TAXONOMY AND SPECIES

ARTICLE OF THE ROYAL SOCIETY JOURNAL:

SPECTRALLY TUNED STRUCTURAL AND PIGMENTARY COLORATION OF BIRDWING BUTTERFLY WING SCALES

Bodo D.Wilts, Atsuka Matsushita, Kentaro Arikawa, Doekele G.Stavenga

The colourful wing patterns of butterflies play an important role for enhancing fitness; for instance, by providing camouflage, for interspecific mate recognition, or for aposematic display. Closely related butterfly species can have dramatically different wing patterns. The phenomenon is assumed to be caused by ecological processes with changing conditions, e.g. in the environment, and also by sexual selection. Here, we investigate the birdwing butterflies,Ornithoptera, the largest butterflies of the world, together forming a small genus in the butterfly family Papilionidae. The wings of these butterflies are marked by strongly coloured patches. The colours are caused by specially structured wing scales, which act as a chirped multilayer reflector, but the scales also contain papiliochrome pigments, which act as a spectral filter. The combined structural and pigmentary effects tune the coloration of the wing scales. The tuned colours are presumably important for mate recognition and signalling. By applying electron microscopy, (micro-)spectrophotometry and scatterometry we found that the various mechanisms of scale coloration of the different birdwing species strongly correlate with the taxonomical distribution of Ornithoptera species.

1. Background

The beauty and brilliancy of this insect are indescribable.

— Alfred Russel Wallace on capturing a birdwing butterfly in Indonesia in 1859, as described inThe Malay Archipelago,1869

The coloured wing patterns of butterflies have attracted the intense interest of scientists, artists and collectors alike for more than a century . Butterfly wings are covered by a lattice of wing scales, which have colours generated by two different mechanisms, structural (physical) and pigmentary (chemical). A structural colour results when a tissue is made up of materials with different refractive indices arranged in periodic structures with distances of the order of the wavelengths of the incident light, and a pigmentary colour occurs when irregularly structured matter contains pigment absorbing in a restricted wavelength range. Structural colours are usually connected to directionally reflected light and are called iridescent when they depend on the direction of illumination and viewing . Pigmentary colours are generally less brilliant than structural colours, due to diffuse scattering by the irregularly arranged material components. Often the two coloration mechanisms are combined, i.e. many structural coloured butterfly wing scales contain pigment that tunes the coloration.

Birdwing butterflies, next to Morpho butterflies, are probably the most amazingly coloured living butterflies and thus are often seen as synonymous for beauty. The Australasian genus Ornithoptera, which includes all birdwings, belongs to the subfamily Troidinae of the swallowtails (Papilionidae ), that evolved from the ancestral troidines .

The genus contains 10 species which are known for their extremely large body size (wing span of up to 28 cm), angular wings and bird-like flight. The Ornithoptera genus includes three distinct subgenera:Schoenbergia,Aetheoptera and Ornithoptera. Each of these subgenera differs in its coloration and the ecosystem they inhabit. The striking wing colorations can range over the whole visible spectrum: from blue-green in the Cape York Birdwing,O. urvillianus, to orange-red in Wallace's Golden Birdwing,O. croesus.

Previous studies have demonstrated that the wing scales of papilionid butterflies harbour a unique class of pigments, the papiliochromes. Notably, the yellow-cream wing colour of the Japanese yellow swallowtail,Papilio xuthus, was shown to be caused by papiliochrome II. This pigment has a narrow-band, violet-peaking absorption spectrum and displays a marked green fluorescence when excited by violet-blue light. In the birdwing butterflies, anatomical studies of the wing scales suggested, however, that their beautiful colours are created by photonic structures.

In which way and to what extent the morphology of the birdwing scales determines the spectral and spatial reflection characteristics, or whether pigments play a principal role, has so far remained unresolved. To clarify this, we investigated seven representative examples ofOrnithopteraby applying morphological as well as various optical methods. We thus found that the coloration of this special group of butterflies is due to a unique combination of structural and pigmentary effects, together causing a diffuse optical signal. We discuss the results in light of the evolution and geographical distribution of the butterflies and we also consider the advantages of the unique structural architecture of the wing scales for coloration and display.

2. Material and methods

2.1. Butterflies

Mounted specimens of the birdwing butterfliesOrnithoptera priamus(Linnaeus, 1758),O. urvillianus(Boisduval, 1832),O. croesus(Wallace, 1859) of the subgenusOrnithoptera(Boisduval 1832),O. tithonus(de Haan, 1840),O. goliath(Oberthür, 1888),O. rothschildi(Kenrick, 1911) of the subgenusSchoenbergia(Pagenstecher, 1893), andO. victoriae(Gray, 1850) of the subgenusAethoptera(Rippon, 1890) (1) were purchased from Worldwide Butterflies O victoriae features two different coloured spots on the forewings, one cyan and one green, which are named as two subspecies in the tables and figures.

Figure 1.Investigated ornithopterans and pigment distribution in the different coloured wing scales. The saturation of the colour corresponds to the (relative) amount of pigment scaled to the maximum absorbance observed in all investigated. 

2.2. Photography

The purchased butterflies were photographed with a

Canon EOS 550D digital camera using a ring flash.

Some butterflies were photographed in the collection

of the Western Australian Museum (Perth, Australia)

using a Canon EOS 7D with a Passport II macro

photography system (Dun Inc., Palmyra, VA, USA).

For UV-photographs (electronic supplementary material,

figure S2), the specimens were illuminated with a Wood's

lamp and photographed with a Nikon D70 digital camera

fitted with a UV-transmission filter

(combined Schott glasses UG3 and BG17). Details of the scale lattice on the wing surface were photographed with a Zeiss Axioscope-A1 Pol microscope, applying white-light epi-illumination and using a Point Grey Grasshopper 3 GS3-U3-50S5C-C or a Mueller DCM310 digital camera. For fluorescence pictures, we used the Zeiss Axioscope with 450–490 nm excitation light and a greater than 520 nm barrier filter.

2.3. Spectrophotometry

The absorbance spectra of the wing scales' pigments were measured on single wing scales immersed in a fluid with refractive index 1.55 (Series A; Cargille Labs, Cedar Grove, NJ, USA) with a microspectrophotometer . Reflectance spectra of the intact wing were measured with an integrating sphere connected to an AvaSpec-2048-2 photodiode spectrometer. The light source was a deuterium–halogen lamp (Avantes D(H)-S). A white standard (Avantes WS-2) served as the reference.

2.4. Scanning electron microscopy

The ultrastructure of the wing scales was investigated using a Philips XL30-ESEM or a Tescan Mira 3 LM scanning electron microscope (SEM). Samples were sputtered with palladium or gold to prevent charging effects prior to imaging.

2.5. Transmission electron microscopy

For transmission electron microscopy (TEM) of the scales, wing parts were prefixed in 2% paraformaldehyde and 2.5% glutaraldehyde in 0.1 mol l⁻¹sodium cacodylate buffer (CB, pH 7.3) for approximately 45 min. After dehydrating with a graded series of ethanol and infiltration with propylene oxide, the tissues were embedded in Spurr's resin. The tissues were cut into 50 nm ultrathin sections, double-stained with uranyl acetate and lead citrate and observed using a Hitachi H7650 (Tokyo, Japan) TEM (as outlined).

2.6. Imaging scatterometry

The hemispherical far-field light scattering pattern of single scales was visualized with an imaging scatterometer . The scatterometer is built around an ellipsoidal mirror, which collects light from a full hemisphere around its first focal point, where the sample is positioned. A small piece of magnesium oxide served as a white diffuse reference object. Images were acquired with an Olympus DP-70 camera and were subsequently corrected for geometrical distortions using a MATLAB routine.

3. Results

3.1. Wing coloration and morphology

The dorsal wing surfaces of male birdwing butterflies feature brightly coloured patches on both wings, marked within a jet-black framing that can be superposed with yellow patches on the hindwing.(2,column I) shows the wing patterns of a few typical ornithopterans, where the colour range varies from blue in O. urvillianus, via green-yellow in O. priamus and O. tithonusto orange-red inO. croesus. All animals have strongly coloured cover scales overlapping a dense lattice of jet-black ground scales (2, column II; electronic supplementary material, figure S1). To investigate whether the coloration of the scales is due to structural effects, we used TEM. The cover scales of the different species all appeared to have a very similar architecture. The scales have a unique layer of pointed ridges covering a lumen existing of an extensive multilayer (2,column III). We derived the average layer distance by performing a fast-Fourier transform (FFT) of the anatomical figures, yielding for instance for the blue and orange wing scales thicknesses of approximately 170 nm and approximately 215 nm, respectively (1). The difference in layer spacing between the differently coloured scales suggests that the scale-specific multilayer properties principally determine the scales' colour. We note here that the multilayers are rather disordered and chirped (i.e. the thickness of adjacent layers varies significantly from layer to layer) when compared with the lumen multilayers of lycaenid butterflies, for instance

Figure 2.Characteristic birdwing butterflies. (a)O. croesus; (b)O. tithonus; (c)O. priamus; (d)O. urvillianus. Column I: picture of the animal; II: scale lattice of the colour patch on the dorsal wing surface; III: TEM cross-section images of structural coloured wing scales; IV: scatterograms of single wing scales. Scale bars for photos in columns I–III are given in rowd; column I, 1 cm; II, 100 µm; III, 2 µm. Column IV: red circles indicate angles 5°, 30°, 60° and 90°.

Table1.

Structural parameters of the wing scales extracted from TEM images.

The average layer distance has been determined by FFT of the TEM

images of figure1 and thus shows the ensemble average over the

full-scale picture.

A classical multilayer reflects light very directionally, and to test if this

was also the case for the birdwing scales we measured the spatial

distribution of the light scattered by single wing scales with an imaging

scatterometer. The obtained scatterograms (2, column IV)

demonstrated that the wing scales of the ornithopterans reflect incident

light somewhat diffusely, arguing against structural coloration and

rather favouring pigmentary coloration. We, therefore, decided to explore the possibility that the scales were coloured by wavelength-selective absorbing pigments.

3.2. Pigmentary coloration

The pigments of papilionids belong to the class of papiliochromes, which can be distinctly fluorescent . To investigate whether the scales contained papiliochrome pigments, we therefore applied fluorescence microscopy. Fig 3 shows results obtained from the wing scales of the Tithonus Birdwing,O. tithonus. Using blue excitation light, the green-yellow coloured wing scales (fi3a) appeared to display a distinct, green emission, strongly suggesting that the scales contained papiliochrome pigemnt (3b). To uncover the pigment's localization, we observed cross-sectioned single wing scales with the fluorescence microscope. The whole wing scale body appeared to be fluorescent (3b), but the fluorescence emerged predominantly from the upper structured layer with structured ridges (R,3c,d), which is a location well suited for a spectral filter.

