932), is more prominent than the obscure white scales

similarly placed on P. papuana kerri. Females of P. diana are very variable

in the extent of white areas on the upperside of both wings but the greyish

violet suffusion on both wings is much more restricted than in P. papuana.

The pale greyish violet suffusion on the upperside of the forewings may be

obscure or absent in P. diana.

The male genitalia of P. diana were figured by Tite (1963). The valvae are

similar to those of P. papuana kerri but the curved apical projection on the

right valva (posterior view) is shorter (Fig. 53) than that of P. papuana (Fig.

54) and the short projection on the right valva of P. diana is broad basally,

rather than the subtubular projection of P. papuana.

Sands (1981a) proposed the diana species-group for Philiris species with

relatively large adults in which both sexes are without a black spot on the

inner margin of the hindwing underside and valvae of the male genitalia are

asymmetrical. The diana species-group was subsequently accepted by

Parsons (1998).

Variation. FWL: $4, 15.3-16.4 mm; 99, 16.7-18.6 mm. In males, the white

patches on the dull purple forewing and the white hindwing costa are variable

(Waterhouse 1932). The forewing median patch between the bases of

M3, CuA; and CuA» occasionally extends between CuA» and 1A+2A

(Waterhouse and Lyell 1914, plate 15) and, on the hindwing, the white area

on the costa may extend from Rs towards M;. Females of P. diana vary in the

extent of blue basal scales and white on the upperside of both wings; the

central area of white on the forewing may be confined to the median and

postmedian region or extend to the subterminal region or inner margin, while

on the hindwing the white area may be limited to the costa or may extend to

postmedian and subterminal regions between M; and Ms.

Distribution. Northern Queensland: Lake Eacham (Atherton Tableland),

Kuranda, Cairns and Paluma.

Biology. A pupa was found near Kuranda on the upperside of a leaf of Litsea

Sp., possibly L. leefeana (F. Muell.) Merr. (Lauraceae) (Braby 2000), and this

is a probable larval food plant for P. diana.

Philiris papuana papuana Wind & Clench, 1947, stat. n.

(Figs 9-14)

Philiris diana papuanus Wind & Clench, 1947: 6; Tite 1963: 235; Kerr 1967: 49-51;

d'Abrera 1971: 370; McCubbin 1971: 72; Common amd Waterhouse 1972: 425;

Sands 1979: 131; Parsons 1998: 362; Edwards 1996: 250; Orr and Kitching 2010:

236.

Philiris diana: Parsons 1991: 136-137; misidentification.

Type. Holotype 8, PAPUA NEW GUINEA: Wau, Morobe District, in MCZ.

Australian Entomologist, 2015, 42 (4) 223

Figs 9-14. Philiris papuana papuana Wind & Clench: (9-10) SS, (11-14) 99; (9, 11,

13) uppersides, (10, 12, 14) undersides. Localities: (9-10) Musgrave River, Central

Province, PNG; (11-14) Madang, Madang Province, PNG.

Diagnosis. The description by Wind and Clench (1947) was based on a

comparison of the type specimen from Papua New Guinea with figures of

male P. diana in Waterhouse and Lyell (1914) and Waterhouse (1932).

Parsons (1998) figured the adults of both sexes.

The male genitalia of P. p. papuana were figured by Tite (1963) and Parsons

(1998); the asymmetrical valvae were noted by Tite (1963) and figured by

Sands (1979). Tite (1963) compared the male genitalia of P. p. papuana with

224 Australian Entomologist, 2015, 42 (4)

those of P. diana, describing the former as ‘larger and stouter' than those of

P. diana.

Sands (1981a) and Parsons (1998) included P. papuana in the diana species-

group proposed by Sands (198 1a).

Variation. FWL: ĉĉ. ca 16.0-183 mm; 99, 16.0-19.0 mm. Males of P.

papuana papuana show some variation in the area of obscure white scales on

the forewing, whereas females vary in the extent of white on the costa and the

distribution of blue scales on the hindwing. In localities in southern Papua

New Guinea, females from higher altitudes have more extensive areas of blue

on the hindwing than in specimens from near the coast. However, these blue

areas in P. p. papuana are not as extensive as those in P. p. kerri.

Distribution. Mainland Papua New Guinea and Papua Province, eastern

Indonesia.

Biology. In Papua New Guinea, Parsons (1998) located larvae of P. papuana

papuana feeding on Litsea guppyi (F. Muell.) F. Muell. ex Forman

(Lauraceae).

Philiris papuana kerri subsp. n.

(Figs 15-20, 54)

Philiris diana papuana: Kerr 1967: 49-51; d'Abrera 1971: 370; McCubbin 1971: 2;

Common and Waterhouse 1972: 425; Sands 1979: 131, Braby 2010: 34-35, Orr

and Kitching 2010: 236; misidentifications.

Types. Holotype 3, QUEENSLAND: labelled "Claudie River Nth. Qld. 26.xii.71,

D.P. Sands’, ‘Figured in Butterflies of Australia, CSIRO Publishing, M.F. Braby’, in

ANIC. Paratypes: | &, Claudie River, | May 1966, J.F.R. Kerr; 1 9, same data with

additional label *Metallotype Philiris diana papuana’, | &, West Claudie River e.p.,

30 August 1999, D.P.A. Sands; 1 4, Lockerbie Scrub, Cape York, 9 Oct 76, D. Sands,

ANIC database No. 32, 043995, Barcode of Life DNA voucher specimen: 11ANIC-

05827 BOLD Proc. ID ANIC1827-11; 1 8, near Mt Lamond, Iron Range, 11 April

1971, AA, all in ANIC; 1 9, ABRS Area 2, 142°45’E, 11?40'S, Dividing Range, 15

km. W. Captain Billy Creek, Cape York Pen., 5-12.11.1976, G.B. Monteith, in OM; 1

&, West Claudie R, Iron Range, XP, 27.ix.2000, R Mayo; 1 £, West Claudie R, CYP,

XP, 4.ix.1999, R Mayo; 1 ĝ, Phillip Hill, Iron Range, XP, 2.x.2003, R Mayo; 1 d,

West Claudie R, XP, 14.1x.2001, R Mayo; 1 9, West Claudie R, Iron Range, XP,

1.x.2000, R Mayo; 1 9, West Claudie R, Iron Range, XP, 2 x 2000, R Mayo; 1 9,

West Claudie R, Iron Range, XP, 31.vii1.2000, R Mayo; 1 9, Gordon Ck, Iron Range,

22.vii1.2001, R Mayo, all in Russell Mayo Collection, Pomona, Old.

Kerr (1967) described a female specimen as a metallotype of P. papuana; the same

specimen is here designated a paratype of P. p. kerri.

Description. Male. Antennal length (of holotype) 9.0 mm, shaft black with

narrow white segmental bands, club dorsally black, apex dorsally edged and

ventrally orange; eyes dark grey, edged white; frons white; thorax and

abdomen dorsally dark grey, ventrally white; palpus white, apical segment

Australian Entomologist, 2015, 42 (4) 225

grey; tibiae and tarsi white with narrow black bands. Forewing length (of

holotype) 15.0 mm, termen almost straight; upperside mauve-purple, an

obscure postmedian whitish suffusion from cell vein and between M; and

CuA, to postmedian area; termen narrowly black (ca 0.2 mm) from apex to

tornus; cilia from apex to CuA» grey and from CuA» to tornus narrowly

white. Hindwing upperside mauve-purple, costa from base, Sc+R;, Rs to

apex, grey; termen narrowly (ca 0.2 mm) black, wider from CuA; to tornus

(ca 0.4 mm); inner margin fold grey-white; cilia narrowly white. Underside

of both wings silvery white, forewing with obscure greyish median patch

between CuA; and CuA»; termen white, narrowly tipped black at vein ends

M; to tornus, hindwing broadly black at vein ends CuA;, CuA» and 1A+2A;

inner margin without black spot; cilia white, except black at vein ends CuA ,

CuA» and 1A+2A.

Male genitalia (Fig. 54). Unmounted: vinculum-tegumen ring narrowly oval

in posterior view. Slide mounted: sociuncus and vinculum broad, saccus

expanded, socii stout, separated by a shallow U-shaped sinus, margins

straight, obtusely angled at 2/3 edge, clothed with fine setae; brachia stout,

inwardly curved, apically broad, tip inwardly tapered to a point; valvae

relatively long, curved, unequal in length (posterior view: left valva longer

than right), both basally broad, cone-shaped, subtriangular, with mid section

long, slender, apices expanded and shapes asymmetrical with fine setae, apex

of left valva (posterior view) with outwardly curved, tapered, sharply pointed

beak-shaped projection, apex of right valva (posterior view) blunt, with

single peg-shaped projection; juxta subcylindrical, with slerotised ring

surrounding aedeagus; aedeagus subcylindrical, prezonal sheath broadened,

retracted cornuti with two slender strips of finely slerotised spines; anal tube

slender, moderately slerotised.

Female. Antennal length (of paratype) 9.0 mm, shaft black with narrow white

segmental bands, club dorsally black, apically tipped orange, ventrally

orange; eyes dark grey, edged white; frons grey-brown; palpus white, apical

segment grey; thorax and abdomen dorsally dark grey, ventrally white; tibiae

and tarsi white with black bands. Forewing length (of paratype) 16.7 mm,

termen bowed, apex obtuse; upperside costa, apex, termen and tornus broadly

dark grey; area from base to cell, to inner margin and postmedian area, pale

blue-mauve; a subcentral white oval patch between bases of M3, CuA; and

CuA», extending to subtermen. Hindwing upperside with apical half of costa

white with greyish suffusion extending to Rs and M;; a broad area from base

and cell to subtermen and subtornus pale blue-violet, crossed by dark grey

veins except at the apical cell bar; termen dark grey-brown (ca 5.0 mm); cilia

white. Underside of both wings silvery white, forewing with small obscure

greyish patch in median region; termen white, narrowly black between CuA ;,

CuA, and 1A+2A and black at vein ends M; to tornus; inner margin white

(without black spot); cilia white, except black at vein ends CuA;, CuA» and

1A+2A.

226 Australian Entomologist, 2015, 42 (4)

Figs 15-20. Philiris papuana kerri subsp. n.: (15-16) SÉ, (17-20) 99; (15, 17, 19)

uppersides, (16, 18, 20) undersides. Localities: (15-16) West Claudie River, Old; (17-

20) Claudie River, Old.

Variation. FWL: ĉĉ. 14.7-16.8 mm; 99, 16.7-19.7 mm. Males are variable

in the density of white scales on the forewing but the upperside of both wings

is otherwise similar to P. p. papuana. Females of P. p. kerri show little

variation in colour of the upperside (Figs 17, 19) but occasionally the blue

areas extend to the termen at M, of the hindwing.

