Research Article |
Corresponding author: Royce T. Cumming ( roycecumming@gmail.com ) Academic editor: Marco Gottardo
© 2020 Royce T. Cumming, Stephane Le Tirant, Sierra N. Teemsma, Frank H. Hennemann, Luc Willemse, Thies H. Büscher.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Cumming RT, Le Tirant S, Teemsma SN, Hennemann FH, Willemse L, Büscher TH (2020) Lost lovers linked at long last: elusive female Nanophyllium mystery solved after a century of being placed in a different genus (Phasmatodea, Phylliidae). ZooKeys 969: 43-84. https://doi.org/10.3897/zookeys.969.56214
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After successful laboratory rearing of both males and females from a single clutch of eggs, the genus Nanophyllium Redtenbacher, 1906 (described only from males) and the frondosum species group within Phyllium (Pulchriphyllium) Griffini, 1898 (described only from females) are found to be the opposite sexes of the same genus. This rearing observation finally elucidates the relationship of these two small body sized leaf insect groups which, for more than a century, have never been linked before. This paper synonymizes the frondosum species group with Nanophyllium Redtenbacher, 1906 in order to create a singular and clearly defined taxonomic group. Five species are transferred from the Phyllium (Pulchriphyllium) frondosum species group and create the following new combinations: Nanophyllium asekiense (Größer, 2002), comb. nov.; Nanophyllium chitoniscoides (Größer, 1992), comb. nov.; Nanophyllium frondosum (Redtenbacher, 1906), comb. nov.; Nanophyllium keyicum (Karny, 1914), comb. nov.; Nanophyllium suzukii (Größer, 2008), comb. nov. The only taxon from this species group not transferred from the frondosum species group to Nanophyllium is Phyllium (Pulchriphyllium) groesseri Zompro, 1998. Based on protibial exterior lobes, this species belongs in the schultzei species group as described in
Nano-leaf insect, Phasmida, Phylliinae, Phyllium, sexual dimorphism, taxonomy, walking leaf, West Papua
The leaf insects (Phylliidae) represent a lineage within the stick insects (Phasmatodea), which are popular as pets and within insect collections due to their impressive leaf-like camouflage. The Phylliidae have an impressive array of morphological adaptations to mimic different types of leaves. Present in many species is the ability for various color forms which allow them to look like living, dying, or dead leaves (Fig.
Included within the Phylliidae are the Nanophyllium Redtenbacher, 1906 a group of small bodied species which for more than 100 years have only been known from rarely collected male specimens. Throughout a majority of those years only the single type species Nanophyllium pygmaeum Redtenbacher, 1906 was recognized. It was not until the last couple of decades that a majority of new Nanophyllium species began to be described (
As it so happens, the breakthrough was not due to the exhaustive review of museum specimens, but instead came as a surprise at the Montreal Insectarium, Quebec, Canada. In April 2018 the Insectarium received eggs of Phyllium (Pulchriphyllium) asekiense Größer, 2002 from Morobe Province, Papua New Guinea with the hopes of raising this beautifully variable species in captivity (Fig.
This pairing of the frondosum species group with the Nanophyllium morphologically was not surprising given numerous morphological similarities between the taxonomic groups (discussed below). From our extensive review of museum specimens, we found males ranging in length from ~ 27 to 40 mm and females ranging from ~ 46 to 71 mm, a difference which is not outside of the norm with the male to female ratio seen throughout the family.
The holotype specimen of Nanophyllium miyashitai sp. nov. was loaned to the Montreal Insectarium (Stéphane Le Tirant, collection manager) from the extensive collection of Tetsuo Miyashita, Japan. See the abbreviations section below for a full list of collections (both institutional and large private) in which relevant material was recovered.
Photos of specimens that were loaned to/held within the Montreal Insectarium were taken by René Limoges of using a Nikon D810 DSLR camera with Nikon Micro-Nikkor 200 mm f/4 lens on Manfrotto 454 micrometric positioning sliding plate. Lighting was provided by two Nikon SB-25 flash units with a Cameron Digital diffusion photo box. Adobe Photoshop Elements 13 was used as post processing software. The Nanophyllium miyashitai sp. nov. holotype specimen is deposited in the Montreal Insectarium (Quebec, Canada) type collection.
Photographs of the female Nanophyllium chitoniscoides (Größer, 1992) comb. nov. (Fig.
The photograph of the female holotype Nanophyllium suzukii (Größer, 2008) comb. nov. (Fig.