Figure 3.Fluorescence of the wing scales of the Tithonus Birdwing,O. tithonus. (a) Epi-illumination light micrograph of the scale lattice on the dorsal side of the forewing. (b) Blue-induced fluorescence (excitation 450–490 nm, emission > 520 nm) of the scale pattern shown in (a). (c) Scanning electron micrograph of a cross-sectioned yellow-green wing scale (cross-section indicated by the dashed line in (a)). The scale structure exists of three parts: a ridge array of tall, peaked ridges (R) on top of a disordered multilayer in the scale lumen (M), with below a flat lower lamina (L). (d) Blue-induced green fluorescence of a cross-section of a single scale. All three parts of the scale contain the pigment. Scale bars: (a,b) 100 µm, (c,d) 2 

To identify the pigments of the scales of the various birdwing species and to determine their spectral characteristics, we immersed single scales in refractive index matching fluid (n= 1.55) and measured absorbance spectra with a

microspectrophotometer (MSP,4a). The scales of the studied birdwing butterflies yielded two classes of absorbance spectra.

The scales of O. urvillianuscontained a predominantly UV-absorbing pigment, with peak absorbance at approximately 375 nm, whereas the scales of O. tithonuscontained a blue-absorbing pigment, absorbing maximally at approximately 460 nm. Some scales, like those ofO. croesuscontained a mixture of both pigments (figures1,4a)

Figure 4.Spectral tuning of the scale reflectance. (a) Absorbance spectra of single wing scales immersed in a refractive index liquid. The wing scales contain different compositions of two different absorbing pigments (see also table 1). (b)

Reflectance spectra of the wings measured with an integrating sphere. Most birdwings are green, with maximal reflectance at approximately 550 nm, but the blue-colouredO. urvillianushas a reflectance peaking at approximately

490 nm and the orangeO. croesusreflects maximally at approximately 660 nm.

To clarify the spectral consequences of the absorbing pigments, we measured the reflectance spectra of the wings of the various birdwing butterflies with an integrating sphere (fig 4b). All reflectance spectra featured pronounced bands in the blue to green wavelength ranges, with a peak value between approximately 0.1 (O. urvillianus) and approximately 0.35 (O. croesus). In addition, the reflectance spectra had characteristic troughs in the shorter wavelength range (4b), strikingly corresponding to the pigments' absorption wavelength range (4a). We conclude, therefore, that the pigments act as spectral filters, suppressing the reflectance in the restricted wavelength range of the pigment absorption bands.

4. Discussion

4.1. The colours of birdwing scales

The scales of birdwings contain distinctly fluorescing pigments, which most likely are intimately related to the papiliochrome pigments of other papilionid butterflies. The violet-absorbing pigment papiliochrome II is evidently amply present in the scales ofO. priamusandO. urvillianus(see also electronic supplementary material, figure S3 for a comparison withTroidesssp.), but another, blue-absorbing pigment exists most prominently in the scales ofO. goliathandO. tithonus, for example.

The wing scales of the relatedP. xuthus, representing the tribe Papilionini, contain the same or very similar blue-absorbing pigments, but their morphological structure is rather irregular. The scales hence act more or less as wavelength-independent, diffuse scatterers, and because the scales' pigment suppresses the reflectance only in the short-wavelength range an overall cream-yellow colour remains. The coloration ofP. xuthuswings hence is fully pigmentary.

Quite differently, the reflectance spectra of the birdwing scales feature clear bands, with at long wavelengths a low reflectance. Therefore, the birdwings are principally structural coloured, due to the multilayers in the scale lumen reflecting only in a restricted wavelength range. Furthermore, the structural coloured cover scales are backed by strongly absorbing, melanin-containing ground scales (fig 2; electronic supplementary material, figure S1). This organization evidently serves to optimize colour contrast, e.g. by the ground scales absorbing straylight.

4.2. Tuning of wing coloration

The multilayers of the brightly reflecting wing scales of the birdwing butterflies are covered by tapered ridges. This organization resembles that of the wing scales of the Emerald-patched Cattleheart,Parides sesostris, also a member of the tribe Troidini, which consist of a highly reflective photonic crystal covered by a ‘honeycomb’ layer acting as both a spectral filter and a diffuser/scrambler [4]. Although the exact anatomy differs, the upper layer of the ornithopteran scales clearly also acts as a spectral filter as well as a strongly scattering medium that causes the broad-angled reflections shown in the scatterograms ofig 2, column IV.

The anatomy of the birdwing scales is very different from that in most other butterfly species, especially concerning its thickness of approximately 10 µm .

Most butterflies, which are usually considerably smaller than the birdwings, feature much thinner scales, hinting at an optimization of the weight of the scales with respect to their colour signalling function. Small butterflies often achieve a strong reflection signal by stacking similar coloured scales, e.g. the Cabbage White,Pieris rapae. The high reflectance achieved by the single scales of the birdwings is presumably sufficient to avoid scale stacking.

The combination of pigmentary and structural coloration to achieve unique optical effects is encountered in the scales of several other butterfly species, and also in bird feathers, e.g. in budgerigars and parrots ]. For instance, the wing scales of thenireusgroup papilionids (Papilio: Papilionini) have inconspicuous ridges above a pigmented layer of irregular cylinders, separated from a flat lower lamina, which acts as a thin film reflector . This principle of coloration is also practiced in the scales of other butterfly families .

The scales of birdwings, on the other hand, have very pronounced ridges above a disordered multilayer in the lumen, consisting of up to 14 chitin layers (2,table1). The large stack of somewhat irregularly tilted layers in the scale lumen together with the highly pointed ridge layer causes light reflection into a wide angular space, which further causes significant shape anisotropy causing a slight polarization-dependent structural colour . The additional filtering pigment enhances the strongly chromatic colour. Preliminary finite-difference time-domain (FDTD) simulations using the TEM ultrastructures of fig.2 results not shown) confirm that the chirped multilayers create broadband, violet-to-green-peaking reflectance spectra and that the papiliochrome pigments effectively suppress the reflectance in the short-wavelength range.

4.3. Colour–phylogeny correlation

The three subgenera of the birdwings show different methods of coloration: (i) the trueOrnithopteraall feature similar wing patterns, which however, strongly vary in colour; (ii) theSchoenbergiaare monocoloured with strongly coloured yellow-green scales and angular (i.e. asymmetric-sized) wing features that further only contain the blue-absorbing papiliochrome pigment leading to strongly chromatic signals (fig 4; electronic supplementary material, figure S2); (iii) theAethoptera, though a small clade, feature different structural-coloured spots with species-dependent pigment expressions (fig 5).

Figure 5.Birdwing phylogeny and geographical distribution. The different coloured squares indicate the colour of the wing patches, similar colours indicate a roughly identical photonic structure, and the blue and black dots the appearance of blue-absorbing and UV-absorbing papiliochromes in the wing scales, respectively. PNG, Papua New Guinea

The different characteristics are also recognizable in the

distribution of the butterflies in the Australo-Indomalayan

region. Weber's line acts as a divider for species having

evolved in Asia from those in Wallacea (the Australian

genera) . Also theOrnithoptera and Schoenbergia

appear to belong to the Australian genera, where as

Aethopterais restricted to small areas to the west of

Weber's line. The origin of this diversification is unknown,

but interestingly the colour mechanisms are distinct between different groups (fig 5).

4.4. Biological importance and butterfly vision

Larval stages ofOrnithopteraare monophagous on the poisonous Aristolochiaceae, the main foodplants.  Both male and female adults have yellow coloured wings, presumably for aposematic warning to predators, e.g. spiders or birds.

As butterfly colour vision extends from the UV into the red , the reflection of UV- and yellow-green light by the birdwing wings into a wide angular space creates a powerful visual signal: a broad-angled, bright ‘butterfly purple’. This is quite in contrast with what is generally assumed to hold for structural coloration, namely that the associated iridescence functions in angle-dependent, directional signalling . In the well-known Neotropical Morpho butterflies (Morphinae: Nymphalidae), multilayer-reflectors in the folded scale ridges create a directional blue-coloured iridescence, which is presumably tuned to the blue-receptors of conspecifics.

Sexual dimorphism is strong inOrnithopteraspecies, where males are black combined with bright green, blue, orange or yellow markings (fig 2), while the larger and less colourful females are overall black or dark brownish with white, pale-brown or yellow markings. The sexual dichromatism functions in mate recognition by using the set of spectral photoreceptors [5]. Most likely theTroidesandOrnithoptera, which are in the same tribe (Troidini) in the family of Papilionidae, have similar sets of spectral receptors. Comparing the photoreceptor spectral sensitivities of the Golden Birdwing butterfly,Troides aeacus formosanus, determined by intracellular recordings, with the measured wing reflectance spectra suggests that the high chromatic contrast created by the pigmentary tuning is well discriminated by the set of different photoreceptors, and thus will facilitate mate recognition in a highly complex visual environment (6a).

Figure 6.Photoreceptor spectral sensitivities and coloration of birdwing butterflies. (a) Normalized spectral sensitivities of the most common photoreceptors of the Golden Birdwing butterfly,Troides aeacus

formosanus. (b) Bar graphs of the relative excitation of the set of photo

receptors for the different coloured wing areas of the butterflies of the subgenera

Ornithoptera(I),Schoenbergia(II) andAethoptera(III). The reflectance spectra

of fig.4b were convoluted with a CIE D₆₅standard illuminant spectrum and each

of the photoreceptor spectral sensitivities. The obtained values were subsequently

normalized to the highest excitation per species. Each bar represents the result of

such a calculation for an individual photoreceptor. Note the strong trend within

OrnithopteraandAethopteraspecies, whereas all excitation profiles are basically

identical for the differentSchoenbergiaspecies.

To show this more directly, we calculated the relative excitation of the different

photoreceptors by the wing reflectances of the different butterflies.

We, therefore, convoluted the spectral reflectance of the various wing areas with

the common daylight spectrum and the photoreceptor spectral sensitivities,

using those ofT. aeacusas the closest approximation.

Comparing the different bar graphs (6b) shows that the different wing colours are

well discriminable by the colour vision system of the birdwings and that especially

the butterflies of the subgenusOrnithopteraexcite strongly different photoreceptors

, as expected from the different colours. Interestingly, species whose photoreceptor spectral sensitivities are sex-dependent, like pierid butterflies, tend to have wings of very different colours between sexes. The wings of papilionid species generally do not feature clear sexual dimorphism, and neither the eyes of papilionid butterflies. In the sister genus of the ornithopterans,Troides, there is weak sexual dimorphism as the two sexes have similarly coloured wings, whereas the wing colours of ornithopteran males and females are quite different (see above). It would, therefore, be interesting to study whether and how photoreceptor spectral sensitivities are sexually dimorphic in ornithopterans. However, the protected status of these species will hamper the necessary experimental studies.

