aunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
Volume 7 ISSN: 2269 - 6016
Numéro 4 Mars 2019 Dépét légal: Mars 2019
Faunitaxys
Revue de Faunistique, Taxonomie et Systématique
morphologique et moléculaire
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Faunitaxys, 7(4), 2019: 1-25.
Review of the Phyllium Mliger, 1798 of Wallacea, with description of a new subspecies from Morotai Island (Phasmatodea: Phylliidae: Phylliinae)
ROYCE T. CUMMING (1), STEPHANE LE TIRANT (2) & FRANK H. HENNEMANN (3)
(1) Entomology Department, San Diego Natural History Museum, POB 121390, Balboa Park, San Diego, California, United States, 92112-1390. Associate researcher for the Montréal Insectarium, Québec, Canada; H1X 2B2 - phylliidae.walkingleaf@gmail.com
- ZooBank : http://zoobank.org/6CA8501F-10BA-4E07-9BF4-65CFCE4E9E92
(2) Collection manager, Montréal Insectarium, 4581 rue Sherbrooke, Montréal, Québec, Canada, H1X 2B2 - sletirant@ville.montreal.qc.ca - ZooBank : http://zoobank.org/A9391F8A-15D7-4D3B-9E3F-7123BA27EA2E
(3) Reiboldstrasse 11, 67251 Freinsheim, Germany- hennemann@phasmatodea.com
- ZooBank : http://zoobank.org/65 1 FCCFA-271B-48A3-A58E-A30FDC739493
Abstract. — The Phylliidae of Wallacea are here reviewed, with notes on species
distributions. Phyllium (Phyllium) tobeloense GroBer, 2007 is also here recorded as having a new subspecies from Morotai Island, Phyllium (Phyllium) tobeloense
bhaskarai Cumming, Le Tirant, and Hennemann ssp. nov. based largely upon the distinct egg morphology. To conclude a key to all known Phyllium from Wallacea is presented with most features illustrated for ease in identification.
Cumming R. T., Le Tirant S. & Hennemann F. H., 2019. — Review of the Phyllium Illiger, 1798 of Wallacea, with description of a new subspecies from Morotai Island (Phasmatodea: Phylliidae:
ZooBank : http://zoobank.org/CEC3E930-3ESD-4B2A-AE4D-51A198FD9E9E
Keywords:
Phasmatodea ; Morotai ;
Phasmida ; tobeloense ;
Phylliidae ; bhaskarai ;
Phylliini ; description ;
Phyllium ; new subspecies ;
Wallacea ; distribution ; Phylliinae). Faunitaxys, 7(4) : 1-25. Indonesia ; biogeography.
Introduction
Members of the genus Phyllium Illiger, 1798 are well known as “Leaf Insects” or “Walking Leaves” and their still poorly understood systematics, phylogeny, and distributions have been studied more comprehensively during the past ten years. It has been shown that the actual biodiversity of these fascinating insects is far larger than previously suggested and several papers by the authors and various colleagues have described 30 new species of Phyllium from throughout Southeast Asia since 2009 (Brock et al., 2018). The Phylliidae of Wallacea are particularly interesting but have not yet been studied in any detail, with only some of the species described before 2009 covered by Hennemann et al. (2009). In addition, two new species from Wallacea have recently been described from the islands of Peleng (Cumming & Teemsma, 2018) and Lombok (Cumming, Valero & Teemsma, 2018). Examination of extensive fresh material that has recently become available from various islands of Wallacea has revealed interesting facts concerning the synonymies and distributions of the species concerned and are summarized in the present paper. This includes a new subspecies from the Island of Morotai north of Halmahera, Maluku Islands, which is described herein. Still however, many questions concerning the true distributional ranges of some species throughout the very complex Wallacea subregion remain unanswered and the authors are aware of further still undescribed species, that will hopefully sometime become available for scientific recognition in order to clarify the biogeographic and phylogenetic facts and help to complete our knowledge of Wallacean Phylliidae.
Biogeography
Wallacea is a group of several thousand islands of various sizes, which cover a total land area of roughly 338,500 km? located between Sundaland in the west, the Philippines in the north as well as New Guinea and Australia in the east and southeast. Recent studies suggest that also the Philippines need to be considered as part of the Wallacea (e.g. Vallejo, 2011), but for covering the Phylliidae of this particular region Wallacea is here treated in its traditional definition that excludes the Philippines. All islands are located almost entirely within the borders of Indonesia and include the island of Sulawesi, the Maluku Islands (also called Moluccas), the Kai Islands, Banda Islands and the Lesser Sunda Islands (except Bali). The largest of these islands is Sulawesi. The Lesser Sunda Islands attributed to the Wallacea are located south of Sulawesi and include Lombok, Sumba, Sumbawa, Flores and Timor, as well as several hundred smaller intervening islands. The Maluku Islands include several hundred islands in the northern portion of Wallacea, the largest being Halmahera in the north and Seram in the south. The most northern islands of the Maluku Islands are the Sangihe Islands and Talaud Islands, the most southeastern ones are the Kai Islands and Banda Islands, situated west of New Guinea.
Principally, Wallacea is a transition zone between the Oriental region and the Australian region with the fauna showing influences from both these regions (Dickerson, 1928). The western border of Wallacea is represented by Wallace’s line (named by Huxley in 1868), which runs between Borneo and Sulawesi in the north and Lombok and Bali in the south (Fig. 1). The eastern border of Wallacea has been disputed by
2 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
geologists and biogeographists and attempts to draw a definite line have produced a number of different lines (Simpson, 1977). The most accepted of these lines is Lyddeker’s line which follows the Sahul shelf, that runs between Seram and the Banda Islands in the west and New Guinea in the east (Fig. 1). Another well-known and much discussed line is Weber’s line, which roughly runs between Sulawesi and the Maluku Islands (Gressitt, 1982). Mayr (1944) regarded Weber’s line as a line of faunal balance, east of which the Papuan faunistic elements have a numerical superiority over Oriental elements.
Wallacea is regarded as a biodiversity hotspot although the subregion contains fewer known species overall compared to e.g. Sundaland, the Philippines, or the Australian region. However, Wallacea contains a very high percentage of endemic species, i.e. species that are only found in Wallacea or on single islands within this subregion, which is explained by several factors. First of all, the subregion contains a large number of different islands and geographic isolation is one of the major factors which leads to diversification and formation of endemic species. Wallace’s line marks the position of deep ocean straits and separates shallow waters in the west from deep waters in the east. The sea-level dropped by an estimated 120 meters during the last ice-age and while the shallow waters in the west disappeared and formed land connections between the Large Sunda Islands (Sumatra and Java), Borneo, and the remaining portions of Sundaland, there were no such land bridge connections towards Wallacea.
Materials and Methods
Photos of specimens were taken by René Limoges of the Montreal Insectarium (IMQC) using a Nikon D810 DSLR camera with Nikon Micro-Nikkor 200mm 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. Measurements of specimens were made to the nearest 0.1 mm using digital calipers. The Phyllium (Phyllium) tobeloense bhaskarai ssp. nov. holotype and allotype specimens are deposited in the Montreal Insectarium Quebec, Canada (IMQC) type collection. Photographs of specimens within the Frank Hennemann personal collection were taken by himself using a Nikon D7000 camera equipped with a Nikon DX AF-S Micro 40mm lens and a wireless Nikon SU-800 dual speed light system. Background lightning was provided by a 18W 6000K LED panel light plate. The photographs of the female holotype Phyllium (Pulchriphyllium) suzukii GroBer, 2008 was taken by Mandy Schroter under direction of Stephan Blanke at the Senckenberg German Entomological Institute Mtincheberg using a Nikon D7200 digital camera and a Nikon Micro Nikkor 105mm f/2.8 G ED objective. Lightning was from the Yongnuo Digital Speedlight YN 560 IV reflected by the inner surface of a styrofoam box set up around the specimen. A grey card was used for white balance. Composite images with an extended depth of field were created using the software StackShot Macro Rail Package (Cognisys Inc., U.S.A.) and Zerene Stacker (release November 7, 2017; Zerene Systems LLC, U.S.A.).
Abbreviations
— AMSA: Australian Museum, Sydney, NSW / Australia. —BPBM: Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A.
— BYU: Monte L. Bean Life Science Museum, Brigham Young University, Provo, Utah / U.S.A.
—CASC: Califomia Academy of Sciences, San Francisco, California / U.S.A.
—IMQC: Insectarium de Montréal, Montréal, Québec / Canada.
—IRSN: Institut Royal des Sciences Naturelles, Brussels / Belgium.
—LEMQ: Lyman Entomological Museum, McGill University, Québec, Canada. —MNHN: Museum National d'Histoire Naturelle, Paris / France.
— MNHU: Museum fiir Naturkunde der Humboldt-Universitét, Berlin / Germany.
—MZSF: Museum Zoologique, Université de Strasbourg, Strasbourg/ France. —NHMUK: The Natural History Museum, London / United Kingdom. —NHMW: Naturhistorisches Museum Wien, Vienna / Austria.
—NMR: Natuurhistorisch Museum, Rotterdam / Netherlands.
—RMNE: Nationaal Natuurhistorisch Museum Naturalis, Leiden / Netherlands. — SDEI: Senckenberg Deutsches Entomologisches Institut, Miincheberg / Germany.
—SDNHM: San Diego Natural History Museum, San Diego, California / U.S.A. —SMEM: Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt a. M. / Germany.
—SMNS: Staatliches Museum ftir Naturkunde, Stuttgart / Germany.
—UCRC: University of California, Dept. of Entomology, Riverside, California / USA.
— UUZM: Uppsala University Zoological Museum, Linnean collection, Uppsala / Sweden.
—ZSMC: Zoologische Staatssammlung, Munich / Germany. — AB: Private collection of Alexander Banko / Canada.
— EB: Private collection of Edy Bhaskara / Indonesia.
— FH: Private collection of Frank H. Hennemann / Germany. —MO: Private collection of Maxime Ortiz / France.
— OC: Private collection of Oskar V. Conle / Germany. —RTC: Private collection of Royce T. Cumming / U.S.A.
— SLT: Private collection of Stéphane Le Tirant / Canada. —ST: Private collection of Sierra Teemsma / U.S.A.
—TM: Private collection of Tetsuo Miyashita / Japan.
Note on Locality Spelling Changes
With a number of historic data labels listed within, it is important to note the various spellings of localities which have changed throughout the years. Although not all those listed below are used within this work, we have noted additional spellings from throughout Wallacea to assist others who may be reviewing historic Wallacean specimens.