Comments. Recognition of P. papuana kerri as a separate subspecies 1s based

on morphological differences in both sexes. The upperside of males of both

Australian Entomologist, 2015, 42 (4) 227

subspecies is similar, but the wingspans of P. p. kerri are usually slightly

smaller (FWL ca 15.00-17.20 mm) than those of P. p. papuana (FWL ca

16.00-18.30 mm). The variable and often obscure whitish suffusion on the

upperside of the forewing of males is usually more prominent on P. p. kerri

than on P. p. papuana, where it is sometimes absent, and the hindwing costa

on the upperside of P. p. kerri is paler than in P. p. papuana. The grey

subbasal suffusion on the forewing underside of males of the two subspecies

differs, with the median patch between CuA; and CuA» of P. p. papuana

more extensive than that of P. p. kerri and reaching the postmedian region.

The hindwing underside termen of P. p. kerri is mostly white with only black

cilia at the margin, whereas in P. p. papuana the termen is narrowly black.

The male genitalia of P. p. kerri do not differ significantly from those of P. p.

papuana, figured by Tite (1963).

Differences between females of P. papuana from Australia and Papua New

Guinea were referred to by Sands (1979), who noted that the blue areas on

both wings [of P. p. kerri] were more extensive (Figs 17, 19) when compared

with females of P. p. papuana (Figs 11, 13) from New Guinea. The

wingspans of females of P. p. kerri are similar (FWL 17.0-19.5 mm) to those

of P. p. papuana. The extent of blue on the upperside of both wings differs:

in P. p. kerri (figured by Kerr 1967) blue areas extend to the postmedian and

inner margin of the forewing and the subterminal area of the hindwing, but in

P. p. papuana blue areas are limited to the basal half of the forewing and

postcellular region of the hindwing.

Etymology. Named in honour of Emeritus Professor J.F.R. Kerr, in

recognition of his many contributions to the study of Australian butterflies.

Distribution. Cape York Peninsula, northern Queensland: Lockerbie Scrub,

Captain Billy Creek, Claudie River and Iron Range.

Biology. The life history of P. papuana kerri was described by Wood (1984).

On Cape York Peninsula, the larvae fed on Litsea breviumbellata C.K. Allen

(Lauraceae) and are not attended by ants (Valentine and Johnson 1997).

Males of P. p. kerri frequently settle on the upperside of leaves up to eight

metres above ground level, on trees growing at the edge of rainforest or in

clearings near streams.

Philiris fulgens fulgens (Grosse-Smith & Kirby, 1897)

Holochila fulgens Grose-Smith & Kirby, 1897: 8.

Candalides fulgens (Grose-Smith & Kirby): Grunberg in Seitz 1921: 854.

Philiris fulgens fulgens (Grose-Smith & Kirby): Wind and Clench 1947: 8; Tite 1963:

242, d'Abrera 1971: 372; Parsons 1998: 365.

Type. Lectotype 2. INDONESIA: Amboina, Maluku, in BMNH. A male was

described and illustrated by Grose-Smith and Kirby (1897) and the male referred to by

Tite (1963) from Amboina (Ambon), bearing a Hewitson label, is probably one of the

syntypes. This specimen was designated the Lectotype by Parsons (1998).

228 Australian Entomologist, 2015, 42 (4)

Male genitalia. Figured by Tite (1963) and Parsons (1991, 1998).

Distribution. Indonesia: Ambon. Other subspecies have been described from

Aru, Biak, mainland New Guinea and Papua New Guinea islands.

Philiris fulgens kurandae Waterhouse, 1903

(Figs 21-26, 55)

Philiris kurandae Waterhouse, 1903a: 651; Waterhouse 1903b: 189-190; Waterhouse

and Lyell 1914: 76; Waterhouse 1932: 138; Barrett and Burns 1951: 147.

Philiris fulgens kurandae Waterhouse: Tite 1963: 242; Common 1964: 122; d' Abrera

1971: 372; Common and Waterhouse 1972: 426-427.

Philiris fulgens (Grose-Smith & Kirby): Edwards 1996: 250; Orr and Kitching 2010:

236.

Types. Lectotype 3 (here designated), QUEENSLAND: labelled ‘Kuranda N.Q. Apr.

1902, 24^ RE. Turner’, ‘Philiris kurandae. G.A. Waterhouse, Type ĉ KL21421’,

“G.A. Waterhouse collection’, in AM. Paralectotypes (here designated): 4 88, 5 99,

with same locality and collector as lectotype but with dates and registration numbers

as follows: 4 S&, 1 9, Dec. 1901-Jan. 1902, KL21426; 1 9, Dec. 1901, KL21422;

3 99, Mch-April 1902, KL21427, all in AM.

When referring to Philiris kurandae in a register of specimens at AM, G.A.

Waterhouse noted: ‘Types É? Kuranda in Australian Museum’. The male referred to

in the Register as the ‘HOLOTYPE MALE is here designated as the Lectotype.

Diagnosis. FWL: ĉĉ. 14.2-15.1 mm; 99, 14.0-16.3 mm. In all geographical

populations of P. fulgens the extent of variation in both sexes is considerable.

It is difficult to distinguish the nominotypical P. f. fulgens from Amboina,

Indonesia, P. f. bicolorata Wind & Clench from mainland New Guinea and

P. f. kurandae from Queensland. Further morphological studies are needed to

validate the subspecific names for P. fulgens.

The characteristic ‘two-tone’ colour of the upperside of males of P. fulgens —

dull purple forewings and sky blue hindwings — is unusual for Lycaenidae

from the Australian Region (Waterhouse 1932) and elsewhere, but this

pattern also occurs in males of Hypochrysops thesaurus (Grose-Smith) from

mainland New Guinea.

Sands (1981a) proposed a fulgens species-group for P. fulgens and related

species, accepted by Parsons (1998). The species and subspecies included in

this species-group require taxonomic reassessment and consideration of the

related species from Papua New Guinea referred to by Parsons (1998).

Variation. FWL: 88, 13.3-15.0 mm; 99, 15.0-16.7 mm. In males, slight

variation occurs in the width of the dull black apex of the forewing of P.

fulgens kurandae and the black termen edging the blue hindwing may also

vary in width. Females are most variable in the extent of dull blue on the

upperside, with these areas often restricted to the base of the wings, reaching

the cell or, occasionally, extending to most of the wing and the subterminal

regions of both wings.

Australian Entomologist, 2015, 42 (4) 229

Figs 21-26. Philiris fulgens kurandae Waterhouse: (21-22) SS, (23-26) 99; (21, 23-

25) uppersides, (22, 26) undersides. Localities: (21-22) Palmerston, Qld; (23) Alice

River, Old; (24) Palmerston, Qld; (25-26) Kuranda, Qld.

Distribution. Northern Queensland: Lockerbie Scrub, Claudie River, Alice

River, Rocky River, Cooktown, Cape Tribulation, Cairns, Palmerston,

Ingham, Innisfail, Mission Beach, Paluma, Bluewater State Forest.

Biology. Eggs are deposited on foliage of the food plants (Wood 1984) and

larvae skeletonise leaves when feeding on the underside. Pupae are attached

between the bases of leaf veins of the food plants, which include Cryptocarya

mackinnoniana f. Muell., C. murrayi F. Muell., Litsea breviumbellata C.K.

Allen, L. leefeana (F. Muell.) Merr. and Endiandra hypotephra F. Muell.

(Lauraceae).

230 Australian Entomologist, 2015, 42 (4)

Philiris ziska ziska (Grose-Smith, 1898)

(Figs 27-30)

Holochila ziska Grose-Smith, 1898: 11.

Candalides pratti Bethune-Baker, 1908: 122; syn. Tite 1963: 243.

Philiris ziska (Grose-Smith): Tite 1963: 243-244; Kerr 1967: 49.

Philiris ziska ziska (Grose-Smith): d'Abrera 1971: 373; Parsons 1998: 371.

Types. Holochila ziska: Lectotype 3, designated by Parsons (1998), INDONESIA:

labelled “Type”, “Kapaur”, ‘Type HT, Ziska Gr.Sm. Kapaur. Types A2". ‘Gen. 1961-

245 G.E.T.’, with genitalia slide, in BMNH.

Male genitalia. Figured by Tite (1963) and Parsons (1998).

Variation. Males of P. z. ziska (FWL ca 14 mm) are larger than males of P. z.

titeus (FWL ca 12 mm) and the hindwing costa on the upperside, whilst

usually white (noted by Grose-Smith 1898), is very variable and sometimes

light brown, rarely as prominently white as the costa of male P. z. titeus.

Females of P. z. ziska are also larger than those of P. z. titeus and the

upperside of both wings has much smaller areas of white than those of P. z.

titeus. Grose-Smith's (1898) description of Holochila ziska refers to a male

but the female he described as P. ziska is a female of another species (Tite

1963).

Distribution. Mainland Papua New Guinea; Papua Province and

neighbouring islands of eastern Indonesia.

Philiris ziska titeus d’ Abrera, 1971

(Figs 31-34, 56)

Philiris ziska (Grose-Smith): Kerr 1967: 49; McCubbin 1971: 72; Common and

Waterhouse 1972: 427; Orr and Kitching 2010: 237.

Philiris ziska titeus d' Abrera, 1971: 373; Edwards 1996: 250; Braby 2009: 119-121.

Types. Lectotype 3, designated by Braby (2009), QUEENSLAND: labelled (as per

Braby 2009) *CLAUDIE R. CAPE YORK, 1 MAY 1966’, ‘Specimen photographed

by B. D'Abrera, 1970’, “B.M. Reg. No. 1966-587’, “253”, ‘Holotype’, “Philiris ziska

titeus, D'Abrera, det. RI. Vane-Wright, HOLOTYPE A" Paralectotypes: l d,

labelled “CLAUDIE R. CAPE YORK, 2 MAY 1966’, ‘Specimen photographed by B.

D'Abrera, 1970’, “B.M. Reg. No. 1966-587’. ‘255’, ‘Paratype, “Philiris ziska titeus,

D'Abrera, det. RI. Vane-Wright, PARATYPE d"; 1 &, labelled ‘CLAUDIE R.

CAPE YORK, 2 MAY 1966’, ‘Specimen photographed by B. D'Abrera, 1970’, *B.M.

Reg. No. 1966-587’, “255”, ‘Paratype’, “Philiris ziska titeus, D' Abrera, det. R.I. Vane-

Wright, PARATYPE A": 1 9, labelled ‘CLAUDIE R. CAPE YORK, 3 MAY 1966’,

‘Specimen photographed by B. D'Abrera, 1970’, “B.M. Reg. No. 1966-587’, “254”,

‘Paratype’, “Philiris ziska titeus, D’ Abrera, det. R.I. Vane-Wright, PARATYPE 9’;

1 9, labelled “CLAUDIE R. CAPE YORK, 26 APR. 1966’, “B.M. Reg. No. 1966-

587° (types collected by J.F.R. Kerr), all in BMNH.

D'Abrera (1971) did not contrast P. z. titeus with the nominotypical P. z. ziska when

describing the Australian subspecies.

Australian Entomologist, 2015, 42 (4)

Figs 27-34. Philiris ziska subspecies: (27-30) P. z. ziska (Grose-Smith), (31-34) P. z.

titeus d' Abrera; (31, 32, 35, 36) SS, (33, 34, 37, 38) 9 9; (31, 33, 35, 37) uppersides,

(32, 34, 36, 38) undersides. Localities: (27-28) Lae, Morobe Probvince, PNG; (29-30)

Sogeri, Central Province, PNG; (31-32) Claudie River, Qld; (33-34) Rocky River,

Qld.