Photographs of the types of Nanophyllium daphne sp. nov. (Fig.
All other photographs were taken by unknown photography equipment/by simple camera phone images, and where photo credit/equipment is known it is stated within the figure caption. Egg orientation terminology follows
The following institutional abbreviations are used:
IMQC Insectarium de Montréal, Montréal, Québec, Canada
RMNH Naturalis Biodiversity Center, Leiden, Netherlands
SMTD Staatliches Museum für Tierkunde, Dresden, Germany
Coll FH Private collection of Frank H. Hennemann, Bad Homburg, Germany
Coll RC Private collection of Royce T. Cumming, California, U.S.A.
Coll SLT Private collection of Stéphane Le Tirant, Montreal, Canada
In January 2018, Stéphane Le Tirant of the Montreal Insectarium applied for a live insect import permit from the
Of these five nymphs, two nymphs refuse to eat any of the host plants offered entirely and died within a few days. The other three nymphs thankfully accepted bramble and during the first few instars all three individuals looked very similar in morphology (Fig.
The female lived as an adult for about nine months before dying, producing a total of 245 eggs of many different colors (Fig.
Despite the failed attempt to bring this species into culture, the successful rearing of both sexes from a batch of eggs from a single female has allowed the much needed definitive evidence to back up the morphological observations that the females of the frondosum species group and the males of Nanophyllium are simply one and the same. Therefore, the frondosum species group of Phyllium (Pulchriphyllium) is here transferred to the genus Nanophyllium (except for Ph. groesseri, which instead belongs to the Phyllium (Pulchriphyllium) schultzei species group based in the protibial lobes and is therefore left within the Phyllium (Pulchriphyllium) subgenus). The genus Nanophyllium is here revised with most species illustrated and a general description for the male, female, and egg morphology after more than 100 years is now summarized together.
Type species. Nanophyllium pygmaeum Redtenbacher, 1906, by monotypy.
Distribution. East of Weber’s biogeographic line of faunal balance (Gressitt, 1982), primarily on the island of New Guinea and surrounding islands, as well as on the northern tip of Australia (Fig.
pygmaeum species group
Nanophyllium adisi Zompro & Größer, 2003 (Male)
Nanophyllium asekiense (Größer, 2002) comb. nov. (Female, Egg, Male)
Nanophyllium australianum Cumming, Le Tirant, & Teemsma, 2018 (Male)
Nanophyllium chitoniscoides (Größer, 1992) comb. nov. (Female, Egg)
Nanophyllium frondosum (Redtenbacher, 1906) comb. nov. (Female)
Nanophyllium hasenpuschi Brock & Größer, 2008 (Male)
Nanophyllium keyicum (Karny, 1914) comb. nov. (Female, Egg)
Nanophyllium pygmaeum Redtenbacher, 1906 (Male)
Nanophyllium rentzi Brock & Größer, 2008 (Male)
Nanophyllium suzukii (Größer, 2008) comb. nov. (Female, Egg)
Nanophyllium daphne Cumming, Willemse, Le Tirant, Teemsma, Hennemann and Büscher, sp. nov. (Female)
stellae species group
Nanophyllium stellae Cumming, 2016 (Male)
Nanophyllium larssoni Cumming, 2017 (Male)
Nanophyllium miyashitai Cumming, Le Tirant, Teemsma, Hennemann, Willemse and Büscher, sp. nov. (Male)
1. Antennae: consisting of nine or ten segments (Fig.
2. Posteriormedial tubercle of the head capsule, split into two points (Fig.
Head capsule posteriomedial tubercles compared between select genera/species A Nanophyllium species female
3. Thorax: mesopleurae on their anterior end are notably wider than the prescutum anterior width. Mesopleurae always with prominent tubercles. Prescutum length to width ratio from 1 : 1.8 (more typical) to 1 : 3.4 (in the more extreme width like which is seen in N. suzukii) (Fig.
4. Tegmina: within the Nanophyllium there are three primary venation patterns (Fig.
Three primary venation patterns seen in female Nanophyllium tegmina, dorsal view A Nanophyllium chitoniscoides Coll FH B Nanophyllium suzukii holotype,
a. In the smallest species (individuals ~ 56.0 mm or less), the radial bend occurs before the splitting of the first radial and the radial sector, therefore the radial sector is straight (for example see N. chitoniscoides comb. nov.; Fig.
b. For the larger species, the bend in the radial vein happens on the radial sector after the splitting of the first radial from the radius (Fig.
c. The final venation pattern is found in Nanophyllium keyicum comb. nov., the only species in this genus which has a wide gap between the media and cubitus veins which persists throughout the entire length of the media, this gap is several times wider than a single vein width. This feature is only seen on this species from Kei Island, Indonesia (Fig.
