On seven undescribed leaf insect species revealed within the recent "Tree of Leaves" (Phasmatodea, Phylliidae).

With the recent advance in molecular phylogenetics focused on the leaf insects (Phasmatodea, Phylliidae), gaps in knowledge are beginning to be filled. Yet, shortcomings are also being highlighted, for instance, the unveiling of numerous undescribed phylliid species. Here, some of these taxa are described, including Phylliumiyadaonsp. nov. from Mindoro Island, Philippines; Phylliumsamarensesp. nov. from Samar Island, Philippines; Phylliumortizisp. nov. from Mindanao Island, Philippines; Pulchriphylliumheraclessp. nov. from Vietnam; Pulchriphylliumdelisleisp. nov. from South Kalimantan, Indonesia; and Pulchriphylliumbhaskaraisp. nov. from Java, Indonesia. Several additional specimens of these species together with a seventh species described herein, Pulchriphylliumanangusp. nov. from southwestern India, were incorporated into a newly constructed phylogenetic tree. Additionally, two taxa that were originally described as species, but in recent decades have been treated as subspecies, are elevated back to species status to reflect their unique morphology and geographic isolation, creating the following new combinations: Pulchriphylliumscythe (Gray, 1843) stat. rev., comb. nov. from Bangladesh and northeastern India, and Pulchriphylliumcrurifolium (Audinet-Serville, 1838) stat. rev., comb. nov. from the Seychelles islands. Lectotype specimens are also designated for Pulchriphylliumscythe (Gray, 1843) stat. rev., comb. nov. and Pulchriphylliumcrurifolium (Audinet-Serville, 1838) stat. rev., comb. nov. from original type material.

Leaves that walk and eggs that stick: comparative functional morphology and evolution of the adhesive system of leaf insect eggs (Phasmatodea: Phylliidae).

Phylliidae are herbivorous insects exhibiting impressive cryptic masquerade and are colloquially called "walking leaves". They imitate angiosperm leaves and their eggs often resemble plant seeds structurally and in some cases functionally. Despite overall morphological similarity of adult Phylliidae, their eggs reveal a significant diversity in overall shape and exochorionic surface features. Previous studies have shown that the eggs of most Phylliidae possess a specialised attachment mechanism with hierarchical exochorionic fan-like structures (pinnae), which are mantled by a film of an adhesive secretion (glue). The folded pinnae and glue respond to water contact, with the fibrous pinnae expanding and the glue being capable of reversible liquefaction. In general, the eggs of phylliids appear to exhibit varying structures that were suggested to represent specific adaptations to the different environments the eggs are deposited in. Here, we investigated the diversity of phylliid eggs and the functional morphology of their exochorionic structure. Based on the examination of all phylliid taxa for which the eggs are known, we were able to characterise eleven different morphological types. We explored the adhesiveness of these different egg morphotypes and experimentally compared the attachment performance on a broad range of substrates with different surface roughness, surface chemistry and tested whether the adhesion is replicable after detachment in multiple cycles. Furthermore, we used molecular phylogenetic methods to reconstruct the evolutionary history of different egg types and their adhesive systems within this lineage, based on 53 phylliid taxa. Our results suggest that the egg morphology is congruent with the phylogenetic relationships within Phylliidae. The morphological differences are likely caused by adaptations to the specific environmental requirements for the particular clades, as the egg morphology has an influence on the performance regarding the surface roughness. Furthermore, we show that different pinnae and the adhesive glue evolved convergently in different species. While the evolution of the Phylliidae in general appears to be non-adaptive judging on the strong similarity of the adults and nymphs of most species, the eggs represent a stage with complex and rather diverse functional adaptations including mechanisms for both fixation and dispersal of the eggs.

Analysis of plant-derived carotenoids in camouflaging stick and leaf insects (Phasmatodea).

