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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)
Abstract
Thirteen species of Phyllium (Phyllium) Illiger, 1798 are studied and (re)described in detail with emphasis on those species which exhibit more or less well developed alae in the females and those occurring in the Philippine Islands and on Palawan. Amongst these five new species are described and illustrated from both sexes and the eggs: Ph. (Ph.) ericoriai Hennemann, Conle, Gottardo & Bresseel n. sp. from the Philippine Islands of Luzon, Marinduque and Batan, Phyllium philippinicum Hennemann, Conle, Gottardo & Bresseel n. sp. from the Philippine Island of Luzon, Phyllium mindorense Hennemann, Conle, Gottardo & Bresseel n. sp. from the Philippine Island of Mindoro, Phyllium mabantai Bresseel, Hennemann, Conle & Gottardo n. sp. from the Philippine Island of Mindanao and Ph. (Ph.) gantungense Hennemann, Conle, Gottardo & Bresseel n. sp. from Palawan. Ph. (Ph.) celebicum de Haan, 1842 is re-described with the male and egg described and illustrated for the first time. It is shown to be restricted to Sulawesi and Ambon with all records from continental Asia based on misidentifications mostly relating to Ph. (Ph.) westwoodii Wood-Mason, 1875. All Philippine records of Ph. (Ph.) celebicum de Haan relate to Ph. (Ph.) ericoriai Hennemann, Conle, Gottardo & Bresseel n. sp.. Both sexes and the eggs of Ph. (Ph.) westwoodii Wood-Mason, 1875 are re-described and illustrated and a survey is provided of its intraspecific variability. This species was misinterpreted by most former authors and is here shown to be widely distributed in southern continental Asia having so far been recorded from the Andamans, Myanmar, Thailand, Laos, Kamputchea, S-China, N-Vietnam, Sumatra and the Riouw Archipelago. The holotype of Phyllium (Ph.) siccifolium (Linné, 1758) is described in detail for the first time with illustrations provided. This, the type-species of the entire family Phylliidae, is shown to have been misinterpreted by almost all previous authors and the distribution to be in fact restricted to the Moluccas (Ambon, Ceram, Halmahera, Sula Islands and Banggai). Ambon is shown to be most likely the type-locality of Ph. siccifolium. Records from Peninsular Malaysia have proven to relate to Ph. (Ph.) hausleithneri Brock, 1999 and Philippine material erroneously referred to as "Ph. siccifolium" by various authors is Ph. (Ph.) philippinicum Hennemann, Conle, Gottardo & Bresseel n. sp.. Ph. (Ph.) tobeloense Größer, 2007 from Halmahera (Moluccas) is shown to represent a junior synonym of Ph. siccifolium (n. syn.). Comparison of the Malayan Ph. (Ph.) hausleithneri Brock, 1999 with Malayan specimens previously referred to as "Ph. siccifolium" has revealed these to be the same species which shows considerable variation concerning to the shape of the abdomen in females. Ph. (Ph.) hausleithneri is characteristic for the conspicuous blue interior marking on the meso- and metacoxae. Both sexes and the eggs as well as the remarkable variation of females are illustrated. Similarly strong variation is recorded and illustrated for females of the Javanese Ph. (Ph.) jacobsoni Rehn & Rehn, 1933. A brief discussion of its variability and distribution as well as a summary of the diagnostic features and illustrations of the females and eggs are presented. The Philippine Ph. (Ph.) bilobatum Gray, 1843 is only known from the unique female holotype and all subsequent records appear to have been based on misidentified material. Subsequent records from Peninsular Malaysia relate to Ph. (Ph.) hausleithneri Brock, 1999 and records from Java have all proven to represent Ph. (Ph.) jacobsoni Rehn & Rehn, 1933. The male allotype of Ph. (Ph.) woodi Rehn & Rehn, 1933 from the Philippine island of Mindanao is specifically distinct from the female holotype from Sibuyan Island and here designated as a paratype of Ph. (Ph.) mabantai Bresseel, Hennemann, Conle & Gottardo n. sp.. The diagnostic features of Ph. (Ph.) woodi, a species so far only known from the island of Sibuyan, are briefly summarized. With emphasis on the Philippine fauna, a checklist and keys are provided for the nine species of Phyllium Illiger, 1798 presently known to occur in the Philippine Islands and Palawan. Critical notes are presented on the current intra-generic systematization of Phyllium Illiger, 1798 along with an extended and more detailed distinction between the two subgenera contained, Phyllium Illiger, 1798 and Pulchriphyllium Griffini, 1898. Based on morphological features of the insects and eggs species-groups are suggested within both subgenus. Phyllium (Phyllium) is proposed to include the siccifolium species-group and celebicum species-group, whereas Phyllium (Pulchriphyllium) subdivides into the bioculatum species-group, schultzei species-group, frondosum species-group and brevipenne species-group. The latter two groups are shown to differ considerably from other members of the subgenus and do not belong in Pulchriphyllium (sensu stricto). Keys are provided for the distinction of the speciesgroups here proposed. The celebicum species-group of Phyllium (Phyllium) is discussed in more detail and provisionally contains all those species in which females have developed alae, a fact overlooked for several species by previous authors. Eight species are here attributed to the celebicum species-group and keys are provided to distinguish these. Five species are transferred from one subgenus to the other. Phyllium drunganum Yang, 1995 and Ph. tibetense Liu, 1993 from S-China are removed from the subgenus Pulchriphyllium and transferred to Phyllium (Phyllium) (n. comb.). Ph. chitoniscoides Größer, 1992 and Ph. frondosum Redtenbacher, 1906 from New Guinea as well as Ph. keyicum Karny, 1914 from they Key-Islands are removed from Phyllium (Phyllium) and transferred to the frondosum species-group of Phyllium (Pulchriphyllium) (n. comb.). Ph. insulanicum Werner, 1922 from the Key Islands is removed from synonymy with the New Guinean Ph. frondosum Redtenbacher, 1906 and synoynmised with Ph. keyicum Karny, 1914; differences between Ph. frondosum and Ph. keyicum are presented. The Philippine Phyllium (Phyllium) pusillulum Rehn & Rehn, 1933 is removed from the genus Phyllium Illiger, 1798 and transferred to Microphyllium Zompro, 2001, hence the valid name now is Microphyllium pusillulum (Rehn & Rehn, 1993 n. comb.). Some taxonomically important features traditionally used for distinguishing the genera and species in the family Phylliidae are critically discussed. The present distinction of Chitoniscus Stål, 1875 and Phyllium Illiger, 1798 is shown to be problematic since research on the length relation of the meso-praescutum (anterior portion of the mesonotum in front of the tegmina) has revealed several species in Phyllium Illiger, 1798 that violate the generic description by having this clearly transverse and actually keying out to Chitoniscus Stål, 1875. The prosternal projection characteristic for Chitoniscus Stål, 1875 is shown to be also present in several members of Phyllium Illiger, 1798. Although the entire family Phylliidae was traditionally diagnosed by females having the antennae with nine segment, six species of Phyllium (Phyllium) Illiger, 1798 are here shown to have in fact ten antennomeres. Another interesting fact are the distinctly pectinate ungues (= claws) seen in Ph. (Ph.) gantungense n. sp. which have so far only been known to occur in the Old World areolate family Aschiphasmatidae.
... All three of the herein described genera have previously been recognized (fully or partially) at one time or another as distinct to some degree. Within the substantial phylliid revision by Hennemann et al. (2009) their intrageneric systematizations recognized several Melanesian clades, namely the schultzei species group, frondosum species group, and the brevipenne species group (all of which at the time were nested within the Pulchriphyllium Griffini, 1898 sensu lato which was thought to be a subgenus of Phyllium Illiger, 1798). Most of the frondosum species group has since been found to be the previously unrecognized females to the Nanophyllium (suspected by Brock and Hasenpusch (2003); confirmed by Cumming et al. (2020a)) but the other clades have not yet been extensively reviewed and are therefore the focus of this work. ...
... Additionally, in reviewing the Nanophyllium sensu lato, this distinct clade was recovered as bifurcate with significant divergence distance between two internal sister clades. These sister clades correspond to two morphologically distinct groups which have been recognized by past authors as significant (Hennemann et al. 2009;Cumming et al. 2020a). Essentially taxonomic ranks above the species level are arbitrarily assigned in order to facilitate communication about natural groups, therefore these two morphologically distinct sister clades based upon the current treatment of genera within the phylliids clearly align themselves with the genus concept well. ...
... This new genus has been recognized as unique in the past, as this clade was designated as the schultzei species group within Hennemann et al. (2009) based upon the shorter tegmina and the two lobes on the exterior protibiae. While these features are helpful to differentiate this genus from others, the below noted autapomorphic features allow differentiation from all phylliid genera. ...
With the first large-scale Phylliidae molecular phylogeny recently published adding a great deal of clar- ity to phylliid diversity, several of the rarer species which could not be included were methodically and morphologically reviewed. This review resulted in identification of numerous substantial morphological features that suggest there are Melanesian clades that create polyphyletic groups within the phylliids which should instead be taxonomically recognized as unique. These rarer Melanesia species have historically been considered to be southern representatives of the Pulchriphyllium Griffini, 1898 sensu lato. However, there are notable morphological differences between the Pulchriphyllium sensu stricto and the “schultzei” group. Therefore, two new genera are erected, Vaabonbonphyllium gen. nov. from the Solomon Islands and Papua New Guinea and Rakaphyllium gen. nov. from New Guinea and the Aru Islands. Erection of these two new genera warrants the following new combinations: Rakaphyllium schultzei (Giglio-Tos, 1912), comb. nov., Rakaphyllium exsectum (Zompro, 2001b), comb. nov., and Vaabonbonphyllium groesseri (Zompro, 1998), comb. nov. Additionally, while reviewing material an undescribed Vaabonbonphyllium gen. nov. specimen was located and is herein described as Vaabonbonphyllium rafidahae gen. et sp. nov. from Mt. Hagen, Papua New Guinea. Additionally, a morphologically unique clade of several species recovered as sister to the Nanophyllium sensu stricto was recognized and their numerous unique morphological features and monophyly leads the authors to erect the new genus Acentetaphyllium gen. nov. which warrants the following new combinations: Acentetaphyllium brevipenne (Größer, 1992), comb. nov., Acentetaphyllium larssoni (Cumming, 2017), comb. nov., Acentetaphyllium miyashitai (Cumming et al. 2020), comb. nov., and Acentetaphyllium stellae (Cumming, 2016), comb. nov. With the addition of several new genera, a key to phylliid genera is included for adult males and females.
... We quantified the effects of variation in morphology (body size and shape) on flight and substrate attachment performance in male leaf insects (Phyllium philippinicum, Hennemann, Conle, Gottardo & Bresseel, 2009, Phylliidae, Phasmatodea). In this solitary canopy-dwelling species, large, sedentary adult females are outstanding leaf-mimics due to lateral 'leaf-like' expansions of the abdominal segments and legs [24][25][26]. Adult males are nine times lighter and almost two times slenderer than females, and have relatively longer antennae (i.e., strong size [SSD] and shape dimorphism; Fig. 1). Adult females lack hindwings but have extended forewings that lie flat on their dorsum, aiding in camouflage [24]. ...
... Adult males are nine times lighter and almost two times slenderer than females, and have relatively longer antennae (i.e., strong size [SSD] and shape dimorphism; Fig. 1). Adult females lack hindwings but have extended forewings that lie flat on their dorsum, aiding in camouflage [24]. These wings cannot flap but can contribute to parachuting if falling. ...