5. Alae: absent.
6. Genitalia: gonapophyses protruding from abdominal segment VIII as long as the terminal abdominal segment, gonapophyses protruding from abdominal segment IX thinner and shorter, not exceeding the terminal abdominal segment, and subgenital plate short and moderately broad with the point just reaching the anterior margin of the terminal abdominal segment (Fig.
The Nanophyllium based on male morphology can be separated into two distinct species groups, the pygmaeum species group (Fig.
Easily observed morphological features which differentiate the species groups are the femoral lobes. The pygmaeum species group has profemoral interior lobes which are angular (Fig.
1. Profemoral interior lobe, in both species groups with most often three small teeth, but occasionally four teeth (rarely), never more than four teeth (Fig.
Front legs and lobes of males of the stellae species group (A–C) and the pygmaeum species group (D–H) A Nanophyllium stellae B Nanophyllium miyashitai sp. nov. C Nanophyllium larssoni D Nanophyllium australianum E Nanophyllium asekiense comb. nov. F Nanophyllium rentzi Coll SLT G Nanophyllium hasenpuschi H Nanophyllium adisi line drawing based on holotype.
2. Posteromedial tubercle of the head capsule split into two points (Fig.
3. Thorax: mesopleurae on their anterior end are moderately wider than the prescutum anterior width (Fig.
4. Tegmina short, length never exceeding the posterior of the metathorax. The venation of the tegmina appears to be rather simple/too sclerotized to identify details of venation. From a review of several different species it appears as though the subcostal vein is lost within an area that is highly sclerotized. The radial vein is moderately present and runs along the edge of the highly sclerotized patch. The medial vein is the most prominent and runs through the center of the tegmina and occasionally has a weak vein splitting from it, but for most observed specimens the medial vein was not prominently branched. The cubitus and first anal are only moderately formed and give stability to the other half of the tegmina and are not notably branched.
5. A unique feature of the alae which appears to be a symplesiomorphy for the Nanophyllium is that the radius splits into the first radial and the radial sector on the distal half of the wing and these two veins run separately to the wing margin without fusing to others (the only other group which also has the radial split on the distal half is the Pulchriphyllium, but their radial sector fuses with the media anterior, media posterior, and the cubitus instead of running by itself to the wing margin; all other leaf insect genera have the radial split happening on the proximal half of the wing). Within the Nanophyllium the alae venation differs between the two species groups.
a. In the pygmaeum species group the key differences are that the media posterior fuses back to the media anterior before reaching the wing margin, and then the fused media runs on its own to the wing margin without fusing with the radial sector (Fig.
Alae wing venation of males for the two species groups, dorsal A Nanophyllium rentzi (pygmaeum species group) B Nanophyllium stellae (stellae species group). Abbreviations used: C (costa); Sc (subcosta); R (radius); R1 (radius 1); Rs (radial sector); M (media); MA (media anterior); MP (media posterior); Cu (cubitus); Cu+1AA (cubitus and first anterior anal); 1AA–7AA (first through seventh anterior anal); 1PA–6PA (first through fifth posterior anal).
b. In the stellae species group the media anterior and the media posterior do not fuse, instead they both run to the wing margin, and the cubitus after splitting from the first anterior anal fuses with the media posterior near the wing margin and then they run fused to the margin as one (Fig.
6. Vomer long and slender with a single apical hook (Fig.
Fortunately, there are several Nanophyllium species with the egg morphology known.
These species include: Nanophyllium asekiense (Figs
1. Cross-section is roundly pentagonal (Fig.
2. Surface is roughly textured, with pitting of various sizes throughout the capsule surface. Surface lacks pinnae. Pits on the capsule surface depending on the species can either have significant depth to them or in other species can have shallow pits.
3. Operculum has distinct pitting surrounding the central gently raised apex (Fig.
4. Micropylar plate is elongate, nearly reaching from end to end of the capsule and with an approximately uniform width throughout except for around the micropylar cup where it is slightly wider. Running parallel along the micropylar plate margin are pits, which vary in number from species to species (Fig.
5. Lateral surface with irregular pitting in no detectable pattern, with some pits very near each other or touching to form wider irregular shapes (Fig.