Present Common Name = Known Historic Names
—Ambon Island = Ambonia / Amboina
—Bacan Island = Bachans / Bachians / Batchians
—Banggai Islands = Bangai / Bankei / Bangkei
—Buru Island = Bouru / Boro / Boeroe
—Buton Island = Button / Boeton / Butung
—Halmahera = Jilolo / Gilolo / Jailolo
~Kai / Kei Islands (both commonly used at present) = Key Islands
— Maluku Province = Moluccas / Muluku Province
—Sanana Island = Sula Besi / Xulla Besi
—Seram Island = Ceram / Seran / Serang
— Sulawesi = Celebes
—Taliabu Island Regency = Taliaboe
Treatment of Species
Subgenus Phyllium (Pulchriphyllium) Griffini, 1898
Type species: Phyllium pulchrifolium Audinet-Serville, 1838, 292, by original designation.
Remarks. — Only two species currently in this subgenus and attributed to the frondosum species group defined by
Faunitaxys, 7(4), 2019: 1-25.
-Wallace Line “
_ Weber Line
; Lydekker Line
BORNEO
NEW GUINEA
Fig. 1. Area defined as Wallacea (in green), bordered to the northwest by Wallace’s line and to the southeast by Lydekker’s line.
Morotai
Tobelo. Galela {rondano Mtlbu
QPalolo
Sanana
Cungadial 5 P tly
Mamasa
eran @imbon
P re y BP. kovicum SP. sp. “Suton” BP. letirand BP. mamasnense >, tobaloanse bhaskare Ys Krayicunn “ay SP ¥ eiranti ¥ in ¥ tobdlosnse bhaskaral
4
Fig. 2. Distribution map of the Phyllium species currently known within the boundaries of Wallacea. (Google Earth: Image Landsat/ Copernicus: Data SIO, NOAA, U.S. Navy, NGA, GEBCO: Accessed December 13th, 2018.
4 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Hennemann et al. (2009) are known from Wallacea at present. One is only known from a unique specimen collected on the island of Bacan west of Halmahera in the northern Maluku Islands and the other species is not uncommon on the Kai Islands in the very southeastern portion of Wallacea. Additionally, both species are the only ones of the frondosum species group that are found outside of mainland New Guinea, and currently only known from female specimens.
Phyllium (Pulchriphyllium) suzukii GroBer, 2008 (Fig. 5, 7,9 & 61)
Phyllium (Pulchriphyllium) suzukii GréBer, 2008: 137, fig. 171-172. Holotype, 2: Phyllium (Pu.) suzukii det. GréBer, Molukken, Ins. Bacan, 9.2006; Phyllium (Pu.) suzukii GréBer sp. nov. Holotypus, det. D. GréBer 9.2006; DEI Hemimetabola #100121 [SDEI].
Distribution.
— INDONESIA: North Maluku Province, South Halmahera Regency, Bacan Island [SDEI].
Discussion. — Phyllium suzukii is one of only two species from the frondosum species group found outside of mainland New Guinea (the other being Phyllium keyicum Karney, 1914 discussed below) and is currently the only known species of Phyllium recorded from Bacan Island. This species is currently only known from the single holotype female held within the SDEI. This species is most characterized by the unique mesopraescutum which is 3.7 times wider than long (Fig. 9, the most extreme width to length ratio of known Phyllium females) as well as an abdomen with prominent, posteriorly directed projections on segments VI-VIII giving the abdomen a unique overall shape (Fig. 5); the outer margins of segments VII and VII posteriorly project beyond the apex of the anal segment. With a body length of only 57.6 mm it is the smallest of all know representatives of the Phylliidae within the boundaries of Wallacea. The egg was illustrated by GréBer (2008: 137, fig. 171) and is easily recognized by the unusually ovoid overall shape and large roughly circular impressions of the capsule.
Phyllium (Pulchriphyllium) keyicum Karny, 1914 (Fig. 3, 4, 6 & 8)
Phyllium keyicum Karny, 1914: 7, fig. 5. Holotype, 9: Kei Islands, Toeal Island [depository unknown].
= Phyllium insulanicum Werner, 1922: 126. Holotype, 2: Toeal, Kei- eilanden, J.H. Jurriaanse 1920 [RMNH]. (Synonymised by Hennemann et al., 2009: 65)
= Phyllium indicum Ginther, 1929: 614 (Misspelling of insulanicum).
Distribution.
— INDONESIA: Maluku Province, Southeast Maluku Regency, Kai Islands [NHMUK, NHMW, NMR, RMNH, SMNS, FH].
Discussion. — A detailed description of the females of this species and the synonymous Ph. insulanicum was presented by Willemse (1947). Since the holotype female of Ph. keyicum is not traced and presumed lost (Otte & Brock, 2005: 274), it may become necessary to designate a neotype. The egg was illustrated by Gr6Ber (2008: 123, fig. 146).
Despite the Kai islands being rather remote, there have been a number of female specimens collected over the years. Unfortunately despite the female being not uncommon, the male Ph. keyicum is still unknown. This species is most notable for its profemoral exterior lobe with a posterior margin that is straight or very nearly so (Fig. 6, black line), this is the only
species within the frondosum species group with this feature. Ph. keyicum also has the unique morphology of a single posteromedial tubercle on the head capsule versus other members of the frondosum species group which have the tubercle split into two. Their most variable morphological feature is the terminal abdominal segments which can range from nearly straight across (Fig. 4) to more pronounced lobes (Fig. 3) and the full range between these two extremes. This intraspecific variability led to the description of Phyllium insulanicum Werner, 1922, based on the more strongly lobed females. Willemse (1947) extensively described and compared a series of specimens from the Kai Islands and from his thorough review it appears as though the variation in the abdominal lobes is the only notable feature between the two taxa. It was not until Hennemann et al. (2009: 65) correctly synominized the Ph. insulanicum with Ph. keyicum, leaving there only a single species of Phyllium known from the Kai Islands. Variation in abdominal shape is now well known as a common occurrence in many species of Phyllium and not a feature that should singularly be relied upon for differentiation. With a body length of 70-79 mm females of this species are considerably larger than the related Ph. suzukii from Bacan Island.
Subgenus Phyllium (Phyllium) Iliger, 1798
Type species: Gryllis (Mantis) siccifolius Linnaeus, 1758: 425, by monotypy.
Remarks. — The seven species known to occur throughout Wallacea belong to two species groups defined by Hennemann et al. (2009). While only the Sulwesian Ph. celebicum de Haan, 1842 belongs to the celebicum species group all other known species fall within the siccifolium species group.
Phyllium (Phyllium) celebicum de Haan, 1842 (Fig. 10-16, 81 & 82)
Phasma (Phyllium) celebicum de Haan, 1842: 111. Holotype, 9: Celebes, Tondano [RMNH - not traced].
Distribution.
— INDONESIA: North Sulawesi Province, Minahasa Regency, Tondano [RMNH - type locality].
— Central Sulawesi Province, Sigi Regency, Palolo [RC].
— South Sulawesi Province, North Luwu Regency, Bungadidi [FH, OC, SLT].
— South Sulawesi Province, North Luwu Regency, Tiulapolu [FH, RC]. — Southeast Sulawesi Province, Buton Regency, Buton Island [OC].
Discussion. — The female holotype was first reported lost in Hennemann et al. (2009: 15) after their extensive search within the RMNH collection in April 2006. Examination of the presumed holotype in the same collection (Bragg, 1996: 110) has revealed a female clearly not celebicum as originally described by de Haan (1842: 111) or illustrated by Willemse (1947: 319, fig. 3). The concerned specimen has features much more agreeing with Ph. tobeloense based on the thorax and profemoral lobes and does not represent the lost Ph. celebicum holotype.
True Ph. celebicum is discussed in depth in Hennemann et al. (2009: 11) where the authors provided illustrations, a detailed redescription of the female, descriptions of the previously unknown males and eggs, summarized the intraspecific variability, and discussed the numerous misidentifications over
Faunitaxys, 7(4), 2019: 1 — 25.
10 mm
10mm
Fig. 3-9. Females, frondosum species group, dorsal view. - 3: Phyllium keyicum [FH 0083-1]. - 4: Phyllium keyicum [NHMUK 012496951 (Kei Is., 1923)]. - 5: Phyllium suzukii GréBer, 2008 holotype [SDEI]. - 6: Left profemor Phyllium keyicum; line illustrating the straight profemoral exterior lobe proximal edge, a unique feature to this species [NHMUK 012496951 (Kei Is., 1923)]. - 7: Phyllium suzukii holotype; line illustrating the curved profemoral exterior lobe proximal edge, and the arrow pointing out the protibial exterior spur, both features clearly contrasting Ph. suzukii from Ph. keyicum [SDEI]. - 8: Mesopraescutum and mesopleurae Phyllium keyicum [NHMUK 012496951 (Kei Is., 1923)]. - 9: Mesopraescutum and
mesopleurae Phyllium suzukii holotype [SDEI].
the years. Phyllium celebicum is currently the only known member of the celebicum species group from Wallacea. This species group is characterized by males with profemoral lobes which are broader than the interior lobes (Fig. 16), and females which have developed alae (length 26.0-32.0 mm, fig. 11), both features which easily separate them from any other Phyllium of Wallacea.
Phyllium celebicum is currently only known from throughout Sulawesi and from nearby Buton Island. The single female specimen from the island of Ambon in the MNHU listed in Hennemann, ef al., (2009: 11) has been reexamined and found to not represent Ph. celebicum. Although very similar, this female specimen has several features which differentiate it from Ph. celebicum such as: a subgenital plate that is long and slender almost reaching the apex of the abdomen (in PA.
celebicum the subgenital plate it short and stout, only just reaching to terminal segment X); mesopleurae which reach nearly to the anterior of the mesopraescutum (in Ph. celebicum the mesopleurae are distinctly shorter reaching only half way through the mesopraescutum); and alae which are notably longer than any examined Ph. celebicum females (49.0 mm in the MNHU specimen versus 26.0-32.0 mm in Ph. celebicum, see Hennemann, ef al., 2009: 16). The vague historic collection data of “Amboina” unfortunately cannot be trusted as truly coming from the island of Ambon and must be taken with little confidence. Realistically this celebicum species group member likely represents one of the many poorly described species from mainland Asia.
Body lengths: 9 9 75.5-92.0 mm, 34 62.0-66.0 mm.