... Colors correspond to different body sizes relative to the original model Table 4 The effects of body size and shape on various aerodynamic variables, as predicted by the CFD models Results of type I ANOVA from linear models contrasting the effects of body length and body aspect ratio on various aerodynamic variables (Fig. 6) their static safety factors for both adhesion and friction forces were still relatively lower than those of males. Male and female leaf insects also exhibit a spectacular interspecific variation in body shape related to leaf mimicry [24,26,61]. This variation is likely driven by masquerade and the advergence of the insect appearance to resemble the size and shape of its host plants' leaves. ...
Background
In most arthropods, adult females are larger than males, and male competition is a race to quickly locate and mate with scattered females (scramble competition polygyny). Variation in body size among males may confer advantages that depend on context. Smaller males may be favored due to more efficient locomotion leading to higher mobility during mate searching. Alternatively, larger males may benefit from increased speed and higher survivorship. While the relationship between male body size and mobility has been investigated in several systems, how different aspects of male body morphology specifically affect their locomotor performance in different contexts is often unclear.
Results
Using a combination of empirical measures of flight performance and modelling of body aerodynamics, we show that large body size impairs flight performance in male leaf insects ( Phyllium philippinicum ), a species where relatively small and skinny males fly through the canopy in search of large sedentary females. Smaller males were more agile in the air and ascended more rapidly during flight. Our models further predicted that variation in body shape would affect body lift and drag but suggested that flight costs may not explain the evolution of strong sexual dimorphism in body shape in this species. Finally, empirical measurements of substrate adhesion and subsequent modelling of landing impact forces suggested that smaller males had a lower risk of detaching from the substrates on which they walk and land.
Conclusions
By showing that male body size impairs their flight and substrate adhesion performance, we provide support to the hypothesis that smaller scrambling males benefit from an increased locomotor performance and shed light on the evolution of sexual dimorphism in scramble competition mating systems.
... While Bradler and Buckley 10 noted that Phylliidae only account for <2% of the phasmatodean diversity with about 50 known species, the number of described species has now doubled within just a few years. Misidentification, overestimation of species' distributions and the unreliability of the highly variable morphological traits 35 had resulted in a chaotic taxonomy that only recently started to be overcome by extensive morphological examinations (e.g., Cumming et al. 36,37 ). Captive breeding and molecular analysis have further helped to shed light on the phylogenetic relationships and to match up males and females of leaf insects [38][39][40] . ...
... According to the most recent studies, Phylliidae currently includes six genera (Chitoniscus, Cryptophyllium, Microphyllium, Nanophyllium, Phyllium and Pseudomicrophyllium) with most species pertaining to Phyllium, which is further divided into four subgenera (Comptaphyllium, Phyllium, Pulchriphyllium and Walaphyllium). Both Phyllium and Pulchriphyllium have undergone further intra-generic systematisation and were classed in several species groups by Hennemann et al. 35 . One of these species groups was recently revealed to be distinct to the remaining phylliids and was therefore transferred to the newly erected genus Cryptophyllium 40 . ...
... Two species originally assigned to Phyllium were not found to belong to any of the former Phyllium groups: Phyllium (Pulchriphyllium) brevipenne and Phyllium (Phyllium) geryon. The former had previously been suggested to be closely related to the frondosum species group 35 , a clade that was recently revealed to belong to Nanophyllium 39 . As our phylogenetic inferences recover P. brevipenne as the sister taxon to Nanophyllium, the species is hereby transferred to Nanophyllium as Nanophyllium brevipenne comb. ...
The insect order Phasmatodea is known for large slender insects masquerading as twigs or bark. In contrast to these so-called stick insects, the subordinated clade of leaf insects (Phylliidae) are dorso-ventrally flattened and therefore resemble leaves in a unique way. Here we show that the origin of extant leaf insects lies in the Australasian/Pacific region with subsequent dispersal westwards to mainland Asia and colonisation of most Southeast Asian landmasses. We further hypothesise that the clade originated in the Early Eocene after the emergence of angiosperm-dominated rainforests. The genus Phyllium to which most of thẽ 100 described species pertain is recovered as paraphyletic and its three non-nominate subgenera are recovered as distinct, monophyletic groups and are consequently elevated to genus rank. This first phylogeny covering all major phylliid groups provides the basis for future studies on their taxonomy and a framework to unveil more of their cryptic and underestimated diversity.
... Taxonomically, Phyllium is further divided into four subgenera: Phyllium, Pulchriphyllium, and the recently described Comptaphyllium and Walaphyllium (Cumming et al. 2019;Cumming et al. 2020b). For the two traditional subgenera Phyllium and Pulchriphyllium, an intra-generic systematization had been proposed in order to facilitate differentiation in a taxonomical context (Hennemann et al. 2009). The subdivision of the most diverse subgenus Phyllium (Phyllium) into the siccifolium and celebicum species groups was mainly based on the presence of developed alae in females of the latter, a problematic character in a group with strong sexual dimorphism and several species only known from single sexes (e.g., Liu 1993;Yang 1995). ...
... Currently, there are two divisions of leaf insects within the singular Phylliinae, the Phylliini Brunner von Wattenwyl, 1893 (which contains the bulk of genera and species) and the Nanophylliini Zompro & Groesser, 2003 (which only contains the Nanophyllium Redtenbacher, 1906) (Brock et al. 2020). Our molecular and morphological review has warranted the description of a novel genus of leaf insect to accommodate the former celebicum species group (sensu Hennemann et al. 2009), which is erected herein. ...
... We chose this name with a double meaning in mind. First, this clade of leaf insects has been "hidden" within the true Phyllium, and only recently have they been noted as unique and placed within the celebicum species group by Hennemann et al. (2009) of Phyllium, but still not completely recognized as separate. Secondly, these insects are masters of disguise and well-hidden within their canopy habitat ( Fig. 1) which means only rarely are individuals found. ...
While the leaf insects (Phylliidae) are a well-supported group within Phasmatodea, the genus Phyllium Illiger, 1798 has repeatedly been recovered as paraphyletic. Here, the Phyllium (Phyllium) celebicum species group is reviewed and its distinctiveness from the remaining Phylliini genera and subgenera in a phyloge- netic context based on morphological review and a phylogenetic analysis of three genes (nuclear gene 28S and mitochondrial genes COI and 16S) from most known and multiple undescribed species is shown. A new genus, Cryptophyllium gen. nov., is erected to partially accommodate the former members of the celebicum species group. Two species, Phyllium ericoriai Hennemann et al., 2009 and Phyllium bonifacioi Lit & Eusebio, 2014 morphologically and molecularly do not fall within this clade and are therefore left within Phyllium (Phyllium). The transfer of the remaining celebicum group members from Phyllium Illiger, 1798 to this new genus creates the following new combinations; Cryptophyllium athanysus (Westwood, 1859), comb. nov.; Cryptophyllium celebicum (de Haan, 1842), comb. nov.; Cryptophyllium chrisangi (Seow-Choen, 2017), comb. nov.; Cryptophyllium drunganum (Yang, 1995), comb. nov.; Cryptophyl- lium oyae (Cumming & Le Tirant, 2020), comb. nov.; Cryptophyllium parum (Liu, 1993), comb. nov.; Cryptophyllium rarum (Liu, 1993), comb. nov.; Cryptophyllium tibetense (Liu, 1993), comb. nov.; Crypto- phyllium westwoodii (Wood-Mason, 1875), comb. nov.; Cryptophyllium yapicum (Cumming & Teemsma, 2018), comb. nov.; and Cryptophyllium yunnanense (Liu, 1993), comb. nov.
The review of specimens belonging to this clade also revealed 13 undescribed species, which are described within as: Cryptophyllium animatum gen. et sp. nov. from Vietnam: Quang Nam Province; Cryptophyllium bankoi gen. et sp. nov. from Vietnam: Quang Ngai, Thua Thien Hue, Da Nang, Gia Lai, Quang Nam, and Dak Nong Provinces; Cryptophyllium bollensi gen. et sp. nov. from Vietnam: Ninh Thuan Province; Cryptophyllium daparo gen. et sp. nov. from China: Yunnan Province; Cryptophyllium echidna gen. et sp. nov. from Indonesia: Wangi-wangi Island; Cryptophyllium faulkneri gen. et sp. nov. from Vietnam: Quang Ngai and Lam Dong Provinces; Cryptophyllium icarus gen. et sp. nov. from Vi- etnam: Lam Dong and Dak Lak Provinces; Cryptophyllium khmer gen. et sp. nov. from Cambodia: Koh Kong and Siem Reap Provinces; Cryptophyllium limogesi gen. et sp. nov. from Vietnam: Lam Dong, Dak Lak, and Dak Nong Provinces; Cryptophyllium liyananae gen. et sp. nov. from China: Guangxi Province; Cryptophyllium nuichuaense gen. et sp. nov. from Vietnam: Ninh Thuan Province; Cryptophyllium phami gen. et sp. nov. from Vietnam: Dong Nai and Ninh Thuan Provinces; and Cryptophyllium wennae gen. et sp. nov. from China: Yunnan Province. All newly described species are morphologically described, il- lustrated, and molecularly compared to congenerics.
With the molecular results revealing cryptic taxa, it was found necessary for Cryptophyllium westwoo- dii (Wood-Mason, 1875), comb. nov. to have a neotype specimen designated to allow accurate differen- tiation from congenerics. To conclude, male and female dichotomous keys to species for the Cryptophyl- lium gen. nov. are presented.
... Zompro & Grösser (2003): A generic revision. Hennemann et al. (2009): A revision of species from the Philippines and reclassification, including clarification of several identifications. ...
... Sexual dimorphism: Female not known, but from the author's evaluation of other taxa, is expected to have a very rounded abdomen. Diagnosis: Closely related to Phyllium (Phyllium) siccifolium (Linnaeus, 1758) and placed in the Phyllium siccifolium group [the distribution range of P. siccifolium is Moluccas: Halmahera, Ambon, Seram, Sula Islands, Banggai Island and Buru (Hennemann et al. 2009), although further research may be needed on the distribution range as this involves Indonesian islands separated by deep water straights. Grösser (2011)] has already disputed the arrangement (see 'Discussion' section). ...
... There is disagreement over taxonomy of certain Phyllium species, with a history of misidentifications, repeated descriptions of the same species and general confusion (there are 52 species of Phyllidae, many described from a single specimen, yet there are another 21 invalid species names (Brock 2013)). For example, Phyllium (Phyllium) tobeloense Grösser, 2007 was regarded as a synonym of Phyllium (Phyllium) siccifolium by Hennemann et al. (2009) but returned to valid species status by Grösser, 2011, although doubts still remain over the correct placement, with a convincing discussion by Hennemann et al. (2009). For the time being the species are separated in the online Phasmida Species File (Brock 2013), although this only reflects the most recent taxonomic change, pending further evaluation by specialists. ...
A new leaf insect species Phyllium (Phyllium) telnovi is described based on a single male specimen from the Tamarau Mountains, West Papua, Indonesia. An overview is also provided on leaf insect (Phylliidae) taxonomy. In: Telnov [Ed.]. Biodiversity, Biogeography and Nature Conservation in Wallacea and New Guinea, volume II
... Phyllium philippinicum Hennemann, Conle, Gottardo and Bresseel, 2009 (Phylliidae) is a species of leaf insect commonly bred in labs and private cultures ( Figure 1A). However, most of the literature on the species revolves around taxonomic and phylogenetic classification and is mainly based on adult morphology [3,[40][41][42]. Leaf insects in general are reported to drop or catapult their egg for deposition from the canopy tops of their host tree [2,13]. ...
... Only very few taxonomic studies hypothesized the function of these pinnae. Hennemann et al. [3] described the unfolding of the pinnae after their contact with water and suggested an adhesive function of this system, however did not further elucidate this idea. Additionally curious, the oviposition technique employed by the females, does not involve active gluing of the eggs, which begs the question of whether there is Insects 2020, 11, 400 3 of 20 a presence or absence of accessory reproductive glands in P. philippinicum. ...