232 Australian Entomologist, 2015, 42 (4)

Diagnosis. Males of P. z. titeus (FWL ca 12 mm) from Queensland are

smaller than those of P. z. ziska (FWL ca 14 mm) from Papua New Guinea,

and the upperside of the hindwing costa of P. z. titeus is more broadly white

than in P. z. ziska. The shining purple-blue ground colour of P. z. ziska and P.

z. titeus and width of the black apex are similar. Median white scales on the

forewing upperside of males of P. z. titeus may be absent, obscure or

occasionally prominent (the late Murdoch De Baar pers. comm.) and, when

present, more noticeable than in P. z. ziska. The male genitalia of P. z. titeus

(Fig. 56) are similar to those of P. z. ziska but the characteristic projection

from the valva is more rounded in P. z. titeus, the median sociuncus lobes are

more produced and the aedeagus is longer and narrower. When compared

with P. azula johnsoni, the purple-blue ground colour of male P. z. titeus is

brighter than the dull violet-blue of P. azula johnsoni and the black forewing

apex of P. z. titeus is narrower. The hindwing costa of P. z. titeus 1s white,

whereas that of P. azula johnsoni is light brown.

Females of P. z. titeus (FWL ca 13 mm ) are smaller than P. z. ziska (FWL ca

15 mm) and the areas of white are more extensive, often merging with

greenish blue scales on the forewing, whereas on the hindwing of P. z. ziska

the white area rarely extends beyond Rs. Females of P. z. titeus are very

similar to those of P. /ucina but may be distinguished by the position of the

white area on the hindwing, that of P. z. titeus extending from the base to the

postmedian region of M3. Moreover, in P. z. titeus the white area near the

base and from the costa to the cell often extends beyond the postmedian

region, whereas in P. /ucina the white area (although variable) is absent from

the grey-black subbasal area of the wing and extends from beyond the cell to

the postmedian region,

Sands (1981a) proposed a ziska species-group for P. ziska and related

species. This proposal was subsequently accepted by Parsons (1998).

Variation. FWL: Ĉe, 12.2-14.0 mm; 99, 12.3-13.7 mm. The median areas

of white scales on the forewing upperside of males of P. z. titeus may be

absent (resembling P. z. ziska), obscure or, occasionally, prominent (Samson

and Johnson 2009) and located post-cell between the bases of Mı, M»; and

CuA, and rarely reaching the subterminal region. The white costa of the

hindwing may also vary in width, sometimes extending from the wing base to

the base of M; at the apex of the cell.

Distribution. Cape York Peninsula, northern Queensland: Iron Range,

Claudie River and Rocky River.

Biology. The life history of P. z. titeus, described by Samson and Johnson

(2009), is very similar to that of nominotypical P. z. ziska from Papua New

Guinea (Parsons 1984). Larvae feed on the vine Trophis scandens (Lour.)

Hook. & Arn. (Samson and Johnson 2009); they are not attended by ants.

Pupation takes place under the base of leaves of the food plant.

Australian Entomologist, 2015, 42 (4) 233

Philiris azula azula Wind & Clench, 1947

(Figs 35-38)

Philiris azula Wind & Clench, 1947: 8-9; Tite 1963: 241; Parsons 1998: 371;

Edwards 1996: 250.

Type. Holotype É, PAPUA NEW GUINEA: Wau, Morobe District, in MCZ.

Diagnosis. The pale, greyish blue colour of males of P. azula azula is

distinctive and not seen in other Philiris spp. of similar wingspan from Papua

New Guinea. The females of P. a. azula are similar to several other female

Philiris spp. from Papua New Guinea, but the upperside is much darker than

the single known female of P. a. johnsoni from Australia. The hindwing costa

of females of P. z. ziska is usually prominently white, when compared with

the greyish white costa of females of P. azula azula.

Sands (1981a) proposed that P. azula be included in a ziska species-group

together with other related species. This was subsequently accepted by

Parsons (1998).

Figs 35-38. Philiris azula azula Wind & Clench: (35-36) ĉĉ. (37-38) 99; (35, 37)

uppersides, (36, 38) undersides. Localities: (35-36) Lae, Morobe Province, PNG; (37-

38) Wau, Morobe Province, PNG.

Variation. FWL: 33, 15.0-15.5 mm; 99, 14.5-16.0 mm. In males of P. a.

azula the width of the black apex on the upperside may vary slightly.

234 Australian Entomologist, 2015, 42 (4)

Females also vary slightly and may be similar to those of several other

Philiris spp. from Papua New Guinea, including P. ziska ziska.

Distribution. Mainland Papua New Guinea and Papua Province, eastern

Indonesia.

Biology. The life history of P. a. azula is not recorded. Adult females were

observed near Lae, Morobe Province, Papua New Guinea, resting on foliage

of Streblus brunonianus (Endl.) F. Muell. (Moraceae), but it is not known if

this 1s a food plant for the larvae of of P. a. azula.

Philiris azula johnsoni subsp. n.

(Figs 39-44, 57)

Philiris azula Wind & Clench: Johnson and Johnson 1984: 89-90.

Types. Holotype 6, QUEENSLAND: labelled ‘Gordon Creek, Iron Range Qld, 1-

10.v11. 1981, LR. & A.J. Johnson’, ‘Figured in Butterflies of Australia (1999), CSIRO

Publishing, M.F. Braby’, ‘Photog. Spm PS 2130’, “GENITALIA SLIDE No. 042’, in

QM. Paratypes: 1 9, labelled “Mt. White Summit, Coen, N. Qld, 6 July 1976, G.B. &

S.R. Monteith’, in QM; 1 ĝ, labelled “Iron Range, Gordon Creek, 15-6-90, Bill

Graham’, in ANIC.

Description. Male. Antenna length (of holotype) 7.4 mm, shaft black with

segmental bands white, club black, tipped orange dorsally, orange 2/3

apically; eyes brown, edged with white cilia; frons white; thorax and

abdomen dorsally dark grey-black, ventrally white; palpus white, apical

segment light brown; tibiae white with brown bands at apex of segment, tarsi

banded white and brown. Forewing length (of holotype) 13.4 mm, apex

acute, termen and inner margin almost straight; upperside base to subtermen

and inner margin dull violet-blue, apex broadly brown-black to ca 1/3

forewing of costa; termen brown-black, broad at apex, narrower at tornus;

cilia from apex to CuA; brown, CuA, to tornus narrowly white. Hindwing

upperside dull violet-blue from base to median region and subtornus; costa,

Rs and apex light brown; base of inner marginal fold light brown, dark brown

towards tornus; termen narrowly dark brown, broader CuA; to tornus;

Underside both wings silvery white, forewing unmarked except brown vein

ends at termen; hindwing inner margin with brown-black spot on inner

margin; termen white with brown flecks at vein ends CuA;, CuA» and

1A+2A.

Male genitalia. Unmounted: vinculum-tegumen ring oval in posterior view.

Slide mounted: sociuncus and vinculum narrow, saccus expanded, socii

tapered with pointed median lobes, separated by a deep U-shaped sinus,

margins weakly convex, clothed with fine setae; brachia slender, inwardly

curved, apically more slender, tip inwardly tapered to a blunt point; valvae

almost symmetrical, subtriangular, longer than wide with blunt subsquared

apices, clothed apically with fine setae; juxta slender, subcylindrical, with

slerotised ring surrounding aedeagus; aedeagus subcylindrical, prezonal

Australian Entomologist, 2015, 42 (4) 235

sheath broadly expanded anteriorly, postzonal sheath with retracted cornuti

with two groups of finely sclerotised spines; anal tube slender, weakly

sclerotised. The male genitalia were also figured by Johnson and Johnson

(1984).

Figs 39-44. Philiris azula johnsoni Sands: (39-42) SS), (43-44) 99; (39, 41, 43)

uppersides, (40, 42, 44) undersides. Localities: (39-42) Gordon Creek, Iron Range,

Qld; (43-44) Mount White, Coen, Qld.

Female. Antennal length (of paratype) 7.3 mm, shaft grey-black with narrow

white segmental bands, club dorsally dark brown-black, apically tipped

orange-brown, orange 2/3 apically; eyes grey-brown edged white; frons grey-

brown, edged white; palpus dorsally brown, ventrally white; thorax and

abdomen dorsally light grey with long grey setae, ventrally white; tibiae and

236 Australian Entomologist, 2015, 42 (4)

tarsi white, broadly banded brown. Forewing length (of paratype) 14.5 mm,

termen bowed, apex obtuse; upperside costa, apex, termen and tornus broadly

grey-brown; area from base to cell, to inner margin and postmedian, light

silvery blue; a subcentral white oval patch between bases of Ms, CuA; and

CuA;, extending to subtermen. Hindwing upperside with apical half of costa

white and with greyish brown suffusion extending from apex to M;; an area

from base and cell to subtermen and subtornus pale blue-violet, greyish

towards termen, crossed by dark grey veins; termen bowed, broadly (ca 1.5

mm) dark grey-brown; cilia white. Underside both wings silvery white;

forewing with small obscure greyish patch in median region, termen and cilia

white; hindwing white, inner margin with prominent submedian black spot;

cilia white, except black at vein ends CuA;, CuA» and 1A+2A.

Comments. The distinctive, weakly iridescent, lilac-blue colour on the

upperside of both wings of P. a. johnsoni males distinguishes it from other

Australian Philiris spp. When males of P. a. johnsoni are compared with P. z.

ziska, the forewings of P. a. johnsoni are slightly narrower and the dark apex

and termen is broader. The male genitalia of P. a. johnsoni do not differ

significantly from those of nominotypical P. a. azula. Identification of the

female specimen as a paratype of P. a. johnsoni is based on the antennae and

the presence of a black spot on the underside of the inner margin of the

hindwing. On the upperside, P. a. johnsoni differs from females of most other

Australian Philiris spp. by the forewing white patch and more extensive areas

of pale blue on both wings. The distribution of white and blue areas on the

upperside resembles that of a small female of P. papuana kerri, but the shade

of blue is different, the hindwing costa is grey rather than white and the black

spot on the inner margin of the underside of the hindwing on P. azula

johnsoni distinguishes it from P. papuana kerri, a species in which the

underside hindwing spot is always absent.

Etymology. Named in honour of Stephen Johnson, in recognition of his many

contributions to the study of Australian butterflies, especially in northern Qld.

Distribution. Cape York Peninsula, northern Qld: Iron Range, Claudie River

and Coen. Only two males and one female have been collected to date.

Biology. The life history and food plant(s) of P. a. johnsoni are unknown.

Philiris innotata (Miskin, 1874)

(Figs 45-52, 58)

Philiris ilias Felder, 1860: Waterhouse 1903b: 652; misidentification.

Pseudodipsas innotatus Miskin, 1874: 165.

Philiris ilias innotatus (Miskin): Waterhouse and Lyell 1914: 76; Waterhouse 1932:

137; Waterhouse 1937: 115.

Philiris innotatus evinculis Wind & Clench, 1947: 11; syn. n.

