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Cryptophyllium, the hidden leaf insects – descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae)
Abstract and Figures
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.
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... The characters of the male vomer are frequently used when describing or differentiating stick insect taxa (Bradler 2009, Bresseel and Constant 2018a, Cumming et al. 2021. The nomenclature proposed here for the dissected vomer can be used in nearly all species, and the structure of the vomer provides useful differential characters when studied from different angles. ...
... The apical portion of the body of the vomer is often upcurved and armed with one to multiple spines. In many species the vomer ends in a single apical spine (Fig. 4, as), referred to as hook(s) or prong(s) by several authors (Bradler 1999, Bradler et al. 2014, Cumming et al. 2021. The apex can sometimes be more complex, bearing two (Neooxyartes Ho, 2018, Cryptophyllium) or more (Spinohirasea Zompro, 2002, Paramenexenus Redtenbacher, 1908 sometimes blunt, spines (Hennemann 2007, Bresseel and Constant 2018a, Bradler et al. 2014, Cumming et al. 2021. ...
... In many species the vomer ends in a single apical spine (Fig. 4, as), referred to as hook(s) or prong(s) by several authors (Bradler 1999, Bradler et al. 2014, Cumming et al. 2021. The apex can sometimes be more complex, bearing two (Neooxyartes Ho, 2018, Cryptophyllium) or more (Spinohirasea Zompro, 2002, Paramenexenus Redtenbacher, 1908 sometimes blunt, spines (Hennemann 2007, Bresseel and Constant 2018a, Bradler et al. 2014, Cumming et al. 2021. The body of the vomer can be symmetrical (Neooxyartes) to asymmetrical (Cryptophyllium) depending on the size and direction of the spine(s) Constant 2018a, Cumming et al. 2021). ...
A new species of Presbistus Kirby, 1896, Presbistus vitivorus sp. nov., is described from Cambodia based on both sexes, nymphs, and eggs. Male genitalia and vomer are described and figured. Illustrations of adults, nymphs, specimens in situ, host plants, a distribution map and records on biology and breeding in captivity are provided. The host plants of the species belong to the family Vitaceae. The genus Presbistus and the family Aschiphasmatidae are recorded from Cambodia for the first time. The species diversity and the distribution of the genus are discussed, and it is shown that the genus is restricted to Sundaland. A nomenclature for the morphology of the dissected vomer is proposed and tries to homologize the previously used terms.
... 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). ...
... 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 . Molecular analyses preceding this study had already repeatedly demonstrated that Phyllium (and Chitoniscus) are paraphyletic and that the Phylliidae are in need of revision 24,25,29,40 . ...
... [2][3][4][5]. However, Chitoniscus and Phyllium are recovered as paraphyletic, which was already shown in previous studies based on molecular data 24,25,29,40,41 . The Chitoniscus spp. ...
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.
... Molecular species delimitation has been successfully used in phasmids in several instances (Glaw et al., 2019;Bank et al., 2021;Cumming et al., 2021). Velonà et al. (2015) first applied DNA barcoding on 16 Australian stick insect taxa, and retrieved a high differentiation among three putative morphospecies of the genus Candovia Stål, 1875: Candovia spp. ...
The Phasmida genus Candovia comprises nine traditionally recognized species, all endemic to Australia. In this study, Candovia diversity is explored through molecular species-delimitation analyses using the COIFol gene fragment and phylogenetic inferences leveraging seven additional mitochondrial and nuclear loci. Molecular results were integrated with morphological observations, leading us to confirm the already described species and to the delineation of several new taxa and of the new genus Paracandovia. New Candovia species from various parts of Queensland and New South Wales are described and illustrated (C. alata sp. nov., C. byfieldensis sp. nov., C. dalgleishae sp. nov., C. eungellensis sp. nov., C. karasi sp. nov., C. koensi sp. nov. andC. wollumbinensis sp. nov.). New combinations are proposed and species removed from synonymy with the erection of the new genus Paracandovia (P. cercata stat. rev., comb. nov., P. longipes stat. rev., comb. nov., P. pallida comb. nov., P. peridromes comb. nov., P. tenera stat. rev., comb. nov.). Phylogenetic analyses suggest that the egg capitulum may have independently evolved multiple times throughout the evolutionary history of these insects. Furthermore, two newly described species represent the first taxa with fully developed wings in this previously considered apterous clade.