Figures
INDONESIA
North Maluku Province:
Batjan Island (N. suzukii:
Maluku Province:
Buru Island (Nanophyllium sp. Undetermined female)
Kei Island (Nanophyllium keyicum:
West Papua Province:
Fak Fak (Nanophyllium rentzi:
Aiduma Island (Nanophyllium sp. Undetermined male, Fig.
Papua Province:
Biak Numfor Regency, Biak Island (N. daphne sp. nov., RMNH, Fig.
Nabire Regency (N. hasenpuschi:
Dogiyai Regency, Mapia (Nanophyllium sp. Undetermined female: Coll SLT, Fig.
Mimika Regency, Utakwa River (Nanophyllium sp. Undetermined male, Wollaston Expedition,
Nduga Regency “Hoofdbivak” (N. adisi, Stirling Expedition, SMTD)
Nduga Regency “Kloofbivak” (Third South New Guinea Expedition, Nanophyllium sp. Undetermined nymph,
Central Mamberamo Regency, Kobakma (Nanophyllium sp. Undetermined female,
Jayapura Regency, Cyclops Mts. (N. stellae (Fig.
PAPUA NEW GUINEA
Western Province:
Katau (N. pygmaeum:
Daru Island (Nanophyllium sp. Undetermined female,
Chimbu Province:
Kerowagi District (N. frondosum, Coll RC, Fig.
Eastern Highlands Province:
Mt. Otto (Nanophyllium sp. Undetermined female nymph, Sixth Archbold Expedition,
Buntibasa dist., N. Guinea, Kratke Mts, 4,000–5,000’, February 1933. (F. Shaw Mayer), (Nanophyllium sp. Undetermined male nymph,
Herowana Village (observational record for N. asekiense by Daniel Levitis, USA)
Gulf Province:
Kerema (N. asekiense, Coll RC)
Morobe Province:
Wau (N. miyashitai sp. nov., IMQC, Fig.
Lae (N. asekiense, Coll RC)
Sattelberg (N. frondosum,
Menyama District, Aseki (N. frondosum and N. asekiense, Coll RC; N. chitoniscoides, Miyashita Private collection, and Coll FH Fig.
Watut (N. chitoniscoides, paratype in the Detlef Größer private collection)
Central Province:
Vanama River (Nanophyllium sp. Undetermined female, Coll RC, Fig.
Port Moresby (Nanophyllium sp. Undetermined female,
Northern Province:
Popondetta (Nanophyllium sp. Undetermined female,
Milne Bay Province:
Normanby Island (Nanophyllium sp. Undetermined female)
AUSTRALIA
Queensland:
Iron Range National Park (N. australianum,
Lockhart (N. australianum, Fig.
Female N. asekiense comb. nov. is most often confused with the sympatric species N. frondosum comb. nov., but N. asekiense comb. nov. can be differentiated by the presence of small exterior pro- and meso-tibial spurs, which N. frondosum comb. nov. lacks.
Only two male N. asekiense comb. nov. are known at present, but their morphology is consistent between them for all features except for the abdominal shape (see Fig.
Nanophyllium asekiense (Größer, 2002), comb. nov. males and females, all originating from Papua New Guinea, Morobe Province. Note the variation in the male abdominal shape and the female abdominal, color, and femoral lobe variability A yellow form female, Morobe Province, Aseki, Oiwa Village, July, 2016, Coll RC 16-268 B green form female, Papua New Guinea, Morobe, Aseki (Oiwa), Nov. 2000, Coll SLT C brown form female, Morobe Province, Aseki, Oiwa Village, July, 2016, Coll RC 16-264 D red/brown form female, Papua New Guinea, Morobe, Aseki (Oiwa), Nov. 2000, Coll SLT E serrate abdominal male bred by the Montreal Insectarium, IMQC F spade shaped male bred by the Montreal Insectarium, Coll RC 19-055.
Nanophyllium rentzi males dorsal, note the variation in abdominal shape A holotype in the
Male. Coloration. Each antennal segment with dark brown and tan coloration. The rest of the body and legs are of a yellow-green based color, with variable brown margins. In the two males bred by the Montreal Insectarium the individual with the undulating abdomen has minimal brown markings, with only brown along the leg margins and the abdomen margins (Fig.
Morphology. Head. Head capsule about as long as wide, with a vertex that is lumpy without notable granulation, two posteromedial tubercles are not notably large but are present with a slight furrow between them (Fig.