6 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 10-16. Phyllium celebicum dorsal. - 10-11: Females; note the well developed alae which help to characterize the celebicum species group [FH 0634-5 (left) and 0634-1 (right)]. - 12: Male [FH 0634-4]. - 13-15: Female left profemor showing intraspecies variability [FH 0634-6, 0634-2, 0634-1]. - 16: Male left profemor; note the exterior lobe with its strong angle and notably larger than the interior lobe, a feature which defines
the celebicum species group [FH 0634-4].
Phyllium (Phyllium) conlei Curmming, Valero, & Teemsma, 2018 (Fig. 23 & 44)
Phyllium (Phyllium) conlei Cumming, Valero, & Teemsma, 2018: 3, fig. 3a-i. Holotype, 4: Indonesien, Lombok, XII.2012 [ZSMC].
Distribution.
— INDONESIA: West Nusa Tenggara Province, Lombok Island [ZSMC].
Discussion. — Currently only known from the unique holotype male deposited in the State Zoological Collection of Munich (ZSMC) originally from the collection of Oskar Conle (Germany). Cumming, Valero, & Teemsma (2018) discuss the lack of knowledge for the Phylliidae of the Lesser Sunda Islands
and described Ph. conlei as the first unique species to the island chain. With the unique male at only 47.5mm in length, Ph. conlei is one of the smaller species of Phyllium. Nearby species also with males of a similar size range are Phyllium jacobsoni Rehn & Rehn, 1934: 42.5-50.4 mm, Java (Hennemann et al., 2009), Phyllium hausleithneri Brock, 1999: 51.4-57.8 mm, West Malaysia and Sumatra (Cumming, Le Tirant, & Teemsma, 2018), and Phyllium chengiae Seow-Choen, 2017 (53 mm, Borneo). From all species of a similar size, Ph. conlei can be differentiated by the slender abdomen with a width only about 30% of the total abdominal length versus the others with broad spade shaped abdomens 40-55%.
Faunitaxys, 7(4), 2019: 1 — 25.
Fig. 17-22. Females, dorsal. - 17: Phyllium mamasaense holotype [SDEI]. 18-19: Phyllium letiranti green and brown colorforms [SLT coll.]. - 20: Phyllium siccifolium holotype [UUZM]. - 21: Phyllium siccifolium from Seram Island [FH 1042-1]. - 22: Phyllium tobeloense tobeloense from
Halmahera Island, Mount Ibu [FH 0657-7].
Phyllium (Phyllium) sp. “Buton” (Fig. 24 & 45)
Distribution.
— INDONESIA: Southeast Sulawesi Province, Buton Regency, Buton Island [FH].
Discussion. — Known only from a single male within the Hennemann private collection (FH 0673-1) this specimen was
the most difficult to place, with a near perfect match within the intraspecies morphological range of Ph. hausleithneri Brock, 1999 and Ph. jacobsoni Rehn and Rehn, 1934, both species found outside of Wallacea. Finding either species on Buton (far from their known distributions outside of Wallacea) would be very surprising and would challenge the well-defined biogeographic boundary lines of Wallacea. The authors are confident in the collection data, but, mistakes by collectors or specimen dealers are a possibility and we cannot ignore the fact that this could simply be a clerical error. Additionally, the
8 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 23-29. Males, dorsal. - 23: Phyllium conlei holotype [ZSMC]. - 24: Phyllium sp. “Buton” [FH 0673-1]. - 25: Phyllium tobeloense tobeloense from Halmahera Island, Mount Ibu [FH 0657-9], note the antennae not reaching the apex of the resting tegmina. - 26: Phyllium tobeloense tobeloense captive reared from Halmahera Island, Galela [FH 0657-13]. - 27: Phyllium letiranti from Peleng Island [SLT coll.]. - 28: Phyllium letiranti from Sanana Island, Waifara [FH 1008-10] note the antennae exceeding the apex of the resting tegmina. - 29: Phyllium siccifolium from Ambon Island [NHMUK].
Faunitaxys, 7(4), 2019: 1 — 25.
Fig. 30-45. Profemora, dorsal. - 30: Female Phyllium siccifolium holotype [UUZM]. - 31: Female Phyllium siccifolium [FH 1042-1]. - 32: Male Phyllium siccifolium [NHMUK]. - 33: Female Phyllium mamasaense holotype [SDEI]. - 34-36: Female Phyllium letiranti showing intraspecies variation in color and lobe serration. - 37: Male Phyllium letiranti [SLT coll.]. - 38-41: Females Phyllium tobeloense bhaskarai ssp. nov. paratypes, showing remarkable intraspecies variation in color and lobe serration [FH 1077-3, 1077-1, 1077-2. 1077-6]. - 42: Male Phyllium tobeloense bhaskarai ssp. nov. allotype [IMQC]. - 43: Male Phyllium tobeloense bhaskarai ssp. nov. paratype [FH 1077-21]. - 44: Male Phyllium conlei holotype [ZSMC]. - 45: Male Phyllium
sp. “Buton” [FH 0673-1].
female and egg from this “Buton” species are not yet known, leaving the possibility that clear morphological features identifying it as a new taxon might simply only be visible in the female or egg morphology. With Ph. tobeloense bhaskarai ssp. nov. only easily separated from the nominate subspecies by the unique egg morphology, it is very possible that this male could represent a new species or subspecies, only recognizable by female or egg morphology. Therefore, with a lack of significant morphological features to base a new taxon on, we simply refer to this species as Phyllium (Phyllium) sp. “Buton” throughout this work. Body length 48.5 mm.
Phyllium (Phyllium) letiranti Cumming & Teemsma, 2018 (Fig. 18, 19, 27, 28, 34-37, 50, 51, 54, 87, 88, 91 & 96-98)
Phyllium (Phyllium) letiranti Cumming & Teemsma, 2018: 2, fig. 1A- D, 2B-D, F-G, 3A-E, 4A-D, 5A-D, 6A-C. Holotype, 9: Indonesia, Peleng Island, Tinagkung Utara, near Luksagu Village: December, 2016: ~60 m. elv. 1°17’ S 123°25.4' E [IMQC]; Paratypes [IMQC, SDNHM, CASC, SDEI, RC, ST, SLT, and FH].
Phyllium (Phyllium) siccifolium, Klante, 1976: 65, 76, fig. 10, (in part).
Phyllium (Phyllium) siccifolium, Hennemann et al., 2009: 58 (in part).
10 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 46-50. Females, ventral view of coxae. - 46: Phyllium mamasaense holotype, showing black interior mark on meso- and metacoxae
[SDEI]. - 47: Phyllium tobeloense tobeloense, showing metacoxae only with a black interior marking, mesocoxae lacking colored mark [FH 0657-3]. - 48: Phyllium tobeloense bhaskarai ssp. nov. paratype, showing metacoxae only with a black interior marking, mesocoxae lacking colored mark [FH 1077-2]. - 49: Phyllium siccifolium, mesocoxae with black interior marking [NHMUK]. - 50: Phyllium letiranti live female bred by Maxime Ortiz (France) showing lack of black coxae markings and the vibrant orange coxae color (© Maxime Ortiz).
Distribution.
— INDONESIA: Central Sulawesi Province, Banggai Regency, Peleng Island [RC, FH, ST, SLT, CASC, IMQC, SDNHM]; Banggai Islands [MNHU].
— INDONESIA: North Maluku Province, Sula Islands Regency, Sanana Island [FH - distribution expansion].
Discussion. — Only recently collected (2016) and described (2018), this species has become widely available for sale as dried specimens and has entered the phasmid breeding community. In captivity it has been described as a relatively easy species to raise with a trend to produce a wide variety of female colorforms (Fig. 96-98). Originally described from Peleng Island, the authors have received photos of Phyllium from the nearby coast of Sulawesi (Central Sulawesi Province, East Peninsula) which appear very similar in morphology to Ph. letiranti, but without high resolution photos or specimens to review, a confident identification is not possible at this time. It would not be surprising to find Ph. letiranti on the Sulawesi coast near Peleng Island but with the similar species Ph.
mamasaense also found on Sulawesi, from photos alone an identification cannot be confirmed. Hopefully future collecting on Sulawesi can bring to light the distribution of Ph. letiranti with more clarity.
A range expansion can however be made for Sanana Island (Sula Islands) as represented by a male specimen from the Hennemann private collection (Fig. 28, FH 1008-10). With the island of Sanana known to have PA. letiranti, the female from “Taliaboe” (Taliabu) in the RMNH illustrated in Klante (1976) and the female from the MNHU from “Bangkei” (Banggai) discussed in Hennemann, et al., (2009) likely also represents Ph. letiranti. Examination of the specimen within the MNHU does reveal a female which matches well with slender PA. letiranti females from Peleng Island.
The two easiest diagnostic features to differentiate female Ph. letiranti from Ph. siccifolium are the ventral color of the coxae (Ph. siccifolium having distinct black markings and Ph. letiranti without) and the number of antennal segments (nine in Ph. siccifolium and ten in Ph. letiranti). Unfortunately, in Klante (1976) neither feature was illustrated clearly and
Faunitaxys, 7(4), 2019: 1 — 25. 11
therefore a thorough examination of the specimen in question must eventually be done to confirm this identification.
With the Sula Islands situated just to the east of the Banggai Islands, this distribution expansion is not surprising and opens the possibility that this species can also be found on the other Banggai or other Sula Islands.
Body lengths: 9 9 86.6-91.2 mm, 33 57.6-62.2 mm.
Phyllium (Phyllium) mamasaense GroBer, 2008 (Fig. 17, 33, 46, 68, 89 & 90)
Phyllium (Phyllium) mamasaense Groer, 2008: 124, fig. 147-148. Holotype, 2: Phyllium (Ph.). mamasaense det. GrofBer, Sulawesi, Mamasa, 9.2006; Phyllium (Ph.) mamasaense GrdBer sp. nov. Holotypus, det. D. Gréer 9.2006; DEI Hemimetabola #100116 [SDE]].
Distribution.
— INDONESIA: West Sulawesi Province, Mamasa Regency, Mamasa District [SDEI].
Discussion. — Only known from very few female specimens collected over the last ten years from the type locality of Mamasa, Sulawesi (Fig. 2). This species has the unique characteristic of having meso- and metacoxae with black interior markings on the ventral surface (Fig. 46) a feature within Wallacea currently only known from Ph. siccifolium (Fig. 49), or only partially seen in Ph. tobeloense (Fig. 47, only metacoxae with black marking). Interestingly, all females observed (although not a wide sample size) have had a very steady morphology with abdominal segments VII and VIII consistently roundly angular (Fig. 68) and the profemoral interior lobe with two large prominent teeth (as well as several small minor teeth) (Fig. 33). These features are variable among certain other Phyllium females and we look forward to hopefully reviewing more material in the future as well as describing the currently unknown male once it is identified. The large eggs are very distinctive (Fig. 89 & 90) and differ from those of all other Wallacean Phylliidae by the feather-like fringes of the capsule forming several parallel longitudinal ridges and the the very large cup-shaped operculum.