... For the outgroup, we further included two species of Embioptera, which were repeatedly recovered as the sister group of Phasmatodea [4,30,56,121]. Our gene sampling comprised three nuclear (18S, 28S, H3) and four mitochondrial loci (12S, 16S, COI, COII) of which data for numerous specimens were already available on GenBank [30,54,55,71,80,89,90,101,[122][123][124][125][126][127][128][129][130][131]. Additionally, we generated new data for 194 specimens following the protocol given by Bank et al. [80] with primers adopted from Buckley et al. [124] and Robertson et al. [55] (see Bank et al. [80]), and deposited the sequences in GenBank under accession numbers OK314533-OK314857, OK324156-OK324324, OK333379-OK334012 (Additional file 3: Table S7). ...
Background
The re-evolution of complex characters is generally considered impossible, yet, studies of recent years have provided several examples of phenotypic reversals shown to violate Dollo’s law. Along these lines, the regain of wings in stick and leaf insects (Phasmatodea) was hypothesised to have occurred several times independently after an ancestral loss, a scenario controversially discussed among evolutionary biologists due to overestimation of the potential for trait reacquisition as well as to the lack of taxonomic data.
Results
We revisited the recovery of wings by reconstructing a phylogeny based on a comprehensive taxon sample of over 500 representative phasmatodean species to infer the evolutionary history of wings. We additionally explored the presence of ocelli, the photoreceptive organs used for flight stabilisation in winged insects, which might provide further information for interpreting flight evolution. Our findings support an ancestral loss of wings and that the ancestors of most major lineages were wingless. While the evolution of ocelli was estimated to be dependent on the presence of (fully-developed) wings, ocelli are nevertheless absent in the majority of all examined winged species and only appear in the members of few subordinate clades, albeit winged and volant taxa are found in every euphasmatodean lineage.
Conclusion
In this study, we explored the evolutionary history of wings in Phasmatodea and demonstrate that the disjunct distribution of ocelli substantiates the hypothesis on their regain and thus on trait reacquisition in general. Evidence from the fossil record as well as future studies focussing on the underlying genetic mechanisms are needed to validate our findings and to further assess the evolutionary process of phenotypic reversals.
... 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.
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.
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.
Paraphasma Redtenbacher, 1906 is a genus of fully-winged stick insects occurring in central and northern South America. We carried out a morphology-based taxonomic revision of this genus with emphasis on the phallic organ, a structure that has been poorly explored for taxonomic purposes in Phasmatodea. Additionally, pairwise genetic distances between mitochondrial COI gene sequences were calculated for ten Paraphasma specimens representing six species. We recognize nine valid species in the genus plus one nomen dubium, Paraphasma fasciatum Gray, 1835. We redescribe Paraphasma and the species previously assigned to it, describe Paraphasma indistinctum Chiquetto-Machado sp. nov., Paraphasma sooretama Chiquetto-Machado sp. nov. and Paraphasma spinicauda Chiquetto-Machado sp. nov., and provide a key to the species in the genus. The male of Paraphasma minus Redtenbacher, 1906 is described for the first time, as well as the eggs of six species. We transfer Paraphasma amabile Redtenbacher, 1906 to Pseudophasma Kirby, 1896 (comb. nov.) and synonymize Pseudophasma xanthotaenidium Günther, 1930 under this species (syn. nov.). In addition, Phasma perspicillaris Stoll, 1813 is removed from the synonymy of Paraphasma laterale (Fabricius, 1775) and synonymized under Parastratocles xanthomela (Olivier, 1792) (syn. nov.). The examination of the phallic organ was essential for species delimitation, as most species of Paraphasma are very similar in the external morphology of both sexes. The analysis of the COI sequences supported the species delimitation, resulting in remarkably lower pairwise distances between conspecific individuals (p-distance ≤ 2.0%) than between different species (p-distance 6.9–17.5%). We hope that this paper will stimulate further studies exploring the taxonomic and phylogenetic potential of the internal male genitalia of stick insects.