... This masquerade, imitating parts of their environment, is particularly striking in the lineage Phylliidae (leaf insects). Consequently, these insects are commonly called "walking leaves" [3][4][5]. Leaf insects extraordinarily imitate the leaves of plants and visually merge with their environment. The first fossil records of Phylliidae date back 47 mya with Eophyllium messelensis Wedmann, Bradler and Rust 2006 as the oldest known representative of this lineage [6]. ...
Leaf insects (Phasmatodea: Phylliidae) exhibit perfect crypsis imitating leaves. Although the special appearance of the eggs of the species Phyllium philippinicum, which imitate plant seeds, has received attention in different taxonomic studies, the attachment capability of the eggs remains rather anecdotical. We herein elucidate the specialized attachment mechanism of the eggs of this species and provide the first experimental approach to systematically characterize the functional properties of their adhesion by using different microscopy techniques and attachment force measurements on substrates with differing degrees of roughness and surface chemistry, as well as repetitive attachment/detachment cycles while under the influence of water contact. We found that a combination of folded exochorionic structures (pinnae) and a film of adhesive secretion contribute to attachment, which both respond to water. Adhesion is initiated by the glue, which becomes fluid through hydration, enabling adaption to the surface profile. Hierarchically structured pinnae support the spreading of the glue and reinforcement of the film. This combination aids the egg’s surface in adapting to the surface roughness, yet the attachment strength is additionally influenced by the egg’s surface chemistry, favoring hydrophilic substrates. Repetitive detachment and water-mediated adhesion can optimize the location of the egg to ensure suitable environmental conditions for embryonic development. Furthermore, this repeatable and water-controlled adhesion mechanism can stimulate further research for biomimeticists, ecologists and conservationalists.
... After reviewing congenerics, we found that not only was this specimen an undescribed species but that there were two other well-known species which formed a unique clade separate from other Phyllium (Phyllium). We here transfer these two species (Phyllium monteithi Brock & Hasenpusch, 2003 and Phyllium zomproi Größer, 2001), from the siccifolium group of Phyllium (Phyllium) as described in Hennemann et al. (2009), and include the herein described Phyllium lelantos sp. nov. ...
... Female Phyllium monteithi can be differentiated from Phyllium zomproi by several morphological features. One is the number of teeth on the stridulatory file of the third antennal segment with 27 to 29 teeth on P. monteithi (Fig. 10C) and 48 to 50 teeth noted on P. zomproi (Hennemann et al. 2009). There is a notable difference in body size between the two species, with P. monteithi a medium sized species ranging from 75.0-76.0 ...
... First, the two previously described species we here transfer to Walaphyllium (P. monteithi and P. zomproi) have historically been placed within the morphologically diverse siccifolium species group of the Phyllium subgenus, which concealed their unique combination of features (Hennemann et al. 2009). It wasn't until we tabulated the morphological diversity of the Phyllium subgenera (see Table 1), that we realized that the Walaphyllium clade essentially contained a mixture of morphological features which are shared between the Pulchriphyllium, Comptaphyllium, and/or the Phyllium subgenera. ...
A new subgenus, Walaphylliumsubgen. nov. , is described within Phyllium Illiger, 1798 to accommodate three leaf insect species. One of the species included is newly described herein as Phyllium (Walaphyllium) lelantos sp. nov. from Papua New Guinea. This new subgenus of Phyllium can be diagnosed by a following combination of features. This new species is compared to the two additional new subgenus members, Phyllium zomproi Größer, 2001 and Phyllium monteithi Brock & Hasenpusch, 2003. Also for the first time the male morphology of Phyllium zomproi is described and illustrated. To conclude, a brief biogeographical view of the leaf insects on either side of the Torres Strait is presented, as well as a key to species and a distribution map to the known species of Phyllium (Walaphyllium) subgen. nov.
... Two of the three Phyllium species on Palawan illustrate this close biogeographic relationship as their morphological sister species can be found on Borneo. Phyllium gantungense Hennemann, et al., 2009 is sister species to Phyllium arthurchungi Seow-Choen, 2016, and Phyllium cummingi Seow-Choen, 2017, both currently only known from Northern Borneo. And Phyllium palawanense Grösser, 2001 is sister species to Phyllium chenqiae Seow-Choen, 2017 from Northern Borneo. ...
... Comparatively, Phyllium saltonae n. sp. appears to be more morphologically related to known species of Phyllium from the Philippines such as Phyllium philippinicum Hennemann, et al., 2009, Phyllium mindorense Hennemann, et al., 2009, and Phyllium mabantai Bresseel, et al., 2009, than it does to any known Bornean species. ...
... This species is remarkable for its extreme variation observed within females, with abdominal shape ranging from nearly perfectly smooth (Fig. 2A) to intricately lobed (Fig. 2C). As noted by Hennemann, et al., (2009) and Cumming (2017) the abdominal shape (ranging from smooth to strongly lobed) directly corresponds to the size of the profemoral interior lobe teeth (small serrate teeth to large triangular teeth, . When viewed in a large set, these variations fall within a continuous series, not in discrete forms. ...
Prompted by the discovery of an undescribed species of Phyllium Illiger, 1798 from within the Natural History Museum, London, United Kingdom collection, the Phylliidae (Leaf Insects) of Palawan, Philippines are here reviewed. The new species Phyllium (Phyllium) saltonae Cumming, Baker, Le Tirant, & Marshall n. sp. is currently only known from female and egg specimens and represents the third species of leaf insect on the island. Taxonomically this new species belongs to the siccifolium species group of Phyllium (Phyllium) as per Hennemann, et al., 2009. To conclude, a key to species of Palawan is included and all species are illustrated.
... Review of leaf-insects collected in Northeastern Laos has revealed a previously unnamed species, which we here describe as Phyllium (Phyllium) oyae species nova. This new species falls within the celebicum species group as described in Hennemann et al. (2009) due to the presence of well developed alae in females and males with a wide exterior profemoral lobe. Upon a review of congenerics, we find that Phyllium (Phyllium) drunganum Yang, 1995 is misplaced and transfer it from the siccifolium species group to the celebicum species group based on the presence of developed alae in females. ...
... After reviewing in detail the material we received from Laos however, it became apparent that the structure of the antennae was consistently different from Phyllium drunganum, and the Laos population Keywords : instead represented an undescribed species. This species is compared below to all members within the celebicum species group as described by Hennemann, et al., 2009 and is named Phyllium oyae n. sp. after Steeve Collard's wife as he first sent us specimens of this new species and has been incredibly helpful in supplying fresh material and photographs of the habitat (Fig. 2). ...
... -Since the original description, where Phyllium drunganum was not placed within a particular subgenus (Yang, 1995& Chen, 1999, there has been significant confusion about the taxonomic placement of this species. Some authors have placed Phyllium drunganum within Phyllium (Pulchriphyllium) based on all tibiae with small exterior lobes (Zompro, 2004;Größer, 2008) and others in the Phyllium (Phyllium) subgenus (Otte & Brock, 2005;Hennemann, et al., 2009;Chen & He, 2008). Most recently with the arrangement of species groups by Hennemann et al. (2009), Phyllium drunganum was returned to the Phyllium (Phyllium) subgenus, but unfortunately it was placed within the siccifolium species group without morphological justification. ...
Review of leaf-insects collected in Northeastern Laos has revealed a previously unnamed species, which we here describe as Phyllium (Phyllium) oyae species nova. This new species falls within the celebicum species group as described in Hennemann et al. (2009) due to the presence of well developed alae in females and males with a wide exterior profemoral lobe. Upon a review of congenerics, we find that Phyllium (Phyllium) drunganum Yang, 1995 is misplaced and transfer it from the siccifolium species group to the celebicum species group based on the presence of developed alae in females. With so many of the species in this group only known from a single sex, we differentiate Phyllium oyae n. sp. from all species. The species name Phyllium rayongii Thanasinchayakul, 2006 is determined to be a nomen nudum and therefore unavailable according to ICZN Article 16.4.1. To conclude, we present dichotomous keys to males and females known within the celebicum species group.
... Review of leaf-insects collected in Northeastern Laos has revealed a previously unnamed species, which we here describe as Phyllium (Phyllium) oyae species nova. This new species falls within the celebicum species group as described in Hennemann et al. (2009) due to the presence of well developed alae in females and males with a wide exterior profemoral lobe. Upon a review of congenerics, we find that Phyllium (Phyllium) drunganum Yang, 1995 is misplaced and transfer it from the siccifolium species group to the celebicum species group based on the presence of developed alae in females. ...
... After reviewing in detail the material we received from Laos however, it became apparent that the structure of the antennae was consistently different from Phyllium drunganum, and the Laos population Keywords : instead represented an undescribed species. This species is compared below to all members within the celebicum species group as described by Hennemann, et al., 2009 and is named Phyllium oyae n. sp. after Steeve Collard's wife as he first sent us specimens of this new species and has been incredibly helpful in supplying fresh material and photographs of the habitat (Fig. 2). ...
... -Since the original description, where Phyllium drunganum was not placed within a particular subgenus (Yang, 1995& Chen, 1999, there has been significant confusion about the taxonomic placement of this species. Some authors have placed Phyllium drunganum within Phyllium (Pulchriphyllium) based on all tibiae with small exterior lobes (Zompro, 2004;Größer, 2008) and others in the Phyllium (Phyllium) subgenus (Otte & Brock, 2005;Hennemann, et al., 2009;Chen & He, 2008). Most recently with the arrangement of species groups by Hennemann et al. (2009), Phyllium drunganum was returned to the Phyllium (Phyllium) subgenus, but unfortunately it was placed within the siccifolium species group without morphological justification. ...
... With the description of the very morphologically similar Phyllium gardabagusi sp. nov., also from Java, the numerous localities noted within Hennemann et al. (2009) cannot be confirmed as Phyllium jacobsoni. This is because the only clear morphological feature to differentiate between these two populations is the coxae color, which can fade in specimens which were not well preserved. ...
... Unconfirmed distributions: Malaysia: Selangor State, Bukit Kutu. This record was noted as Phyllium siccifolium in Brock (1999) and treated as a smooth abdominal form of P. hausleithneri in Hennemann et al. (2009), however, without examining the specimen in question it is best to leave this as an unconfirmed record. This is because the morphologically similar Phyllium rubrum Cumming, Le Tirant, & Teemsma, 2018 is also found in Peninsular Malaysia and has in the past been considered a form of P. hausleithneri. ...
... This species has been in the phasmid breeding community for many years ( Figure 6A-E), with most of those years being sold erroneously as Phyllium siccifolium which has led to much of the confusion surrounding this species. To help clear up confusion, the morphological variations and numerous misidentifications were presented in the discussion of Phyllium hausleithneri in Hennemann et al. (2009). Phyllium hausleithneri from Peninsular Malaysia was morphologically compared extensively to P. nisus sp. ...
Within the last two years, the leaf insects of the genus Phyllium of both the islands of Java and Sumatra have been reviewed extensively based on morphological observations. However, cryptic species which cannot be differentiated morphologically may be present among the various populations. Since it has frequently been demonstrated that analyses based on molecular data can bring clarity in such cases, we conducted a phylogenetic analysis based on three genes (nuclear gene 28S and mitochondrial genes COI and 16S) from the Phyllium species of these islands. The results show distinct molecular divergence for several populations and suggest the presence of two new cryptic species, morphologically inseparable from Phyllium hausleithneri Brock, 1999. From Sumatra, the population originally thought to be a range expansion for Phyllium hausleithneri, is now here described as Phyllium nisus sp. nov., with the only consistent morphological difference being the color of the eggs between the two populations (dark brown in P. hausleithneri and tan in P. nisus sp. nov.). Further, an additional population with purple coxae from Java was morphologically examined and found to have no consistent features to separate it morphologically from the other purple coxae species. This cryptic species from Java was however shown to be molecularly distinct from the other purple coxae populations from Sumatra and Pen-insular Malaysia and is here described as Phyllium gardabagusi sp. nov. In addition, Phyllium giganteum is here officially reported from Java for the first time based on both historic and modern records of male specimens.