Body length 93.5-105.0 mm.
Phyllium (Phyllium) siccifolium (Linnaeus, 1758) (Fig. 20, 21, 29, 30-32, 49, 52, 55, 63 & 69) Gryllus (Mantis) siccifolium Linné, 1758: 425, No. 3. Holotype, 9:
Indes, Gustav IV Adolph; Gryllus (Mantis) siccifolius type; Coll. n° 1818 [UUZM].
= Phyllium brevicorne Latreille, 1806: 272. (Synonymised by Gray, 1835: 30).
= Phasma chlorophylla Stoll, 1813, 69, pl. 23: 89. (Synonymised by Redtenbacher, 1906: 176).
= Phasma citrifolium Lichtenstein, 1796: 78. (Synonymised by Gray, 1835: 30).
= Phasma donovani Gray, 1835: 31. (Synonymised by Redtenbacher, 1906: 176).
= Phasma foliatus Perry, 1810: plate 24. (Synonymised by Westwood, 1859: 172).
= Phasma gorgon Gray, 1835: 31. (Synonymised by Westwood, 1859: 172).
= Phyllium stollii Le Peletier de Saint Fargeau & Serville, 1827: 115. (Unnecessary replacement name for Phasma chlorophylla Stoll, 1813). [Not: Phyllium (Phyllium) tobeloense GréBer, 2007: 15. Erroneous synonym of Hennemann et al., 2009: 53].
Distribution.
— INDONESIA: Maluku Province, Ambon [MNHU, NHMUKkK, RMNH, UUZM], Seram [NHMUK, FH] and Buru Islands [NHMW].
Discussion. — As the type species for the family, this species is one of great importance but unfortunately it is also one of the many poorly known species because of the lack of available material in modern times. The holotype female (Fig. 20) was redescribed in Hennemann ef al. (2009: 55) as well as a male specimen from the island of Ambon from NHMUK (Fig. 29). These authors also discussed the numerous synonyms and provided information of the intraspecific variability. The exact type-locality of Ph. siccifolium originally stated to be “Habitat in Indiis” by Linnaeus (1758: 425) was discussed in detail in Hennemann et al. (2009: 57). These authors stated the holotype female in the UUZM collection to be most likely from the island of Ambon.
In addition to the island of Ambon, various collections examined over the years have revealed numerous females from the islands of Seram and Buru which match well with the morphology of the holotype. The likely misidentification by Klante (1976) discussed above from Taliabu Island is probably the morphologically similar Ph. letiranti known from the Banggai and Sula Islands, leaving true Ph. siccifolium restricted to east of Weber’s Line (Fig. 1) and only known from Ambon, Seram, and Buru. Unfortunately, the male of this species is little known and the eggs are still unknown for Ph. siccifolium. With egg morphology so drastically different between PA. letiranti and Ph. tobeloense hopefully future fresh material will reveal the eggs of Ph. siccifolium and their morphology should shed light on the relationship to the other similar species.
Body lengths: 9 89.0-94.0 mm, 34 67.0-73.0 mm.
Phyllium tobeloense Phyllium siccifolium
Ventral coxae color
Profemoral exterior lobe*
2 (Fig. 38-41)
Antennae Length
Metacoxae with distinct black interior marking, Q mesocoxae slightly orange or light green, no black interior mark (Fig. 47 & 48)
Thin and smoothly arcing, greatest width only 1.5 to 2 times the width of the profemora shaft
Short, not projecting beyond the apex of the oes resting tegmina (Fig. 25)
Mesocoxae and metacoxae and with distinct black interior marking (Fig. 49)
Broad and slightly angular, greatest width 2.5 to 3 times the width of the profemora shaft (Fig. 30 & 31)
Long, reaching or projecting beyond the apex of the resting tegmina (Fig. 29)
Table 1. Summary of comparison between Phyllium tobeloense and Phyllium siccifolium females and males. * Measured at the proximal edge of the profemoral interior lobe. ** When antennae are held at rest back along the body.
12 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
twos [2 [ws [nme [wen [—tenmorane fp esp fam
Table 2. Measurements of Phyllium (Phyllium) tobeloense bhaskarai ssp. nov., holotype female and allotype male. For the paratype material only the smallest and largest in the series were measured to give a range. All measurements made
to the nearest 0.1mm. * Including cerci and head, excluding antennae.
** Only measured on paratype specimen RC 18-421 which measures 101.2 mm in length.
Phyllium (Phyllium) tobeloense tobeloense Groer, 2007 (Fig. 22, 25, 26, 47, 64, 66, 83, 84, 92, 99 & 100)
Phyllium (Phyllium) tobeloense Gréer, 2007: 15, fig. 1-4. Holotype, ®: Phyllium (Ph.) tobeloense det. GréBer, Molukken, Halmahera, Tobelo, 8.1998 [SDE]T].
Distribution.
— INDONESIA: North Maluku Province, North Halmahera Regency, Tobelo [SDEI, FH, OC - type locality], Mount Ibu [FH, RC, SLT] and Galela [FH].
Discussion. — Originally collected in Tobelo, Halmahera, this species has been widely bred by enthusiasts and like PA. letiranti has yielded a number of beautiful female colorforms (Fig. 99 & 100). This species was briefly disputed in Hennemann et al. (2009) where it was presented as a junior synonym for Ph. siccifolium until GréBer (2011) where it was reestablished. Here we agree with the reestablishment of Ph. tobeloense as a valid species and discuss the morphological reasoning for our decision.
Although very similar in morphology, when a review of a significant series is examined these two species can be separated by the following characteristics. The first feature for differentiating females is the ventral color of the coxae, with Ph. tobeloense having a black marking only on the metacoxae, not on the mesocoxae (Fig. 47 & 48) versus Ph. siccifolium with both meso- and metacoxae with black markings. Also for females, the width of the profemoral exterior lobe is consistently different with Ph. tobeloense having a more slender lobe, no more than two times the width of the profemoral shaft (Fig. 38-41), as compared to Ph. siccifolium
which always has a profemoral exterior lobe that is notably wider (2.5 to 3 times) than the width of the profemoral shaft (Fig. 30 & 31). For males, the antennae immediately separate the two species, with Ph. tobeloense with shorter antennae, not longer than tegmina length when at rest (Fig. 25), versus Ph. siccifolium males which have longer antennae which reach to or past the apex of the folder tegmina. These distinguishing features are presented in summary in table 1. These features have proven to be consistent across all specimens examined of both populations, including extreme variations within each population.
Body lengths: 29 89.0-101.0 mm, 33’ 59.0-62.0 mm.
Phyllium (Phyllium) tobeloense bhaskarai Cumming, Le Tirant, and Hennemann, ssp. nov.
(Fig. 38-41, 48, 53, 56-58, 65, 67, 70-77, 79, 80, 85, 86, 101-103)
ZooBank : http://zoobank.org/4ED84870-BA6D-4546- B518-278B66EEAF4D
Holotype: 9: INDONESIA, Morotai island, North Maluku, Pulau Morotai Regency, Daeo village, XI-XII.2018 [IMQC].
Allotype: 3: INDONESIA, Morotai island, North Maluku, Pulau Morotai Regency, Daeo village, XI-XII.2018 [IMQC].
Paratypes: [103 33, 90 2°, 46 eggs], INDONESIA, Morotai island, North Maluku, Pulau Morotai Regency, Daeo Majiko village, XI-XII. 2018.
Depositions.
-14,2 99, 1 egg [AMSA] -1¢,1 9 [BPBM] -13,2 29, 1 egg [BYU]
Faunitaxys, 7(4), 2019: 1 — 25. 13
-2 88,2 22, 1 egg [CASC] ~5 34, 18 29, 10 eggs [IMQC] ~163,3 29 [IRSN]
~138,19 [LEMQ]
~2 33,2 29, 2 eggs [MNHN] ~13,3 29 [MNHU]
~163,1 9 [MZSF] ~2 33,2 29, 2 eggs [NHMUK]
~2 34,3 299° [NHMW]
~163,3 29 [RMNH]
-2 8,2 29 [SDE]
~2 83,2 29, 1 egg [SDNHM]
-13,3 99 [SMFM]
-13,2 99, 1 egg [UCRC]
-~2 83,3 29, 1 (penultimate instar) [ZSMC]
~13,1 9 [coll. AB]
-1 8,2 2 [coll. EB]
—15 33, 20 29, 12 eggs [coll. FH, No’s 1077-1 to 35, E]
-~1 4,1 [coll. MO]
~5 34,5 29 [coll. OC]
~6 64,7 29, 5 eggs [coll. RTC 18-421, 18-422, & 19-087 to 19-102] ~20 33, 20 99, 5 eggs [coll. SLT]
~1 4,1 9, 1 egg [coll. ST]
~1,1 9 [coll. TM].
Distribution.
— INDONESIA: North Maluku Province, Morotai Island Regency.
Remarks. — Currently only known from Morotai Island, Indonesia. Morotai is separated from northem Halmahera by the Morotai Strait, a stretch of water only about 10 kilometres (6.2 mi) wide. The only other island near Morotai is Bau Island, a small island on the west coast. It is possible that Phyllium (Phyllium) tobeloense bhaskarai ssp. nov. may also be found on Bau Island, due to its close proximity but currently no specimen records exist.
Discussion. — This new population has only recently been discovered (November, 2018 by Alim Yasin) on Morotai and the first impression was that this was likely a range expansion for Phyllium tobeloense until photos of the eggs were received. Immediately it became apparent that this Morotai population was not identical to the Halmaheran population. The decision to classify this Morotai population a subspecies of Phyllium tobeloense and not full species status is tentative but was based on a number of observations. Upcoming genetic analysis will help reveal the extent of the variability between the populations, but our present morphological review suggests a very close relationship. First is the incredible similarity of the adult morphology between the two populations. With the Halmaheran population rather popular in both phasmid breeding communities and commonly bred for commercial export as dried specimens, a wide review of the intraspecies variation has been possible. This has revealed that the females of this species have moderate variability in the profemoral interior lobe, and slight variability in abdominal segment VIII. Captive bred specimens have also resulted in a significant variety of color forms (Fig. 99 & 100) and a variety of major and minor profemoral interior lobes, similar to those seen in exported dried specimens. With this significant view into the intraspecies variation of males, females, and eggs, the egg morphology differences between the Morotai and Halmahera populations can easily be identified as not simply intraspecies variation and is discussed in detail below. With egg
morphology one of the only concrete features to allow differentiation between the populations, and the obvious similarity between the population’s eggs, we consider this Morotai population an offshoot of the mainland Halmahera population and classify it as a subspecies.