Insects of the order Phasmatodea are mainly distributed in the tropics and subtropics and are best known for their remarkable camouflage as plants. In this study, we sequenced three complete mitochondrial genomes from three different families: Orestes guangxiensis, Peruphasma schultei, and Phryganistria guangxiensis. The lengths of the three mitochondrial genomes were 15,896 bp, 16,869 bp, and 17,005 bp, respectively, and the gene composition and structure of the three stick insects were identical to those of the most recent common ancestor of insects. The phylogenetic relationships among stick insects have been chaotic for a long time. In order to discuss the intra- and inter-ordinal relationship of Phasmatodea, we used the 13 protein-coding genes (PCGs) of 85 species for maximum likelihood (ML) and Bayesian inference (BI) analyses. Results showed that the internal topological structure of Phasmatodea had a few differences in both ML and BI trees and long-branch attraction (LBA) appeared between Embioptera and Zoraptera, which led to a non-monophyletic Phasmatodea. Consequently, after removal of the Embioptera and Zoraptera species, we re-performed ML and BI analyses with the remaining 81 species, which showed identical topology except for the position of Tectarchus ovobessus (Phasmatodea). We recovered the monophyly of Phasmatodea and the sister-group relationship between Phasmatodea and Mantophasmatodea. Our analyses also recovered the monophyly of Heteropterygidae and the paraphyly of Diapheromeridae, Phasmatidae, Lonchodidae, Lonchodinae, and Clitumninae. In this study, Peruphasma schultei (Pseudophasmatidae), Phraortes sp. YW-2014 (Lonchodidae), and species of Diapheromeridae clustered into the clade of Phasmatidae. Within Heteropterygidae, O. guangxiensis was the sister clade to O. mouhotii belonging to Dataminae, and the relationship of (Heteropteryginae + (Dataminae + Obriminae)) was recovered.
With every molecular review involving Chitoniscus Stål, 1875 sensu lato samples from Fiji and New Caledonia revealing polyphyly, the morphology from these two distinct clades was extensively reviewed. Morphological results agree with all previously published molecular studies and therefore Trolicaphyllium gen. nov. is erected to accommodate the former Chitoniscus sensu lato species restricted to New Caledonia, leaving the type species Chitoniscus lobiventris (Blanchard, 1853) and all other Fijian species within Chitoniscus sensu stricto. Erection of this new genus for the New Caledonian species warrants the following new combinations: Trolicaphyllium brachysoma (Sharp, 1898), comb. nov., Trolicaphyllium erosus (Redtenbachher, 1906), comb. nov., and Trolicaphyllium sarrameaense (Größer, 2008a), comb. nov. Morphological details of the female, male, freshly hatched nymph, and egg are illustrated and discussed alongside the Chitoniscus sensu stricto in order to differentiate these two clades which have been mistaken as one for decades.
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.
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.
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.
IQ-TREE (http://www.iqtree.org, last accessed February 6, 2020) is a user-friendly and widely used software package for phylogenetic inference using maximum likelihood. Since the release of version 1 in 2014, we have continuously expanded IQ-TREE to integrate a plethora of new models of sequence evolution and efficient computational approaches of phylogenetic inference to deal with genomic data. Here, we describe notable features of IQ-TREE version 2 and highlight the key advantages over other software.
The study was carried out to find the approaches to the assessment of the conservation effectiveness of Red Data insect species in the especially protected natural territories in Belarus (East Europe) and Vietnam (Southeast Asia). A review of the structure of a modern system of protected areas in both countries is provided. The structure is similar and includes the national parks, nature reserves, the special protected areas, and a few other categories of protected areas like, for example, scientific and experimental forest areas in Vietnam. The total number of the protected territories is many times higher in Belarus than in Vietnam-1,285 versus 164, although the area of the protected territories in both countries is different, as well as a total area of the countries. The lection of the criteria to assess the conservation effectiveness of Red Data insect species in the especially protected natural territories is very difficult because their multiplicity and different interpretations. We tried to use the only criterion of presence of the Red Data species in the protected areas. The nature reserves, national parks and 5 the largest special protected territories (zakaznik) were analysed in this way in Belarus. In Belarus, the effectiveness of the protected areas according to this criterion is very high because: 1). More than 80 % of Red Data species are presented in the protected areas, 2). There are at least three Red Data species inhabited every territory and 3).There are species presented almost exclusively in the protected territories. There is a problem to use this criterion in Vietnam because of the poor knowledge of the regional biodiversity and the local faunas, existing of the large number of the endemics and undescribed species, as well as the rarity of many species. Some problems of the conservation of Red Data insect species in the especially protected natural territories are outlined.