... 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 and Lombok (Cumming, Valero & Teemsma, 2018). ...
... Faunitaxys, 7(4), 2019 : 1 -25. 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. ...
... 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 Discussion. ...
The Phylliidae of Wallacea are here reviewed, with notes on species distributions. Phyllium (Phyllium) tobeloense Größer, 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.
... The Phyllium (Pulchriphyllium) bioculatum Gray, 1832 comes under the family Phyllidae. The genus Phyllium Illiger, 1798 is further sub-divided into 2 sub-genus, Phyllium and Pulchriphyllium Griffini, 1898 [4] . The family Phyllidae Redtenbacher, 1908 contains 5 species [2] of both of these sub genera [3] from India. ...
... Based on the external morphological characters and literature by Zompro O, Größer D(2003) [7] , Hennemann F, Conle O, Gottardo M, Bresseel J(2009) [4] , Mandal SK, Yadav K(2010) [3] and Srinivasan G, Surendar C, Chatterjee P, Mukherjee TK(2017) [6] all the species was identified as male Phyllium (Pulchriphyllium) bioculatum Gray, 1832. The detail of the species is discussed below:- ...
... The species is diagnosed under the genus Phyllium Illiger, 1798 for having the prominent and broad exterior lobe of the tibiae, and the mesonotum in front of tegmina which is about quadrate but not remarkably transverse [4,7] . Further, the species is characterized by the subgenus Pulchriphyllium Griffini, 1898 by the presence of interior lobes on the prefemora [7] . ...
Phyllium (Pulchriphyllium) bioculatum Gray, 1832 (Phasmida) is a leaf insect which is the most incredible, attractive and wonderful one. It was reported earlier from some places of India viz., Andaman and Nicobar Islands, West Bengal and Assam. This is for the first time report from Medziphema in Dimapur district of Nagaland, NorthEast India. Three male individuals of this species were collected from the light source. The species has been properly diagnosed and the description has been made based on the morphological characters using morphometric measurement. To know the distribution pattern, abundance, behavior and biology of this amazing species in this region further study is needed.
... Phyllium (Phyllium) yapicum Cumming and Teemsma n. sp., with a broad thorax and parallel sided abdomen, appears to be most morphologically similar to Phyllium (Phyllium) ericoriai Hennemann et al. (2009), Phyllium (Phyllium) bonifacioi Lit and Eusebio (2014), and Phyllium (Phyllium) tibetense Liu (1993). Geographically, of these three species, Phyllium (Ph.) ericoriai, and Phyllium (Ph.) bonifacioi, logically fit as the closest relations, because these two species are native to the Philippines, the island nation to the west of Micronesia. ...
... Differentiation. The exterior lobe of the profemora is wider than the interior lobe and with developed alae, Phyllium (Phyllium) yapicum n. sp. is placed within the celebicum species group, as described by Hennemann et al. 2009. Phyllium (Ph.) yapicum n. sp. ...
... Adapted from the Hennemann et al. (2009) key. Distributional data cited in this key obtained from Hennemann et al. (2009), Cumming et al. (2017), Thanasinchayakul (2006), and Seow-Choen (2017). ...
A new species of leaf insect from the celebicum species group, Phyllium (Phyllium) yapicum Cumming and Teemsma, new species (Phasmida: Phylliidae), is described from a female specimen from the California Academy of Sciences collection, United States. This new species is the first recorded species of Phylliidae from the country of Micronesia and represents a notable range expansion for the family. With Phyllium (Phyllium) yapicum Cumming and Teemsma, new species, currently only known from a female holotype; a key to females is included for the celebicum species group.
... The final feature reviewed on the female specimens was the number of teeth on the pars stridens of antennomere III, a feature that has proven useful in species differentiation when larger features seemed to be difficult to keep separated in morphological keys. Größer in 2008 lists Ph. (Ph.) hausleithneri as having a pars stridens with 44-46 teeth, and the following year Hennemann et al., 2009 then listed it as having 44-48 teeth, a moderately wide range for a species. The number of teeth observed on the Sumatran populations pars stridens was again eye opening to the wide range of morphologies of this species with a range of 37-44 teeth. ...
... rarely offered. The lack of exterior lobes and the rudimentary alae place this new species within the siccifolium species-group within the (Phyllium) subgenus as per Hennemann et al., 2009. It was originally planned to wait until a male specimen could be obtained to describe the species, however, due to the lack of an official name within the phasmid breeding community, a description of at least the female was warranted to help clear up current confusion and it is unlikely that a male will be obtained in the near future due to the species apparent rarity. ...
... (Ph.) bradleri appear more closely related to Ph. (Ph.) jacobsoni, a congeneric which Seow-Choen failed to differentiate both species from. Eggs, morphologically are less similar to Ph. (Ph.) hausleithneri and are instead more similar in their pinnae pattern on the lateral view to those of Ph. (Ph.) philippinicum Hennemann, et al., 2009. Instead of a pinnae pattern on the lateral view with straight marginal rows and a center row that varied in pattern from straight sided to intricate, Ph. (Ph.) philippinicum eggs by the micropylar plate which is narrow and covers approximately 5/6 th of the dorsal view ( in Ph. (Ph.) philippinicum the micropylar plate only covered around 2/3rds of the dorsal view and is wider with each side from the micropylar cup tapering evenly). ...
Abstract. – The Phyllium diversity of Peninsular Malaysia and Sumatra, Indonesia is discussed with records of recent collection and from past literature. The previously unknown female Phyllium (Pulchriphyllium) abdulfatahi Seow-Choen, 2017a. is described along with a discussion of the range in which the species can be found and figures of the female. A range expansion into Sumatra and additional regions of Borneo for Phyllium (Pulchriphyllium) giganteum Hausleithner, 1984 is discussed along with figures of corresponding specimens.
Phyllium (Phyllium) hausleithneri Brock, 1999 is also newly recorded from Sumatra and a discussion on intraspecies variation with corresponding figures of females and eggs. From Peninsular Malaysia, a new species Phyllium (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. is described and illustrated from Perak, Malaysia. Phyllium (Pu.) mannani Seow-Choen, 2017a. and Phyllium (Pu.) rimiae Seow-Choen, 2017a. ranges are expanded to include Peninsular Malaysia, and Phyllium (Pu.) bioculatum Gray, 1832 is expanded to include
Sumatra, with discussion on the intraspecies variation of males of all three species included. To conclude, with Phyllium (Ph.) rubrum Cumming, Le Tirant & Teemsma n. sp. described from a female, a key to females of all currently described species of Phyllium from Sumatra, Singapore, and West Malaysia is included.
... With the lack of exterior lobes of the tibiae and simple antennae this new species is placed within the subgenus Phyllium (Phyllium), species-group placement however is tentative. Hennemann et al., 2009 describe the males of the siccifolium species-group as having an exterior lobe of the profemora which is always distinctly narrower than the interior lobe, a feature only weakly accurate for Ph. (Ph.) antonkozlovi sp. ...
... Upon closer examination, several differences became clear. Like Ph. (Ph.) philippinicum, this new species falls under the siccifolium species-group of Hennemann, et al., 2009 because the female lacks developed alae and the male has an exterior lobe of the profemora that is more slender than the interior lobe. It is possible that future genetic analysis will find that these two populations are not conspecific, but the marked differences in the female genitalia led us to here describe them with species level status. ...
... Gray's original distribution note of "Inhabits the Philippine Islands", has led to much confusion over the years and erroneous records. Several authors over the years have mistakenly noted Ph. (Ph.) geryon from localities such as Java or New Caledonia (see Hennemann et al., 2009 for a thorough explanation of these erroneous localities). ...
Examination of unidentified Phylliidae specimens revealed a number of undescribed species from the island of Luzon, Philippines. Morphological and molecular study of specimens from the obscure phasmid genus Microphyllium Zompro, 2001, revealed a new species, which we describe as Microphyllium haskelli Cumming sp. nov.. It is here described and differentiated from the two other species in the genus, both currently only known from adults of a single sex. Pseudomi-crophyllium Cumming gen. nov. is described as a new genus within Phylliidae with the type species Pseudomicrophyllium faulkneri Cumming gen. et sp. nov. as the sole known species in the genus. As is unfortunately often the case in the leaf-mimicking family Phylliidae, this new genus and species is only known from a single specimen. In addition to the new genus, two new Phyllium (Phyllium) species from the siccifolium species-group are named and described as Ph. (Ph.) an-tonkozlovi Cumming sp. nov. and Ph. (Ph.) bourquei Cumming & Le Tirant sp. nov.. In addition to the newly described species, Phyllium (Phyllium) geryon Gray, 1843 is redescribed from a nearly perfect specimen, completing some of the morphological knowledge gaps currently missing because of the severely damaged holotype specimen. A key to all known species of Phylliidae from Luzon is included. Holotype specimens for all four new species will be deposited in the National Museum of the Philippines type collection and paratype specimens will be deposited into the San Diego Natural History Museum collection or retained within the first author's collection.
... The eggs of leaf insects were examined e.g. by Clark (1978), Viscuso & Longo (1983) and Hausleithner (1984). Hennemann et al. (2009) published an extensive survey of the genus with a revision of Philippine species, which questioned the validity of the so far established systematic groups and provided a reclassification as well as clarification of several identifications. This paper describes and discusses a tiny new leaf insect found in West Papua (New Guinea), which is in accordance to the classification presented by Hennemann et al. (2009) placed in the siccifolium species-group of the subgenus Phyllium (Phyllium). ...
... Hennemann et al. (2009) published an extensive survey of the genus with a revision of Philippine species, which questioned the validity of the so far established systematic groups and provided a reclassification as well as clarification of several identifications. This paper describes and discusses a tiny new leaf insect found in West Papua (New Guinea), which is in accordance to the classification presented by Hennemann et al. (2009) placed in the siccifolium species-group of the subgenus Phyllium (Phyllium). The description is based on a single adult female and four eggs. ...
... Eggs differ by the raised hairy, umbrella-or feather-like appendages, lack of longitudinal lamellae of the capsule and flat operculum. For full details, as well as diagnoses of the genus and subgenus and keys see Hennemann et al. (2009). ...
The female and egg of the new Papuan leaf insect Phyllium (Phyllium) riedeli n. sp. are described and illustrated. The species belongs in the siccifolium species-group of the subgenus Phyllium and, with a body length of 56.3 mm, represents the smallest leaf insect so far described for the genus. The type-specimens are stored in the State Museum of Natural History Karlsruhe, Germany (SMNK). A checklist and key is provided for the New Guinean representatives of Phyllium (Phyllium) Illiger, 1798.
... Such a mechanism with a similar functionality in two organisms from phylogenetically distant lineages is represented by the adhesive mechanism of the eggs of the Philippine leaf insect Phyllium philippinicum Hennemann, Conle, Gottardo & Bresseel, 2009 (Phasmatodea, Phylliidae) and the seeds of the ivy gourd Coccinia grandis (L.) Voigt (Cucurbitaceae). Both systems include fibrillary surface structures on the surface and a glue component that is applied to the substrate. ...