Philiris imotatus innotatus (Miskin): Tite 1962: 247; Common 1964: 124; McCubbin

1971: 72; d'Abrera 1971: 374.

Australian Entomologist, 2015, 42 (4) 237

Philiris innotatus evinculis Wind & Clench: Tite 1963: 248; Common 1964: 124;

d'Abrera 1971: 374; McCubbin 1971: 72; Common and Waterhouse 1972: 428;

Edwards 1996: 250.

Philiris innotata innotata (Miskin): Common and Waterhouse 1972: 427-428.

Philiris innotatus (Miskin): Edwards 1996: 250; Braby 2000: 683.

Philiris innotata (Miskin): Parsons 1998: 375-376; Orr and Kitching 2010: 236.

Types. Philiris innotata: Miskin’s syntypes of Pseudodipsas innotatus from Brisbane

have not been located (Hancock 1995) and specimens lodged in the Queensland

Museum are not part of the syntypic series (S. Wright pers. comm.). In a register of

specimens at AM, G.A. Waterhouse noted, when referring to Philiris ilias: “Types

ĉ Brisbane in Miskin Coll, Queensland Museum”. G.A. Waterhouse examined most

of Miskin’s types in July, 1910 (unpublished records, Australian Museum, Sydney;

Hancock 1995) and referred to two males and one female of Philiris innotata but did

not attach any labels to these specimens.

Philiris innotatus evinculis: Holotype 3, QUEENSLAND: labelled “Redlynch, North

Queensland, Australia, August 14, 1938 (R.G. Wind)’; allotype 9 with same locality

and collector but September 17, 1938; 2 paratype ĉĉ, l paratype 9, same locality

and collector but dated October 1, 1938, October 3, 1938 and August 14, 1938

respectively. Holotype, allotype and one paratype 2 in Cornell University collection,

one paratype &, one paratype 9 in collection of R.G. Wind, Berkeley, California.

Diagnosis. The description of both sexes of P. innotata by Miskin (1874) was

adequate and the male genitalia were figured by Tite (1963).

Both sexes of P. i. evinculis were described from specimens collected at

Redlynch, Cairns, by R. G. Wind (Wind and Clench 1947). However,

consistent differences (in colour and width of termen) between males from

Cairns, Townsville and Magnetic Island, and those from Brisbane, are

insufficiently distinct to justify separate subspecific rank. Not recognised as

distinct in recent literature (and informally listed in synonymy by Braby

2010), P. i. evinculis 1s here formally synonymised with P. i. innotata.

Sands (1981a) proposed the innotata species-group to include several related

species and this was subsequently accepted by Parsons (1998).

Variation. FWL: Ĉĉ, 11.7-13.55 mm; 99, 11.7-14.2 mm. Males of P.

innotata from Brisbane to Cooktown are variable in wingspan, shape of the

forewing, extent of purple on the upperside and width of the black termen at

the apex of the forewing. Males of P. innotata from the Claudie River are

mostly smaller than those from Cairns and Townsville, the apex of the

forewing is often more acute and the termen almost straight. In females, the

extent of blue on the upperside of both wings is very variable and sometimes

absent from the hindwing. When the blue area is extensive on the forewing,

the veins of the forewing are sometimes white. Some females from northern

inland Queensland have the most extensive areas of blue on both wings,

possibly a result of extremes in temperature experienced during immature

development.

Australian Entomologist, 2015, 42 (4)

Figs 45-52. Philiris innotata (Miskin): (45-48) SS, (49-52) 99; (45-47, 49-52)

uppersides, (48) underside. Localities: (45, 48-50) Innot Hot Springs, Old; (46) Coen

River, Qld; (47) Brisbane, Old; (51) Davies Creek, Qld; (52) Burleigh, Qld.

Australian Entomologist, 2015, 42 (4) 239

Figs 53-58. Philiris spp., male genitalia: (53) P. diana Waterhouse & Lyell; (54) P.

papuana kerri Sands; (55) P. fulgens kurandae Waterhouse & Lyell; (56) P. ziska

titeus d' Abrera, (57) P. azula johnsoni Sands; (58) P. innotata (Miskin). Images

clockwise: lateral view, unmounted; slide mounted, posterior view, sociuncus, valvae

& aedeagus. Scale bar = ca 1 mm for each species.

240 Australian Entomologist, 2015, 42 (4)

Distribution. Queensland: Cape York, Portland Roads, Claudie River, Coen

River, Mcllwraith Range, Cooktown, Cairns, Kuranda, Townsville, Magnetic

Island, Great Palm Island, Mackay, Rockhampton, Gayndah, Maryborough,

Kin Kin, Brisbane, Mt Tamborine, Burleigh Heads; New South Wales:

Richmond River, Coffs Harbour, Port Macquarie. An undescribed subspecies

of P. innotata has been recorded from the Eastern and Western Highlands of

Papua New Guinea (Sands and Fenner 1978, Parsons 1998).

Biology. The life history is well known and the larval food plants are mostly

sandpaper figs (Ficus spp: Moraceae), including F. coronata Spin, F. fraseri

Miq. and F. opposita Miq. Larvae also feed on introduced figs, including the

Indian F. benghalensis L. and Asian F. carica L. The feeding patterns

produced by larvae of P. innotata on Ficus are distinctive and similar to those

of several other Philiris spp. on other food plants, including P. intensa Butler

in Papua New Guinea (Parsons 1998). In subtropical eastern Australia, larvae

frequently compete for suitable leaves with larvae of the chrysomelid beetle

Ponerida semipullata (Clark) (Sands and House 1990). In some areas, such

as Magnetic Island and near Brisbane, the abundance of P. innotata may

increase locally if the exotic F. benghalensis has been cultivated and where

competitive herbivory by the larvae of Ponerida semipullata is absent.

Philiris nitens (Grose-Smith, 1898)

(Figs 59-66, 73)

Holochila nitens Grose-Smith, 1898: 107.

Philiris kamerungae Waterhouse, 1903a: 650; Waterhouse 1903b: 189; syn. Sands

1980: 82.

Philiris nitens (Grose-Smith): Waterhouse 1903b: 190; Edwards 1996: 250, Braby

2010: 34; Orr and Kitching 2010: 237.

Philiris kamerungae kamerungae Waterhouse: Waterhouse and Lyell 1914: 77;

Waterhouse 1932: 137; Tite 1963: 240; Common 1964: 124; d'Abrera 1971: 372;

McCubbin 1971: 72; Common and Waterhouse 1972: 425.

Candalides kamerungae (Waterhouse): Grunberg in Seitz 1921: 853.

Philiris nitens nitens (Grose-Smith): Tite 1963: 240; Common and Waterhouse 1972:

426; Sands 1980: 82.

Philiris nitens restricta Tite, 1963: 241; Common and Waterhouse 1972: 426; syn.

Sands 1980: 82.

Types. Holochila nitens: Holotype 6, QUEENSLAND: labelled ‘nitens Grose-Smith

Type 3’, ‘N. Queensld.”, “Gen. 1962. 436. G.E.T.”, ‘Rothschild bequest BMNH 1939-

P with genitalia slide, in BMNH.

Philiris kamerungae: Lectotype É (designated by Peters (1971) ‘by inference of

holotype’), QUEENSLAND: labelled ‘Kuranda, N.Q., Mar. 1902, R.E. Turner’,

“Philiris kamerungae type £, G.A. Waterhouse, KL21485', in AM. This specimen

bears a red label and was listed as the holotype, in the handwriting of G.A.

Waterhouse, in a register of specimens at AM. It was also listed as the holotype by

Peters (1971) in his catalogue of types in the Australian Museum.

Australian Entomologist, 2015, 42 (4) 241

Philiris nitens restricta: Holotype 6, QUEENSLAND: labelled ‘Cedar Bay, S. of

Cooktown (Meek)’, “Rothschild bequest BM. 1939-1’, “Gen. 1962 440. G.E.T.”,

“Type HT’, “Philiris nitens restricta Tite, Holotype 2. BMNH Type No. RL

16843’ with genitalia slide; allotype 9, labelled “Cedar Bay, S. of Cooktown (Meek)’,

“Type AT’, ‘Philiris nitens restricta Tite’, “ALLOTYPE 9”, “BM Type No. RL.

16844’, both in BMNH.

Diagnosis. The wingspans of males of P. nitens (FWL ca 12 mm) are similar

to those of P. sappheira manskiei (FWL ca 12.5 mm) but slightly greater than

those of P. /ucina (FWL 11.5 mm). In both sexes the ventral surface of the

antennal club of P. nitens is weakly tipped orange, but in P. sappheira

manskiei and P. lucina the ventral apical half of the club is orange. Males of

P. nitens can be distinguished from males of P. sappheira manskiei by the

blue to purple-blue upperside areas, while the forewings are narrower and the

hindwings weakly bowed; in males of P. sappheira manskiei the upperside

areas are always bluish green, the forewings broader and the hindwings more

rounded than in P. nitens. The blue areas of P. nitens males are also more

extensive than the blue-green areas of P. sappheira manskiei males. The

upperside of both fore and hindwings in male and female P. nitens usually

have areas of white, unlike both sexes of P. sappheira manskiei, which are

without any areas of white on the upperside. However, when areas of white

are occasionally absent in males of P. nitens, the costa of the hindwing has

whitish suffusions (Ring and Olive 1997) that are absent in P. sappheira

manskiei. The hindwing black terminal band of P. nitens 1s subparallel to the

termen, but in P. sappheira manskiei this band extends from the costa and

from Rs to 1A+2A. The upperside of females of P. nitens has variable areas

of white that are not present on the upperside of females of P. sappheira

manskiei.

Areas of white on both fore and hindwings of P. nitens and P. lucina are

often similar in the two species (Common and Waterhouse 1972) and females

may be difficult to distinguish. Males of P. nitens can be distinguished from

P. lucina by the darker blue or purple-blue areas on both wings in P. nitens,

compared with the paler sky-blue areas on both wings in P. /ucina. The

forewing of males of P. nitens usually, but not always, has a suffusion of

white scales or a prominent oval patch of white scales beyond the cell

between the bases of M; and CuA,, whereas in males of P. /ucina there is a

narrow strip of white scales from the bases of Rs and M, between cell and

tornus and below CuA;. On the hindwing of P. nitens males, the clear purple-

blue area is bounded by a narrow grey-black terminal band, with or without a

variable white area extending from mid costa to the apex and occasionally the

termen at Mı, whereas in males of P. /ucina the greenish blue area is crossed

by dark veins. The grey-black terminal band of P. /ucina is broad from Rs to

the tornus, as well as on the apical half of the inner margin between 1A+2A.

The basal half of the inner marginal fold of P. /ucina is more prominent and

more broadly white than that of P. nitens.

242 Australian Entomologist, 2015, 42 (4)

Figs 59-66. Philiris nitens (Grose-Smith): (59, 61, 63, 65) SS, (60, 62, 64, 66) 99;

(59-64) uppersides, (65-66) undersides. Localities: (59) Bloomfield, Qld; (60-61)

Kuranda, Qld; (63, 65) Innisfail, Qld; (64) Cairns, Qld; (62, 66) Josephine Falls, Qld.