An additional reason for our decision to classify the Morotai population as a subspecies is the significant number of other insect species which follow this trend. Frequently, the Morotai populations are subspecies with sibling subspecies from the other northern Maluku Islands. For example, Wallace’s golden birdwing butterfly Ornithoptera croesus Wallace, 1859 is endemic to the northern Maluku islands, with five total subspecies distributed throughout these islands. In particular, from Halmahera there is Ornithoptera croesus lydius (Felder & Felder, 1865) and from Morotai Ornithoptera croesus toeantei Parrot & Schmid, 1984 . Interesting, these butterfly subspecies parallel our own Phyllium tobeloense subspecies as the females are more easily morphologically separated than males are, with males of these two-island populations near impossible to differentiate (pers. comm. Gilles Delisle, birdwing butterfly expert). As discussed in more detail below, we were able to find consistent morphological differences between the females and eggs of these two populations of Phyllium tobeloense but between males no reliable features could be found.
Differentiation. — Phyllium tobeloense bhaskarai ssp. nov. may be separated from the nominate subspecies from Halmahera by only a few subtle morphological features. This Morotai population is the first subspecies described for Phyllium tobeloense GréBer, 2007 and is only easily separated from the nominate by the egg morphology.
The most easily seen morphological feature separating the eggs of the two subspecies is the overall shape when viewed laterally. The nominate subspecies has eggs that are asymmetrical with a reniform (kidney shaped) (Fig. 83) whereas the new subspecies has eggs which are ovoid in shape and almost symmetrical (Fig. 85). Also, in the nominate subspecies the long pinnae along the margins are not fully formed around the entire capsule with the area opposite the micropylar plate and below the operculum bald (Fig. 83). In the new subspecies the long pinnae along the margins are fully formed around the entire capsule, with no bald portion opposite of the micropylar plate below the operculum (Fig. 85).
Adults of the two subspecies are much more difficult to differentiate morphologically with only subtle differences found when examining a large series of both populations. The only consistent feature observed on the females were the length of the subgenital plate. In the nominate subspecies the subgenital plate was stouter, protruding only about half the length of sternite X, leaving a significant amount of the gonapophyses exposed (Fig. 64), and for the new subspecies the subgenital plate was longer, about three-quarter the length of sternite X, leaving only the tips of the gonapophyses exposed (Fig. 65).
After an extensive search for a consistent feature to differentiate males of these two populations, none was found that was reliable enough to be useful in a morphological key. With this species so common thus allowing the intraspecies variation to be well known, our conclusion is that it is likely that only molecular comparison can confidently differentiate these two populations males.
Habitat and Host Plant. — The local collectors were kind enough to share a number of photos of live specimens in the wild on their host plant (Fig. 78-80). The photos of this host plant were shared with Binjie Ge of the Shanghai Chenshan
14 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 51-58. Mesothorax - 51: Female Phyllium letiranti, dorsal view [SLT coll.]. - 52: Female Phyllium siccifolium, dorsal view [NHMUK]. - 53: Holotype female Phyllium tobeloense bhaskarai ssp. nov. dorsal view [IMQC]. - 54: Male Phyllium letiranti, dorsal view [SLT coll.]. - 55: Male Phyllium siccifolium, dorsal view [NHMUK]. - 56: Allotype male Phyllium tobeloense bhaskarai ssp. nov., dorsal view. - 57: Holotype female Phyllium tobeloense bhaskarai ssp. nov. lateral view of thorax [IMQC]. - 58: Allotype male Phyllium tobeloense bhaskarai ssp. nov. lateral view of thorax [IMQC].
Plant Research Center who identified them as likely a species of Macaranga (Euphorbiaceae) but without a specimen to examine a definitive identification is not possible at this time. Macaranga would not be surprising as this has been recorded as a phasmid host plant in the past for several other phasmids (Seow-Choen, 1997). The native people of Morotai call the tree by the common name “Liwiwi”.
Also accepted as a host plant was Psidium guajava (Myrtaceae) by individuals collected in the wild and kept in captivity by Edy Bhaskara (Fig. 101).
Morphological description of the female.
Coloration. — Coloration descriptions are largely based on photographs of live material sent to the authors from collectors on Morotai. Preserved specimens with color that was decently retained were also examined to help describe the spectrum of observed colors.
Two main color forms have been observed, a green form (Fig. 103) and a yellow/orange form (Fig. 102). In both color forms the antennae are of a slightly darker color than the rest of the body and with the terminal three segments with more dense setae and a general darker color than the other antennae segments. Compound eyes of both forms are generally of a similar color to that on the head capsule or slightly lighter in color (in some dried specimens the eyes were a reddish brown but that is possibly from the drying of the specimen). In the green form, the
serration of the femoral lobes, the margins of the mesothorax, and the venation of the tegmina are lighter in color than the remaining surfaces, generally a pale green/yellow or cream color.
In the yellow/orange form, the wing venation is a darker orange than the tegmina ground color which can be more yellow than orange. In both color forms the ventral surface of the metacoxae have a clear black marking, mesocoxae without, cream, yellow, or pale green only.
Morphology
Head. — Capsule \onger than wide (1.1-1.2 times). — Vertex nearly smooth, with only slight texture and no notable granules except the prominent posteriomedial tubercle. Frontal convexity stout, only slightly shorter than the length of the first antennomere, and with a moderate number of clear setae throughout. — Antennae consisting of nine segments, with the terminal two segments (VIII and IX) approximately the same length, and segments IV through VII shorter than the terminal two, and IV through VII of a uniform length (Fig. 76). — Antennomeres I-VII sparsely marked with transparent setae (segments I through III with longer setae than the following segments), and the terminal two antennomeres are more densely covered in brown setae with fewer transparent setae. — Compound eyes snug to the head capsule, not strongly protruding from the head capsule. — Ocelli absent. Antennal fields wider than the first antennomere but not protruding back farther than the frontal suture.
Faunitaxys, 7(4), 2019: 1 — 25. 15
Thorax. — Pronotum with anterior and lateral margins that are nearly straight, lateral margins converging to a straight posterior margin that is about 60 percent the width of the anterior rim. The pronotum surface lacks nodes but has prominent furrows giving it a lumpy surface. The surface has a prominent pit in the center and one along the sagittal plane near the anterior rim, the remaining furrows are prominent and clearly visible. The pronotum has a moderate anterior rim, lateral rims are weak, and the posterior lacks a rim. — Prosternum and the anterior quarter of the mesosternum with slight granulation. — Metasternum rather smooth, without prominent granulation. — Mesopraescutum approximately as long as wide, with lateral rims with five prominent tubercles rather uniform in size, with occasionally one to two small nodes between the anterior most tubercles. — Mesopraescutum anterior rim prominent with a large tubercle in the center with the remainder of the rim without clear nodes but slightly lumpy, not perfectly smooth (Fig. 53). — Mesopraescutum crest (excluding the tubercle of the anterior rim) nearly flat, not prominent, with four small nodes unevenly spaced along the sagittal plane (Fig. 53). — Mesopraescutum surface lumpy, but lacking clear nodes. — Mesopleurae straight and evenly diverging; lateral margin with five to seven major tubercles which are almost always even in size; occasionally those with five tubercles have one to two smaller tubercles present (Fig. 53). Face of the mesopleurae smooth, with two divots, one more prominent divot on the anterior third and one closer to the center that is faint. — Tegmina slightly variation in length, at the shortest only reaching the anterior margin of segment VIII and at the longest reaching just over three quarters the way through VIII.
Alae. — Rudimentary, only about 6mm in length.
Abdomen. — Abdominal segments IT through the anterior two thirds of IV diverging, posterior third of IV through VII gently converging. — Segments VII-X converge more acutely than the previous segments, converging uniformly to a terminal segment that is about as wide as it is long with a broad rounded apex. — Subgenital plate starts at the anterior margin of segment VIII and extends three quarters of the way into segment X ending in a fine point, leaving only the tips of the gonapophyses exposed. — Gonapophyses long, reaching the posterior margin of segment X (Fig. 65). — Cerci relatively flat not strongly cupped, with a prominently granular surface and very few short setae near the apex only.
Legs. — Profemoral exterior lobe smoothly arcing from end to end without an angle, exterior lobe approximately the same width as the interior lobe, or slightly thinner and with a granular surface. — Profemoral interior lobe obtuse angled and marked with five to seven serrate teeth with generally four to five of these teeth larger than the other smaller teeth and not perfectly evenly spaced. — Mesofemoral interior lobe smoothly arcs from end to end and has five to seven serrate teeth distributed on the distal half. — Interior and exterior lobes are of a similar width, but the exterior lobe is clearly angled, not smoothly arcing like the interior. — Mesofemoral exterior lobe with generally only a single small tooth on the distal half of the arc, rarely an additional smaller tooth is present. — Metafemoral interior lobe wider than exterior lobe, and arcs end to end and has nine to eleven serrate teeth pointing distally. — Metafemoral exterior lobe is thin and smooth, hugging the metafemoral shaft, lacks teeth. — Protibiae lacking an exterior lobe. — Protibial interior lobe spans the entire length in an evenly weighted triangle and is about twice the width of the protibiae shaft itself: — Mesotibiae and metatibiae lacking exterior and interior lobes.
Morphological description of the male.
Coloration. — Most of the body is of a lime green color with little variation throughout. Ventral of coxae a cream color, without black markings. The antennae are of a slightly darker green/brown than that found throughout the body. Protibial interior lobe occasionally with slight brown markings.
Morphology
Head. — Capsule slightly longer than wide, with a weakly granular vertex. Frontal convexity stout with a dull point and sparsely marked with transparent setae. — Antennae consisting of 25 segments (including the scapus and pedicellus which are marked with few transparent setae), all other antennal segments with a dense short covering of darker setae relatively uniform in length throughout all segments. — Compound eyes large and notably protruding away from the head capsule. — Ocelli developed clearly between compound eyes.