5. Conclusion

The showy coloration of birdwings is controlled by two mechanisms: diffuse reflection of incident light by chirped multilayers and spectral filtering by the embedded pigments. Changes in the multilayer dimensions and the pigment absorption spectrum will modify the colour. This is especially interesting as the different birdwing species cover the full range of visible colours, ranging from deep-blue (O. urvillianus) to orange-red (O. croesus) . The characterization of the novel combination of structural and pigmentary coloration in birdwings expands our insight into biophotonic coloration, especially in insects. This may provide biomimetic inspiration, e.g. for adjusting the colour of photonic materials or for improvement of the viewing angle of displays.

Authors' contributions.

B.D.W. and D.G.S. designed the study, all authors performed experiments, collected and analysed data, B.D.W. and D.G.S. wrote the paper with input from all authors.

© 2015 The Author(s)

Published by the Royal Society. All rights reserved.

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SOME BOOKS ABOUT BIRDWING BUTTERFLIES:

1."Outstanding Birdwing butterflies" 

    two volumes, Ornitho Press, 2015 , 1642 pp.           GILLES DESLISLE

    Probably the most correct and complete work about birdwings... but expensive.

2."Ornithopterae"                    vol. 1 :  O. priamus, races et formes                                             B. THIERRY
                                                                         vol. 1  : O. priamus, races et formes
                                                                         vol. 2 : O. aesacus et O. croesus, races et formes

                                                                          vol. 3 : O. goliath, races et formes
                                                                         vol. 4 : O. tithonus, races et formes
                                                                         vol. 5 : O. chimaera et O. rothschildi, races et formes
                                                                         vol. 6 : O. paradisea et O. meridionalis, races et formes
                                                                         vol. 7 : O. victoriae, races et formes
                                                                        vol. 8 : O. alexandrae, formes et variations
                                                                        vol. 9 : Ornithoptera, formes hybrides                             Blurb.com, 2016
 2."Birdwing butterflies of the world"                                                                                         D'ABRERA
                                                                    Melbourne, Lansdowne, 260 pages

3."A new subgenus in the genus Ornithoptera  for the species alexandrae"  

(Lepidoptera, Papilionidae) 2001, pp. 288-321                                                                                   GILLES DESLISLE
4."Outstanding Birdwing butterflies"

2 volumes, Ornitho Press, 2015 , 1642 pp.                                                                                                GILLES DESLISLE

SEEN AS THE MOST COMPLETE WORK ABOUT BIRDWING BUTTERFLIES

5."A monograph of the birdwing butterflies", 6 parts in 2 volumes.          HAUGUM & LOW
Klampenborg, Scandinavian Science Press, 1978/85, 664 pp., also 28 color plates
6."A list of the type-specimens of Ornithoptera (Lepidoptera : Papilionidae) 

British Natural History Museum"                                                                                 THOMAS HOWART
London, Bull.Br.Mus.nat.Hist., 1977

7."Ornithoptera, Schönbergia, Aetheoptera"                                           BELLA VON KNOETGEN
Wangen, M.G.G. Verlag, 1997, xx, 291 pp.,

8."Natural history of birdwing butterflies"                                                                      H. MATSUKA
Tokyo, Matsuka, 2001, 367 pp.
9."Catalogue of birdwing butterflies 1758-2008"                                                TAKASHI OHYA
 published 2009,  288 pp.
10."Fluttering encounters in the amazing Archipelago"                              JAN PASTERNAK
 published 2000, 135 pp.
11."A monograph of the Papilionine tribe Troides of Hübner

Ornithoptera (bird-wing butterflies) of Boisduval",                     2 vol.                H.F. RIPPON
12."Butterflies of the world"                           K. RUMBUCHER , B. VON KNÖTGEN  , O. SCHAEFFLER
Keltern, Goecke & Evers, 1999, with color plates

13

Sumiyoshi, Kaoru.Birdwing butterflies of the genera Trogonoptera and Troides (Lepidoptera: Papilionidae). Osaka: Osaka Museum of Natural History, 1994.

14

New,TimR., and DonaldP.A.Sands.Conservation of the Richmond Birdwing Butterfly in Australia. Springer, 2013.

15

Stidworthy, John.Queen Alexandra's Birdwing: The World's Largest Butterfly (Supersized!). Bearport Publishing, 2007.

16

Haugum,J., and Archibald Montgomery Low.A Monograph of the Birdwing Butterflies, The Systematics of Ornithoptera, Troides and Related Genera: Volume 2. The Genera Trogonoptera, Ripponia & Troides. E. J. Brill, 1985.

17

Sands,DonaldP.A., and TimR.New. "Birdwing Butterflies and Their Conservation Needs." InConservation of the Richmond Birdwing Butterfly in Australia, 1–27. Dordrecht: Springer Netherlands, 2013. http://dx.doi.org/10.1007/978-94-007-7170-3_1.

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STORIES ABOUT BIRDWING BUTTERFLIES:

Wallace discovery of Ornithoptera Croesus

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Wallace wrote in the Malay Archipelago,

pages 257-258 ofO. croesus"

" On taking out of my net and opening those

glorious wings my heart began to beat violently,

the blood rushed to my head, and I felt much

more like fainting than I had done when in

apprehension of immediate death.

I had a headache the rest of the day, so great

was the excitement produced by what appear

to most people a very inadequate cause".

Wallace had in fact written a similar account of the discovery of the new Birdwing on Batchian (now usually spelt Bacan) a decade before, not long after he caught a specimen visiting a yellow flowered Mussaenda shrub in early January 1859. In a letter dated 28 January 1859, written to Samuel Stevens (1817– 1899) his natural history agent, Wallace wrote in this extract "I had determined to leave here about this time, but two circumstances decided to prolong my stay-first, I succeeded at last in taking the magnificent new Ornithoptera and, secondly I obtained positive information here of a second species of Paradisea, apparently more beautiful and curious than the one I obtained.

You perhaps imagine my excitement when, after seeing it only two or three times in three months, I at length took a male Ornithoptera. When I took it out of my net, and opened its gorgeous wings, I was near fainting with delight and excitement than I have ever been in my life ; my heat beat violently, and the blood rushed to my head, leaving a headache for the rest of the day.

The insect surpassed my expectations, being allied to Priamus,perfectly new, distinct, and one of the most gorgeous and unique colour ; it is a fiery golden orange, changing when viewed obliquely, to opaline-yellow and green.

It is, I think, the finest of the Ornithoptera, and consequently the finest butterfly in the world. Besides the colour it differs much in the markings from all the Priamus group. Soon after I took it I sent one of my men to search for it daily, giving him a premium on every specimen, good or bad, he takes, he consequently works hard from morn to dewy eve, and occasionally brings one ; unfortunately several of them are in bad condition.

I also occasionally take the lovely Papilio Telemachus, n.s.

I have sent a small box containing four males, one female, and one young bird of the new Batchian Paradisea, besides an ink red-ticketed private specimen ; six males and five females of the new Ornithoptera, and seven Papilio Telemachus.

Tell Mr Gray and Mr Gould that the Paradisea had better not be described yet, as I am making great exertions to get the second species.

For the Ornithoptera I propose Croesus as a goodname. "

Stevens read the whole of Wallace's letter to the members of the Entomological Society at a meeting held on 6 June 1859 and extracts (some of which are above) were published in theTransactions of the Entomological Society of London1859, 5:70, andThe Zoologist : a monthly journal of natural history, 1859. 17: 6621. Surprisingly John Obadiah Westwood (1805 –1893), who had visited the Leyden Museum, and had seen the holotype ofOrnithoptera tithonusDe Hann 1840, which then, was the only one known to science of that species, expressed his opinion at the Society meeting that the male of theOrnithopteraWallace described in his letter belonged to that species and his females were those ofOrnithoptera victoriaedescribed by G. R. Gray and figured in theProceedings of the Linnean Society, from a specimen take by Mr M'Gillivary on Guadalcanal . It was then suggested by a member that Wallace, before visiting the Far East, had carefully investigated all the available Natural History literature and was aware of De Hann's figure ofO.tithonuson the first plate of theInsects of the Dutch Settlements.
At an Entomological Society meeting in London on the 4 July 1859, it was recorded that "Stevens exhibited both sexes the new Ornithoptera croesus,and both sexes of the beautiful Papilio allied to P. ulysses, for which Wallace proposed the specific name of *telemachus which had arrived since the last meeting".

Stevens told the members at the meeting, "although Wallace's new birdwing was allied to O, tithonus it was by no means identical with that insect, as had been conjectured by Westwood, from the description given in Mr Wallace's letter." *Papilio ulysses telemachus.Thespecific nametelemachushad been used previously for a subspecies ofPapilio ulysseson Woodlark Island by Montrouzier, 1856, and the Bachan butterfly found by Wallace is now known asP. ulyssestelegonusC. & R. Felder, 1860. See this link for Wallace's specimens ofPapilio ulysses.

Stevens who was on very friendly terms Wallace, importance to the British naturalist travelling in the Far East should never be underestimated. Not only did Stevens very efficiently dispose of all of Wallace's specimens that included many duplicates that he did not want for his private collection, he exhibited many of them at the meetings of the Entomological Society, where he mixed with many the elite of the scientific establishment in Britain.

He sent Wallace all his equipment, clothes, boots and many other much needed things, and carried out small, but important tasks for his friend, such as taking his favourite pair of broken spectacles to the makers, and returning them to him in his next parcel. Stevens was an acknowledged expert on his native Coleoptera and Lepidoptera, being an excellent field worker, and thus had much in common with Wallace. Stevens was often the first person to read to Wallace and Bates accounts from the Amazon, and later Wallace reports from the Far East. He other than them, was the first to look on their natural history treasures, and give some of them to the taxonomists who sought to describe them. He knew that both Bates and was Wallace were doing very important scientific work, and with their wish, published many extracts of their travels in theAnnals and Magazine of Natural Historyand later in theZoologist, so many readers were able to follow the excitement and discovery of their journeys, not too long after they happened.

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D'Abrera versus Raymond Straatman

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In 2003 Bernard D' Abrera published his new edition of his 'Birdwing Butterfies of the World' and in this work, the author is very critical of Raymond Straatman, who having passed away in 1987 was unable to counter these accusations and it was left to Jan Pasternak to come to his defense in his CD ROMA Naturalist in Birdwing Paradise. The D' Abrera and Straatman intrigues are legendary among lepidopterists and there was a great rivalry between the two. D' Abrera having produced lavish volumes on lepidoptera and Straatman with his unrivaled field knowledge of butterflies.