Australian Entomologist, 2015, 42 (4) 243

In females of P. /ucina the postmedian areas of white on the hindwing are

crossed by dark postcellular veins, whereas the veins of P. nitens are not

prominently darker than the ground colour. A darker apical cell bar on the

hindwing of P. /ucina at the bases of Mj, M and M; assists in distinguishing

females of P. /ucina from those of P. nitens. Sands (1981a) proposed a nitens

species-group to include P. nitens and related species, subsequently accepted

by Parsons (1998). The male genitalia were figured by Tite (1963).

Variation. FWL: dd, 12.2-13.0 mm; 99, 12.3-14.8 mm. Males and females

of P. nitens vary considerably in the extent of areas of white on the upperside

of both fore and hindwings, due in part to seasonal effects and the

temperatures experienced during immature development. For example, a

female of P. nitens reared in Brisbane in June (Fig. 60) has much more

extensive areas of white than specimens reared or collected during warmer

months (J.F.R. Kerr pers. comm.). In males, the pale blue forewing usually

has a variable small white median patch between the bases of M3, CuA; and

CuA», but the forewing is occasionally entirely pale blue or dark blue,

without any areas of white. On the hindwing, the white costa of males also

varies in extent and may be just visible or extend as far as Mı.

Distribution. Northern Queensland, from near Cooktown south to Bluewater

Range, 24 km north of Townsville: 19 km S. of Cooktown (Ring and Olive

1997), Cedar Bay, Rossville, Mossman Gorge, Kuranda, Lake Placid, Cairns,

Mission Beach, Innisfail, Herbert River, Ingham, Kirrama, Paluma,

Mutarnee, Bluewater Range.

Biology. Larvae feed on Macaranga involucrata var. mallotoides (F. Muell.)

L.M. Perry, and occasionally G/ochidion phillipicum (Cav.) C.B. Rob.

(Common and Waterhouse 1981) and Macaranga tanarius (L.) Muell. Arg.

(L. Ring).

Philiris lucina Waterhouse & Lyell, 1913, stat. n.

(Figs 67-72, 74)

Philiris kamerungae lucina Waterhouse & Lyell, 1914: 77; Waterhouse 1932: 138;

Tite 1963: 240; Common 1964: 124; d'Abrera 1971: 372; McCubbin 1971: 72;

Common and Waterhouse 1972: 425.

Philiris nitens lucina: Common and Waterhouse 1981: 464; Edwards 1996: 250,

Braby 2010: 34.

Types. Lectotype É (here designated), QUEENSLAND: labelled “Cape York, 18 JLY

1910, H. Elgner’, 'KL21517', “G.A. Waterhouse collection’. Paralectotypes (here

designated): 1 A. labelled “Cape York, H. Elgner’, “KL21520”; 1 9, labelled “Cape

York, 22 Oct 1908, H. Elgner’, “Cape York, Q. 3:3:08, G.A. Waterhouse Coll.

Elgner’, and “KL21518”; 1 9, labelled ‘Cape York, 22 Oct 1908, H Elgner’, and

“KL21518”; 1 9, labelled “Cape York, 3.8.10, H. Elgner’, “KL21521”, all in AM.

The Lectotype, here designated, was listed as the holotype, in the handwriting of G.A.

Waterhouse, in a register of specimens at AM.

244 Australian Entomologist, 2015, 42 (4)

Figs 67-72. Philiris lucina Waterhouse & Lyell: (67, 69, 71) do, (68, 70, 72) 99;

(67-70) uppersides, (71-72) undersides. Localities: (67-69, 71) Claudie River, Old;

(70, 72) Iron Range, Qld.

Diagnosis. The pale shining blue areas and white patches on the upperside of

males of P. /ucina are somewhat similar to those of P. scintillata Sands from

Papua New Guinea, but the wingspan of P. /ucina is smaller and the male

genitalia differ considerably from those of P. scintillata (Sands 1981b).

When males of P. nitens are without white on the upperside, they may be

similar to P. sappheira manskiei but are blue rather than blue-green. Philiris

lucina can be distinguished from P. nitens by the more extensive orange on

Australian Entomologist, 2015, 42 (4) 245

the ventral surface of the antennal club and by the shorter valvae of the

genitalia.

Females of P. /ucina and P. nitens are readily distinguished from P.

sappheira manski by the presence of white areas on the upperside of both

species. In females of P. sappheira manski both wings have variable blue-

green areas and are without the white areas that are always present in females

of P. lucina and P. nitens.

The male genitalia of P. /ucina (Fig. 74) and P. nitens (Figs 73) are readily

distinguished from those of P. sappheira manskiei (Fig. 75) by the length and

shape of the valvae (Ring and Olive 1997).

Figs 73-76. Philiris spp., male genitalia: (73) P. nitens Waterhouse & Lyell; (74) P.

lucina Waterhouse & Lyell; (75) P. sappheira manskiei Ring & Olive; (76) P.

sappheira sappheira Sands. Images clockwise: lateral view, unmounted; slide

mounted, posterior view, sociuncus, valvae & aedeagus. Scale bar = ca 1 mm for each

species.

246 Australian Entomologist, 2015, 42 (4)

Variation. FWL: ĉĉ. 10.8-11.4 mm; 99, 10.8-12.4 mm. In males, the extent

of pale shining blue areas and white patches is variable and in females, on the

hindwing, the extent of white may be restricted to the costa or extend almost

to the apex.

Distribution. Cape York Peninsula, northern Queensland: Bamaga and Iron

Range to the Rocky River. Philiris lucina is most abundant near Heathlands

and near the Claudie River. Philiris lucina, P. nitens and P. sappheira

manskiei are allopatric in distribution.

Biology. Larvae of P. lucina feed on Macaranga involucrata (Ring and Olive

1997). The life history is otherwise very similar to that of P. nitens.

Philiris sappheira sappheira Sands, 1980

(Figs 76, 83)

Philiris nitens sappheira Sands, 1980: 82.

Philiris sappheira sappheira Sands: Ring and Olive 1997: 65; Orr and Kitching 2010:

236.

Type. Holotype 8, PAPUA NEW GUINEA: Rouna Falls, Central Province, in ANIC.

Male genitalia (Fig. 76). Very similar to those of P. s. manskiei from

northern Queensland but the valvae are not as prominently ‘club-shaped’ as

in P. s. manskiei.

Variation. FWL: ĉĉ. 12.5 mm; 9, 13.5 mm. Males of P. sappheira may be

distinguished from other species of similar size in the nitens species-group by

the distinctive greenish blue areas and the absence of white on the upperside

of both wings. Only one female (a paratype) of P. s. sappheira is known and

confirmation of its identity will require more material, preferably when

females are reared with males and confirmed to be conspecific, or by using

DNA methods capable of discriminating closely-related taxa.

Distribution. Known from only the southern end of the Kokoda Track, Sogeri

Plateau and Rouna Falls, Central Province, Papua New Guinea.

Biology. Unknown. Adults were collected near the embankments of shallow

watercourses where a Macaranga sp. was abundant.

Philiris sappheira manskiei Ring & Olive, 1997

(Figs 75, 77-82, 84)

Philiris sappheira manskiei Ring & Olive, 1997: 66-71; Braby 2010: 34; Orr and

Kitching 2010: 236.

Types. Holotype 3, QUEENSLAND: labelled *McIvor River Road, 11 May 1994,

L.R. Ring, xp M. involucrata’, ‘HOLOTYPE Philiris sappheira manskiei”, “ANIC

Type Reg. No. 3330, genitalia slide No 3438’, “GART Exemplar und Etik-ketten

dokumentiert specimen and label data documented 22.3.2002’, + pupal shell on point,

in ANIC.

Australian Entomologist, 2015, 42 (4) 247

Y é

L9 D A

uid

TU A

a ~E

83 84 Tn fs

Figs 77-84. Philiris sappheira subspecies: (83) P. s. sappheira Sands, (77-82, 84) P.

s. manskiei Ring & Olive; (77, 79, 81, 83) 33, (78, 80, 82, 84) 9 9; (77-80, 82-83)

uppersides, (81, 84) undersides. Localities: (78) 35 km NW Cooktown, Old; (77, 79-

82, 84) Mclvor River, Qld; (83) Rouna Falls, Central Province, PNG.

248 Australian Entomologist, 2015, 42 (4)

Diagnosis. Males of P. sappheira manskiei differ only very slightly from

males of P. s. sappheira, with the blue area of the hindwing of males

extending to the postmedian region in P. s. manskiei but only to the median

region in P. s. sappheira. The illustrated female in Braby (2004) is a paratype

of P. s. sappheira from Papua New Guinea. This specimen of the

nominotypical subspecies differs from P. s. manskiei by the extensive areas

of white on the upperside of both wings.

The male genitalia were figured by Ring and Olive (1997) and compared

with those of P. nitens nitens and P. nitens lucina.

Sands (1981a) proposed a nitens species-group for P. nitens and related

species including P. sappheira and which now includes P. s. manskiei.

Variation. FWL: oo, 10.8-12.8 mm; 99, 12.7-13.3 mm. Females of P. s.

manskiei may resemble female P. moira Grose-Smith (Forbes 1977) from

Papua New Guinea, particularly in some forms when the forewing area of

green is restricted. The extent of silvery greenish blue areas on the upperside

of the forewing of females varies from an obscure narrow strip of greenish

scales from the base to the cell, not reaching the inner margin (e.g. in Braby

2004), to an extensive oval and silvery green central area, extending from the

base beyond the cell to the inner margin and postmedian region. The

hindwing may be almost uniformly dark grey with paler costa, to light grey

with obscure lighter grey scales in the central areas reaching the cell and

postmedian region.

Distribution. Queensland: 39 km NW of Cooktown, 8 km SW of Mount

Webb, Mclvor River, 3 km NW of Hopevale, Cedar Scrub, Endeavour Falls,

3 km SE of Isabella Falls (ANIC, Ring and Olive 1997).

Biology. Larvae of P. sappheira manskiei feed on Macaranga involucrata

(Ring and Olive 1997).

Discussion

Infraspecific variation in adult morphology, particularly that of females (e.g.

Forbes 1977), has contributed to difficulties in associating sexes in several

Philiris spp. from Papua New Guinea and without any evidence for seasonal

variation. By contrast, variation in Australian species appears to be seasonal,

evident in some species with allopatric distributions. Morphological

variation, particularly the extent of white or blue scales, is thought to be

induced by variation in temperatures (unpublished observations) during

development; for example, females of P. nitens and P. innotata reared from

immature stages during winter months are usually much paler, with more

white or blue on the upperside respectively, than females reared during

summer months. The extent of white areas on the upperside of both sexes of

P. diana may also be temperature dependent.

Australian Entomologist, 2015, 42 (4) 249

Asymmetry in the male genitalia of some Philiris spp., in both shape and

length of the valvae when viewed posteriorly, is known in other members of

the diana species-group (e.g. P. siassi Sands: Sands 1979) and this unusual

morphology is most easily observed when the genitalia are slide mounted.

Moreover, the morphology of the male genitalia, including sclerites of the

prezonal sheath, may vary with age of the specimen (for example, with

freshly eclosed specimens when sclerotisation of the prezonal sheath is

weakly developed) and the position of retractile cornuti in the aedeagus may

also vary according to age.