Holotype : ♂, Papua New Guinea: Morobe Province, Wau: IX, 2000. From the collection of Tetsuo Miyashita, Japan. Deposited in the Montreal Insectarium (Quebec, Canada) type collection.
With the interior lobe of the profemora rounded, not angular, and the mesofemoral interior lobe broad and reaching fully end to end in a rounded triangle, this new species falls within the stellae species group. This is the first species from the stellae species group recorded from Papua New Guinea. The other two species are known from Jayapura, Irian Jaya, Indonesia (very near the border with Papua New Guinea so it is likely they also occur there but to date, we have not confirmed any specimens). This new species can be differentiated from the other two species in the stellae species group by the mesopleurae which have a prominent anterior tubercle followed by four additional small tubercles (only a single anterior tubercle in the other two species) and tegmina that are shorter, only about half the length of the metathorax (almost the length of the metathorax in the other two species).
Like the other members of the Nanophyllium stellae species group, the holotype is a male specimen and the female is unknown. It is expected that the female is larger than other known female Nanophyllium as the stellae species group members are larger than the pygmaeum species group members.
Male. Coloration. Antennae dark brown, a similar brown to that found throughout the head and thorax. The majority of the dorsal aspect throughout the remainder of the body and legs is of a similar lighter brown, but not a light as the stripe of light brown running along the sagittal plane along the head and thorax. Alae and tegmina have a similar dark brown to that found on the antennae. Throughout the ventral surface the coloration is the same as that found on the legs. Granulation on the body is mostly of a lighter brown than the surface it is found on.
Morphology. Head. Head capsule slightly longer than wide, with a vertex that is heavily granulose, which includes the two posteromedian tubercles which are no larger than the surrounding granulation around them (Fig.
[mm]. Length of body (including cerci and head, excluding antennae) 40.0, length/width of head 2.6/2.5, antennae (repaired) 16.4, pronotum 1.8, mesonotum 1.8, length of tegmina 5.9, length of alae 30.5, greatest width of abdomen 8.0, profemora 7.4, mesofemora 6.8, metafemora 7.3, protibiae 3.9, mesotibiae 5.0, metatibiae 6.7.
Currently only known from the type locality of Wau, Morobe Province, Papua New Guinea (Fig.
Patronym. This species is dedicated to Mr. Tetsuo Miyashita (Japan). Miyashita is a major private collector who has amassed one of the largest insect collections in the world. Miyashita and the specimens from his collection have allowed the description of several new beetle taxa over the years with this being the first phasmid described from his collection.
Holotype ♀: Indonesia: Biak. 16/9.54; NNM-Leiden, ex collectie A. Veldhuyzen. In the collection RMNH, Leiden, Netherlands.
This small species (body length of the holotype only 54.0 mm) has several interesting morphological features which differentiate it from other known Nanophyllium females. The tegmina venation places this species most closely aligned to N. chitoniscoides due to the venation pattern having the radial bend occurring before the splitting of the first radial and the radial sector, therefore the radial sector is straight (Fig.
This new species can be differentiated from all other Nanophyllium by several morphological features. First, it is the only species which has exterior profemoral lobes which are obtuse (Fig.
These unique morphological features coupled with the geographic isolation from the mainland makes it unlikely that this female represents the unknown female sex of one of the many species which are only known from males from the mainland (Fig.
Female. Coloration. Presently, only the dried holotype specimen is known, which is fairly well-preserved with only minimal discoloration along the midline due to a lack of gutting. The majority of the body is of a pale light green coloration, with the areas of discoloration (such as the head, thorax, and shafts of the legs) being a pale brown/tan in coloration. Leaf insects are more vibrantly colored in life and it can be assumed that this specimen was a brighter green in life.
Morphology. Head. Head capsule slightly longer than wide, vertex with small granulation throughout the surface and unevenly spaced in no detectable pattern (some right next to each other some with more spacing). The posteromedial tubercle is small, only slightly noticeable and split into two lobes. Frontal convexity stout, not prominently protruding, with a lumpy surface which is marked by numerous pale setae. Antennae. Antennae consisting of nine segments. The terminal segment has a narrower base than segment VIII, instead with a width only about as wide as segments IV or V, and it is about as long as the previous two segments combined length. All segments have setae present; segments I through III have sparse but long pale setae; segments IV through VIII have sparse, stout, tan setae; and the terminal segment IX has dense, stout, dark setae. Compound eyes slender and tightly formed to the head, only reaching across one quarter of the head capsule length. Ocelli absent. Antennal fields approximately the same dimensions as the compound eyes, wider than the base of the first antennomere, and not protruding back farther than the frontal suture. Thorax. Pronotum with anterior margin that is slightly concave and lateral margins that are straight that slightly converge to a broad, slightly convex posterior margin that is about the same width as the anterior rim (Fig.