... Attaching the eggs in higher levels of the forest is a widespread strategy to avoid these parasitoids, which evolved independently in many phasmid lineages [64,65]. (2) Localization of the offspring close to the foodplant could guarantee suitable food for the offspring [14,15,66]. (3) Attachment can be used for dispersal [26], as has been shown for many seeds as well [40]. ...
Plants and animals are often used as a source for inspiration in biomimetic engineering. However, stronger engagement of biologists is often required in the field of biomimetics. The actual strength of using biological systems as a source of inspiration for human problem solving does not lie in a perfect copy of a single system but in the extraction of core principles from similarly functioning systems that have convergently solved the same problem in their evolution. Adhesive systems are an example of such convergent traits that independently evolved in different organisms. We herein compare two analogous adhesive systems, one from plants seeds and one from insect eggs, to test their properties and functional principles for differences and similarities in order to evaluate the input that can be potentially used for biomimetics. Although strikingly similar, the eggs of the leaf insect Phyllium philippinicum and the seeds of the ivy gourd Coccinia grandis make use of different surface structures for the generation of adhesion. Both employ a water-soluble glue that is spread on the surface via reinforcing fibrous surface structures, but the morphology of these structures is different. In addition to microscopic analysis of the two adhesive systems, we mechanically measured the actual adhesion generated by both systems to quantitatively compare their functional differences on various standardized substrates. We found that seeds can generate much stronger adhesion in some cases but overall provided less reliable adherence in comparison to eggs. Furthermore, eggs performed better regarding repetitive attachment. The similarities of these systems, and their differences resulting from their different purposes and different structural/chemical features, can be informative for engineers working on technical adhesive systems.
... Additionally, Brock and Hasenpusch (2003) discuss the subadult female found by G. B. Monteith in 1971 in the Iron Range, Queensland as additional evidence that a "frondosum-like species" is probably the unknown female for the Nanophyllium. As it turns out, the frondosum species group, as defined by Hennemann et al. (2009), are also somewhat of a quandary since they are only known from female specimens, except for Phyllium (Pulchriphyllium) groesseri Zompro, 1998. In light of the speculation by Brock and Hasenpusch (2003) the authors scoured major collections around the world for additional Nanophyllium male specimens and females of the frondosum species group looking for shared morphological features which might support the thoughts of Brock and Hasenpusch (2003). ...
... Egg orientation terminology follows Clark (1978). Species group organizations follow the classification presented in Hennemann et al. (2009) and Cumming (2017). Wing venation terminology follows Burt (1932) and Ragge (1955). ...
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 Hennemann et al. 2009 and is therefore excluded from further discussion here. The rearing of Nanophyllium also yielded the male Nanophyllium asekiense (Größer, 2002), comb. nov. thus, enabling comparison of this male to the other previously known Nanophyllium species. Two new species of nano- leaf insects are described within, Nanophyllium miyashitai sp. nov., from Morobe Province, Papua New Guinea, and Nanophyllium daphne sp. nov., from Biak Island, Papua Province, Indonesia. With such distinct sexual dimorphism in Nanophyllium between sexes, which have only now been matched up via captive rearing, illustrated within are numerous specimens which might represent the unknown opposite sexes of the many currently known species of Nanophyllium. Due to pronounced sexual dimorphism in Nanophyllium, only future captive rearing or molecular analysis will match up the many unknown sexes. To conclude, with the description of two new Nanophyllium species, dichotomous keys to species for known males and females are presented.
... Additionally, Brock and Hasenpusch (2003) discuss the subadult female found by G. B. Monteith in 1971 in the Iron Range, Queensland as additional evidence that a "frondosum-like species" is probably the unknown female for the Nanophyllium. As it turns out, the frondosum species group, as defined by Hennemann et al. (2009), are also somewhat of a quandary since they are only known from female specimens, except for Phyllium (Pulchriphyllium) groesseri Zompro, 1998. In light of the speculation by Brock and Hasenpusch (2003) the authors scoured major collections around the world for additional Nanophyllium male specimens and females of the frondosum species group looking for shared morphological features which might support the thoughts of Brock and Hasenpusch (2003). ...
... Egg orientation terminology follows Clark (1978). Species group organizations follow the classification presented in Hennemann et al. (2009) and Cumming (2017). Wing venation terminology follows Burt (1932) and Ragge (1955). ...
... The orange and yellow L. arcticus morphs may also mimic the colour and shape of an autumn coloured polar willow (S. polaris) leaves. Leaf insects (Phylliidae) and dead leaf butterfly (Kallima sp.) use camouflage to take the appearance of a leaf (Nijhout 2001& Hennemann et al. 2009). Mimesis of leaves is well documented in other taxa (Eigenmann & Allen 1921, Hennemann et al. 2009), but the ability of L. arcticus to mimic a real leaf may be the first documented crustacean leaf mimesis in the aquatic environment. ...
... Leaf insects (Phylliidae) and dead leaf butterfly (Kallima sp.) use camouflage to take the appearance of a leaf (Nijhout 2001& Hennemann et al. 2009). Mimesis of leaves is well documented in other taxa (Eigenmann & Allen 1921, Hennemann et al. 2009), but the ability of L. arcticus to mimic a real leaf may be the first documented crustacean leaf mimesis in the aquatic environment. The mimicry in colour, shape and movements is very close to the polar willow leave, the photographs and drawings of the dead specimens scarcely doing it justice. ...
Lepidurus arcticus (Pallas, 1793) is a keystone species in High Arctic ponds, which are exposed to a wide range of environmental stressors. This thesis provides information on the ecology of this little studied species by paying particular focus on the sensitivity of L. arcticus to acidification and climate change. Respiration, reproduction, olfaction, morphology, salinity and pH tolerance of the species were studied in the laboratory and several environmental parameters were measured in its natural habitats in Arctic ponds.
Current global circulation models predict 2–2.4 °C increase in summer temperatures on Spitsbergen, Svalbard, Norway. The L. arcticus respiration activity was tested at different temperatures (3.5, 10, 16.5, 20, 25 and 30 °C). The results show that L. arcticus is clearly adapted to live in cold water and have a temperature optimum at +10 °C. This species should be considered as stenothermal, because it seems to be able to live only within a narrow temperature range.
L. arcticus populations seem to have the capacity to respond to the ongoing climate change on Spitsbergen. Changes can be seen in the species’ reproductive capacity and in the individuals’ body size when comparing results with previous studies on Spitsbergen and in other Arctic areas. Effective reproduction capacity was a unique feature of the L. arcticus populations on Spitsbergen. L. arcticus females reached sexual maturity at a smaller body size and sexual dimorphism appeared in smaller animals on Spitsbergen than anywhere else in the subarctic or Arctic regions. L. arcticus females were able to carry more eggs (up to 12 eggs per female) than has been observed in previous studies. Another interesting feature of L. arcticus on Spitsbergen was their potential to grow large, up to 39.4 mm in total length. Also cannibalistic behaviour seemed to be common on Spitsbergen L. arcticus populations. The existence of different colour morphs and the population-level differences in morphology of L. arcticus were unknown, but fascinating characteristic of this species. Spitsbergen populations consisted of two major (i.e. monochrome and marbled) and several combined colour morphs. Third interesting finding was a new disease for science which activated when the water temperature rose. I named this disease to Red Carapace Disease (RCD).
This High Arctic crustacean lives in ponds between the Arctic Ocean and glaciers, where the marine environment has a strong impact on the terrestrial and freshwater ecosystems. The tolerance of L. arcticius to increased water salinity was determined by a LC50 -test. No mortality occurred during the 23 day exposure at low 1–2 ‰ water salinity. A slight increase in water salinity (to 1 ‰) speeded up the L. arcticus shell replacement. The observations from natural populations supported the hypothesis that the size of the animals increases considerably in low 1.5 ‰ salt concentrations. Thus, a small increase in water salinity seems to have a positive impact on the growth of this short-lived species.
Acidification has been a big problem for many crustaceans, invertebrates and fishes for several decades. L. arcricus does not make an exception. Strong acid stress in pH 4 caused a high mortality of mature L. arcticus females. The critical lower limit of pH was 6.1 for the survival of this acid sensitive species. Thus, L. arcticus populations are probably in danger of extinction due to acidification of three ponds on Spitsbergen. A slight drop (0.1–1.0) in pH values can wipe out these L. arcticus populations. The survival of L. arcticus was strongly related to: (1) the water pH, (2) total organic carbon (TOC) and pH interaction, (3) the water temperature and (4) the water salinity. Water pH and TOC values should be monitored in these ponds and the input of acidifying substances in ponds should be prevented.
... The Philippine archipelago is considered the second ''hottest hotspot'' of biodiversity on Earth [17], and represents an area with potential high levels of euphasmatodean diversity and endemicity. Motivated by this assumption, we have undertaken a series of taxonomic investigations in order to improve the knowledge of the stick and leaf insect fauna of the Philippines [18][19][20][21][22][23]. In our latest contribution we documented an interesting representative of the family Aschiphasmatidae, Dallaiphasma eximius Gottardo, 2011, from Mount Halcon on the island Mindoro in the western part of the archipelago [23]. ...
... The enigmatic stick insect described in this article further underpins the high degree and uniqueness of biodiversity in general, and also in Phasmatodea, reported from Philippine archipelago [17][18][19][20][21][22][23]. At this time, the taxonomic position of C. enigma gen. ...
A new genus and species of stick insect is described and figured from Mount Halcon, on the Philippine island of Mindoro. Conlephasma enigma gen. et sp. n. is a stout, flightless, and apparently ground-dwelling species with vivid integumental colors. When disturbed, specimens spray a defensive secretion from the prothoracic exocrine glands. The systematic position of Conlephasma within Euphasmatodea is unclear. The elongated galealobulus and the trichome area located laterally in the galea, represent unusual apomorphic characters of the maxilla that could indicate affinities with Necrosciinae or Pseudophasmatinae. All tibiae exhibit the anareolate condition. Euplantulae are of two types: those of tarsomeres I–IV feature a nubby microstructure, whilst the one on the ventral side of the pretarsus is smooth. Males are characterized by the presence of a well-developed vomer on the tenth abdominal segment. A distinctive and apomorphic trait of female terminalia is represented by the elongated tenth abdominal tergum. Conlephasma can represent an interesting taxon for studies on the evolution of the stick and leaf insects.
... Gottardo [7] summarized the few published reports, recording four genera and five species from the islands of Luzon, Mindanao, Palawan, and Panay. During the work on ongoing project ''Systematics of Philippine Phasmatodea'' [7][8][9][10][11], the author examined new samples of micropterous Aschiphasmatidae characterized by a set of peculiar apomorphic features which allowed recognition of a new genus and species. Also, the specimens of the new taxon were collected on Mindoro island, where members of this group were not yet reported. ...
... et sp. n., Mount Halcon on the eastern half of Mindoro island, represents the third highest mountain in the Philippines, but so far is known to harbour only few other phasmatodean taxa, specifically: four members of Lonchodinae: [11,[20][21][22]. Most of these species appear to be Mindoro endemic, and future studies on this locality will certainly raise the number of taxa represented. ...
Based on characters of both sexes, a new genus and species of the basal euphasmatodean lineage Aschiphasmatidae is described and figured from the Philippines. Dallaiphasma eximius gen. et sp. n. displays interesting features for the group, including: a cone-shaped vertex, which is notably raised above the pronotum; the tibial area apicalis represented by a depressed membranous posterior lateral region, and a strongly sclerotized central apical region; the euplantulae consisting of smooth-type attachment pads; the pretarsal claws pectination reduced to minute denticulations; and the well-differentiated boundary between the metanotum and the first abdominal tergum. The phylogenetic information content of the new findings is discussed. Furthermore, as a result of this study, the Aschiphasmatidae are newly recorded from Mindoro island, and now include five genera and six species of the Philippines.