A common way to avoid predators is by use of camouflage, a strategy which the stick and leaf insects (Phasmatodea) have refined by appearing as leaves, sticks, lichen, and moss. Stick and leaf insects have perfected their camouflage by sequestering diet-based carotenoids within their exoskeleton. Visual and chemical details of such camouflage have likely been influenced through the millennia of co-evolution between these insects and the plants they mimic. It is this evolutionary struggle that has resulted in a plethora of morphological and chemical adaptations across the stick and leaf insect family tree. In this chapter we discuss prior stick and leaf insect carotenoid studies, proper rearing of specimens, and describe methods for preparation of insect exoskeleton and plant samples, carotenoid extraction and analysis.

Three new genera and one new species of leaf insect from Melanesia (Phasmatodea, Phylliidae).

With the first large-scale Phylliidae molecular phylogeny recently published adding a great deal of clarity 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: Rakaphylliumschultzei (Giglio-Tos, 1912), comb. nov., Rakaphylliumexsectum (Zompro, 2001b), comb. nov., and Vaabonbonphylliumgroesseri (Zompro, 1998), comb. nov. Additionally, while reviewing material an undescribed Vaabonbonphyllium gen. nov. specimen was located and is herein described as Vaabonbonphylliumrafidahae 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: Acentetaphylliumbrevipenne (Größer, 1992), comb. nov., Acentetaphylliumlarssoni (Cumming, 2017), comb. nov., Acentetaphylliummiyashitai (Cumming et al. 2020), comb. nov., and Acentetaphylliumstellae (Cumming, 2016), comb. nov. With the addition of several new genera, a key to phylliid genera is included for adult males and females.

A tree of leaves: Phylogeny and historical biogeography of the leaf insects (Phasmatodea: Phylliidae).

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 the ~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.

Resolving a century-old case of generic mistaken identity: polyphyly of Chitoniscus sensu lato resolved with the description of the endemic New Caledonia Trolicaphyllium gen. nov. (Phasmatodea, Phylliidae).

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 Chitoniscuslobiventris (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: Trolicaphylliumbrachysoma (Sharp, 1898), comb. nov., Trolicaphylliumerosus (Redtenbachher, 1906), comb. nov., and Trolicaphylliumsarrameaense (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.

Assemblages of Necrophagous Blow Flies (Diptera: Calliphoridae) in Four Contrasting Ecoregions of Los Angeles County, California, United States.

An ecological study to evaluate necrophagous species richness, abundance, and diversity was conducted in four contrasting ecoregions of Los Angeles County, California, United States: a highly anthropized area, two moderately populated areas, and a sparsely populated area. Our hypothesis states that there will be higher abundance of exotic species in greatly anthropized ecoregions and higher diversity of native calliphorid flies in sparsely populated sites. A total of 2,883 blow flies belonging to 12 species were captured from the four ecoregions. The exotic Chrysomya rufifacies (Macquart) (Diptera: Calliphoridae) was the most abundant species, representing 72.84% of all Calliphoridae collected. The highest abundance was in the highly anthropized ecoregion. Differences of the blow fly assemblages across environmentally and anthropically different ecoregions were found, and that the relative abundance of the exotic species increased with anthropization. Our results suggest that both anthropization and elevation alter the structure of blow fly assemblages, modifying the abundance and species richness of Calliphoridae. The results of this study support the hypothesis of biotic homogenization, since we found a higher proportional abundance of exotic species and lower diversity of native blow flies in ecoregions heavily modified by human action. Additionally, the hypothesis of intermediate disturbance was supported as the highest diversity was found in the ecoregion with moderate anthropization mixed with natural elements. This is the first study to conduct an intensive survey of the Calliphoridae assemblages in Los Angeles County, which can be potentially used as a tool in human and veterinary health and forensic sciences.

Drawing the Excalibur bug from the stone: adding credibility to the double-edged sword hypothesis of coreid evolution (Hemiptera, Coreidae).