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.
A record is provided for the Pink Orchid Mantis, Hymenopus coronatus Olivier, 1792 (Insecta: Mantodea: Hymenopodidae) from the Queen Sirikit Botanical Garden, Chiang Mai, Chiang Mai Province, northern Thailand on 1 July 2017. Brief notes are provided on the specimen which was noticed resting motionless on the top of two flowers of a 2.5 m high specimen of the tropical Perfume Tree, Cananga odorata (Lam.) Hook.f. & Thomson (Annonaceae). The biology, "mimicry" system of the various life stages and previously published literature on the species are reviewed. We have postulated the view that these mantids do not normally occupy orchids for foraging purposes and that they never evolved in the first place to mimic orchids. The supposed orchid mimicry is simply coincidence. The mantids are extremely rare in the field. Our observations indicate that the mantid forages on insects attracted to fragrant flowers where at least in the adult stage and the mantid coloration acts as like a break up pattern (i.e. camouflage) in relation to the environment. The vernacular name of the Pink Orchid Mantis is thus a serious misnomer.
Research in the Indo-Pacific region has contributed massively to the understanding of speciation. White-eyes (Aves: Zosteropidae: Zosterops), a lineage containing both widespread ‘supertramp’ species and a high proportion of island endemics, have provided invaluable models. Molecular tools have increased speciation research, but delimiting species remains problematic. We investigated the evolutionary history of Zosterops species in south-east Sulawesi using mitochondrial DNA, morphometric, song and plumage analyses, to draw species limits and assess which techniques offer best resolution. Our investigation revealed a novel Zosterops species, >3000 km from its closest relative. Additionally, we demonstrated unanticipated diversity in the alleged ‘supertramp’ Zosterops chloris and propose the Wakatobi Islands subspecies (Z. c. flavissimus) to be given full species status. Furthermore, we provide the first molecular and phenotypic assessment of the Sulawesi endemic Zosterops consobrinorum. While local populations of this species vary in either genetics or morphometrics, none show consistency across measures. Therefore, we propose no change to Zosterops consobrinorum taxonomy. This study gives insight into one of the great Indo-Pacific radiations and demonstrates the value of using multiple lines of evidence for taxonomic review.
Stick and leaf insects (Phasmatodea) are large, tropical, predominantly nocturnal herbivores, which exhibit extreme masquerade crypsis, whereby they morphologically and behaviorally resemble twigs, bark, lichen, moss, and leaves. Females employ a wide range of egg-laying techniques, largely corresponding to their ecological niche, including dropping or flicking eggs to the forest floor, gluing eggs to plant substrate, skewering eggs through leaves, ovipositing directly into the soil, or even producing a complex ootheca. Phasmids are the only insects with highly species-specific egg morphology across the entire order, with specific egg forms that correspond to oviposition technique. We investigate the temporal, biogeographic, and phylogenetic pattern of evolution of egg-laying strategies in Phasmatodea. Our results unequivocally demonstrate that the ancestral oviposition strategy for female stick and leaf insects is to remain in the foliage and drop or flick eggs to the ground, a strategy that maintains their masquerade. Other major key innovations in the evolution of Phasmatodea include the (1) hardening of the egg capsule in Euphasmatodea; (2) the repeated evolution of capitulate eggs (which induce ant-mediated dispersal, or myrmecochory); (3) adapting to a ground or bark dwelling microhabitat with a corresponding shift in adult and egg phenotype and egg deposition directly into the soil; and (4) adhesion of eggs in a clade of Necrosciinae that led to subsequent diversification in oviposition modes and egg types. We infer at minimum 16 independent origins of a burying/inserting eggs into soil/crevices oviposition strategy, 7 origins of gluing eggs to substrate, and a single origin each of skewering eggs through leaves and producing an ootheca. We additionally discuss the systematic implications of our phylogenetic results. Aschiphasmatinae is strongly supported as the earliest diverging extant lineage of Euphasmatodea. Phylliinae and Diapheromerinae are both relatively early diverging euphasmatodean taxa. We formally transfer Otocrania from Cladomorphinae to Diapheromerinae and recognize only two tribes within Diapheromerinae: Diapheromerini sensu nov. and Oreophoetini sensu nov. We formally recognize the clade comprising Necrosciinae and Lonchodinae as Lonchodidae stat. rev. sensu nov.
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.