... Alae of female is not developed. Eggs have lamellated longitudinal carinae; without hair-like structures; operculum conically (Hennemann et al., 2009;Bank et al., 2021b). Diagnosis. ...
The leaf insect Pulchriphyllium giganteum (Hausleithner, 1984) is recorded in Thailand for the first time. The diagnosis, measurements, photographs, and short notes of the specimen are also provided.
... In the present study, as the first step to dissect de novo reference genome of one representative leaf insect, we used Illumina next-generation sequencing (NGS) data to generate the nearly complete mitogenome (with the exception portions of the control region) of Phyllium westwoodii (Wood-Mason 1875). This species is one member in Phylliidae and mainly distribute in countries of Southeast Asia, including India (South Andaman Island), Myanmar, China (Yunnan), Thailand, Laos, Vietnam, Sumatra, and Singapore etc. (Hennemann et al. 2009). Male and female adults were collected from Muang Fuang, Nang Ha, Laos (102 7 0 3 00 E; 18 39 0 12 00 N) during June 2017 by Zhiwei Dong and local villagers, and bred in the greenhouse with host plant Rubus sp. ...
The nearly complete mitochondrial genome (mitogenome) of Phyllium westwoodii, a typical leaf mimic insect in Phasmatodea, was obtained in this study. This mitogenome is 17,222 bp in length and contains 13 protein-coding genes (PCGs), 22 transfer RNA genes (tRNAs), two ribosomal RNA genes (rRNAs) and almost complete control regions. All PCGs initiate with ‘ATN’ except for NAD4L that uses ‘TTG’ as the start codon, and terminate with ‘TAA’ except for COX2 that uses a single ‘T’ residue as the stop codon. The phylogenetic analysis based on the concatenated sequences of 13 PCGs and two rRNAs shows that P. westwoodii is closer to Phyllium tibetense than Phyllium giganteum.
... Very well-known examples for this aspect of phasmatodean evolution are the Walking Leaves (Phyllidae). These insects perfectly imitate leaves of plants and blend with their environment [11][12][13]. This is substantially made possible by their abdominal lateral extensions and their green coloration (see, e.g., in [14][15][16][17][18]). ...
Leaf insects (Phylliidae) are well-camouflaged terrestrial herbivores. They imitate leaves of plants almost perfectly and even their eggs resemble seeds—visually and regarding to dispersal mechanisms. The eggs of the leaf insect Phyllium philippinicum utilize an adhesive system with a combination of glue, which can be reversibly activated through water contact and a water-responding framework of reinforcing fibers that facilitates their adjustment to substrate asperities and real contact area enhancement. So far, the chemical composition of this glue remains unknown. To evaluate functional aspects of the glue–solvent interaction, we tested the effects of a broad array of chemical solvents on the glue activation and measured corresponding adhesive forces. Based on these experiments, our results let us assume a proteinaceous nature of the glue with different functional chemical subunits, which enable bonding of the glue to both the surface of the egg and the unpredictable substrate. Some chemicals inhibited adhesion, but the deactivation was always reversible by water-contact and in some cases yielded even higher adhesive forces. The combination of glue and fibers also enables retaining the adhesive on the egg, even if detached from the egg’s surface. The gained insights into this versatile bioadhesive system could hereafter inspire further biomimetic adhesives.
... With the description of this new species, and review of its closest relatives, Phyllium caudatum Redtenbacher, 1906 andPhyllium riedeli Kamp &, these species share a unique set of characteristics that sets them apart from all other Phylliidae. We here transfer these three species from the siccifolium species group of Phyllium (Phyllium) as defined by Hennemann et al. (2009) into their own subgenus Phyllium (Comptaphyllium) subgen. nov., based on the below discussed characteristics. ...
A large species of leaf insect from Obi Island, Indonesia is here described as Phyllium regina n. sp. which is currently only known from a single female specimen. With the description of this new species, and review of its closest relatives, Phyllium caudatum Redtenbacher, 1906 and Phyllium riedeli Kamp & Hennemann, 2014, these species share a unique set of characteristics that sets them apart from all other Phylliidae. We here transfer these three species from the siccifolium species group of Phyllium (Phyllium) as defined by Hennemann et al. (2009) into their own subgenus Phyllium (Comptaphyllium) subgen. nov., based on the below discussed characteristics. In addition to the description of the new species and subgenus, the unknown male Phyllium riedeli morphology is here described. To conclude, a distribution map for the three species is presented, as well as a morphological key to the three species.
... Differentiation. Phyllium (Phyllium) conlei new species morphologically fits within the siccifolium species-group as described by Hennemann et al., 2009 with the exterior lobe of the profemora which is thinner than the interior lobe. Phyllium (Phyllium) conlei new species is most morphologically similar to Phyllium (Phyllium) brossardi Cumming et al., 2017, and Phyllium (Phyllium) caudatum Red tenbacher, 1906 because of the slender abdomen with a maximum width of only about 30% of the overall abdominal length (Fig. 4a-c). ...
The Phylliidae (Phasmatodea) diversity of the Lesser Sunda Islands, Indonesia is preliminarily examined, and revealed to be notably lacking in completeness with only two species currently recorded. Of the nine islands/ island groups within the Lesser Sunda Islands, only the westerly islands (Bali and Lombok) have single species recorded: Phyllium (Pulchriphyllium) pulchrifolium Audinet-Serville, 1838, from Bali, and Phyllium (Phyllium) conlei Cumming, Valero, and Teemsma, new species, from Lombok. The latter species is herein described and differentiated from congenerics. To conclude, with so few species recorded from the Lesser Sunda Islands, a key to species for Java and the Lesser Sunda Islands is presented for males.
... Males have an exterior lobe of the profemora which is thinner than the interior lobe, females have tibiae lacking lobes and rudimentary alae, and eggs have long pinnae on the margins and an operculum which is flat. Features of both sexes agree that Phyllium (Phyllium) letiranti new species falls within the siccifolium species-group as described by Hennemann et al., (2009). Phyllium (Phyllium) letiranti new species morphologically is most similar to Phyllium (Ph.) mamasaense Größer, 2008, Phyllium (Ph.) siccifolium (Linnaeus, 1758), Phyllium (Ph.) philippinicum Hennemann et al., 2009, and Phyllium (Ph.) bourquei Cumming and Le Tirant, 2017. ...
A new species of leaf insect, Phyllium (Phyllium) letiranti Cumming and Teemsma, new species (Phasmida: Phylliidae), is described from a series of males, females, and eggs from Peleng Island, Indonesia. This new species is the first record of the family Phylliidae on the island and is here differentiated from congeners. Keys to males, females, and eggs of the Phyllium species of Sulawesi and Peleng islands are included within.
... The exterior lobe of the protibia is reduced almost to the status of the Phyllium (Phyllium) subgenus, however the other prominent lobes of the meso-and metatibiae, and serrate antennae clearly place the new species in the Phyllium (Pulchriphyllium) subgenus and the bioculatum species-group as per Hennemann et al., 2009. Phyllium (Pulchriphyllium) shurei n. sp., is morphologically similar to Phyllium (Pulchriphyllium) mannani Seow-Choen, 2017, which unfortunately is also only known from male specimens, with the female of both species currently unknown. ...
A new species, Phyllium (Pulchriphyllium) shurei n. sp. is described and
illustrated from Java, Indonesia. Phyllium (Pulchriphyllium) shurei n. sp. is currently
only known from a male specimen, therefore only a key to known Phyllium males
from Java is included. Nomenclature note are added about the name Phyllium
(Pulchriphyllium) lambirense Seow-Choen, 2017.
... HENNEMANN et al.33–35 teeth on the pars stridens of antennomere III; four teeth on interior lobe of profemora with a larger distinct gap between the 2nd and 3rd; V & VI abdominal lateral margins consistently parallel; abdominal segment VII with lobes variable, if strongly lobed, profemoral teeth prominent, if lobe reduced or absent the teeth of the profemoral lobe are also reduced in size; presence of a median spine on the transverse anterior ridge of the mesopraescutum; (Palawan) ................ gantungense HENNEMANN et al. * The range of P. mabantai Bresseel et al., 2009 is here noted to be more extensive than noted by the original authors. Fresh material from both Leyte and Samar islands have since been recovered belonging to P. mabantaiBresseel et al., 2009. ...
A new species of leaf insect, Phyllium (Phyllium) fallorum n. sp. is described and illustrated. In addition a key to the current known species of Phyllium Illiger, 1798 from the Philippines is provided. This species is currently only known from the single unique holotype, which will be deposited in the San Diego Natural History Museum. Measurements of anatomical figures were made to the nearest 0.1 mm. Photos were taken using a Canon 5D Mark II and a MP-E 65mm macro lens and stacked using Zerene photo stacking software, version 1.04, 64-bit. Currently, ten species of Phyllium Illiger, 1798 have been described from the Philippines, with Phyllium (Phyllium) fallorum n. sp. the eleventh. Of the eleven known species from the Philippines, five are only known from female specimens.
... Currently there are 53 species of Phylliidae described, 41 of which belong to the genus Phyllium Illiger, 1798 (Brock, 2015). The genus Phyllium is divided into the two subgenera Phyllium and Pulchriphyllium Griffini, 1898(HenneMann et al., 2009. Phyllium maethoraniae n. sp. ...
A new species of leaf insect is described from Thailand, Phyllium (Pulchriphyllium) maethoraniae n. sp., very close to P. (Pulchriphyllium) sinense Liu, 1990, but differs by various distinct characters such as the armature of the mesonotum, shape of the anterior legs, abdomen and abdominal apex. Like P. sinense, P. maethoraniae n. sp. is only known from the female. A table is given to distinguish between the two species.
... In the female, important phasmatodean features are represented by the specialized sternum VIII (=operculum), which covers the ovipositor (Bradler & Buckley 2011;Wang et al. 2014). Characters of the male and female terminalia of Phasmatodea represent a useful tool in taxonomic research (Hennemann et al. 2009;Gottardo 2011a;Buckley et al. 2014) as well as in phylogenetic and evolutionary research (Bradler 2001(Bradler , 2009Wedmann et al. 2007;Buckley et al. 2009;. ...
Aschiphasmatinae is a small group of stick insects from the Oriental region whose genital morphology has been rarely described in detail. The subfamily is of particular interest, as phylogenetic studies have shown Aschiphasmatinae to be the sister group to the remaining Euphasmatodea. In this paper, the male and female terminalia are described for the first time in Dajaca napolovi Brock, a little known aschiphasmatine species from Vietnam. In the male, the transversally undivided abdominal sternum IX and gently incurved cerci with a conspicuous apical tooth represent apomorphies of Aschiphasmatinae. Thorn pads on the hind margin of abdominal tergum X consist of only a single row of 6–7 ventrally oriented teeth. The simple thorn pad structure of Aschiphasmatinae can represent an ancestral condition for Euphasmatodea. The vomer on venter X is well-developed and features two unusually large basal apodemes and a strongly developed apical spine showing a specialized streaked surface micropattern. Female terminalia are characterized by an unkeeled abdominal sternum VIII covering the reduced primary ovipositor. Gonapophysis VIII does not extend beyond the tip of gonapophysis IX. An asymmetry concerning the size of the paired gonapophyses is reported for the first time in Phasmatodea.