Very little is known of the ecology of Australian Philiris spp. except for

some larval food plants. The distribution of members of the nitens species-

group is enigmatic, with all three species being allopatric in Australia and

utilising the same food plant, Macaranga involucrata.

Acknowledgements

Sincere thanks to Peter Valentine, David Hancock, Graham Forbes, Russell

Mayo and Ted Edwards for helpful discussions and for providing access to

specimens for study and thanks are due to Bill Graham for depositing a

paratype of P. azula johnsoni in the Australian National Insect Collection,

Canberra. Special thanks to Albert Orr, John Kerr and Tim New for

constructive reviews of the manuscript and to Susan Sands for preparing

many of the genitalia illustrations. The late Murdoch De Baar provided much

valuable information during earlier preparations for this review and loan of

his specimens for study.

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papuana Wind & Clench (Lepidoptera: Lycaenidae). Australian Entomological Magazine 10:

81-83.

Australian Entomologist, 2015, 42 (4): 253-255 253

A FURTHER RECORD OF DANAUS CHRYSIPPUS CRATIPPUS

(C. FELDER, 1860) (LEPIDOPTERA: NYMPHALIDAE: DANAINAE)

FROM THE NORTHERN TERRITORY, AUSTRALIA

MICHAEL F. BRABY

Research School of Biology, Australian National University, Canberra, ACT 0200

Abstract

A female Danaus chrysippus cratippus (C. Felder) is recorded from the Darwin suburb of

Wanguri, Northern Territory, on 13 April 2015. This record represents only the third location for

the species within Australian limits, the two others being the Black Point and Smith Point areas

of Cobourg Peninsula, Northern Territory, and Thursday Island in Torres Strait, Queensland.

Introduction

The Plain Tiger, Danaus chrysippus (Linnaeus, 1758), 1s a polytypic species

distributed widely in the Afrotropical and Oriental Regions (Smith 2014). In

Australia, however, it is rarely encountered, being known previously only

from the Black Point and Smith Point areas of Cobourg Peninsula, Northern

Territory [a male collected in 1977 (Common and Waterhouse 1981) and 13

specimens collected in 2007 (Braby 2014, Braby ef al. 2015)] and from

Thursday Island in southern Torres Strait, Queensland [a female collected in

1995 (Lambkin 2009)]. All of these specimens were collected from January

to April, which is typically the monsoon or wet season in northern Australia,

and all specimens have been assigned to the subspecies D. chrysippus

cratippus (C. Felder, 1860), which is restricted to the eastern half of Wallacea

(Braby et al. 2015). Permanent breeding populations of the species do not

appear to be established on the Australian mainland, with all previous records

representing either vagrants from Maluku and/or the Lesser Sunda Islands, or

progeny from temporary colonisations that failed to establish.

The species is taxonomically and phylogenetically distinct from the well

known Lesser Wanderer, D. petilia (Stoll, 1790) (Lushai et al. 2005, Braby et

al. 2015), and is distinguished from that species by the following six

phenotypic characters (see Braby ef al. 2015): (1) the upperside ground

colour is uniformly tawny brown or dark orange-brown, with the forewing

discal cell and basal area of cell M; diffusely contrasted in darker brown; (2)

the underside ground colour of the forewing is uniformly rusty brown or dark

orange-brown, with the discal cell and basal area of cell M3 a darker shade of

rusty brown; (3) the length of the white subapical spot in cell M» of the

forewing is significantly longer; (4) the white subapical spot in cell M3 of the

forewing is always present and significantly larger; (5) the width of the black

marginal band on the upperside of the hindwing is significantly narrower,

often enclosing a few white submarginal spots, with the inner edge of the

band usually lacking the rusty brown suffusion; and (6) the black marginal

band on the underside of the hindwing is narrower and usually encloses only

a single row of white spots or, if two rows of spots are present, the inner row

of spots is smaller and generally less distinct and adjacent to the band.

254 Australian Entomologist, 2015, 42 (4)

Observations

At Wanguri, an outer northern suburb of Darwin, NT (12.373°S, 130.886°E),

a female D. chrysippus cratippus was observed feeding on flowers of Tridax

procumbens L. (Asteraceae) at 1130 hrs on 13 April 2015. The specimen

(Figs 1-4) was in worn condition and was located near a shopping centre at

the intersection of Gsell Street and Vanderlin Drive. At the time of

observation I had no collecting equipment with me, but I was able to

photograph and collect the specimen by hand. The female was kept alive for

48 hrs, fed with diluted honey-sugar solution and placed inside a plastic bag

supplied with fresh cuttings of Cynanchum carnosum (R.Br.) Schltr.

(Apocynaceae) in a shade house, but did not oviposit during this period.

Figs 1-4. Female Danaus chrysippus cratippus from Darwin, NT, showing: (1)

upperside, (2) underside, (3-4) feeding from flowers of Tridax procumbens. Scale bar

for Figs 1 and 2 = 20 mm.

Discussion

The specimen agrees with the diagnosis of Braby et al. (2015) in that it

possesses all six phenotypic characters of D. chrysippus cratippus that

distinguish it from D. petilia. The specimen was collected at the end of the

wet season and presumably had arrived following the summer monsoon trade

winds from Indonesia, which typically occur during December-March.

Darwin is located approximately 200 km south-west of Black Point on

Australian Entomologist, 2015, 42 (4) 255

Cobourg Peninsula and a straight line drawn between these two locations is

perpendicular to the direction of the monsoon trade winds, suggesting that the

specimen was unlikely to have originated from Cobourg Peninsula.

References

BRABY, M.F. 2014. Remarks on the spatial distribution of some butterflies and diurnal moths

(Lepidoptera) in the Top End of the Northern Territory, Australia. Northern Territory Naturalist

25: 29-49.

BRABY, M.F., FARIAS QUIPILDOR, G.E., VANE-WRIGHT, R.I. and LOHMAN, D.J. 2015.

Morphological and molecular evidence supports recognition of Danaus petilia (Stoll, 1790)

(Lepidoptera: Nymphalidae) as a species distinct from D. chrysippus (Linnaeus, 1758).

Systematics and Biodiversity 13: 386-402.

COMMON, I.F.B. and WATERHOUSE, D.F. 1981. Butterflies of Australia. Revised Edition.

Angus and Robertson, Sydney; xiv + 682 pp.

LAMBKIN, T.A. 2009. A record of Danaus chrysippus cratippus (C. Felder) (Lepidoptera:

Nymphalidae: Danainae) from Thursday Island, Torres Strait. Australian Entomologist 36: 33-

36.

LUSHAI, G., ZALUCKI, M.P., SMITH, D.A.S., GOULSON, D. and DANIELS, G. 2005. The

lesser wanderer butterfly, Danaus petilia (Stoll 1790) stat. rev. (Lepidoptera: Danainae),

reinstated as a species. Australian Journal of Entomology 44: 6-14.

SMITH, D.A.S. 2014. African Queens and their kin: A Darwinian odyssey. Brambleby Books,

Taunton, UK; 848 pp.

256 Australian Entomologist, 2015, 42 (4)

BOOK NOTICE

All about butterflies of Australia, by Garry Sankowsky. Paperback; 167 pp;

ISBN: 9781921517433; $29.99. Available from: Attp://www.publish.csiro.au/

nid/18/pid/7532.htm

This delightful book draws on the enormous experience of the author with

butterflies in the field and is filled with hundreds of photographs of living

butterflies and their life histories and food plants. Included are many snippets

of information about living butterflies, their behaviour, breeding habits and

preferences, seasonal quirks and migrations, mimicry and defences. Also

discussed is the decline in butterflies due to factors such as clearing of the

brigalow belt, decline in dry vine scrubs due to annual burn-offs and the

steady invasion of natural habitats by aggressive introduced pasture grasses

that are rapidly replacing the natural vegetation, including butterfly food

plants, throughout Australia.

For further details see the News Bulletin of the Entomological Society of

Queensland, Volume 43, Part 5, page 88 (August 2015).

Australian Entomologist, 2015, 42 (4): 257-263 257

TELICOTA SP. NR KEZIA KEZIA EVANS, 1949 (LEPIDOPTERA:

HESPERIIDAE: HESPERIINAE) FROM DAUAN ISLAND, TORRES

STRAIT, QUEENSLAND

TREVOR A. LAMBKIN

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

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

Abstract

Four specimens of a species of Telicota Moore, 1881 new to Australia are recorded from Dauan

Island, Torres Strait: a male and female collected from blossom of Citharexylum spinosum L.

and Melaleuca L. sp. respectively, and another two males collected while flying amongst tall

grass in semi-shade under Melaleuca trees. Both sexes are illustrated, including the genital

armature of a male that is closest in structure to that of 7. kezia kezia Evans, 1949 from Papua

New Guinea and 7. ohara ohara (Plótz, 1883) from Papua New Guinea and Australia, while the

external facies, except the male sex brand, best fit T. k. kezia. In addition, the specimens possess

on the hindwing underside a postmedian band that is divided by black veins, a character shared

with 7. o. ohara. The male sex brand of the Dauan Island specimens is broader and more

centrally placed in the dark central band of the forewing than that of 7. k. kezia. Because the

Dauan Island specimens do not match any described species, and due to their external similarity

to T. k. kezia, they are referred to here as 7. sp. nr kezia kezia. It is proposed that because of their

similarities to 7. o. ohara, they might share some relationship with this species and thus could

share the same larval hostplant, i.e. Flagellaria indica L.

Introduction

Telicota Moore, 1881 is a genus of predominantly tropical skippers (Evans

1949) often referred to as darters or darts (Corbet and Pendlebury 1992,

Braby 2000). The genus is wide ranging, occurring from Sri Lanka and India

across south-east Asia, through Indonesia and New Guinea, to the Solomon

Islands and Australia (Evans 1949, Corbet and Pendlebury 1992, Parsons

1998, Bascombe et al. 1999, Braby 2000, Tennent 2002). Species in the

genus are characterised by their black and orange colouration, distinctive

forewing sex brands in males and rounded forewings in females. They are

moderate-sized skippers, with forewing lengths ranging from 14-17 mm in

males and 15-19 mm 1n females (Evans 1949, Corbet and Pendlebury 1992,

Parsons 1998, Braby 2000).

The genus is diverse with more than 38 species recognised by Parsons

(1998), who suggested that there were 25 species in Papua New Guinea

alone. Nine species are currently recognised in Australia (Braby 2000). The

type species is 7. colon Fabricius, 1775 from India (Evans 1949, Edwards et

al. 2001).

Telicota species are found in a variety of habitats but predominantly occur in

damp, swampy or estuarine areas, or along the margins of monsoon forest,

rainforest or riparian forest (Parsons 1998, Braby 2000), although some

species can occur in more open forest habitats (Parsons 1998, Braby 2000).

Larval host plants of Te/icota in Australia and Papua New Guinea include

grasses (Poaceae), Flagellaria indica L. (Flagellariaceae) and sedges

(Cyperaceae). In Queensland, several species are known from the islands of

258 Australian Entomologist, 2015, 42 (4)

Torres Strait (Waterhouse and Lyell 1914, Braby 2000), but little is

documented on the species diversity of Telicota in the far north of the Strait,

particularly from the islands close to the southern coastline of Papua New

Guinea, viz. Dauan, Saibai and Boigu Islands.