Perhaps the best known of these intrigues are the events in 1973 on Kokoda Trail at the Goldie River near Straatman's home at Sogeri. Ray had taken D' Abrera to photograph theO. goliathlarvae, where they arrived late. D' Abrera, who would not use flashlight, complained of poor light conditions and wanted to move the larvae into a more open space to take his photographs. This Ray would not allow because he did not wish to disturb the larvae. Furious and returning the next day on his own, D' Abrera says that he found the larvae gone and the hostplant savagely cut down. Straatman gave Jan Pasternak his version of these events. Ray told Jan that theO. goliathlarvae had devoured and then destroyed theAristolochiavines. Ray had them moved early to a new location and to stop any interference of them by D' Abrera. In spite of D' Abrera opinion of Raymond Straatman he was a highly respected lepidopterist who was very popular with many people.

source: collector -secrets.proboards .com

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BUTTERFLIES WITH BULLET HOLES

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By Lisa Hendry

The holes in this specimen show the passion butterflies have inspired

for centuries.

In 1906, English naturalist and explorer Albert Stewart Meek was

two days deep into the rainforests of New Guinea when he spotted an

enormous butterfly. With little hesitation, he shot it out of the sky.

He wasn't the only collector in the 1800s and early 1900s to capture large flying insects in this way. Obsessed with possessing spectacular specimens, they would sometimes shoot them, using special ammunition to limit the damage caused.

World-record wingspan

Meek presented his exciting find to the renowned zoologist Walter Rothschild, who prepared a scientific description of the butterfly - the first known example of its kind. Rothschild named the new species Queen Alexandra's birdwing,Ornithoptera alexandrae, in honour of Britain’s queen at the time.

The specimen is now housed in the Museum’s Lepidoptera collection along with several other shot birdwings. It has an astounding wingspan of nearly 20 centimetres across.

Today, Queen Alexandra’s birdwing is recognised as the world’s biggest butterfly. The largest specimen, which has a wingspan of 27.3cm, is also cared for by the Museum.

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THE BUTTERFLY EFFECT

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article by Jo Chandler

"Stalking a giant in Papua New Guinea's Ranges"

Sometime in 1906, butterfly hunter Albert Stewart Meek disembarks from an old pearler namedHeklaon the northeast coast of New Guinea. He unloads his provisions and tools of trade: killing bottles with cyanide of potassium for small insects, syringes with acetic acid for larger ones, non-rusting pins for setting his trophies, cork-lined collecting cases. He waves off the boat with instructions to the skipper to return for him in three months.

He has high hopes of claiming discoveries in a wilderness still largely unexplored by Europeans. But things are not going so well.

By his own account —A Naturalist in Cannibal Land—Meek is the swashbuckling, superior Edwardian opportunist from central casting. The son of a naturalist, but with no formal scientific training, he’d travelled from London to Queensland at seventeen to work as a jackaroo, with a sideline in collecting and trading antipodean specimens. At eighteen he had his first commission from Lionel Walter Rothschild, 2nd Baron Rothschild, heir to the Rothschild fortune and a zoologist, to venture into Queensland’s central ranges and bag three pairs of every kind of insect, bird or animal he could find.

Fifteen years later, Meek is being bankrolled by Rothschild to explore the Pacific, capturing exotic butterflies and moths to add to the baron’s natural history collection. It’s the adventurous life he yearned for, and a handsomely profitable one, but it is not without its travails. The islanders he has recruited as labourers have bolted into the bush. He recovers them, punishing the ringleader to persuade them back into reluctant service. By the time they have hauled his kit inland, though, seven are down with disease. It’s fair to say their welfare is not his paramount concern.

But amid this chaos is a glimpse of something beguiling. Setting up camp in the ranges on the Mambare River — likely within sight of the village of Kokoda — he sees an enormous and unfamiliar butterfly. She’s flying so high he brings her down with a shotgun armed with special ammunition. She’s brown with pale yellow markings, her wingspan measuring almost twenty centimetres.

Over the next month, Meek pushes about 130 kilometres inland and high into the formidable Owen Stanley Range — country where, thirty-six years later, the Japanese push down to Port Moresby would be defeated and the Australian legend of Kokoda born. The butterfly eludes him. Meek is “unfortunate enough to lose a couple of boys” — his carriers attacked and murdered. “The collecting was good, but the natives made it practically impossible for me to stay there any longer.” He retreats to Queensland for a spell.

A year later he returns, pulling up in Oro Bay, about thirty kilometres from where he shot the female butterfly. More misfortune: this time he’s laid up himself with terrible sores and raging fever. Then, somewhere near Popondetta, the present-day capital of Oro Province, he stumbles into the butterfly’s garden. He captures males splashed with iridescent turquoise and blue, and more females, some measuring twenty-eight centimetres wingtip to wingtip. Aided by villagers he rewards with mirrors and knives, he finds velvety black caterpillars with ruby spines. He plucks pupae from vine leaves and witnesses — “to my great joy” — a butterfly emerge. The species feeds on an “entirely different vine to other butterflies,” a hitherto unknown species ofAristolochia.

The butterfly specimens are dispatched to Rothschild, who names the subgenus Queen Alexandra’s birdwing,Ornithoptera alexandrae, in honour of the wife of Edward VII.

Meek’s faded trophies, complete with bullet holes, still reside under glass at Rothschild’s estate in Hertfordshire, now part of the British Natural History Museum. In Oro, their elusive kind, the world’s biggest butterfly, has largely vanished, along with much of its nourishing garden.O. alexandrae’s prospects for survival are now entwined with those of the 22,000 people who are the owners of its remnant habitat on the remote Managalas Plateau.

After thirty-three years of negotiations with and between 152 clans, the people of the plateau last year declared a conservation area over their 360,000 hectares of country, putting it out of reach of encroaching loggers, miners and oil palm plantations. It’s only the second, and by far the largest, conservation area in the country. They have resolved to find other ways in which the land might support their livelihoods.

Because such wild places are the last strongholds for so much fast-vanishing biodiversity, and critical to buffering the effects of climate change, we all have a stake in how this crazy-brave gamble turns out. And there are other intriguing dimensions to this story. In an era of seismic corruption and the fracturing of fragile services and infrastructure across Papua New Guinea, the preservation of the Managalas Plateau looks like something almost as elusive asO. alexandrae —good news. As a grassroots initiative, might it signal a turn of the tide in the narrative of external plunder — so much of it unapologetically rapacious; some of it, lately, dressed in finer ambitions but wearing the same old soiled colonial attitudes, blind and deaf to indigenous needs and desires?

After a decade travelling in and out of PNG, collecting too many bleak stories of violence, disease and dysfunction (yes, I too am a plunderer), I’m a little reluctant to chase this apparition for fear it will vanish. Up close, things are sometimes not as they appear from a distance. Yet here I am, stumbling around with my butterfly net on other people’s country.

At Oro Bay, where theHeklahad moored 111 years earlier, we pull off the coast road from Popondetta to pick up cold drinks and provisions at a cavernous tin-shed trade store before heading bush. To have any chance of seeing a Queen Alexandra birdwing butterfly (or QABB, as the locals shorthand it), I will have to travel deeper into the country than Meek likely did. Blessedly, not on foot, though at times it seems like that might be quicker.

The store sits on an inlet, a tide choked with garbage lapping at the shore. Stallholders do steady trade inbuai, betel nut, for chewing and there’s a window offering liquor. Disconcertingly, one of the police escorts we collected back in town — pistol waving in his hand and thongs on his feet — makes a purchase. His young offsider, in a neat shirt and black lace-ups buffed to a mirror finish, waits in the back of our Toyota 4WD ute. My guide — veteran environmental champion, lawyer and local son Damien Ase — works his phone before we drive beyond network coverage.

Wilting in hot, wet shade, I slip the sandwich I have no appetite for to an appreciative hound. Eventually we are on our way.

We pass towering tanks full of crude palm oil awaiting shipment to become cakes and cosmetics around the world. They are fed by the harvest of red nuts from the squat, bristling oil palms that line the road. Palm oil production accounts for more than half of PNG’s agricultural export earnings. The cost had been plain to see through the window on my flight into Popondetta, over the ranges from Port Moresby: wild jungle canopy gobbled up by a meticulously machined green industrial landscape.

Before hitting the road to the plateau, I met Malchus Kajai, chair of the Managalas Conservation Foundation, in Popondetta — from whereO. alexandraehas disappeared. “Because of the oil palm, the feeding grounds have been destroyed,” he explained. “We’re fortunate to have it up there.”

Kajai, who is fifty-seven, has spent more than half his life campaigning to preserve his birthright. He was studying for the Anglican priesthood when he began to worry there was more harm than good coming from development in Oro — the logging, the mining, the explosion of oil palm. His reading of the Bible was that “we have been entrusted to manage the forest. I started to take up the responsibility to speak and protect… The forest which is still virgin contains a lot of fauna, flora, a lot of species that have yet to be discovered.”

He had other concerns too. The fading of culture. Failing schools and health services. Diminishing income from crops like coffee and cocoa — not for lack of effort, but because without functional roads and communications, farmers could not market their produce. There’s been coffee on the plateau for sixty years, but lately it hasn’t been worth picking because of the obstacles getting it to market and the price, now less than $1 a kilo in a fifty-kilo bag. (Meanwhile, in Melbourne, I pay $50 for a kilo of PNG beans when I can find them.)

Such hardships underlie the willingness of many people — including some on the plateau — to sell rights to their country, but Kajai was one who led the resistance. Urbanised landowners were particularly keen for the pay-off, but then they wouldn’t have to live with the consequences. “We had a lot of conflicts, a lot of problems, especially with the elites — educated people who have been to town and were lucky enough to come down and get employment,” he said of the decades wrangling the conservation push.

To understand the obstacles the project had to navigate requires a few insights. First, land in PNG is still largely held under customary ownership. Second, PNG’s population is a diverse patchwork, with over 850 languages, so negotiations over one region may involve a multitude of tongues. (There are five across the Managalas.) Third, land is beyond price in a nation where the state is still so absent that country is all that can be relied on for survival.

These hurdles mean that any number of conservation efforts by marquee conservation outfits and other international non-government organisations — so-called BINGOs — have crashed and burned in the gulf between what distant donors expect and what local people need. Meanwhile, the country’s wilderness is being devoured by logging, much of it illegal. Exports of tropical timber have doubled over the past decade, making PNG the world’s largest exporter of round logs.