Thorax. — Pronotum with anterior margin slightly concave and lateral margins that are straight and gently converging to the posterior margin that is about 60 percent the width of the anterior rim. Anterior and lateral margins with moderate rims, posterior margin without a rim and smooth. Face of the pronotum with moderate furrow on the anterior half and the surrounding surface without granulation and smooth. — Prosternum with slight granulation throughout, not highly noticeable. — Mesosternum surface with more noticeable granules on the anterior. — Metasternum surface mostly smooth. — Mesopraescutum about as wide as long, with lateral rims with six to seven tubercles of varying sizes spread nearly evenly throughout. — Mesopraescutum anterior rim prominent with a large tubercle in the center with the remainder of the rim smooth (Fig. 56). — Mesopraescutum crest (excluding the tubercle of the anterior rim) with three to four small nodes unevenly spaced along the sagittal plane (Fig. 56). — Mesopleura diverging weakly on the anterior but more prominently on the posterior half; lateral margin with six to seven major tubercles throughout, with clear gaps between each tubercle with at most two small nodes in the gaps. — Mesopleural face with two faint divots, one on the anterior third and one on the posterior third, anterior and posterior of the face smooth, center of mesopleural face with a wrinkled texture. — Tegmina reaching half way through abdominal segment I'V or to the posterior margin on IV.
Alae. — Well developed, reaching segment IX or partly into the anal abdominal segment.
Abdomen. — Ovular, with segments II through IV diverging, V and VI parallel or slightly convex, VII through X steadily converging to a blunt rounded apex. — Anal abdominal segment longer than wide with lateral margins that converge slowly throughout the majority of the length and then converge prominently at the terminal third. — Poculum starts in abdominal segment VIII, with parallel lateral margins, ending in a broad apex that slightly reaches under segment X. — Cerci are slightly cupped and exceed the length of segment X, lateral margins are straight and the surface of the cerci are covered in nodes and short brown setae throughout. — Vomer broad with nearly straight margins converging to the apical hook which points upwards into the paraproct.
Legs. — Profemoral exterior lobe lightly granular and _ slightly thinner than the profemoral shaft, lobe gently arcing along the entire profemoral length. — Profemoral interior lobe about as wide as the shaft and the exterior lobe combined, situated on the distal two thirds and obtuse, not sharply angled. Profemoral interior lobe with four to five serrate teeth of similar sizes or with those in the center only slightly larger. The spacing is uniform except for the middle space which is slightly greater than the spacing on the sides. — Mesofemoral exterior lobe about as wide as the shaft, arcing from end to end with the widest portion weighted slightly off center on the distal end and with two to four small serrate teeth. — Mesofemoral interior lobe slightly thinner than the exterior lobe and with less of a bend, and six to eight serrate teeth on the distal half of the lobe. — Metafemoral exterior lobe thin and lacking distinct dentition but with a slight rough texture, hugging femoral shaft. — Metafemoral interior lobe slightly wider than exterior lobe with ten to twelve serrate teeth throughout the distal two-thirds of the length. No exterior protibial lobe, interior lobe extends the entire length in a rounded triangle, with the widest portion in the center or slightly off center on the distal end and slightly wider than the shaft width. — Meso- and metatibia simple, lacking lobes.
Description of Egg (Fig. 85 & 86).
The lateral surfaces are flattened and the dorsal and posterior surfaces are slightly convex, which gives the egg a slightly rounded ovular appearance. When viewed from the lateral aspect, the anterior and posterior widths of the capsule are similar, with the center of the capsule the widest point. When viewed from the lateral aspect, the anterior, posterior, and lateral margins are set with numerous rope-like pinnae, each with a split, slightly hooked apex giving them a barbed appearance (Fig. 92). Lateral surfaces with textured surface of circular markings, with 16 to 25 of those circular markings developed into deeper pits. These pits are in no detectable pattern and vary from egg to egg. Area around all pits lacking
16 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Keys to known species of Phyllium from Wallacea
Keys are broken into female, male, and egg morphology due to the marked sexual dimorphism. Female and egg Ph. (Ph.) conlei and Ph. (Ph.) sp. “Buton” unknown. Egg of Ph. (Ph.) siccifolium unknown. Male of Ph. (Ph.) mamasaense, Ph. (Pu.) keyicum and Ph. (Pu.) suzukii unknown. Males of Ph. (Ph.) tobeloense tobeloense and Ph. (Ph.) tobeloense bhaskarai ssp. nov. cannot be morphologically separated and are simply listed together as Ph. (Ph.) tobeloense in the below key.
Female
1. Mesopraescutum distinctly wider than long (>1.7 times) (Fig. 8 & 9) — Mesopraescutum about as long as wide, or longer than wide (Fig. 51-53)
2. Protibia with an exterior spur near the center (Fig. 7 arrow); profemoral posterior margin distinctly recurved (Fig. 7 line); length to width ratio of the mesopraescutum approximately | : 3.4 (Fig. 9) Ph. (Pu.) suzukii
— Protibiae lacking exterior lobe; profemoral posterior margin straight (Fig. 6 line); length to width ratio of the mesopraescutum approximately 1 : 1.8 (Fig. 8) Ph. (Pu.) keyicum
3. Alae developed (Fig. 11) — Alae rudimentary
4. Mesocoxae or metacoxae with distinct black interior marking (Fig. 46-49); antennae with nine segments — Mesocoxae and metacoxae without distinct black interior marking (Fig. 50); antennae with ten segments
5. Mesocoxae and metacoxae with distinct black interior marking; exterior lobe of the profemora wide and gently angled with
the greatest width 2.5 to 3 times the width of the profemora shaft (measured at the promixal edge of the interior lobe of the
profemora) (Fig. 30-31)
— Metacoxae with distinct black interior marking, mesocoxae slightly orange or light green, no black interior mark; exterior lobe of profemora thin and smoothly arcing with the greatest width only 1.5 to 2 times the width of the profemora shaft (measured at the promixal edge of the interior lobe of the profemora) (Fig. 38-41)
6. Three to five teeth of approximately even size on the interior lobe of the profemora (Fig. 30-31); abdominal segment VIII margin always almost perfectly straight (Fig. 69 line ) Ph. (Ph.) siccifolium
— Two large teeth and smaller minor teeth between on the interior lobe of the profemora (Fig. 33); abdominal segment VIII margin always with a smooth bend giving a slight lobe to the segment (Fig. 68 line) Ph. (Ph.) mamasaense
7. Subgenital plate stout, protruding about half the length of sternite X, leaving a significant amount of the gonapophyses exposed Ph. (Ph.) tobeloense tobeloense
— Subgenital plate long, about three-quarter the length of sternite X, leaving only the tips of the gonapophyses exposed Ph. (Ph.) tobeloense bhaskarai ssp. nov.
Male
1. Profemoral exterior lobe wider than the interior lobe, strongly angled (Fig. 16) — Profemoral exterior lobe thinner than the interior lobe, arcing end to end, not strongly angled (Fig. 42-45)
2. Interior lobe of the protibia thin and arcing from end to end without a distinct triangular shape and slender, only about as wide as the protibial shaft; abdomen long and slender, at its greatest width only about 30% of the total abdomen length (Fig. 23) Ph. (Ph.) conlei
— Interior lobe of the protibia with a distinct triangular shape and wider than the protibial shaft; abdomen broad, with a greatest width of about 37-50% of the length of the total abdomen
3. Abdomen spade shaped with abdominal segments V through the apex distinctly converging (Fig. 24) Ph. (Ph.) sp. “Buton”
— Abdomen ovoid, with abdominal segments V and VI with parallel margins or only slightly subparallel, only segments VII through the apex distinctly converging (Fig. 25)
4. Antennae when resting back along the dorsal surface are similar in length or longer than the folded tegmina (Fig. 28) — Antennae when resting back along the dorsal surface are shorter than the folded tegmina (Fig. 25) Ph. (Ph.) tobeloense
5. Large species 67.0-73.0 mm; antennae apex when resting back along the dorsal surface reaching apex of tegmina or only exceeding length by one to two segments; mesopraescutum stout (length to maximum width ratio of about 0.8-0.9) and with an anterior that is distinctly broader than the posterior (Fig. 55) Ph. (Ph.) siccifolium
— Medium species 57.6-62.2 mm; antennae apex when resting back along the dorsal surface exceeding apex of tegmina by four to five segments; mesopraescutum long and thin (length to maximum width ratio about 1.0-1.2) and with anterior and posterior margins approximately the same width (Fig. 54) Ph. (Ph.) letiranti
Faunitaxys, 7(4), 2019: 1 — 25. 17
Egg
1. Capsule entirely lacking distinct pinnae
— Capsule with distinct pinnae (most prominent along the operculum rim, if not prominent along the rim, then with short distinct
pinnae along the longitudinal margins of the capsule itself)
2. Lateral margins of the micropylar plate with four to five large pits throughout the length — Lateral margins of the micropylar plate with eight to nine small pits throughout the length
Ph. (Pu.) suzukii Ph. (Pu.) keyicum
3. Operculum raised, not flat, lacking long pinnae along the rim (Fig. 81) — Operculum flat, with long straight pinnae along the rim (Fig. 83-90)
4. Opercular rim with pinnae that are broad and featherlike (Fig. 91), broadest throughout their length
— Opercular rim with pinnae that are thin, ropelike, broadest at the apex (Fig. 92)
5. Lateral surface with short pinnae, notably shorter than the pinnae of the posterior and anterior ends (Fig. 88) — Lateral surface with rows of long pinnae, of a similar length to the pinnae of the posterior and anterior ends (Fig. 90)
Ph. (Ph.) letiranti Ph. (Ph.) mamasaense
6. Overall shape somewhat reniform (kidney shaped) (Fig. 83); long pinnae along the margins not fully formed around the entire
capsule, area opposite the microplylar plate and below the operculum bald (Fig. 83)
Ph. (Ph.) tobeloense tobeloense
— Overall shape more symmetrical, ovoid in shape (Fig. 85); long pinnae along the margins fully formed around the entire capsule,
no bald portion opposite of the microplylar plate below the operculum (Fig. 85)
long pinnae, leaving the lateral surfaces without significant projections. Micropylar plate covering approximately half of the overall length, situated near the center, not weighted towards either end of the capsule. Micropylar plate in a thin teardrop shape, with most of the width thinner than the micropylar cup. Micropylar cup of moderate size and placed on the posterior third of the capsule. Operculum nearly circular, with around ten round impressions encircling the center, and the outer margin of the operculum has a row of pinnae similar in length to those found along the other edges of the capsule. Overall color yellow, tan, or light brown.
Measurements [mm].