[mm]. Length of body (including cerci and head, excluding antennae) 54.0, length/width of head 5.7/5.1, antennae 2.9, pronotum 4.0, mesonotum 2.7, length of tegmina 36.0, greatest width of abdomen 28.0, profemora 10.0, mesofemora 8.3, metafemora 9.9, protibiae 5.7, mesotibiae 6.4, metatibiae 8.2.
Noun. Named for the nymph Daphne of Greek mythology who was pursued tirelessly by the god Apollo and was eventually after pleading with her father for a way to escape the relentlessness of Apollo, was turned into a laurel tree. Derived from Greek, Δάφνη.
Currently only known from Biak Island, Papua Province, Indonesia.
One male, observed and collected by Mike Wild (USA/Indonesia) in 2015. Indonesia: Papua Province, Puncak Jaya Regency, Mokndoma, around 2,180 meters elevation.
This individual was observed and photographed by Mike Wild, who notes that despite living in the area for more than 14 years, and actively observing and collecting insects there the entire time, this is the only leaf insect he has ever seen. This species has highly reduced exterior profemoral lobes, which places it morphologically most similar to N. australianum (Fig.
Originally proposed by
One male, observed by Achmad Rian Dietra (Indonesia) in May of 2017. Indonesia: West Papua Province, Kaimana Regency, Aiduma Island.
Discussion. This is only known from photographs of a live individual taken by Achmad Rian Dietra (Indonesia). Based on pro- and mesofemoral lobes being strongly angular and not smoothly arcing from end to end, this individual belongs to the pygmaeum species group. This species group only has males known for six species: N. pygmaeum Redtenbacher, 1906, N. asekiense (Größer, 2002), comb. nov., N. adisi Zompro & Größer, 2003, N. rentzi Brock & Größer, 2008, N. hasenpuschi Brock & Größer, 2008, and N. australianum Cumming, Le Tirant & Teemsma, 2018.
Based on the profemoral exterior lobe that is wider than the shaft width and not larger than the interior lobe, that rules out N. australianum (exterior lobe of profemora same width as shaft width; Fig.
Nanophyllium australianum specimens are exceedingly rare (likely due to a lack of extensive collecting in the area they are found in), with only four collections/observations known to the authors to date and all known from in/near Iron Range National Park of Northern Queensland, Australia (Fig.
The first known record is a subadult female discovered by G. B. Monteith in June 1971 while he was collecting along the edge of the rainforest of Iron Range. Monteith recognized this individual as a second species for Australia and likened the species to a specimen from Popondetta, Papua New Guinea (Fig.
Likely N. australianum subadult females A preserved specimen which was collected by G. B. Monteith in June 1971 on the edge of Iron Range, Queensland (photograph by Susan Wright, collections manager, Queensland Museum) B dorsal, live individual photographed by Chien C. Lee (Malaysia) in July 2014; Lockhart, Queensland C same individual as in B but viewed from the lateral aspect.
The second collection record we are aware of is an early instar nymph collected by G. B. Monteith in February 1976, near Gordon’s Mine Area, Iron Range (see
The third record was the holotype male which was collected as a nymph by D. C. F. Rentz near Mt. Tozer within the Iron Range in December 1986 and which matured to adult in January 1987 (see
The only other individual we are aware of is a subadult female which was photographed by Chien C. Lee in July 2014 in Lockhart, Queensland (Fig.
Nanophyllium sp. Female NYMPH (35 mm): Papua New Guinea: Eastern Highlands District LJBrass, Coll. Sixth Archbold Exped. To Papua New Guinea. No.7, Kotuni, south slopes Mt.Otto, 2200m. Aug. 4–20, 1959. (
Nanophyllium females and their collection data which likely represent the unknown female Nanophyllium rentzi, Nanophyllium hasenpuschi, Nanophyllium pygmaeum, Nanophyllium adisi, or possibly undescribed species, scaled to relative size A Eastern Highlands District LJBrass, Coll., Sixth Archbold Exped. To Papua New Guinea, No. 7, Kotuni, south slopes Mt. Otto, 2200m. Aug.4–20.1959,
This subadult could be N. asekiense or N. frondosum based on the geographic proximity to N. asekiense and N. frondosum known localities and how large it might be if it had reached adulthood. It is likely too large to have been one of the smaller species like N. pygmaeum.