The profits from this trade have failed to improve life for most people on the ground, said Kajai. He’s the father of eight grown children. “The system has failed them… The system has failed us. But we have land. Land will not fail you. It is only when you are not creative that you’ll fail yourself.”

There’s bitumen carpeting the routes of the oil palm trucks, but it disappears after the turn-off to Afore village on the Managalas Plateau. Afore is only sixty kilometres away, but it will take around four hours to get there because the road is so bad. A dozen people are piled in the back of our ute. Most are locals who pay K50 (A$21) for the ride — around half the annual income earned by many households on the plateau.

I’m sitting up front with driver Colin Fred, who lives in Afore with his schoolteacher wife and three children. Grinding two sets of gears, playing the pedals like a 4WD virtuoso across the range from full throttle to light staccato, he somehow extricates the overburdened ute from dry ruts and muddy bogs. We plough through a wide river where the bridge was taken out by Cyclone Guba a decade ago. The road is ruined, but then so are pretty much all the routes relied on by the 80 per cent of PNG’s eight million–plus population who live in rural and remote areas. For them, this reality defines all else. Without a functional road you can’t bring teachers and medicines in or send crops and emergency cases out.

We lurch up onto the plateau, a shallow basin that sits between 650 and 850 metres above sea level, encircled by mountain ranges pushing up to over 2000 metres. A breeze flushes out the vehicle’s stifling cabin.

The road winds through stands of rainforest and wild banana trees, rows of coffee and cocoa, swathes of grassland, huts planted on stilts in scrupulously kept gardens of flowers and vegetables. The lushness is fed by rich volcanic soils. A scientist working here twenty years ago on an AusAID-funded research program theorised that the QABB probably gained its monumental size from the vigorous health of the single rare species ofAristolochiavine on which it lays its eggs, and the nectar of the hibiscus and ixora flowers it cruises. Dozens of eruptions had scattered layers of phosphate-rich ash across the plateau, providing “ample nutrients to sustain the caterpillars of such a large butterfly.” And it’s not the only extraordinary creature nurtured by these conditions: a billboard celebrating the conservation project lists half a dozen other flagship species, among them the Raggiana bird of paradise and Doria’s tree kangaroo.

The crucible for the project was a Tok Pisin literacy program back in 1984, enlisting academics and students from the University of PNG — among them aspiring lawyer Damien Ase. A central figure in the PNG conservation movement nationally and locally, Ase hails from a village on the other side of the plateau. “I saw all the destruction that was going on in those places where cash crops like palm oil and cocoa were taking over the forest,” he recalls. “I didn’t want my people to go through that… so I played my part.”

The literacy program evolved into a non-government organisation called Partners with Melanesians, which over the next decade shifted into conservation and development, securing funding from the Rainforest Foundation of Norway, which has supported the project from concept to realisation last year.

The Managalas declaration doesn’t entirely lock up the forests. Rather, it lays out a program of sustainable use of the landscape. Every part of the plateau has been mapped and zoned for one of five purposes: village life, subsistence gardening, larger-scale cropping, hunting grounds and no-go conservation areas. The hope is that this portfolio will generate a mix of activities and attract a variety of players — including researchers, tourists and produce buyers — to support local livelihoods. In Kajai’s vision of the future, farmers will find markets for their organic coffee and spices, village houses will have electric light, schools will plug into the internet, and students will become teachers, tour guides, scientists and health workers employed on the plateau, raising their own families, and sowing an ongoing connection to land and culture.

This model is what American anthropologist and PNG specialist Paige West describes as “conservation-as-development.” Such projects assume that environmental conservation can provide a flow of cash income, and that “development needs, wants and desires, on the part of rural peoples, could be met by the protection of biodiversity on their lands.” West has spent years closely observing the dynamics of such projects, which turn on contracts between villagers and outsiders — maybe a big non-government organisation, maybe research scientists. She has seen how much gets lost in translation: rural people don’t always understand the outsider notion of “conservation” and outsiders don’t always understand what villagers think of when they imagine “development.”

These days, West collaborates with John Aini, a PNG conservationist, to spotlight these failures among specialists, scholars and practitioners, and challenge them to find strategies for “decolonising conservation.” They describe how, time and again, they have seen outsiders come into communities with their own well-formed ambitions but little capacity to understand the links between local livelihoods and healthy biodiversity. Donor-driven projects almost inevitably fail, often leaving behind a volatile mess of failed expectations.

A big part of the problem, according to Vojtech Novotny, a Czech ecologist who has been working in PNG for decades — including running a modest sustainable livelihoods program in the villages around his field sites — is that many people who donate money and effort to saving faraway forests are afflicted by a crippling romanticism. He explored this in a provocatively titled 2010 paper, “Rain Forest Conservation in a Tribal World: Why Forest Dwellers Prefer Loggers to Conservationists,” arguing that “the global machinery of nature conservation remains, regrettably, remarkably inept at presenting indigenous owners of tropical forests with a decent offer in exchange for their continued management and conservation of a substantial amount of the world’s biodiversity.”

Forest people need income and services, and he’s seen little sign of improvement from the BINGOs in delivering on these. “They need a stream of new projects to excite donors, and that doesn’t really work here.”

But he’s “cautiously optimistic” about the prospects for the Managalas Plateau. He and others credit the realisation of the project last year to its organic roots, the engagement of local participants throughout, and the long endurance of both the homegrown Partners with Melanesians and its Norwegian benefactors. None of this guarantees it will deliver what is hoped, but “the advantage is that after thirty years, they have already been through the cycle of hope and disappointment,” says Novotny. They are perhaps well placed to ride it out a bit longer yet.

Malchus Kajai is banking on it. “It’s almost a year now, and people are asking us, ‘When is the service going to be delivered? When are we going to have a coffee mill? When are we going to secure a market for our coffee and our vanilla? When are we going to have better roads?’” People are anxious, he says. “I’m anxious.”

We arrive in Afore at nightfall, pulling up at a pair of spartan shacks that serve as headquarters of the conservation project. We settle in by torchlight, talking late by the cooking fire, eating rice and tinned fish beneath undiluted stars. We sleep under mosquito nets, serenaded by the chorus of forest creatures and a choir of mothers just returned from a church retreat.

I’m up at 5am, slip-sliding down a dark, muddy path to a pit toilet, praying our crew will soon be en route to a stand of forest where — they promise — I will findO. alexandrae. It’s my only shot. I’m booked on a flight out of Popondetta that afternoon, so I’m desperate to get this show on the road. But by the time the fire’s awakened, and pots of tea and rice brewed for breakfast, it’s gone 7am.

In the rear-vision mirror, Afore is otherworldly, an island in the sky encircled by rivers of ethereal morning mist. The track into the forest is barely discernible. I feel myself breathing in to help Colin Fred squeeze the ute between close trees and across too-narrow improvised bridges. Households sprinkled through the bush are surprised by the rare traffic, children chasing and laughing. We pass women hauling up pots of water from streams. The bone-weary gaze of one of them, as I wave, wipes the smile off my face.

Raynold Pasip leans in from the back of the ute to tap my shoulder. “This is my land!” he shouts over the revving engine as we pass through some invisible jungle boundary. Pasip is a wiry elder and a member of the Managalas Conservation Foundation board. We spoke at length the night before about the land, his history with the project, his hopes.

“When I walk to my own bush, I see the bird of paradise. I see a cassowary. I see a wallaby. I see all these things, I feel proud,” he said. “If I want to kill them for my meat for the dinner, it doesn’t cost me money. I can kill some of them, and then come and cook. The feathers for my dancing, [the] tails for makingbilum[bag], and the bones I use [to] make a needle, different things. Our young children are taught, when they go they have a certain time for hunting, and a reason to kill birds and a reason to catch animals. They are not careless in killing birds or cutting trees.”

Pasip talked about the destruction he had witnessed elsewhere in PNG. He describes how that persuaded him to join forces with the elders of 151 other clans, deciding “this plateau should be declared for conservation… so our young generation, the children, will have benefit from their own resource.” They should not be mere labourers on their own land. But without a good road, without airstrips, “people struggle. They carry their own food, cargo, from their shoulders… they walk down to town and they do their marketing.”

He also talked about the peacefulness of the plateau, in the bush and even in the villages, where people were not disturbed by the fighting that has become part of life in so many communities. He spoke of places where you could find birds of paradise or great waterfalls or tiny frogs. The secret sites where, in old times, his people would take their dead. “They didn’t bury them in the ground. They have to go and wrap them around with a mat, and they make a little house on the tree, and they leave them there.”

We pull up at the tiny village of Dareki. The phone network on the plateau hasn’t worked for over a month, so Damien Ase couldn’t send word ahead of our coming. We surprise the man we have come to see — Conwell Nukara, the butterfly whisperer — at home with his small children under a verandah of palm leaves, drying off after a bath in the Pongani River. Briefed on our hurried mission, he leads the way, on foot, deeper into the forest.

We climb over logs and under a tangle of trailing vines including, Nukara points out, theAristolochiafavoured byO. Alexandrae. To us it’s poisonous, he warns. We wade through cloying air and a cool, shallow creek. Pasip and June Toneba, the women’s representative on the conservation board, walk with me. She points out cultivated plots mixed in with the wild growth of the forest: plantings of corn, peanuts, chillies. One of her objectives under the project is to bring in teaching programs for the women. They are such accomplished gardeners, but they struggle to turn this into profitable business, and their children are malnourished because they don’t have the knowledge or resources to feed them a sustaining diet.

Fixing the road is also, for her and other women, truly a matter of life and death. Because when their labours go wrong they can’t get to hospital in Popondetta, many mothers die delivering their babies. Toneba is still mourning the recent death of her own daughter, Imelda, after an asthma attack.

My gaze is on the ground, picking through the labyrinth of tree roots, when Toneba cries out. A flash of movement, a disturbance in the ether. A butterfly the size of a small bird, swooping and dancing around us. “It’s a Queen Alexandra… a QABB!” Toneba declares, and I’m swivelling about with excitement, or perhaps delirium. It circles close, but juggling camera, recorder and notebook I fail to get a fix, and then it’s vanished. When we catch him up, butterfly whisperer Nukara says that we were almost certainly mistaken, tricked by a similar but smaller birdwing.

Finally, we arrive in a clearing where a sign declares: WELCOME TO MISU — QUEEN ALEXANDRA BIRDWING BUTTERFLY FARM: A COMMUNITY INITIATIVE. Nearby is a green shade house, about half the size of a tennis court, which Nukara built last July. Inside are rows of saplings sprouting broad leaves.