— length (including operculum) : 3.8 — length (excluding operculum) : 3.5 — height : 2.1
— width : 2.4
— length of micropylar plate : 2.2
Etymology. — This new subspecies is dedicated to Edy Bhaskara from Malang, Indonesia who was the first to send us specimens for this description. May he be thanked for his numerous collaborations to discover new taxa.
Note on Phyllium female genitalia in relation to egg morphology
As discussed in depth above, the only easily observed distinguishing features between the two subspecies of Phyllium tobeloense are egg morphology and the female subgenital plate length. This difference in subgenital plate length is likely linked to the morphological variation observed between the eggs of the two populations, with subgenital plate length reflecting the ideal arrangement to hold the egg before it is flung away.
Within the Phylliidae in general, the ratio of subgenital plate and gonapophyses lengths help to reveal general features of egg morphology. Members of the bioculatum species group with eggs characterized by five distinct ribs along the egg (with the exception of Phyllium giganteum Hausleithner, 1984 which has 3 rounded ribs) have females with shorter subgenital plates and
Ph. (Ph.) tobeloense bhaskarai ssp. nov.
shorter gonapophyses, likely because the eggs can easily be held with this arrangement or the shorter features aid in the action of flinging the egg away from the female (Fig. 59). In contrast, the frondosum species group have short subgenital plates and longer gonapophyses (Fig. 61 & 62), with eggs which lack these distinct ribs. In this case the longer gonapophyses are likely needed to help hold the egg in place before flinging it. This arrangement of a short subgenital plate and long gonapophyses is also seen in Phyllium giganteum (Fig. 60) likely for the same reason, as Ph. giganteum eggs have stout rounded ribs much like the rounded ribs of the frondosum species group. Members of the Phyllium (Phyllium) subgenus generally all have longer gonapophyses and a range of subgenital plate lengths (presumed to be according to the ideal length for their specific egg shapes). With eggs of varying morphologies, the subgenital plate appears to be the feature most variable among females with differing egg morphologies, as observed by the two Ph. tobeloense subspecies and their
eggs.
Biogeography and discussion
With nine species of Phyllium currently known from throughout Wallacea (Fig. 2), several biogeographical trends can clearly be seen. Particularly the Weber Line which separates the Papuan subregion species (Ph. suzukii and Ph. keyicum within the frondosum species group, which is only found additionally in the Papuan subregion) from the Oriental elements, particularly illustrative Ph. celebicum, the only member of the celebicum species group found this far south, with the others either father north in the Philippines, Micronesia, or on mainland Asia. The Weber line also illustrates the likely delineation between Ph. letiranti (west of the line: Banggai and Sula Islands) from Ph. siccifolium (east of the line: Buru, Ambon, and Seram Islands).
Of the Lesser Sunda Islands within Wallacea, currently only Phyllium conlei from Lombok Island is known, with all other islands devoid of literature and specimen records (Cumming, Valero & Teemsma, 2018).
To date Phyllium mamasaense has only been recorded from the type locality of Mamasa, Sulawesi, but with the other
18 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 59-69. Fig. 59-67. Genitalia ventral view. - 59: Phyllium abdulfatahi Seow-Choen, 2017 [RC16-039], short subgenital plate and short gonapophyses. - 60: Phyllium giganteum [RC16-044], short subgenital plate and long gonapophyses. - 61: Phyllium suzukii holotype [SDEI] female short subgenital plate and long gonapophyses. - 62: Phyllium keyicum [NHMUK 012496951 (Kei Is., 1923)]. - 63: Phyllium siccifolium [FH 1042-1] Seram Island. - 64: Phyllium tobeloense tobeloense [FH 0657-4] Halmahera Island, Tobelo. - 65: Phyllium tobeloense bhaskarai ssp. nov. holotype female [IMQC]. - 66: Phyllium tobeloense tobeloense [FH 0657-6] Halmahera Island, Tobelo. - 67: Phyllium tobeloense bhaskarai ssp. nov. allotype male [IMQC]. Fig. 68-69. Dorsal view of female terminal abdominal segments; lines show the curve of segment VIII, illustrating a consistent feature found between Ph. mamasaense and Ph. siccifolium. - 68: Holotype Phyllium mamasaense [SDEI]. - 69: Phyllium siccifolium [FH 1042-1].
Sulawesian species Ph. celebicum found throughout the island it would not be surprising if Ph. mamasaense were also more widely distributed on the island.
Within Wallacea there are a number of islands currently devoid of Phylliidae records, some of the largest being from the eastern Lesser Sunda Islands: Sumbawa, Sumba, Flores, and Timor. With the island of Timor a rather large island situated just south of the Phylliidae rich island of Sulawesi, the authors expected there to be records from Timor as well. However, after speaking with Chris Reid, Principal Research Scientist at the Australian Museum in Sydney he is skeptical Phylliidae can be found on the island. The Australian Museum has been conducting field work on Timor since 2012 and _ his observations were as follows, “Most of Timor is dry woodland on limestone dominated by either Eucalyptus, Casuarina, or
Acacia (now mostly introduced spp), with a bit of sandalwood etc. Plus a lot of pasture and a lot of weedy arable. The remaining rainforest is almost entirely dry vine thicket — the wetter stuff is either converted to coffee or too remote for easy access or we just didn't get into it. So in general I’d say Phyllium is unlikely” (pers. comm. Chris Reid).
Undoubtedly, our current knowledge of Wallacean Phylliidae is still fractional. Of the nine species that have so far become known from throughout Wallacea, five are only known from a single sex: Ph. conlei and Ph. sp. ‘Buton’ are both only known from unique males, and Ph. mamasaense, Ph. keyicum and Ph. suzukii are only known from females. There is another questionable celebicum-like species only known from an old female specimen labelled “Amboina” in the collection of MNHU, whose identity deserves clarification by additional
Faunitaxys, 7(4), 2019: 1 — 25. 19
Fig. 70-77. Phyllium tobeloense bhaskarai ssp. nov. - 70: Holotype female dorsal [IMQC]. - 71: Paratype female dorsal [FH 1077-1]. - 72: Paratype female ventral [FH 1077-1]. - 73: Paratype male [FH 1077-22]. - 74: Paratype male [FH 1077-23]. - 75: Dorsal view of terminal abdominal segment (= tergum X) and cerci, holotype [IMQC]. - 76: Dorsal view of antennae, holotype female [IMQC]. - 77: Dorsal view of antennae, allotype male [IMQC].
CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
s3) ~
— ~
-
Fig. 78-80. Macaranga sp. (Euphorbiaceae) with wild Phyllium (Phyllium) tobeloense bhaskarai ssp. nov. on Morotai Island (© Alim Yasin). - 78: Macaranga sp., host plant. - 79: Subadult female on Macaranga sp. - 80: Adult male on Macaranga sp.
material (see discussion on PA. celebicum). Also, the authors are aware of at least one further still undescribed Phyllium species which is only known from a photograph. It is very much hoped that the present paper will encourage collectors to look out for Phylliidae throughout Wallacea to obtain fresh specimens that can help broaden our knowledge of these fascinating insects from this particular region so well known for its high biodiversity and high degree of endemism.
Acknowledgments
The authors thank René Limoges, entomological technician at the Montreal Insectartum for taking many photos for this work, as well as for many professional courtesies. Thank to Vadim Voitsekhovskii (Bali, Indonesia) for sending a series of specimens for this study. A special thanks to Alim Yasin (Morotai, Indonesia) who sent to us an extra 50 males and 50
Faunitaxys, 7(4), 2019: 1 — 25. 21
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micropylar plate. - 81 & 82: Phyllium celebicum. - 83 & 84: Phyllium tobeloense tobeloense. - 85 & 86: Phyllium tobeloense bhaskarai ssp. nov. paratype. - 87 & 88: Phyllium letiranti paratype. - 89 & 90: Phyllium mamasaense (© Gré8er D.). Fig. 91-92. Pinnae from the opercular rim. - 91: Featherlike pinnae, frills throughout length, wide throughout length, apex not widest portion, Phyllium letiranti. - 92: Ropelike pinnae, stalk without frills, broadest portion the apex, Phyllium tobeloense tobeloense. Scale bar = 2 mm.
females that allowed us to deposit many paratypes in various museums and who also sent us photos of wild specimens and the habitat. Thanks to Maxime Ortiz (France) who sent us eggs that have been used in this work. Thanks also to Binjie Ge (Shanghai Chenshan Plant Research Center and Gilles Vincent, Special advisor of the Executive President of the Chenshan Botanical Garden-Shanghai, China) for the identification of the host plant. Thank you to Jim Berrian (SDNHM, San Diego) for years of support and access to the museum’s camera. Thank you to Tim Van molle (Rupelmonde, Belgium) for allowing us to use photos of his beautiful Phyllium tobeloense color forms and Detlef GroBer (Berlin, Germany) for allowing us to use his photos of Phyllium mamasaense eggs in this work. Sigetake Suzuki (Hokkaido, Japan) kindly provided various specimens from Sulawesi and helpful information. We thank Edy Bhaskara (Malang, Java, Indonesia) for providing the specimens of Phyllium tobeloense bhaskarai ssp. nov. and Ph. tobeloense tobeloense for description as well as pictures of live
specimens of the new subspecies. Thanks are also due to the curators of various museums for access to the corresponding collections that contain material listed in the present paper and/ or providing photographs of specimens: Dr. Michael Ohl and Ulrike Dorandt (MNHU, Berlin), Dr. Susanne Randolf and Prof. Ulrike Aspé6ck (NHMW, Vienna), Dr. Caroline Pepermans (RMNH, Leiden) and Hans Mejlon (UUZM, Uppsala). Mandy Schréter and Stephan Blanke (SDEI, Senckenberg German Entomological Institute Miincheberg) for photographs of specimens used in this work. Judith Marshall and Benjamin Price (NHMUK, London) for processing the loan of the Phyllium keyicum specimen despite the lack of an Orthopteroid collection curator. To conclude we of course thank our two peer reviewers who gave us valuable feedback and suggestions on how to improve this text, thank you Philip E. Bragg (Nottinghamshire, United Kingdom) and Dr Allan Taylor (Quebec, Canada).
CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 93-95. Live Phyllium celebicum captive reared F1-generation. - 93: Male/female pair. - 94: Male. - 95: Female.
Faunitaxys, 7(4), 2019: 1-25.
Fig. 96-98. Live Phyllium letiranti captive reared (© Maxime Ortiz). - 96: Yellow form female. - 97: Yellow form female, detail of head, thorax, and profemoral lobes. - 98: Phyllium letiranti Males.