... The older divisions of Areolatae and Anareolatae (Stål, 1875;Brunner von Wattenwyl and Redtenbacher, 1906, 1907, 1908, are not natural (Bradler, 2001(Bradler, , 2009Tilgner, 2002), and therefore of no practical use, except to note that the presence of the areola apicalis is putatively plesiomorphic for Phasmatodea (Bradler, 2009). We concur with Hennemann et al. (2009) and Conle et al. (2011), that the systems, largely of the 'areolates' but also of other groups, of Zompro and Gr€ oßer (2003) and Zompro (2004aZompro ( , 2004b are confused and do more to crimp work than advance it. Accordingly, we generally follow the various reorganizations of the order as advocated by Hennemann and Conle (2008), Bradler (2009), andBradler et al. (2014). ...
A new fossil stick-insect (Phasmatodea) is described and figured from a male preserved in Cretaceous amber from Myanmar. Echinosomiscus primoticus Engel and Wang, gen. et sp. nov., is a robust, somewhat-compressed stick-insect bearing abundant acanthae over the head and body, and remarkably lacks an area apicalis on the tibiae. The species is described and assigned to a new, extinct subfamily of Phasmatidae s.l., as Echinosomiscinae Engel, subfam. nov. Brief remarks are made regarding the dating of phasmatodean lineages, with E. primoticus providing the first reliable evidence for Euphasmatodea and even Neophasmatodea in the Cenomanian.
... As part of our research on the Philippine stick and leaf insect fauna (Gottardo 2007;Gottardo 2008;Hennemann et al. 2009;Bresseel 2012), a field expedition was carried out to the Benguet province of Luzon island during which several specimens of Orthomeria were found. After a careful examination of the general anatomy, the specimens were assumed to represent a new species of the subgenus Orthomeria showing an original set of morphological traits, including a distinctive red colouration on the wings which was not previously reported within the genus (see Bragg 2001). ...
A new stick insect of the genus Orthomeria Kirby, 1904 (Phasmatodea, Aschiphasmatidae) is described from the Philippines. Orthomeria (O.) kangi sp. n. is readily distinguished from all other congeners by the distinctive blood red colouration of the costal region of the hind wings. Major features of the external morphology of adults, eggs, and first-instar nymphs are illustrated. Locomotory attachment pads are of the smooth type with irregular microgrooves on the contact surface. An unusual condition of male terminalia is the absence of tergal thorn pads on segment 10. The male clasping organs are represented by an elongated vomer terminating in a prominent spine, and by incurved cerci featuring a bilobed apex equipped with a sharp blade-like ridge. Intraspecific variation in body colouration and hind wing length occurs in females. The new species lives at 400-650 m elevation in the surroundings of the Sablang and Tuba regions, in the Benguet Province of Luzon island. Host plants include Ficus spp. (Moraceae), and Pipturus spp. and Leucosyke spp. (Urticaceae). Observations on the mating and defensive behaviour are presented. Orthomeria (O.) catadromus (Westwood, 1859) is recognised as a junior synonym of O. (O.) pandora (Westwood, 1859), syn. n. A lectotype is designated for both species. Finally, an updated identification key to the species of the subgenus Orthomeria is provided.
... Recently, while rearing Phyllium westwoodii Wood-Mason (Phasmatodea: Phylliidae) (Fig. 1), a mid-sized leaf insect widely distributed in Southeast Asia (Hennemann et al. 2009), we noticed that the aperture used for oviposition, the distance that the eggs were propelled and the color of the eggs appeared to differ between fertilized and unfertilized eggs. As far as we are aware, this situation has not been reported previously in phasmatids or in other insects. ...
We report that in a leaf insect, Phyllium westwoodii Wood-Mason (Phasmatodea: Phylliidae), two differing apertures can be used for oviposition, the color of eggs being affected by which aperture is used. Eggs which are forcibly propelled from the internal space within the valvulae of the abdomen are brown, whereas white eggs emerge slowly from the opening between the eighth sternite and the valvulae, and are deposited close to the ventral surface of the female. This unusual oviposition system does not appear to have been previously reported in phasmatids or in other insects.
... Daug nuostabių jų nuotraukų pateikiama užsienio in- terneto svetainėse. Retkarčiais publikuojamos monografijos (Hennemann, Conle, 2008;Hennemann et al., 2009) ir kitos mokslo studijos (Kristensen, 1975;Tilgner et al., 1999;Bradler, 1999;Thomas, 2001;Tilgner, 2003;Gillott, 2005;Conle, Hennemann, 2005;Conle et al., 2009). Deja, leidinių apie gyvalazdes lietuvių kalba iki šiol dar nebuvo. ...
... The results are discussed with respect to phasmatodean phylogeny, and the placement of H. leytensis within Stephanacridini. This article is also a contribution to a project aimed at studying the systematics of Philippine stick and leaf insects [41][42][43][44]. ...
External morphological features of adult males are described in the stick insect Hermarchus leytensis from the Philippines, a species belonging to the little-known euphasmatodean lineage Stephanacridini. Mouthparts are characterized by some likely specialized features: (1) a dentate dorsal cutting edge on the mandibles; (2) distinct differences in size and shape between the galeae; (3) absence of an apical field of trichomes on the galeae; and (4) lacinial setae not protruding from the mesal margin, which features three bearing-like protuberances. The latter character state represents a very unusual condition, not known in other phasmatodeans or even in polyneopteran insects. A distinctive characteristic of attachment devices is that each euplantula is divided into two separated pads with a smooth microstructure, as it also occurs in some members of the clade Schizodecema. Male terminalia exhibit character states previously unknown in Stephanacridini, including (1) a symmetrical type of vomer and (2) claspers equipped with ∼70 very minute ventral teeth on each thorn pad. Potential implications for the systematic placement of H. leytensis are discussed. The results also underline the importance of microanatomical investigations as source of substantial characters for future analyses on phasmatodean systematics.
... In the present study, an additional distinctive species is described from Panay Island as new for science, along with an emended diagnosis of the genus, and a summary of the Philippine taxa of Platycraninae. This article is a contribution to the series ''Systematics of Philippine Phasmatodea'' [7][8][9][10]. ...
A distinctive new species of the phasmatodean genus Ophicrania Kaup, 1871 (Phasmatidae: Platycraninae) is described and figured from the Philippines. Ophicrania conlei n. sp. (from Mount Madja-as, on Panay island) is characterized by the bicoloured anal region of the male hind wing, divided into an inner whitish patch, and an outer brownish area. The species is also distinguished from its most similar congeners on the basis of integumental coloration, features of the antennae, thoracic nota, wings, legs, and terminalia. The present study also provides an emended diagnosis of Ophicrania, and an updated checklist of the taxa of Platycraninae recorded from the Philippine archipelago.
The subfamily Platycraninae Brunner v. Wattenwyl, 1893 is polyphyletic in the traditional treatment. None of the genera attributed to the subfamily is closely related to the type-genus Platycrana Gray, 1835, which has already been suggested by morphological studies. The main key feature of Platycraninae traditionally used to characterize the subfamily, the remarkably large head and strongly enlarged genae ("cheeks"), is not true for Platycrana. Consequently, all other genera, the true "Palm Stick Insects" need to be removed from Platycraninae and are here accommodated in Megacraniinae subfam. nov.. Megacrania Kaup, 1871 is chosen as the type-genus of this new subfamily, because the name very well describes the main characteristic of this clade. Previous molecular studies have shown Megacraniinae subfam. nov. to be a member of the Old World family Phasmatidae s. str. (= Lanceocercata Bradler, 2001), which is characterized by the enlarged, laterally flattened and foliaceous or lanceolate cerci and by the male anal segment longitudinally split into two movable, interiorly dentate hemi-tergites that serve as a clasping apparatus to grasp the female abdomen during copulation and makes lacking a vomer. Neither character is true for Platycrana and Platycraninae sensu nov. respectively. Molecular studies have revealed Platycraninae sensu novo as the sister group of Phasmatidae s. str. (= Lanceocercata). The tribe Stephanacridini Günther, which has previously been without a subfamilar affiliation is transferred as a subordinate taxon of Platycraninae sensu novo. Platycraninae is characterized and distinguished from Phasmatidae s. str. (= Lanceocercata) by the small cylindrical cerci, simple anal segment and presence of a vomer in males, as well as the strongly keeled subgenital plate and remarkably elongated, filiform gonapophysis VIII of females, which greatly project over the apex of the abdomen. Characterizations are presented for Megacraniinae subfam. nov. as well as Platycraninae sensu nov. and its two tribes Platycranini and Stephanacridini. Lists of genera are provided for the three taxa. Three genera are removed from Platycraninae and Megacraniinae subfam. nov.: The Australian Echetlus Stål, 1875 is transferred to Phasmatinae: Acanthomimini and Redtenbacherus Özdikmen & Darilmaz, 2008 and Elicius Günther, 1935 are transferred to Lonchodidae: Necrosciinae. The male of Megacrania spina Hsiung, 2007 as well as the eggs of Megacrania brocki Hsiung, 2002, Erastus apalamnus Rehn, 1904, Ophicrania apterus (Redtenbacher, 1908), Ophicrania nigroplagiatus (Redtenbacher, 1908), Platycrana viridana (Olivier, 1792) and Macrophasma lyratus (Redtenbacher, 1908) are illustrated for the first time. Xenomaches moderata (Kirby, 1884) from the island of Ambon is synonymised with Platycrana viridana (Olivier, 1792) (syn. nov.).
Published in Zoologica — Original Contributions to Zoology (ISSN 0044-5088):
Abstract
Attachment devices with different microstructure and surface microstructure evolved on the tarsi of hexapods. In biomechanical studies it was shown for a few species that different types of structure have different attachment properties. However, it is yet unclear if these structures evolved in correlation with the species’ ecology. Stick and leaf insects (Phasmatodea) is a speciose insect taxon with several different ecological preferences. We therefore chose 116 species from all subfamilies currently recognised within this taxon, to uncover correlations between the euplantular microstructure and the ecological preferences or oviposition techniques of the examined species. Twelve different types of attachment microstructures have been found using scanning electron microscopy (SEM), seven of them previously unknown. The correlation of the microstructures with the ecology, habitat and phylogeny of stick and leaf insects is discussed. Two different hypotheses are possible based on this analysis: (1) Smooth euplantular structures may be a ground plan in Phasmatodea and structures with different degrees of elevations evolved convergently in many groups starting from this type of microstructures, or (2) elongated acanthae are the plesiomorphic state which is conserved in some taxa and convergently reduced to smooth types of microstructures in other groups. These assumptions should be tested in subsequent phylogenetic studies based on the use of extensive molecular data. In addition, one species of Embioptera (Hexapoda), which is assumed to be the sister group of the Phasmatodea, has been examined, and its rather smooth attachment structures might be a result of their extraordinary mode of living within galleries.
URL: https://www.schweizerbart.de/publications/detail/isbn/9783510550517/Zoologica_Vol_164
Abstract. – A new species of nano-leaf insect, Nanophyllium australianum Cumming,
Le Tirant, and Teemsma, n. sp. (Phasmida, Phylliidae), is described from a single male
specimen from Queensland, Australia. This single Australian record was originally
considered as a range expansion for Nanophyllium pygmaeum Redtenbacher, 1906
(Rentz, 1988). Reexamination of this specimen as well as the intraspecies variation
within other Nanophyllium specimens revealed this Australian record as morphologically
distinct and geographically isolated. With this species now distinguished from N.
pygmaeum and additional specimens within the genus examined, a more comprehensive
key to species is included.
Type specimens of 437 Phasmida taxa have been located in the Natural History Museum, London (NHMUK, formerly BMNH), including 480 primary types of 372 taxa. Taxa with types are listed alphabetically by their specific or subspecific name, and the number of specimens, sex and locality data are given.