He gently turns one over. Stapled underneath is a portion of another leaf he has plucked from the surrounding forest. Stuck to this is the brown and yellow pupa ofO. alexandrae. Nukara turns another leaf, revealing another pupa. The one cocooning a female is as long as his index finger but plumper, the male specimen a little smaller.

Nukara says he roams the forest every morning looking for pupae. When he finds them he brings them into the shade house, where they stay up to eight weeks. This keeps them safe from birds and spiders. A day after they hatch, he opens the door and releases them, he says. So far, he’s waved out twenty-five: fifteen females, ten males.

“This is the largest butterfly we have in the world,” he says, “so that is our pride… It is also endangered, and in the future this will bring people from outside, tourists and other people who are interested, so we can make a small income from that.”

Nukara is wearing a t-shirt with the emblem of New Britain Palm Oil, one of the biggest producers in the country. He’s not being ironic, merely pragmatic. Decades of campaigning by environmentalists against rapacious habitat destruction by the industry, enlisting the orangutan as the poster child of the devastation through Sumatra and Borneo, has put producers under intense pressure to improve their sustainability credentials. In Oro, New Britain Palm Oil has recently announced it will bankroll a captive breeding program for the QABB to try to rescue its precarious population. Its experts have been visiting Nukara’s butterfly farm, talking to him about collaborating on what may be the last chance to save the vanishing species.

Nukara keeps a close watch on his growing flock ofO. alexandrae. He checks in on the pupae and patrols the glade every morning. Butterfly poachers, modern-day Meeks, remain a real threat. Prized QABBs sell for thousands on the black market. There is an argument — including from some of the species’s passionate champions — that its best protection might be to permit landowners to trade a limited quota of specimens: killing butterflies to save them.

What happens, I ask Nukara, when you release these freshly hatched spirits into the wild? Do they flap away? “They hang around for a while,” Nukara says. “They even come back to me… then they fly up.” He saw half a dozen here just this morning. “You should have come early,” he admonishes. “Every morning, I am always filled with joy.”

To my great joy.That’s what A.S. Meek wrote on witnessing his prize emerge from its cocoon. At which point he pulled out his kit and killed it, securing the trophy inside one of his airtight japanned containers.

I scan the enveloping green one last time. Nothing. I shut my eyes. Birdsong, the chirp of lizards, the cacophony of unseen tiny creatures, the fall of fruit on the forest floor. The pulse of fecund energy. The silence of the ancestors perched in their trees. The more disturbing ghost of my own kind, Meek.

I’ve read somewhere thatO. alexandraeflaps through the high canopy with the power and thrust of a bat. It’s not registering on my poorly tuned radar. Which is not to say it isn’t there. •

This essay appears inGriffith Review 63: Writing the Country, edited by Ashley Hay.

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THE QABB (QUEEN ALEXANDRAE BIRDWING BUTTERFLY) BREEDING PROGRAM:

source: Article by New Britain Palm Oil Limited

Conservation of the world’s largest butterfly
‍With a wingspan of 19–30cm, Queen Alexandra’s Birdwing Butterfly (QABB) (Ornithoptera alexandrae) is the largest globally and one of the rarest. Endemic to Papua New Guinea’s Northern Province (also known as Oro Province), it can only be found in two locations: the forest areas of the Managalas Plateau, 1,000 metres above sea level, and the coastal lowlands of the Popondetta Plains. QABB numbers in the Managalas Plateau are relatively healthy and thriving, but the Popondetta Plains are small and patchy.

The QABB is one of only three butterflies listed in Appendix I of the Convention on International Trade in Endangered Species (CITES) and is classified as endangered in the International Union for Conservation of Nature’s Red List of Threatened Species. On a positive note, the Papua New Guinea government declared the Managalas Plateau a conservation area in 2017, and it is now a protected National Park. However, much more needs to be done to ensure the survival of the species.

‍The NBPOL QABB captive breeding programme
‍We are privileged that part of our Higaturu operations is home to the QABB species. Consequently, we have designated certain forest areas in the company’s lease area to preserve its habitat so that the butterfly can thrive in a legally protected environment.

To further strengthen our conservation efforts, with a long-term aim that the QABB will no longer be an endangered species, NBPOL has built and equipped a new laboratory, flight cages and foodplant nurseries. This has been established at our secure residential and operations compound in an attempt to breed the QABB in captivity. Our objective is to eventually release them into previously inhabited areas enriched with additional foodplants.

Some fundamental questions need to be addressed as the breeding programme continues to develop. We know now there is little genetic variation between the sub-populations. However they may still have different ecological requirements, even in their specific food plants. This is vital information for breeding success. Before any releases can be contemplated, past surveys of existing populations need to be consolidated and possibly repeated so that a conservation baseline is established to measure any future success. NBPOL has engaged a birdwing butterfly expert from France to conduct genetic analysis across the two populations. Although the results have been delayed because of the COVID-19 pandemic, we are advancing with other aspects of the trials.

Given that the QABB species is so rare, we have started developing breeding protocols with a closely related but common species – the Common Birdwing Butterfly (Ornithoptera priamus). We will only begin the QABB breeding programme when we have demonstrated two successful Common Birdwing Butterfly egg-to-adult life cycles. NBPOL recorded successful results for the first round of breeding in September 2020. We noted the butterflies were naturally feeding, courting, and mating – a development that bodes well for the second cycle.

‍Inside the lab
‍Construction of the breeding laboratory began in February 2018 and was completed in May 2019. We also built staff accommodation and hired full-time researchers, ground staff, and a Lead Entomologist. The laboratory contains two primary structures: the breeding room and the flight cage. Our breeding room hosts the caterpillars where they are fed until they pupate. We grow the vines that the butterflies feed on in specially designated onsite nurseries. When the pupae are close to emerging as butterflies, we move them to cool dark boxes to enable the adult butterflies to harden their wings for 24-48 hours after emergence. They are then transferred to a flight cage designed to replicate the QABB’s natural environment: a large tunnel-shaped enclosure. We also developed a bespoke ventilation system to recreate the temperature and humidity of the natural forest habitat. Throughout 2020, we worked on improving the flight cage conditions and enhancing our knowledge of butterfly behaviour within the captive rearing environment.

The results are promising, and we plan to build a more extensive flight cage to house more butterflies.

‍Working with communities, organisations and partners
A significant component of this project involves working with local communities, such as the Hombareta Butterfly Conservation Reserve. It is hard to overstate the importance of the QABB to local communities. The Oro provincial flag includes the QABB (representing wildlife) on a green background (for vegetation). QABB sightings in this region are rare. Given the local community’s proximity to the QABB’s natural habitat, we will seek their help in locating and nurturing the QABB. Higaturu Oil Palm Limited has supported this programme by building a guesthouse in one of the villages near a birdwing butterfly farm to house researchers and visitors.

The QABB have long been targeted by poachers and can fetch up to US$5,000–10,000 each on the black market. Unfortunately, some community members have been coerced to help meet this growing demand. NBPOL is actively encouraging local stakeholders to shift their attention towards a conservation focus. NBPOL also works with local community-based organisations, such as Partners with Melanesians (PWM), to help in butterfly conservation efforts in the Managalas Plateau Conservation Area.

NBPOL is grateful for support from the Conservation and Environment Protection Authority (CEPA). They closely tracked the project’s progress and approved our permit to handle and breed the birdwing butterflies in captivity.

The project is financed by Sime Darby Foundation, whose representatives conduct regular visits and assist in the programme’s implementation. The project will run from August 2020 to December 2022, and we hope to raise independent funding to ensure that the project is maintained into the foreseeable future.

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PICTURES:

(*) Most specimens pictures and data comes from my own collection and from friend entomologists, who I bought many butterflies from. Contacts on request. Further sources of information from the Natural History Museum and references below:

• Wikispecies: taxonomy of Ornithoptera, Troides and Trogonoptera. Species an subspecies.
• Boisduval In:M. J. Dumont d'Urville
• Deslisle, G.Straatmania: A replacement name for the subgenusZeuneraDeslisle2001
• Deslisle, G. A new local form ofOrnithoptera (Schoenbergia)ssp.t. tithonusde Haan, 1840
• Deslisle, G.&Sclavo, J.P. New individual forms of the genusOrnithopteraBoisduval, 1832
• Deslisle, G.&Sclavo, J.P. Outstanding Birdwing Butterflies. Volume 1: 1–813.
• Deslisle, G.&Sclavo, J.P. Outstanding Birdwing Butterflies. Volume 2: 814–1642.
• Hanafusa, H. On a new subspecies ofOrnithoptera goliath.
• Haugum, J.&Low, A.M. A monograph of the birdwing butterflies. Vol. 1, part 1. Introduction, Ornithoptera
• (Aetheoptera). Scandinavian Science Press Ltd., Denmark.
• von Knötgen, B. Eine extreme Variante des VogelflüglersOrnithoptera (Ornithoptera) priamus urvillianus
• Schäffler, O. Papilionidae VI, Ornithopthera inBauer & Frankenbach, Butterflies of the World. part 12.
• So, H.&Sato, S. A new subspecies of Ornithoptera paradisea Staudinger, 1893 (Lepidoptera, Papilionidae).Lepidopteral society of Japan.
• Hübner, J.Verzeichniß bekannter Schmettlinge2–11: 17
• Okano,M&J,Ohkurat, 1979: The geographical races of Troides minos (Cramer)
  
• Rumbucher, K.&Schäffler, O. Butterflies of the world 19: Papilionidae X. Troides III. haliphron-species gr.
• Rumbucher, K.&Schäffler, O. Butterflies of the world 21: Papilionidae XI. Troides IV. helena-group.
• --------------------------------------------------------------------------------------

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THE MAN WHO SHOT BUTTERFLIES

John Tennent

Storm Entomological Publications, Oxford

2021

ISBN: 9780954204525

Reviewed by Richard Harrington

I’ve just reached for my Brooke Bond tea card album “Butterflies of the World”, which came out in 1964. The reason for doing this is that I recall from my youth reading that Queen Alexandra’s Birdwing,Ornithoptera alexandrae, the world’s largest butterfly, was first described from a female killed by gun shot. The tea card, it turns out, was indeed the source of that knowledge. The shooter was Albert Stewart Meek (1871–1943), naturalist and explorer, and this 603-page giant of a book is all about him, his exploits and his discoveries.