A new genus and species of exaggerated antennae Coreidae is described from Myanmar amber of the Late Cretaceous (Cenomanian stage). Ferriantenna excalibur gen. et sp. nov. appears related to another Cretaceous coreid with exaggerated antennae, Magnusantenna Du & Chen, 2021, but can be differentiated by the fourth antennal segment which is short and paddle-like, the undulating shape of the pronotum and mesonotum, and the shorter and thicker legs. The new coreid, with elaborately formed antennae and simple hind legs instead of the typical extant coreid morphology with simple antennae and elaborately formed hind legs, begs the question: why were the elaborate features of the antennae lost in favor of ornate hind legs? Features that are large and showy are at higher risk of being attacked by predators or stuck in a poor molt and subjected to autotomy and are therefore lost at a higher rate than simple appendages. We hypothesize that because elaborate antennae play an additional significant sensory role compared to elaborate hind legs, that evolutionarily it is more costly to have elaborate antennae versus elaborate hind legs. Thus, through the millenia, as coreid evolution experimented with elaborate/ornate features, those on the antennae were likely selected against in favor of ornate hind legs.

Cryptophyllium, the hidden leaf insects - descriptions of a new leaf insect genus and thirteen species from the former celebicum species group (Phasmatodea, Phylliidae).

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 phylogenetic 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 ericoriaiHennemann 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.; Cryptophyllium 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.; Cryptophyllium 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 Vietnam: 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, illustrated, and molecularly compared to congenerics. With the molecular results revealing cryptic taxa, it was found necessary for Cryptophyllium westwoodii (Wood-Mason, 1875), comb. nov. to have a neotype specimen designated to allow accurate differentiation from congenerics. To conclude, male and female dichotomous keys to species for the Cryptophyllium gen. nov. are presented.

Lost lovers linked at long last: elusive female Nanophyllium mystery solved after a century of being placed in a different genus (Phasmatodea, Phylliidae).

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.

Walaphyllium subgen. nov., the dancing leaf insects from Australia and Papua New Guinea with description of a new species (Phasmatodea, Phylliidae).

A new subgenus, Walaphyllium subgen. 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) lelantossp. 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.

Notes on the leaf insects of the genus Phyllium of Sumatra and Java, Indonesia, including the description of two new species with purple coxae (Phasmatodea, Phylliidae).

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 Peninsular 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.

Addendum to "Leaf insects from Luzon, Philippines, with descriptions of four new species, the new genus Pseudomicrophyllium, and redescription of Phyllium (Phyllium) geryon Gray, 1843, (Phasmida: Phylliidae)".

At the time Cumming et al. (2017) was published, the GenBank accession numbers for partial cytochrome c oxidase subunit I (COI) DNA barcode sequences from Microphyllium specimens sampled in the phylogenetic analysis were not yet available. This information is provided in Table 1.

A second new species of Nanophyllium Redtenbacher, 1906 from the northern coast of New Guinea (Phasmida, Phylliidae).

A new species of leaf insect, Nanophyllium larssoni n. sp. is described and illustrated bringing the known species in the genus to a total of six. Like all species of Nanophyllium Redtenbacher, 1906, the new species is named from a single male specimen, which is deposited in the San Diego Natural History Museum. With this newly identified species, two clear species-groups emerge and are described, in detail, in the species key. Measurements of anatomical figures were made to the nearest 0.1 mm.

Leaf insects from Luzon, Philippines, with descriptions of four new species, the new genus Pseudomicrophyllium, and redescription of Phyllium (Phyllium) geryon Gray, 1843, (Phasmida: Phylliidae).

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. Pseudomicrophyllium 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.) antonkozlovi 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.

A new species of Nanophyllium Redtenbacher, 1906 from the northern coast of New Guinea (Phasmida, Phylliidae).

A new species of leaf insect, Nanophyllium stellae n. sp. is described and illustrated. As for the majority of the current taxa of Nanophyllium Redtenbacher, 1906, the new species is named from a single male specimen, as no other specimens are currently known to exist. A key to the current species of Nanophyllium Redtenbacher, 1906 is provided. The holotype was photographed using a Canon 5D Mark II and a MP-E 65mm macro lens and will be deposited in the San Diego Natural History Museum. Photos were stacked using Zerene photo stacking software, version 1.04, 64-bit. Measurements of anatomical figures were made to the nearest 0.1 mm. Currently, only four species of Nanophyllium Redtenbacher, 1906 have been described, all from males, with N. stellae n. sp. marking the fifth species. To date, there have been no females described, save one possible N. hasenpuschi Brock & Grösser, 2008 specimen illustrated from Nabire, Irian Jaya (Brock & Hasenpusch, 2003).