Nanophyllium sp. Female (46.7 mm): Papua New Guinea: Central Province, Along Hiritano Highway, East of Vanapa River crossing. June 21st, 1989. Collected by L. M. Munsey, previously from the collection of Jerri Larsson (California), (Coll RC 16-224). (Fig.
Habitat. From the notes of L. M. Munsey the collector of the two specimens: “Daytime beating in a 1 to 3 acre area of cuttings with few small and large downed trees remaining”.
Nanophyllium sp. Female (54.0 mm): Papua New Guinea: Central Province, 20Km SE Port Moresby “bushes” 26.i.1985 J.W.Ismay. Ex Papua New Guinea DPI-CRIC Konedobu. C.I.E. COLL. A. 17440.
Nanophyllium sp. Female (59.6 mm): Papua New Guinea: Daru, Papua (New Guinea) Mouth of Fly R. VII-1941. Collector RG Wind. Van Dyke Collection. (
These three small adult females possibly represent the unknown female for Nanophyllium pygmaeum as they are the correct size and geographically are from southern Papua New Guinea, with the female from
Nanophyllium sp. Female (70.5 mm): Indonesia: New Guinea: West Irian, Kobakma, North Slope of Central Range N. of Baliem Valley 3500’ October 1971, Robert Mitton, Coll., Presented by H. Vannoy Davis, C.A.S. Accession. (
Based on the size and morphology this female from Kobakma is similar to N. frondosum females which are known from other distant localities in Papua New Guinea. Unfortunately, the holotype N. frondosum has no other locality information other than “Neu-Guinea” (Redtenbacher, 1906) and therefore we cannot determine if N. frondosum is a wide-ranging species or if there is a complex of N. frondosum-like species. Hopefully future molecular comparison of freshly collected material can reveal the extent of N. frondosum.
An additional possibility is that this female could be the opposite sex of Nanophyllium adisi, which is known from nearby this collection location from “Hoofdbivak, 250 m Datum IX” from the Stirling Expedition of 1926 (present day Indonesia: Papua Province, Nduga Regency). This possibility is in our opinion unlikely as these localities are separated by the expansive Maoke Mountains which are most likely a geographic barrier between these populations.
Nanophyllium sp. Female (75.0 mm): Indonesia, West Papua, Mapia, V.1999 (Coll SLT) (Fig.
This female is from very near the type locality of N. hasenpuschi and could possibly represent the unknown female. Hopefully the holotype N. hasenpuschi can be sequenced one day to be compared and possibly matched with this female. Morphologically this female is similar to N. asekiense as it has slight exterior tibial lobes and similar prominent serration of the femoral lobes.
Figures
Phylliidae arose according to
1 | Profemoral interior lobes which are rounded without a sharp angle; mesofemoral interior lobes which are a large rounded triangle, reaching from end to end without prominent spination; (alae) the media anterior and the media posterior do no fuse, instead they both run to the wing margin, and the cubitus after splitting from the first anterior anal fuses with the media posterior near the wing margin and then they run fused to the margin as one; (stellae species group) | 1 |
– | Profemoral interior lobe angular; mesofemoral interior lobes which do not reach from end to end of the shaft and have distinct serrate teeth; (alae) the media posterior fuses back to the media anterior before reaching the wing margin, and then the fused media runs on its own to the wing margin without fusing with the radial sector; (pygmaeum species group) | 4 |
2 | Mesopleurae with a single anterior tubercle, remainder lacking tubercles; tegmina longer (almost the length of the metathorax) | 3 |
– | Mesopleurae with a prominent anterior tubercle followed by four additional small tubercles; tegmina shorter (only about half the length of the metathorax) | N. miyashitai sp. nov. |
3 | Exterior profemoral lobe smoothly rounded with an obtuse angle; abdominal segments with smooth edges creating a clean, spade-shaped abdomen | N. stellae |
– | Exterior profemoral lobe slightly recurved creating an overall acute angle; abdominal segment V with two large clear spots; segments V–VII each with margins that extend and then contract creating a scalloped edge | N. larssoni |
4 | Alae completely transparent, not fully brown in color or with a brown band along the margin; tegmina transparent | 5 |
– | Alae color either completely brown or with a transparent center and brown margin; tegmina completely brown | 6 |
5 | Profemoral interior lobe notably larger than the exterior lobe | N. asekiense |
– | Profemoral interior lobe equal width to the exterior lobe | N. rentzi |
6 | Exterior profemoral lobe distinct, wider than the width of the profemoral shaft | 7 |