Meek was employed by Lionel Walter Rothschild to collect anythingAnimaliafor his museum at Tring. This Meek did in south-east Asia, mainly in New Guinea and the Solomon Islands. Much of what he found was previously unknown to science. John Tennent’s preface hints at Meek’s character and strained family relationships, which may partly explain his wish to get away from it all. There is then a series of maps showing the areas explored and an introduction covering, amongst other things, the history and perils of collecting in Meek’s time, his specimens and his letters. Twenty chapters follow, the first dealing with his father and his dodgy exploits, the rest detailing his life chronologically. Chapter 2 takes us up to 1894 when he was 23 and had carried out his first expeditions, in Australia, and had his first collecting agreement with Rothschild. Chapters 3 to 17 each cover a period of one to three of his most productive years, whilst chapters 18 to 20 cover the 28 years leading up to his death. Helpfully, each chapter has a very short précis in green type. These notes helped me to home in on the shooting ofO. alexandrae(ironically an unusually small female), which happened in late 1905 or early 1906 and is covered in Chapter 11. That holotype is now in London’s Natural History Museum and there is a delightful picture of Meek’s grandson and great grandson holding the drawer in which it resides. As it happens, this event seems not to have greatly excited Meek at the time and is described by him in a 48-word afterthought in a letter to Tring’s Karl Jordan. We learn from John Tennent that several of the large butterflies collected a century ago were brought down by dust shot used primarily for shooting small birds without damaging their plumage. How wrong it all feels now! On page 333 there is a summary of what is known about the butterfly, some illegal trading and the problems encountered in trying to protect it.

Of course, the book covers very much more than this eye-catching story – so much more, that attempting to summarise it here would be futile. The one-volume abridged version of Churchill’sA History of the English-Speaking Peoplesis a little over 600 pages in length. John Tennent uses a similar number of pages on one of them, and brings him out of obscurity. The research that went in to all of this is extraordinary. It involved digging out and sifting through articles, letters, obituaries, collectionsetc., and seeking out numerous people all over the world to talk to about the man. The 21 pages of references give a clue as to what must have been involved.

Weighing in at 3.26 kg, this book is hard to read comfortably from your lap. In spite of its huge size, the great majority is text, although there are some magnificent photographs sprinkled throughout (including – Fig, 6.4 – the cover ofAntenna42(2)!).

Appendix A is a useful summary of the relationships between Meek and some key people mentioned in the book (parents, siblings, wives, children, the Eichhorns and the Barnards). Appendix B presents pictures from his book:A Naturalist in Cannibal Land; others from an article written by Frank Fox in 1913, and 81 photographs from Meek’s photograph album, courtesy of Sir David Attenborough who now has it in his library. Appendix C contains an important chronology of Meek-related events from 1871 to 1944.

John Tennent has been able to write this book with such authority not only because of years of research, but also because he has experienced searching for butterflies in the same remote areas. It is a terrifically engaging read. In his foreword, Dick Vane-Wright says “This is the story of a remarkable naturalist and explorer, by a remarkable naturalist and explorer”. I couldn’t put it better.

TRADE AND THREATS ON ORNITHOPTERA CROESUS (LEPIDOPTERA:THE WALLACE’S BIRDWING BUTTERFLY  IN NORTH MALUKU, INDONESIA
Djunijanti Peggie*
Museum Zoologicum Bogoriense, Research Center for Biosystematics and Evolution,
National Research and Innovation Agency (BRIN),
Ornithoptera croesus, Wallace’s Golden Birdwing butterfly, is one of the most highly-demanded
butterflies in the world. Annual trades of this endemic species have been monitored and mechanisms
to control the trades have been developed to ensure the sustainability of the species. This paper aims
to address and clarify the issues on the trade and threat to the species. The methodology involves the
probe into the CITES Trade Database, into the threat to the species based on observation during field
surveys, and into the possible solution. The results showed that there were some discrepancies in the
data provided by the exporter and importer of the species and that the threats to the species are real.
An approach to improve the condition is discussed. The solution to meet the demand for this species
is offered through captive breeding and ranching for sustainable use.
Key words: birdwing butterfly, Ornithoptera croesus, sustainable use, trade, threats
INTRODUCTION
Ornithoptera croesus Wallace, 1859 can be regarded as the most remarkable butterfly
species in North Maluku, Indonesia. The beauty of this magnificent species stunned A.R.
Wallace when he discovered the species, which he expressed with his very famous and
impressive quote: “The beauty and brilliancy of this insect are indescribable, and none but
a naturalist can understand the intense excitement I experienced when I at length captured
it. – I had a headache the rest of the day, so great was the excitement produced by what will
appear to most people a very inadequate cause.” (Wallace, 1869).
Knowledge of birdwing butterflies (Ornithoptera Boisduval, 1832, Troides Hübner,
[1819], and Trogonoptera Rippon, [1890]) in general is based on information from betterknown common birdwings O. priamus (Linnaeus, 1758) (e.g., Straatman, 1969; Heidelberger
& Heppner, 1999; Parsons, 1999; Sands & New, 2013; Kazama et al., 2017) and T. helena
(Linnaeus, 1758) (e.g., Böhm et al., 2018; Nurjannah, 2010; Peggie et al., 2021b). This
species is one of the endangered butterflies in the world (Collins & Morris, 1985; New &
Collins, 1991; Endo & Ueda, 2004; IUCN, 2022), but further knowledge about O. croesus
is very limited and has only been recently reported (Mas’ud et al., 2016, 2020; Peggie et
al., 2021a). The life history has been reported (Igarashi & Fukuda, 2000) and larvae feed on
Aristolochia spp. so indeed it is possible to breed this species. In a study at Gunung Sibela
Nature Reserve, Bacan, this species was found in very few numbers, only a few individuals
seen each sighting time, at four sites of altitude 20 m, 200 m, 400 m, and 800 m asl. (Mas’ud
et al., 2016). Assessment on the breeding approach of O. croesus was recently presented
(Peggie et al., 2021a).
Treubia, 49(2): 85–96, December 2022
This species has different subspecies on Bacan, Halmahera, and Morotai (Peggie et
al., 2005). The subspecies on Bacan is O. croesus croesus Wallace, 1859, the subspecies on
Halmahera is O. croesus lydius (Felder & Felder, 1865), and the subspecies on Morotai is O.
croesus toeantei Parrott & Schmid, 1984. Some even recognized distinct subspecies from the
small islands of Kasiruta and Mandioli.
Despite the limited knowledge about this species, there has been pressure to this species
due to habitat deterioration and demand for trade. Direct collections of this species from the
natural habitats are prohibited due to the protected status under Indonesian regulation (Lampiran
PP7, 1999; Peggie, 2011b; KLHK, 2018).

Trades can be allowed for specimens derived from captivity.

Butterfly captive breeding and ranching can offer solutions to this matter. A butterfly
ranching operation needs to be in-situ at the natural habitat (see Neville, 1993 for the example;
Peggie et al., 2021a). In addition, captive breeding can be operated ex-situ, outside of the
distributional ranges, but this should be monitored closely to avoid any escape to the outside.
Annual trades of this species have been monitored nationally and internationally, as it
is included as one of Appendix II CITES-regulated butterflies.

Stakeholders have developed schemes and mechanisms to control the trades to ensure sustainable usage of the species.

In recent years, O. croesus has been an issue as a candidate of Review of Significant Trade (RST)
by CITES due to the high volume of trades and the differences in the source code usage (CITES,
2022). UNEP-WCMC and CITES Secretariat have asked Indonesia to pay close attention to this
species since the Conference of the Parties CoP17 of CITES in 2016. The Standing Committee
recommended that Indonesia provides a report on the ranching operations of O. croesus to the
Secretariat and confirms that an NDF will be made prior to authorizing the exportation of any
specimens with source code ‘R’.
MATERIALS AND METHODS
To understand the issues on O. croesus, an approach was pursued to look at the trade data
of O. croesus which was taken from CITES Trade Database (CITES, 2022) and followed the
guide to using it (CITES Secretariat and UNEP-WCMC, 2022). Field surveys were conducted
in April 2018 on Bacan and Halmahera, and in September 2019 on Bacan and Morotai, and
threats to the species were observed.
RESULTS
The distribution of this birdwing butterfly is limited only to Bacan, Halmahera, and
Morotai; and is very likely extinct from Ternate. Observation during the field surveys indicated
that this species is rare, with only very few encounters (Peggie et al., in prep.). On the other
hand, the beauty of this butterfly (Fig. 1) has made it very popular among collectors. The
demand for this species and other CITES-regulated and protected species (Fig. 2) should be
fulfilled through a breeding mechanism as no direct collecting is allowed. In most cases, the
specimens are exported as adults, preserved in a folded position, each inside a glassine envelope
(Fig. 3a), and occasionally it is mounted and placed inside a glass-top frame (Fig. 3b).
P
Trade Data of O. croesus
Based on the trade database of O. croesus (CITES, 2022, see Table 1), there were
specimens exported from Indonesia to various countries with source code ‘R’ (ranched
specimens, specimens reared in a controlled environment taken as eggs or juveniles from the
wild), ‘C’ (captive bred specimens, animals bred in captivity), ‘F’ (animals born in captivity as
F1 or subsequent generations that do not fulfill the definition of “bred in captivity”), and even
‘I’ (confiscated or seized specimens). The data presented here are from the year 2011 until 2017.
The search was allowed to include until 2022 but apparently, there was no trade of this species
recorded after 2017 due to the Indonesian policy to voluntarily suspend the trade since 2017.

The quantities reported by importers and those reported by exporters were in most cases
not the same. The purpose of the transactions was mostly reported as T (commercial), except for
the data in 2012 in which 250 bodies imported by the US were listed as E (educational) purpose,
and data in 2014 in which 6 bodies imported by France were listed as P (personal) purpose. The
source code of the transactions of O. croesus varied from 2011 to 2017. In 2011, the source code
R was used in all transactions. In 2012, source code R was used in most transactions, except
for three transactions which were listed as source code C by the exporter country, and two
transactions were listed as confiscated or seized specimens (source code I), of which most were
used for E (educational) purpose, but 14 bodies of confiscated or seized specimens were used
as T (commercial) purpose. In 2013, most transactions were listed as source code R, except 220
bodies were listed as C by the US as the importer. In 2014, Indonesia as the exporter used the
source code F, but importers used the source code R on 7 transactions and used the source code
C on one transaction. In 2015, source code F was used in all transactions.

In 2016, source code F was used for almost all transactions except one which was listed using source code R by Japan as the importer. In 2017, source code F was used in both transactions.
Threats to O. croesus
Surveys in April 2018 and in November 2019 have shown that O. croesus was indeed
difficult to find (Peggie et al., 2021a) and detailed data on the occurrence of this species will be
presented separately (Peggie et al., in prep.). During the surveys, some extractions of trees in
the forests were seen. In some areas, the land use changes took place in the concession forests
(Fig. 4a), meaning that it was done legally. However, at lower elevations at Gunung Sibela
nature reserve, the trees were also cut down (Fig. 4b).