A complete taxonomic catalogue of the Stick and Leaf-insects (Phasmatodea) recorded or described from the mainland China (excluding Taiwan) is presented. 241 valid species are listed, which are currently attributed to 50 genera, 5 families and 7 subfamilies. Genera and species are listed alphabetically. All available type-data is provided based mainly on literary sources for species described by Chinese workers from 1986 to 2006, including documented depository of type-specimens. The catalogue therefore also provides complete lists of the type-material of Phasmatodea housed in the following Chinese institutions: Administration of Baishuijiang Natural Reserve (ABNR), Beijing Forestry University, Beijing (BFU), China Agricultural University, Beijing (CAU), Geological Museum of China, Beijing (GMC), Inca Science Ltd., Chongqing (INCA), Institute of Zoology, Chinese Academy of Sciences, Beijing (IZCAS), Department of Biology, Nankai University, Tianjin (NKU), Northwest Sci-Tech University of Agriculture and Forestry, Shaanxi (NWSUAF), Institute of Zoology, Shaanxi Normal University, Xi’an (SNU), Institute of Entomology, Sun Yat-sen University (SYSU), Shanghai Institute of Entomology, Academia Sinica, Shanghai (SIES), Tianjin Natural History Museum, Tianjin (TJNHM), Zhejiang Museum of Natural History, Hangzhou (ZJMNH). The known distribution of each species, in means of provinces is provided as well. 14 species are shown to have been recorded from China in error, several of these based on misidentifications. The “Phasmatodea-like” fossil taxa described from the the Late Jurassic Yixian Formation of North Hebei and West Liaoning are listed in a separate section.
Two new generic synonyms are recognized: Arthminotus Bi, 1995 synonymised with Lopaphus Westwood, 1859 (n. syn.) and Dianphasma Chen & He, 1997 synonymised with Parasosibia Redtenbacher, 1908 (n. syn.). The genus Linocerus Gray, 1835 (Type-species: Linocerus gracilis Gray, 1835) was erroneously synonymised with the mediterranean Bacillus St. Fargeau & Audinet-Serville, 1825 and is here re-established in Phasmatidae: Pachymorphinae: Gratidiini (rev. stat.). Relationship to Clonaria Stål, 1875 (= Gratidia Stål, 1875, = Paraclonaria Brunner v. Wattenwyl, 1893), Sceptrophasma Brock & Seow-Choen, 2000 and Macellina Uvarov, 1940 is obvious.
13 species are transferred to other genera (new combinations): Asceles dilatatus Chen & He, 2004 and Asceles quadriguttatus Chen & He, 1996 to Pachyscia Redtenbacher, 1908, Arthminotus sinensis Bi, 1995 to Lopaphus Westwood, 1859, Baculum dolichocercatum Bi & Wang, 1998a and Baculum politum Chen & He, 1997 to Medauroidea Zompro, 1999, Dixippus bilippus Chen & He, 1999, Dixippus hainanensis Chen & He, 2002, Dixippus nigroantennatus Chen & He, 2002, Dixippus parvus Chen & He, 2002 and Entoria bobaiensis Chen, 1986 to Lonchodes Gray, 1835, Sipyloidea obvius Chen & He, 1995 to Sinophasma Günther, 1940, and Gratidia bituberculata Redtenbacher, 1889 and Leptynia xinganensis Chen & He, 1993 to Sceptrophasma Brock & Seow-Choen, 2002. Acrophylla sichuanensis Chen & He, 2001 remains of unknown generic assignment, but is shown to be not a member of the Australian genus Acrophylla Gray, 1835. Furthermore, as Baculum Saussure, 1861 is a neotropical genus and most Old World species previously attributed this genus are now listed in Ramulus Saussure, 1861, all Chinese species described in this genus are consequently transferred to Ramulus Saussure. Other changes of specific placements are based on published literature and concern to the following three synonymies not recognized by Chinese workers: Abrosoma Redtenbacher, 1906 (= Prosceles Uvarov, 1940), Necroscia Audinet-Serville, 1838 (= Aruanoidea Redtenbacher, 1908), Lopaphus Westwood, 1859 (= Paramyronides Redtenbacher, 1908). Megalophasma Bi, 1995 is transferred from Necrosciinae to Lonchodinae.
Four lectotypes are designated and three new specific synonyms revealed. A lectotype is designated for Rhamphophasma modestus Brunner v. Wattenwyl, 1893, the type-species of Rhamphophasma Brunner v. Wattenwyl, 1893, in order to fix this genus and species. The male paralectotype is shown to be a male of Parapachymorpha nigra Brunner v. Wattenwyl, 1893, the type-species of Parapachymorpha Brunner v. Wattenwyl, 1893. Clitumnus porrectus Brunner v. Wattenwyl, 1907 is synonymised with Bacillus ? artemis Westwood, 1859 with a lectotype designated for the former (n. syn.). A lectotype is designated for Oxyartes lamellatus Kirby, 1904. Paracentema stephanus Redtenbacher, 1908 is shown to have been erroneously synonymised with Neohirasea japonica (de Haan, 1842) and here synonymised with Neohirasea maerens (Brunner v. Wattenwyl, 1907) (n. syn.). In order to fix the new synonymy a lectotype is designated for Paracentema stephanus Redtenbacher, 1908.
Finally, a biogeographic analysis of the Chinese phasmid fauna is presented. This includes brief background information on the topography and biogeography of China along with maps showing the seven zoogeographical subregions currently recognized as well as the 4 municipalities, 23 provinces, 5 autonomous regions and 2 special administrative regions of China. A summary of the taxonomic compilation of the fauna is provided and its relationships with neighbouring regions, of both the Palearctic and Oriental realms, are discussed. A study is presented on the distribution of the taxa and species densities of each province / autonomous region.
Studies on the rarely reported Australian leaf insects reveal that Phyllium species from northern Queensland, P. frondosum Redtenbacher 1906 and P. siccifolium (L.), are misidentifications. Consequently, a new species Phyllium (Phyllium) monteithi is described and figured, including the egg. Nanophyllium pygmaeum Redtenbacher 1906 is confirmed as Australian, although it may be a different species, similar in appearance or related to Phyllium (Phyllium) frondosum Redtenbacher. Females are needed to clarify its position. A third Australian species is also reported, Chitoniscus lobiventris (Blanchard 1853), based on a single museum record from 1903. Keys are provided to distinguish Australian leaf insects.
190 species of stick-and leaf-insects in 58 genera are known to occur in New Guinea and the surrounding islands. This paper, based on a literature study, provides keys to identify species up to their genus following the current accepted classification. For each genus a summarised description and where available a schematic reproduction of an existing illustration of a representative is given, as well as a list of species ocurring in the subregion. For each species the reference of the original description including page number, sex of the type material, illustrations, and the location of the type material are given. When applicable, references to first description of opposite sex and synonyms followed by the reference containing the nomenclatural act are included. For each species a general geographic distribution is given. In addition, a gazetteer has been compiled including all known localities of type and non-type specimens reported in the literature from the New Guinea subregion. Technical terms used are described in the glossary.
Im September vorigen Jahres konnte ich zwei Wochen in der entomologischen Sammlung des naturhistorischen Reichsmuseums in Leiden arbeiten und nicht nur das interessante, ältere Mantodeenmaterial von De Haan studieren, sondern auch die eingetroffenen Eingänge bestimmen. Was damals nicht möglich war, aufzuarbeiten, erhielt ich durch die grosse Freundlichkeit von Herrn Direktor Van Oort nach Wien gesandt. Die nachstehende Arbeit beschäftigt sich nur mit diesem Material. Auch Herrn R. van Eecke, der mir während meines Aufenthaltes in Leiden stets in liebenswürdigster Weise behilflich war, möchte ich an dieser Stelle herzlichst danken. Wie man sieht, sind zur Fauna des malayischen Archipels abermals mehrere neue Arten hinzugekommen. Dieser Umstand lässt es ratsam erscheinen, mit der Abfassung eines Katalogs der niederländisch-indischen Mantodeen noch etwas zu warten, bis das noch im Mus. Leiden befindliche Material, sowie die noch in diesem Jahre zu erwartenden Sendungen meiner Freunde Dr. Karny und Dr. Fulmek aus Java und Sumatra bearbeitet sind. Im Nachstehenden sind vorwiegend indomalayische, wenige surinamensiscbe Arten beschrieben. Eine neue Gattung aus Australien habe ich im Material des Mus. Leiden auch gefunden und hier beschrieben, ebenso eine neue Phasmide der Gattung Phyllium. I. EREMIAPHILINAE. Theopompa burmeisteri De Haan. ♂ Smitau, Dr. J. Büttikofer Febr. 1894. ♂ Amboina, Hoedt 1867. ♀ Amboina, Hoedt 1864.
Type specimens of 75 taxa of Phasmida have been identified in the Nationaal Natuurhistorisch Museum, Leiden. The taxa are listed alphabetically by species, with the number of specimens, sex and locality data. The material includes species described by Bragg, Bragg & Chan, Gunther, de Haan, Kirby, Lichtenstein, Olivier, and Stoll. The type of pin used was found to be particularly useful as an aid to identification of older material. There are a number of omissions and discrepancies in the published data, these are clarified. Some background information on the original publications and the described material is included.
An attempt is made to standardize further the descriptive terminology of the phasmid egg capsule by introducing stricter definitions and standard abbreviations. In addition, the various forms of the internal micropylar plate are categorized. Eophasma Sellick is replaced by Eophasmodes nov.n. A key to 131 generic forms of these eggs is provided. Where more than one egg form is associated with a genus, a diagnosis of the subgroups is provided.
The genera of the family Phylliidae (Walking Leaves) (Phasmatodea: Areolatae) are revised and the relationships between them discussed. Nanophyllium Redtenbacher, 1906 differs strikingly from the other genera and is transferred in the Nanophylliini, trib. nov. A key to genera and species is provided. Nanphyllium adisi, spec. nov. from New Guinea is described for the first time. The paper includes a key to the species of Nanophyllium.
Phyllium westwoodii is a phasmid insect (Order Phasmatodea) belonging to the Family Phylliidae (leaf insects). These rather large and ornate creatures are known for their morphological resemblance to plant leaves for camouflage. Pyrazines are a common class of compounds used or produced by a wide variety of organisms, even humans. When an individual of P. westwoodii is disturbed, it sprays an opaque liquid from a pair of prothoracic glands, which are utilized by other phasmid species for defense. The current study has found that this liquid contains glucose and a mixture of 3-isobutyl-2,5-dimethylpyrazine, 2,5-dimethyl-3-(2-methylbutyl)pyrazine, and 2,5-dimethyl-3-(3-methylbutyl)pyrazine. This is the first report of pyrazines found in the defensive gland spray of phasmid insects, and the first chemical analysis of glandular material from family Phylliidae.
Phasmatodea of China
- S C Chen
- Y H He
Chen, S.C. & He, Y.H. (2008) Phasmatodea of China. China Forestry Publishing House, Beijing, 476 pp., 12 plates
Anatomical observations on a variety of Phyllium celebicum (de Haan) from Thailand
- A Deschandol
Deschandol, A. (1991) Anatomical observations on a variety of Phyllium celebicum (de Haan) from Thailand. Phasmid Study Group Newsletter, 47, 10–12
Famille des Phasmides (Phasmidae) In : Voyage de M. Dumont d’Urville au Pôle Sud et dans l’océanie par Hombron et Jaquinot
- E Blanchard
Blanchard, E. (1853) Famille des Phasmides (Phasmidae). In : Voyage de M. Dumont d’Urville au Pôle Sud et dans l’océanie par Hombron et Jaquinot. Zoologie IV, pp. 358–360, pl. 1: 8–9
Preliminary Catalogue of the Orthopterous Insects of the Philippine Islands
- L Bruner
Bruner, L. (1915) Preliminary Catalogue of the Orthopterous Insects of the Philippine Islands. University of Nebraska Studies, 15(2), 195–281