24 CUMMING, LE TIRANT & HENNEMANN. — Phyllium of Wallacea
Fig. 99-100. Live Phyllium tobeloense tobeloense captive reared female (© Tim Van molle). - 99: Red color form. - 100: Yellow and green color forms.
Fig. 101-103. Live Phyllium tobeloense bhaskarai ssp. nov. - 101: Captive adult pair feeding on Psidium guajava (Myrtaceae) (© Alim Yasin). - 102: Orange form female, paratype (© Alim Yasin). - 103: Green form female, paratype (© Edy Bhaskara).
Faunitaxys, 7(4), 2019: 1 — 25. 25
References
Audinet-Serville J. G. A., 1838. — Histoire Naturelle des Insectes. Orthopteéres. Libraire Encyclopédique de Roret, Paris, 18, 776 p.
Bragg P. E., 1996 — Type specimens of Phasmida in the Nationaal Natuurhistorisch Museum, Leiden (Insecta: Phasmida). Zoologische Mededelingen, Leiden, 70(6): 105-115.
Brock P. D., 1999. — Stick and Leaf Insects of Peninsular Malaysia and Singapore. Malaysian Nature Society, Academe Art & Printing Sdn. Bhd., Kuala Lumpur, 222 p.
Brock P. D., Biischer T. H. & Baker E., 2018. — Phasmida SF: Phasmida Species File Version 5.0/5.0. In: Roskov Y., et al., (Eds.), Species 2000 & ITIS Catalogue of Life. Species 2000: Naturalis,
Leiden. Digital resource at www.catalogueoflife.org/col. ISSN 2405-8858. [Accessed January Sth, 2019].
Clark-Sellick J. T., 1997. — Descriptive terminology of the phasmid egg capsule, with an extended key to phasmid genera based on egg structure. Systematic Entomology, 22: 97-122.
Cumming R. T., Valero P. R. & Teemsma S. N., 2018. — Description of Phyllium (Phyllium) conlei, new species, and a first look at the Phylliidae (Phasmatodea) of the Lesser Sunda Islands, Indonesia. Insecta Mundi, 0677: 1-9
Cumming R. T. & Teemsma S. N., 2018. — Phyllium (Phyllium) letiranti sp. nov. (Phasmida: Phylliidae) a new leaf insect from Peleng Island, Indonesia. Jnsecta Mundi, 0618: 1-16.
Cumming R. T., Le Tirant S. & Teemsma S. N., 2018. — On the Phyllium of Peninsular Malaysia and Sumatra, Indonesia, with range expansions for currently known species, description of the previously unknown Phyllium (Pu.) abdulfatahi Seow-Choen female, and description of the new species Phyllium (Ph.) rubrum n. sp. from Peninsular Malaysia (Phasmida: Phylliidae). Faunitaxys, 6(2): 1-21.
de Haan W., 1842. — Bijdragen tot de Kennis der Orthoptera, p. 95-248. Jn: C. J. Temminck (ed.). Verhandelingen over de natuurlijke Geschiedenis der Nederlandsche overzeesche Bezittingen. Vol. 2. Zoologie. S. and J. Luchtmans and C. C. van der Hoek; Leiden, Netherlands, 248 p.
Dickerson R. E., 1928. — Distribution of life in the Philippines. Bureau of Science Monograph, 21: 1-322, plates 1-42.
Gressitt J. L., 1982. — Ecology and biogeography of New Guinean Coleoptera (beetles). In: Gressitt, J. L. [Ed.] Monographiae Biologicae 42, Biogeography and Ecology in New Guinea. Dr W. Junk Publishers, The Hague, pp. 709-734.
Griffini A., 1898. — Intorno al Phyllium geryon Gray. Bollettino dei Musei di Zoologia ed Anatomia comparata della Royal Universita di Torino, 13(312): 1-4.
GréBer D., 2007. — Eine neue Phyllium Art von Halmahera, Nord- Molukken (Phasmatodea: Phylliidae). Arthropoda, 14(3/4), 15-17.
GréBer D., 2008. — Wandelnde Blatter. Ein Katalog aller bisher beschriebenen Phylliinae-Arten und deren Eier mit drei Neubeschreibungen. 2nd Edition. Edition Chimaira, Frankfurt am Main, 175 p.
GréBer D., 2011. — New Insights and Critical Remarks on certain species of Walking Leaves. ARTHROPODA Generalis, 3: 1-17.
Resumé
Ginther K., 1929. — Mitteilungen aus dem Zoologischen Museum, Berlin, 14: 614.
Hausleithner B., 1984. — Eine neue Phyllium-Art aus Malaysia (Phasmatodea: Phylliidae). Entomologische Zeitschrift mit Insektenborse, 94(4): 39-43.
Hennemann F. H., Conle O. V., Gottardo M. & Bresseel J., 2009. — On certain species of the genus Phyllium Illiger, 1798, with proposals for an intra-generic systematization and the descriptions of five new species from the Philippines and Palawan (Phasmatodea: Phylliidae: Phylliinae: Phylliini). Zootaxa, 232: 1-83.
Huxley T. H., 1868. — On the Classification and Distribution of the Alectoromorphae and Heteromorphae. Proceedings of the Zoological Society, London, 294-319.
Illiger J. K. W., 1798. — Verzeichnis der Kafer Preussens. Johann Jacob Gebauer, Halle, 510 pp.
Karny 1914. — Jahresber. K. K. Maximilian-Gymnasium Wien, 14(7).
Klante H., 1976. — Die “Wandelnden Blatter”. Eine taxonomische Revision der Gattung Phyllium Ill. (Insecta, Orthoptera, Phasmatoptera). Zoologische Beitrdge, 22: 49-76.
Linnaeus C., 1758. — Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Laurentii Salvii; Stockholm, 824 p.
Mayr E., 1944. — Wallace’s line in the light of recent zoogeographic studies. The Quarterly Review of Biology, 19(1), 1-14.
Rehn J. A. G. & Rehn J. W. H., 1934[1933]. — On certain species of the genus Phyllium (Orthoptera; Phasmidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 85: 411-427.
Seow-Choen F., 1997. — Stick and Leaf Insect (Phasmida: Insecta) Biodiversity in the Nature Reserves of Singapore. Proceedings of the Nature Reserves Survey Seminar. Gardens’ Bulletin Singapore, 49: 297-312.
Seow-Choen F., 2017. — A taxonomic guide to the stick insects of Borneo, Volume II. Natural History Publications (Borneo), Kota Kinabalu, 261 p.
Simpson G. G., 1977. — Too many lines; the limits of the Oriental and Australian zoogeographic regions. Proceedings of the American Philosophical Society, 121(2): 107-120.
Vallejo B., 2011. — The Philippines in Wallacea. In: Telnov, D. M. [Ed.] Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, Vol. 1. Latvian Society of Entomology, Riga, Latvia, 27-42.
Wallace A. R., 1869. — The Malay Archipelago: The land of the Orang-Utan, and the bird of paradise. A narrative of travel, with studies of man and nature. New York Harper + Brothers, Publishers, Franklin Square, 478 p.
Werner F., 1922. — Zur Kenntnis der Mantodeenfauna der niederlandischen Kolonien. Zoologische Mededeelingen Uitgegeven Vanwege's Rijks Museum van Natuurlijke Historie te Leiden, 7(1-2): 115-126.
Willemse C., 1947. — On Phyllium species, known from the Key Islands. (Orthoptera: Phasmidae, Phylliinae). Tijdschrift Voor Entomologie, 88: 316-322.
Cumming R. T., Le Tirant S. & Hennemann F. H., 2019. — Révision des Phyllium Illiger, 1798 de Wallacea (Indonésie), avec la description d’une nouvelle sous-espéce de l’ile Morotai (Phasmatodea: Phylliidae: Phylliinae). Faunitaxys, 7(4) : 1-25.
Les Phylliidae de Wallacea sont révisés. La répartition des espéces est précisée. Une nouvelle sous-espéce de Phyllium (Phyllium) tobeloense Gréker, 2007 est décrite de l’ile de Morotai, Phyllium (Phyllium) tobeloense bhaskarai Cumming, Le Tirant et Hennemann ssp. nov. dont l’identification est basée principalement sur la morphologie distincte des ceufs. Enfin une clé de tous les Phyllium connus de Wallacea est présentée permettant une identification rapide.
Mots-clés. — Phasmatodea, Phasmida, Phylliinae, Phylliini, Phyllium, Wallacea, Indonesia, Morotai, tobeloense, bhaskarai, description,
espéce nouvelle, distribution, biogeographie.
Faunitaxys
Volume 7, Numero 4, Mars 2019
SOMMAIRE
Révision du genre Phyllium Illiger, 1798 de Wallacea, avec la description d’une nouvelle sous- espéce de l’ile Morotai (Phasmatodea: Phylliidae: Phylliinae).
Royce T. Cumming, Stéphane Le Tirant & Frank H. Hennemann
CONTENTS
Review of the Phyllium Illiger, 1798 of Wallacea, with description of a new subspecies from Morotai Island (Phasmatodea: Phylliidae: Phylliinae).
Royce T. Cumming, Stéphane Le Tirant & Frank H. Hennemann
Mlustration de la couverture: Forét tropicale de Morotai.
Crédit: Fig. 2 © (Google Earth : Image Landsat). Fig. 3-10-11-12-13-14-15-16- 31-34-36-38-4 1-43-45-47-48-63-64-66-69-7 | -74-8 1-84-89-9 1-93-95 © Hennemann F.H., Germany. Fig.-29-32- 49-52 © Natural History Museum, United Kingdom.
Fig. 5-7-9-17-21-22-24-25-26-28-33- 46-61-68 © Senckenberg Deutsches Entomologisches Institut, Miincheberg, Germany. Fig. 4-6-8-18-27-37-42-5 1-53-54-56-57-58 -62-65-67-70-75-76-77-85-86-88 © Limoges R., Insectarium de Montréal, Québec, Canada. Fig. 20-30 © Uppsala University Zoological Museum, Linnean collection, Uppsala, Sweden.
Fig. 23-44 © Zoologische Staatssammlung, Munich, Germany.
Fig. 50-96-98 © Ortiz M., France.
Fig. 59-60 © Cumming R.T., U.S.A.
Fig. 78-80-101-102 & couverture © Yasin A., Indonesia.
Fig. 89-90 © GréBer D., Germany. Fig. 99-100 © Van molle, T, Belgium.
Fig. 103 © Bhaskara E., Indonesia.
Publié par |’ Association Frangaise de Cartographie de la Faune et de la Flore (AFCFF)