Besides habitat deterioration, the threat to the species is also due to direct capture, possibly
through illegal operations. Direct capture may be avoided by introducing ranching and captive
breeding operations.
Possible Solution through Ranching or Captive Breeding
A breeding facility at Labuha, Bacan and a few potential sites for ranching operations at
Halmahera and Morotai were visited to assess whether the in-situ semi-natural breeding method
meets the criteria for sustainable use of O. croesus (Peggie et al., 2021a). On other islands
in North Maluku, i.e., on Halmahera and Morotai, there were some areas (Fig. 5) with the
potential occurrence of O. croesus. In the past, before the species is protected, certain villagers
had harvested pupae from host plants in forests nearby the villages.
With the protected status of the species, breeders would need a harvest permit issued by
the Forestry office to take parental stocks from the wild. The eggs or caterpillars were placed
on the leaves of the host plants which had been planted to enrich the habitat. Then the parts of
plants that had eggs or caterpillars were covered with a large net sleeve to avoid any predators
and parasites. Every day, the breeder would check if the caterpillars had enough leaves and
would need to move the caterpillars to other branches if needed. This practice is quite common
among breeders to optimize the usage of host plants (Fig. 5). When the caterpillars reach the
pupation stage, they can be moved to the pupal chamber. After they emerge into adults, they can
be harvested and some are released to the wild for restocking to comply with the Indonesian
regulation.

DISCUSSION
This birdwing butterfly species is urgently in need of serious attention. It was suspected to
become extinct from Ternate since the volcanic eruption of Mount Gamalama in 1983 (Peggie
et al., 2005) as confirmed by a field survey in 2009 (Peggie, 2011a). However, detailed data on
the distribution of this species is lacking, thus further research is needed to confirm the IUCN
status. The beauty and the rarity made this species a highly demanded species. Though schemes
and mechanisms to control the trades are available, the implementation needs improvement to
ensure the sustainable use of the species.

There were discrepancies in the quantities reported by importers and exporters at the Trade
Database for O. croesus, the numbers were in most cases not the same. As noted by CITES
Secretariat and UNEP-WCMC (2022), many of the annual reports submitted to the Secretariat
do not clearly state whether the data were derived from the actual number of specimens traded
or from the quantity for which the permits or certificates were issued. The numbers were often
considerably different. The term used to indicate the piece might be different and this has
somewhat added problems to the data, in this data the exporter country used ‘bodies’ but there
were 5 entries in which the importers used the term ‘specimens’ and ‘trophies’. This difference
in the term used might create a discrepancy in the quantities, for example, the data in 2016 of 90
specimens reported by China as the importer was most likely the same as the data of 90 bodies
reported by Indonesia as the exporter, but somewhat appeared as two transactions.
The usage of different source codes (R, F, and C) has created confusion.

Since 2014,
Indonesia has produced individuals with source codes ‘F’ and ‘C’ (no more with code ‘R’). This
was done in conjunction with the suggestion from the CITES authority to use F source code for
specimens derived from captive breeding operations that have not fully met the requirements.
Indeed, the Indonesian management and scientific authorities had checked the facilities and
confirmed the captive breeding operations but due to the lack of traceability of parental stocks,
which are understandable for fast-growing insects, we agreed upon the usage of F as the source
code for those time periods. It was suggested that the breeders should improve their management
including the tracing of parental stocks. However, when the condition of the documentation
and reporting did not improve, and the confusion of the different source code applications
continued, the Indonesian management and scientific authorities decided to voluntarily suspend
the trade of O. croesus in 2017. This decision was based on the consideration to improve the
management of the species, primarily on the confusion about the usage of source codes.
This species has been classified as NT – near threatened of the IUCN status (Böhm, 2018),
based on the conditions of (1) relatively localized threats of logging, (2 likelihood of occurrence
in more than ten locations, (3) possible fragmentation of the distribution of the species, and (4)
continuing habitat decline.

It has been suggested that the primary threat to wildlife is habitat
deterioration. Habitat alterations as seen during the surveys and as reported by Mas’ud et
al. (2016) indicate the real threat to this endemic species and may have affected the spatial
distribution of this species. Obtaining detailed information in regard to the distribution and the
rarity of this species should be a priority and become a major concern to all stakeholders. Smiet
(1982) reported that the commercial logging operations in northern and central Maluku might
put wildlife including O. croesus in declining populations. Data on the status and trends of the
populations of O. croesus, and the impact of the threats is urgently needed.
Data on life history and other aspects of the species can be obtained through captive
breeding operations (Matsuka, 2001; Daniels et al., 2020). Important data such as the
reproductive capacity of this species needs to be revealed and taken into consideration for the
conservation of this species. An effort has been made to take some pupae of this species to the
Butterfly Research Facility in Cibinong but they were not successful in mating at that time.
As for the few villagers who had harvested pupae from host plants in forests nearby
the villages, rearing or captive breeding operation can be encouraged. Incentive needs to be
given to villagers so that they can breed the butterfly legally with source code R or even C for
sustainable use of the species.

Indonesia is required to have an NDF assessment before the exportation of specimens
with source code ‘R’. Meanwhile, Indonesia has not developed an NDF for any species of
birdwing butterflies, and we will follow the recommendation to prepare an NDF and notify the
NDF result to CITES Secretariat before authorizing the exportation of specimens with source
code ‘R’.
Regarding the question of whether there is a mechanism in place to ensure that exports of
ranched butterfly specimens under the source code “R” consist only of specimens collected at a
very early developmental stage (eggs or the first larval stage), I would like to confirm that this
has been the case. Breeders usually planted many host plants to attract female butterflies to lay
eggs on the leaves of the host plants. The eggs will then be covered with a net sleeve (in the case
of ranching) or protected from natural enemies in the butterfly enclosure (in the case of captive
breeding). The hatched caterpillars will feed on the leaves of the host plants, undergo 5 instars,
and pupate. The chrysalis, or the pupae, will then be selected upon emerging to adults, i.e.,
some will be harvested and some others will be kept alive as the parent stocks of subsequent
generations. In the case of O. croesus, it was difficult to find parent stocks in the wild. Therefore,
it is natural to think that breeders would ensure to keep sufficient individuals for parent stocks
and not harvest too many.

Otherwise, they would need to spend extra resources (efforts, time,
money for transportation, etc.) to go to the forests and obtain parent stocks. Considering the
challenges of walking up the river at Gunung Sibela Nature Reserve that we also experienced
during the field surveys (Peggie et al., 2021a), sparing decent numbers of individuals will be a
viable option that the breeders would take. Therefore, it is recommended that an in-situ seminatural butterfly breeding approach can be developed for butterfly conservation and sustainable
use of O. croesus with “R” source code or with “C” source code for the breeding facility inside
a butterfly enclosure (Peggie et al., 2021a).
Domestic and international trading (Soehartono & Mardiastuti, 2002) may add to the
threats. Excessive trades, mostly uncontrolled direct collecting, may lead to a threat to endemic
and rare species like O. croesus. Much better alternatives such as harvesting through captive
breeding and ranching operations should be encouraged.
In conclusion, the trade and threat issues of O. croesus need to be considered thoroughly
based on the viability to implement conservation strategies. The demand for the trade of O.
croesus can be fulfilled through good understanding and cooperation to achieve the expected
goals of sustainable use. An approach such as ranching and captive breeding practices need to
be encouraged and supported, within the boundaries of international trade compliance. More
importantly, the remaining forests on the islands of North Maluku need to be preserved to spare
the future of this magnificent species and many others.
ACKNOWLEDGMENTS
The field surveys were carried out with DIPA funds from KSK CITES 3400.002.051A,
Research Center for Biology - LIPI. I am grateful to KSK CITES and the director. I also thank
the head of BKSDA Maluku and his staff for the access given to enter the areas on Bacan
Island. I thank Sarino, Sih Kahono, and all those who have assisted during the field surveys. I
appreciate insights provided by reviewers to improve an earlier version of the manuscript.

FIRST DESCRIPTION of... Ornithoptera priamus and tithonus

FIRST DESCRIPTION of... Trogonoptera brookiana

FIRST DESCRIPTION of... Ornithoptera victoriae forma gabrieli

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BREEDING PROJECT RICHMONDS BIRDWING - AUSTRALIA

Attention Attention !!!         RichmondBirdwing from 'Urban Birdwing Butterfly Facebook'
Grafton, Gympie Maryborough and Hervey Bay regions.
Today we are announcing an expansion for our plans with the1 in 100 Project .....
To bring the Richmond Birdwing back to the Northern most & Southern Range for this species.

Attached is a rough map/guide of where according to historical records, the Richmond Birdwing Butterfly's range used to be.

In the far North and the far South of the range, the Birdwing is not found. Let's fix that - together.
The 2nd image shows a Tried & True host plant with Birdwings munching upon the leaves.

Let's look at Grafton...
In less than 2-3 years we could have Richmond Birdwings in Grafton gardens and backyards in a self sustaining colony.
With that as a goal....Does anyone know anyone in the Grafton zone who might be interested in participating? Surely we can find 10-20 people or a couple of school teachers who will want to be a part of this re-introduction project?
20 host plants in two - three years (quicker in warmer zones) will be big enough to support a self sustaining colony of these butterflies. Too Easy.
One of the Butterfly Mentors of mine mentioned today that they grow the Tried & True host without issues in temperatures down to -4C (minus 4C). Obviously this means that these host plants will do absolutely great at Grafton & surounding areas.
Fabulous news right !!??
I am not so sure that Super Swift or Vulcan will cope with temperatures that cool - because those sulk a bit at my place in temperatures of 8 to 14C.
But having Tried & True available in gardens seems a logical choice in a water-wise garden in Grafton.
Of course we can still use Slow Poke in Grafton as well. But now we know that we have a great combo with having the two available as host plants in home gardens.
We also have another one or two varieties that are cold tolerant in the pipeline... Not available as yet, but hopefully within the next 12 months they will be available.
I am happy to bring some Richmond Birdwings to some dedicated gardeners in the Grafton area in order to bring the Birdwing back to your area.

Moving North... Up in Gympie, Maryborough & Hervey Bay.
We absolutely have the capacity to increase the host plant availability for Richmond Birdwings to find them.
In this vain, is there anyone you can refer the 1 in 100 Project to, in order to increase the amount of fodder available for Richmond Birdwings by growing host plants in those areas?
Once again, all we need is about 10-20 people or a couple of schools involved, growing a total of 20 or more host plants and in two years, these individual areas could also have Richmond Birdwings flying around again very soon.

Let's do this thing...pass on the message to all and sundry.