While a study is currently underway documenting the Telicota species from

these northern islands (TAL in prep.), between 2009 and 2015 three males

and one female of a strikingly large species of Telicota were collected at the

western end of Dauan Island. After examination of their wing patterns,

including the male forewing sex brands, and their genital armature, it was

found that the species was hitherto unrecorded from within Australian

territory and best matched 7. kezia kezia Evans, 1949, which occurs

predominantly in New Guinea (Evans 1949, Parsons 1998). Two subspecies

of T. kezia were described by Evans (1949): T. k. kezia from mainland New

Guinea and the islands to the west, including Maluku (Evans 1949, Parsons

1998), and T. k. lenna Evans, 1949 from New Britain, Witu Island, New

Hanover and New Ireland (Evans 1949). The type locality for T. k. kezia is

Mt Mado, Buru, Maluku (Evans 1949). Currently, there appears to be nothing

published on the habits and ecology of 7: kezia (Parsons 1998). Interestingly,

T. k. kezia appears to have a more restricted distribution than Evans (1949)

and Parsons (1998) indicated, as the latter did not record it from the Bulolo-

Wau Valley (Morobe Province) in the north-east of Papua New Guinea,

despite a long period of butterfly sampling in the region (Parsons 1991).

Here I report the details of these specimens and their collection on Dauan

Island, including field observations, and illustrate male and female adults and

the male genital armature. In addition, I discuss the uncertainty surrounding

their identification, together with why observations of this species on Dauan

Island are infrequent despite a relatively robust collecting effort by butterfly

enthusiasts over the last two decades or so.

The following abbreviations refer to repositories from which material has

been examined: ANIC — Australian National Insect Collection, Canberra;

CGMC - Collection of C.G. Miller, Lennox Head; TLIKC — Joint collection

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

Abbreviations of collectors’ names are: CGM - C.G. Miller; EJH- Sir

Edward J.L. Hallstrom; TAL — T.A. Lambkin; WWB — W.W. Brandt.

Material examined

Telicota sp. nr kezia kezia (Figs 1-8)

QUEENSLAND: 1 £, 1 9, Dauan Island, Torres Strait, 6.iv.2009 (4), 13.1.2014 (9),

TAL (TLIKC); 2 GG, same data except 18.iv.2015, 19.iv.2015, CGM (CGMC).

Telicota kezia lenna

PAPUA NEW GUINEA: 1 ĝ, Rabaul, New Britain, 22.viii.1952, collected WWB,

EJH (ANIC); 1 &, Keravat, New Britain, 14.xi.1952, collected WWB, EJH (ANIC).

Australian Entomologist, 2015, 42 (4) 259

Figs 1-8. Telicota sp. nr kezia kezia from Dauan Island, Torres Strait: (1-2) adults:

upperside left, underside right [forewing lengths, in mm, in square brackets]: (1) 3

6.1v.2009 [19 mm] TLIKC, (2) 9 13.1.2014 [20 mm] TLIKC; (3-5) male sex brands

LH forewing [sex brand lengths, in mm, in square brackets]: (3) 6.1v.2009 [5.5 mm]

TLIKC, (4) 19.iv.2015 [5.3 mm] CGMC, (5) 18.1v.2015 [5.4 mm] CGMC; (6-8)

genital armature [lengths of structures, in mm, in square brackets]: 2 6.iv.2009

TLIKC: (6) right valva, dorsal surface uppermost [1.5 mm]), (7) divided uncus dorsal

view [1.5 mm], (8) phallus [1.8 mm].

260 Australian Entomologist, 2015, 42 (4)

Observations

Dauan Island is one of Australia's most northern islands, lying approximately

10 km from the southern coastline of Papua New Guinea. It is roughly 2.5 sq.

km in size, with the bulk of the island being hills composed of exposed large

granite boulders with, amongst these boulders, extensive patches of semi-

deciduous monsoon forest. The highest of these hills is Mt Cornwallis (275

m), which is the dominant feature of the island. The bulk of the village on

Dauan Island is situated at the northwestern end of the island.

The first specimen recorded of this Dauan Island Telicota was a male

collected in April 2009 from blossom of Citharexylum spinosum L.

(Verbenaceae) (fiddlewood) along the main street of the village. This plant is

a common garden tree grown in the village and, at that particular time, the

butterflies Cephrenes moseleyi (Butler, 1884) (Hesperiidae) and Graphium

codrus medon (C. & R. Felder, 1864) (Papilionidae) were collected from the

same blossom. The other three Te/icota specimens were collected towards the

western end of the village, the female in January 2014 from blossom of

Melaleuca L. sp. overhanging the road and the remaining two males in April

2015 at the same location, but in tall grass in semi-shade under Melaleuca.

As in most species of Telicota, the Dauan Island males recorded in April

2015 were observed flying rapidly in sunlit glades and settling on tall grass

stems, and would frequently challenge males of other Telicota species for

territory (C.G. Miller pers. comm.).

The four specimens collected are typical of Te/icota in form, as per Evans

(1949), but they differ from all other Telicota species recorded thus far from

Australia in being much larger and roughly similar in size to C. moseleyi: i.e.

male and female with forewing lengths of around 19-20 mm. The forewing

uppersides of the Dauan Island specimens have almost no extension of the

outer edge of the orange postmedian bands along the veins to the termen

(Figs 1-2) and the hindwing undersides have a fuscous, tawny postmedian

band that is divided by black veins. In addition, the sex brands of the three

Dauan Island males appear slightly broader than in 7. kezia males from Papua

New Guinea (Parsons 1998: figs 391-393 and 395) and are placed more in the

centre of the dark central band of the forewing (Figs 3-5).

The male genital armature of a Dauan Island specimen (in TLIKC) (Figs 6-8)

is close to that illustrated as 7. kezia by Parsons (1998) and, to a lesser

degree, to those depicted by Evans (1949). It is also similar to that illustrated

as T. ohara (Plótz, 1883) from Papua New Guinea (Parsons 1998), Hong

Kong (Bascombe et al. 1999) and Australia (Braby 2000). The valvae (Fig. 6)

and uncus (Fig. 7) of the Dauan Island specimen closely resemble those of

the several specimens of 7: kezia illustrated by Parsons (1998), while only the

uncus matches the single example of 7. kezia depicted by Evans (1949). The

phallus (Fig. 8) was not illustrated by either author. Unfortunately neither

Australian Entomologist, 2015, 42 (4) 261

author indicated whether their drawings of the male genital armature were of

T. k. kezia or T. k. lenna.

Discussion

Evans (1949) placed T. kezia and its allies (T. ohara, T. ternatensis Swinhoe,

1907, T. sadra Evans, 1949) into a group primarily based on the morphology

of the genital armature, in particular on the pointed ends of the divided uncus.

He indicated that the valvae of the four species were variable, even across

subspecies within species, and thus species were difficult to separate using

the structure of the genital armatures alone. Similarly, the genital armature of

the Dauan Island male (Figs 6-8) has some similarities, particularly the

valvae, with the armature of 7. ohara (Evans 1949, Parsons 1998, Bascombe

et al. 1999, Braby 2000).

Within the Telicota group in which Evans (1949) placed 7. kezia, a

distinctive feature of 7. kezia is the shape and placement of the male

forewing upperside sex brand. The sex brand of T. kezia meets vein 1A+2A

approximately midway along its length between termen and base and forms

an almost vertical line, relatively thin and grey-black in colour, lying in the

outer edge of the dark central band, between veins M; and 1A+2A. The sex

brands of the Dauan Island males differ in that they are broader and are

placed centrally in the dark central band of the forewings (Figs 3-5). In

addition, Evans (1949) indicated that the postmedian bands on the hindwing

undersides in both sexes of T. k. kezia are fuscous and narrow, orange in

colour and with distinctive black veins running through them. The

postmedian bands on the hindwing undersides of the Dauan Island males

match those of a 7. k. kezia male illustrated by Parsons (1998) in width but

are broader than those of 7. k. lenna males illustrated by Evans (1949) and

Parsons (1998) and in 2 8 ĝ of T. k. lenna examined in ANIC.

Except for the character and position of the sex brands of the three Telicota

males from Dauan Island, their external facies closely resemble the

illustration of a male 7. k. kezia in Parsons (1998): a ‘pale form’ collected at

Waidoro, Western Province, Papua New Guinea, which is approximately 43

km NE of Dauan Island. In addition, the size of the three Telicota males

reported here from Dauan Island (Figs 1-2) is notable. Evans (1949) and

Parsons (1998) specified that 7. k. kezia males have a forewing length of 16-

18 mm, smaller than the Dauan Island specimens, which have a forewing

length of 19 mm (n = 3), although this might not be significant.

Taking into account size, overall wing pattern and character and placement of

the male sex brands, it remains uncertain whether the Dauan Island material

is T. k. kezia or another, closely related taxon. I cannot, therefore, place the

specimens from Dauan Island confidently in T. kezia and thus refer to them

here as 7. sp. nr kezia kezia.

262 Australian Entomologist, 2015, 42 (4)

Evans (1949) indicated that T. kezia was the only member of his kezia group

that has distinctive black veins running through the postmedian band of the

hindwing underside. He further specified that T. ohara lacked this feature.

Despite this, 7. o. ohara is the only species of Telicota in Australia that does

have predominately black veins running through the postmedian band of the

hindwing underside (Braby 2000). The presence of this character in 7. o.

ohara and T. sp. nr k. kezia from Dauan Island (Figs 1-2) might indicate that

the taxonomy of this Telicota group could be more intricate than Evans

(1949) indicated.

Despite the intensity of butterfly collecting undertaken on Dauan Island over

the past 20 years or so, it is surprising that only four specimens of this taxon

have been collected. Based on Evans’ (1949) assessment that 7. k. kezia was

related to 7. ohara and my assessment that the two taxa share some

similarities in their external facies and genital armature to 7. sp. nr k. kezia, it

is possible that 7. sp. nr k. kezia might share the same hostplant as T. o. ohara

recorded from mainland tropical Queensland (Braby 2000). If this is the case,

then the larvae of T. sp. nr k. kezia might also be dependent on Flagellaria

indica, which is a common component of the semi-deciduous monsoon forest

on Dauan Island (Torres Strait Regional Authority 2013). If F. indica 1s the

host plant of 7. sp. nr k. kezia on Dauan Island, this might explain its apparent

rarity, due at least in part to the butterfly primarily inhabiting the forest

canopy where F. indica predominantly grows and where, in this habitat, F.

indica can form impenetrable dense understories (Jones and Gray 1977,

Williams 1979). Certainly, its southern congener, 7. h. ohara, is also reported

to be uncommon (Common and Waterhouse 1981) and infrequently observed

(Braby 2000), perhaps due to its propensity to also inhabit the forest canopy.

This could mean that 7. sp. nr k. kezia is rarely observed outside the forest,

particularly at ground level, much like 7. o. ohara.