– | Exterior lobe of profemora greatly reduced, not wider than the width of the profemoral shaft | N. australianum |
7 | Exterior profemoral lobe notably tapered on the distal and proximal ends; the interior profemoral lobe can be of the same size as the exterior lobe or larger than the exterior lobe | 8 |
– | Exterior profemoral lobe only notably tapered on the proximal end, with the distal nearly reaching the end of the profemoral shaft; profemoral interior lobe always smaller than the exterior lobe | N. adisi |
8 | Alae almost completely brown, or completely brown in color | N. pygmaeum |
– | Only the alae margin and sclerotized section brown, interior half of the alae transparent | N. hasenpuschi |
1 | Small species (~ 56.0 mm or less); (tegmina) the radial bend occurs before the splitting of the first radial and the radial sector, therefore the radial sector is straight; the radius and medial crossvein is present on the radial bend at or before the splitting of the first radial | 2 |
– | Larger species (> 56.0 mm); (tegmina) the bend in the radial vein happens on the radial sector after the splitting of the first radial from the radius; the radius and medial crossvein occurs after the splitting of the first radial, instead originating on the radial sector | 3 |
2 | Profemoral exterior lobe broad, with a slight recurve, giving the exterior angle an acute or right angle; mesofemoral interior lobe with the widest portion on the proximal half | N. chitoniscoides comb. nov. |
– | Profemoral exterior lobe narrow, smoothly arcing from end to end with the exterior angle distinctly obtuse; mesofemoral interior lobe with the widest portion on the distal half | N. daphne sp. nov. |
3 | (Tegmina) there is a wide gap between the media and cubitus veins which persists throughout their entire length, this gap is several times wider than a single vein width; profemoral exterior lobe proximal margin is straight, not recurved | N. keyicum comb. nov. |
– | (Tegmina) the media and cubitus veins run side by side throughout the entire length either touching or no farther than a single vein width apart; profemoral exterior lobe proximal margin is recurved, not straight | 4 |
4 | Prescutum width more than two times the length; mesofemoral exterior lobe broad, notably wider than the mesofemoral shaft | N. suzukii comb. nov. |
– | Prescutum width less than two times the length; mesofemoral exterior lobe as wide as or thinner than the mesofemoral shaft | 5 |
5 | No protibial exterior lobes and no mesotibial exterior lobes, exteriors simple | N. frondosum comb. nov. |
– | Distinct protibial exterior lobes and mesotibial exterior lobes, present as small spurs | N. asekiense comb. nov. |
Review of a wide number of institution and private collections as well as the successful rearing by the Montreal Insectarium has revealed the identity of a previously unconfirmed female Nanophyllium. This has allowed us to synonymize the frondosum species group (only known from females) with the Nanophyllium (only known from males) into a single genus.
Unfortunately, due to the striking sexual dimorphism in Nanophyllium, this leaves many females and males with unknown opposite sexes and the possibility that some of the presently described species of either group might simply be the opposite sex of an already known species. Hopefully future collections of fresh material from throughout the region will either allow successful rearing of species to elucidate the unknown sex or allow pairing of sexes on a molecular basis. Additionally, we eagerly await extensive molecular analysis for the Phylliidae as a whole to elucidate the higher taxonomy within the family and the placement of the Nanophyllium.
Mr. Tetsuo Miyashita who recognized the importance of describing this new Nanophyllium species from his personal collection. Alexandre Banko for his collaboration and transportation of the holotype Nanophyllium miyashitai specimen. René Limoges, entomological technician at the Montreal Insectarium for taking many of the photos for this work, as well as for many professional courtesies. Paul Harrison, Mario Bonneau, Thierry Boislard, Thomas Théry, Jennifer De Almeida, and Dominic Ouellette, entomological technicians at the Montreal Insectarium for caring for the live Phylliidae cultures and for taking photos of the live Nanophyllium. Lorraine Bluteau, horticulturist at the Montreal Insectarium for growing the host plants for the breeding laboratory for many years. Maxim Larrivée (director of the Montreal Insectarium) and Michel Saint-Germain (head of research and entomological collections at the Montreal Insectarium) for their support to “Team Phyllies”. Stephan Blanke and Mandy Schröter at the Senckenberg German Entomological Institute Müncheberg (