Parsons (1998) noted that W. Brandt, a butterfly worker in Papua New

Guinea in the 1950s (his collection is housed in the ANIC), supposed that the

taxon 7. k. lenna was likely a distinct species. Parsons (1998) also suspected

that, because T. k. kezia varied in its external facies and male genital structure

across Papua New Guinea, there might be several species contained in 7. k.

kezia, which may begin to explain the unexpected size of the specimens

collected on Dauan Island. For now, the four Dauan Island specimens are

tentatively placed as T. sp. nr k. kezia until a more thorough revision is made

of the kezia species-group, particularly in New Guinea.

Acknowledgements

I thank the local community councils and island Elders of Dauan Island,

Torres Strait for allowing entry into their communities and providing

assistance during time spent on their islands. Appreciation is given to E.D.

Edwards (ANIC) and C.G. Miller for allowing access to specimens in their

care. This paper partially fulfils the requirements for a Master of

Australian Entomologist, 2015, 42 (4) 263

Philosophy degree undertaken by the author at The University of Queensland,

Brisbane.

References

BASCOMBE, M.J., JOHNSTON, G. and BASCOMBE, F.S. 1999, The butterflies of Hong

Kong. Academic Press, San Diego, California; xiii + 422 pp.

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

CSIRO Publishing, Collingwood; xxvii + 976 pp.

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

Angus and Robertson, Sydney; xiv + 682 pp.

CORBET, A.S. and PENDLEBURY, H.M. 1992. The butterflies of the Malay Peninsula. 4th

edition, revised by J.N. Eliot. Malayan Nature Society, Kuala Lumpur; 578 pp.

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.

EVANS, W.H. 1949. A catalogue of the Hesperiidae from Europe, Asia and Australia in the

British Museum. Trustees of the British Museum (Natural History), London; xix + 502 pp, 53

pls.

JONES, D.L. and GRAY, B. 1977. Australian climbing plants. A.H. and A.W. Reed, Sydney;

166 pp.

PARSONS, MJ. 1991. Butterflies of the Bululo-Wau Valley. Handbook Wau Ecological

Institute. No 12: 280 pp, 25 pls, 23 figs.

PARSONS, MJ. 1998. The butterflies of Papua New Guinea: their systematics and biology.

Academic Press, London; xvi + 736 pp, 162 pls.

TENNENT, W J. 2002. Butterflies of the Solomon Islands: systematics and biogeography. Storm

Entomological Publications, Dereham, Norfolk; 413 pp, 90 pls.

TORRES STRAIT REGIONAL AUTHORITY. 2013. Profile for management of the habitats

and related ecological and cultural resource values of Dauan Island, January 2013. Prepared by

3D Environmental for Torres Strait Regional Authority Land & Sea Management Unit.

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

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

WILLIAMS, K.A.W. 1979. Native plants Queensland. Vol 1. Cranbook Press, Brisbane; 288 pp.

264 Australian Entomologist, 2015, 42 (4)

BOOK NOTICE

Tropical Queensland Wildlife from Dusk to Dawn, by Buck Richardson.

Hardbound; 348 pp; ISBN: 9780957729018; $65 (including postage within

Australia). Available from: Attp://vww.leapfrogoz.com.au/LeapFrogOz/

Tropical Queensland Wildlife from Dusk to Dawn Science and Art.html

Tropical Queensland

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Buck Richardson

This sumptuous coffee-table-sized book is devoted primarily to photographs

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from beetles, bugs and katydids to snakes, frogs and mammals, all taken at

night of living subjects and properly identified by leading authorities.

Interspersed among the photographs are numerous examples of computer

artwork of these amazing creatures, plus a concluding critique of the

language used by scientists in discussing evolution and its products.

For further details see the News Bulletin of the Entomological Society of

Queensland, Volume 43, Part 5, page 89 (August 2015).

THE

AUSTRALIAN

ENTOMOLOGIST

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2015

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THE AUSTRALIAN ENTOMOLOGIST

Contents: Volume 42, 2015

BRABY, M.F.

First record of Petrelaea tombugensis (Róber) (Lepidoptera: Lycaenidae) from

Western Australia

A further record of Danaus chrysippus cratippus (C. Felder, 1860) (Lepidoptera:

Nymphalidae: Danainae) from the Northern Territory, Australia

BROWN, S.S., MEYER, C.E. and WEIR, R.P.

The first records of Anthene seltuttus seltuttus (Rober, 1886) and Anthene lycaenoides

sutrana (Fruhstorfer, 1916) (Lepidoptera: Lycaenidae: Polyommatini) from northern

Torres Strait, Queensland, Australia

BROWN, S.S., MEYER, C.F., WEIR, R.P. and MILLER, C.G.

Tagiades nestus korela Mabille, 1891 (Lepidoptera: Hesperiidae: Pyrginae) from

Mer Island, Torres Strait, Queensland

CLAYTON, J.

The Lophocoleus group of genera (Lepidoptera: Erebidae: Herminiinae) in

Fiji, with the description of a new genus and species

HALES, D.F., FOOTIT, R.G. and MAW, E.

Sexual morphs of Aphis acaenovinae Eastop, 1961 (Hemiptera: Aphididae:

Aphidinae)

HANCOCK, D.L.

A new subgenus for six Indo-Australian species of Bactrocera Macquart (Diptera:

Tephritidae: Dacinae) and subgeneric transfer of four other species

A review of the tree, fig and fruit-infesting flies of the Aethiothemara, Diarrhegma,

Dirioxa and Themaroides groups of genera (Diptera: Tephritidae: Acanthonevrini)

HANCOCK, D.L. and DREW, R.A.I.

A review of the Indo-Australian subgenus Parazeugodacus Shiraki of Bactrocera

Macquart (Diptera: Tephritidae: Dacinae)

HANCOCK, D.L. and WHITMORE, D.

A new species of Euphranta Loew (Diptera: Tephritidae: Adramini) from Sulawesi

HOFFMANN, B.D.

A new species of Polyrhachis (Hagiomyrma) Wheeler from the Northern

Territory, Australia (Hymenoptera: Formicidae: Formicinae)

LACHLAN, R.B.

Additional polymorhism in females of Hypolimnas bolina pallescens (Butler)

(Lepidoptera: Nymphalidae) from the islands of Taveuni and Viti Levu, Fiji

Polymorphic form pallescens females of Hypolimnas bolina (Linnaeus)

(Lepidoptera: Nymphalidae) from Tonga

LANE, D.A. and MOULDS, M.S.

An annotated list of hawkmoths and butterflies (Lepidoptera) from Hammond

Island, Torres Strait

LAMBKIN, T.A.

Telicota sp. nr kezia kezia Evans (Lepidoptera: Hesperiidae: Hesperiinae) from

Dauan Island, Torres Strait, Queensland

LAMBKIN, T.A. and JOHNSON, S.J.

Cethosia cydippe damasippe C. & R. Felder, 1867 (Lepidoptera: Nymphalidae:

Heliconiinae) in Torres Strait, Queensland, Australia

MEYER, C.E., WEIR, R.P. and BROWN, S.S.

A new subspecies of Hasora hurama (Butler, 1870) (Lepidoptera: Hesperiidae:

Coeliadinae) from the Northern Terrirory, Australia

iii

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253

129

107

149

189

257

133

MILLER, C.G. and JOHNSON, S.J.

Further records of Delias lara (Boisduval, 1836) (Lepidoptera: Pieridae) from

Australia 35

MONTEITH, G.B.

Two new records of Australian Triatoma Laporte (Hemiptera: Reduviidae: Triatominae) 105

MÜLLER, C.J.

The life history of Candalides insanea Müller, 2013 (Lepidoptera: Lycaenidae) and

description of the female 45

The female of Deudorix cleora Miller & Miller, 1986 (Lepidoptera: Lycaenidae) 59

NIELSEN, J.E.

A range extension for Eurema alitha (C. & R. Felder) (Lepidoptera: Pieridae) in

Australia, with notes on the migratory behaviour of Eurema species in south-east

Queensland 15

ORR, A.G.

Additional records of predation on butterflies by breeding Rainbowbirds (Merops

ornatus: Meropidae) 38

PIERCE, F. and WALKER, G.

Photographic record of female A//ora major Rothschild (Lepidoptera: Hesperiidae) at

Iron Range National Park 13

POPPLE, L.W.

A new species of Toxala Moulds (Cicadidae: Cicadettinae: Cicadettini) from north

Queensland, with notes on its calling song and comparisons with a congeneric species 77

RICHTER, R.

Rediscovery of the rare coccinellid Micraspis flavovittata (Crotch, 1874) in western

Victoria (Coleoptera: Coccinellidae) 73

SANDS, D.P.A.

Review of Australian Philiris Rober (Lepidoptera: Lycaenidae), with notes on

variation and descriptions of two new subspecies from Cape York Peninsula,

Queensland 219

SANDS, D.P.A. and SANDS, M.C.

Review of variation in Acrodipsas cuprea (Sands, 1965) and A. aurata Sands, 1997

(Lepidoptera: Lycaenidae), with descriptions of a new subspecies of A. cuprea and

a new species of Acrodipsas Sands from inland southern Queensland 197

STOLARSKI, A.M.P. and MEYER, C.E.

Variability in the adult colour forms of Theclinesthes albocincta (Waterhouse, 1903)

(Lepidoptera: Lycaenidae: Polyommatinae) from coastal South Australia 187

VALENTINE, P.S.

The contribution of Stephen James Johnson to the study of Australian butterflies 161

MISCELLANEOUS NOTES 148

NEW PUBLICATIONS 160

BOOK NOTICES 256, 264

Publication dates: Part 1 (pp 1-52) 28 February 2015

Part 2 (pp 53-104) 15 June 2015

Part 3 (pp 105-160) 2] September 2015

Part 4 (pp 161-264) 14 December 2015

THE AUSTRALIAN

Entomologist

Volume 42, Part 4, 14 December 2015

BRABY, M.F.

A further record of Danaus chrysippus cratippus (C. Felder, 1860) (Lepidoptera:

Nymphalidac: Danainac) from the Northern Territory, Australia

LACHLAN, R.B.

Polymorphic form pallescens females of Hypolimnas bolina (Linnacus) (Lepidoptera:

Nymphalidae) from Tonga

LAMBKIN, TA.

Telicota sp. nr kezia kezia Evans, 1949 (Lepidoptera: Hesperiidae: Hesperiinac) from

Dauan Island, Torres Strait, Queensland

SANDS, D.P.A.

Review of Australian Philiris Rober (Lepidoptera: Lycacnidac), with notes on variation

and descriptions of two new subspecies from Cape York Peninsula

SANDS, D.P.A. and SANDS, M.C.

Review of variation in Acrodipsas cuprea (Sands, 1965) and A. aurata Sands, 1997

STOLARSKI, A.M.P. and MEYER, C.E.

Variability in the adult colour forms of Tbeclinestbes albocincta (Waterhouse, 1903)

(Lepidoptera: Lycaenidae: Polyommatinac) from coastal South Australia

VALENTINE, P.S.

The contribution of Stephen James Johnson to the study of Australian butterflies

BOOK NOTICES

ISSN 1320 6133