Trophobiosis between a new species of Acropyga (Hymenoptera, Formicidae) and new Neochavesia (Hemiptera, Xenococcidae) from Peru, and establishment of the Acropygasmithii species-group.

We describe a new pair of trophobiotic partners from the ant genus Acropyga and the root mealybug genus Neochavesia. A recent field study on Acropyga ants and associated root mealybugs, conducted in the Peruvian Amazon, led to the discovery of Acropygamanuense LaPolla & Schneider, sp. nov. and its root mealybug symbiont Neochavesiapodexuta Schneider & LaPolla, sp. nov. The new root mealybug belongs to the family Xenococcidae, whose members are all obligate associates of Acropyga ants. Providing joint descriptions of new mutualist partners in the same article is a novel approach for this system, and it offers benefits to the ongoing study of mutualism and patterns of association among these symbiotic ants and scales. Here, we also begin to revise the species-group composition of Acropyga by establishing the smithii species-group, and we provide updated information to aid in identifying the new ant species and root mealybug species.

Distinguishing Symbiotic Partners of Acropyga Ants from Free-Living Soil Inhabitants.

The fruitful study of associations between ants and scale insects yields insight into the mechanisms that shape these symbioses. Field collections provide the basic information linking partnered species, and as such it is critical that collection techniques from the field reflect true species-to-species partnerships in the published literature. It is equally critical that such practices limit the potential for mistaking free-living "neighbors" for symbiotic partners and publishing erroneous associations. This article describes a protocol for collecting subterranean scale insects and associated Acropyga Roger ants, which relies upon the activity of worker ants to sort and distinguish symbionts from free-living scale insects that happen to live near the colony. By collecting samples of ants and scales into nest boxes and allowing a resting period of several hours, worker ants will gather symbiotic partners into dense, protected clusters in which symbionts are actively tended. Free-living scale insects neighboring the colony can be collected from soil along with colony samples, but these free-living individuals are excluded from protective clusters and ignored by workers. Following confirmation of ant attendance, true symbiotic partners can be confidently collected, preserved, and recorded for future study. We illustrate the value of employing this collection protocol using a case study from Peru.

A Neotropical complex of Ripersiella species (Hemiptera, Coccomorpha, Rhizoecidae) collected from the nests of Acropyga ants (Hymenoptera, Formicidae).

We describe five new Neotropical species of Ripersiella living in association with Acropyga ants: R.campensis sp. nov., R.illicians sp. nov., R.montanae sp. nov., R.pediandensis sp. nov., and R.telalia sp. nov. We also redescribe R.andensis and R.colombiensis based on type specimens and other collections. Together, these seven species form a morphologically similar group that we informally refer to as the andensis-complex of Ripersiella. All members of the andensis-complex are confirmed or are speculated to be mutualists of Acropyga ants. We discuss the implications of these associations and provide an identification key to the Neotropical species of Ripersiella that are lacking bitubular cerores, including the new species.

Four new species of Aspidiotini (Hemiptera, Diaspididae, Aspidiotinae) from Panama, with a key to Panamanian species.

Four new species of armored scale insect, Clavaspis selvatica sp. nov., Clavaspis virolae sp. nov., Davidsonaspis tovomitae sp. nov., and Rungaspis neotropicalis sp. nov., are described and illustrated from Panama. We also transfer two previously described species of Panamanian Aspidiotini to new genera, Hemiberlesia crescentiae (Ferris) comb. nov. and Rungaspis rigida (Ferris) comb. nov., and report the first record of Selenaspidopsis browni Nakahara in Panama. A key to the species of Aspidiotini occurring in Panama is provided.

Facilitative pollinator sharing decreases with floral similarity in multiple systems.

Investigating the factors that determine whether interactions are competitive or facilitative is essential to understanding community structure and trait evolution. Co-flowering plants interact indirectly through shared pollinators, and meta-analyses suggest that phylogenetic relatedness and floral trait similarity may predict the outcome of these interactions. In a comparative approach, we manipulated the floral community across five focal species to assess how floral similarity and phylogenetic relatedness affect the outcome of interactions. To assess the extent of pollinator-mediated competition versus facilitation, we compared pollen limitation in five focal species growing with floral neighbors (either congeners or neighbors from a different family) relative to a control (growing alone). We measured floral morphology, color, and nectar traits to calculate multivariate floral similarity between species pairs and inferred a phylogeny to calculate phylogenetic distance. Pollinator-mediated interaction values were regressed against floral similarity and phylogenetic distance. We found evidence of pollinator-mediated facilitation in nine of 13 species pairs. Furthermore, floral similarity and phylogenetic distance reduced facilitative interactions, but the latter relationship was not significant when controlling for the identity of the focal species. Our results suggest that facilitative pollinator sharing is more common than reported in the literature, but co-flowering plant species with similar floral traits are less likely to facilitate pollination. A better understanding of the factors that promote facilitation versus competition has important potential applications for managing rare and invasive species.

A mid-Cretaceous female scale insect (Hemiptera: Sternorrhyncha: Coccomorpha) in Burmese amber.

A new genus and species of scale insect (Hemiptera: Coccomorpha) is described from a female specimen in mid-Cretaceous Burmese (Myanmar) amber. Fossil female scales are rare and the present species, described as Paleolepidotus macrocolus gen. et sp. n., has such an unusual assortment of morphological features that it could not be assigned to any particular extant or extinct family. The small, ferruginous specimen exhibits a series of long wax pencils that extend around the body, including the head. The antennae and legs are quite long compared to other extant and extinct scale fossils. Of special interest are the protruding eyes, and a conical-triangular rostrum arising from between the forelegs; the claws with bifid apices are also unique. The ovisac contains immature stages.

Trophobiosis between a new species of Williamsrhizoecus (Hemiptera: Coccomorpha: Rhizoecidae) and Acropyga silvestrii (Hymenoptera: Formicidae) in Tanzania.

A new myrmecophilous species of root mealybug, Williamsrhizoecus udzungwensis sp. n., is described from individuals found living within a nest of Acropyga silvestrii in the Udzungwa Mountains of Tanzania. Acropyga ants are highly specialized, obligate associates of scale insects, typically members of the scale family Xenococcidae. Acropyga are best known for vertically transmitting trophobiotic partners during their nuptial flights and for housing them within brood chambers. This article presents the first record of trophobiosis between a species of Williamsrhizoecus and Acropyga, and only the second record of an association between Acropyga and rhizoecids in the Old World. This discovery contributes important information about the few species of Rhizoecidae confirmed to engage in these unique symbioses, each putatively the result of a past horizontal transmission event from a xenococcid to a rhizoecid lineage. Included is a discussion on the diagnosis of Williamsrhizoecus and an updated key to the species.

Five new species of Aspidiotini (Hemiptera, Diaspididae, Aspidiotinae) from Argentina, with a key to Argentine species.

Five new species of armored scale insect from Argentina are described and illustrated based upon morphological and molecular evidence from adult females: Chortinaspis jujuyensis sp. nov., Clavaspis patagonensis sp. nov., Hemiberlesia ozolita sp. nov., Melanaspis lilloi sp. nov., and Melanaspis targionoides sp. nov. The genera Chortinaspis and Melanaspis are recorded for the first time from this country. An identification key to all recorded species from tribe Aspidiotini occurring in Argentina is provided.

Phylogeny and classification of armored scale insects (Hemiptera: Coccomorpha: Diaspididae).

Armored scale insects (Hemiptera: Coccomorpha: Diaspididae) are major economic pests and are among the world's most invasive species. Here we describe a system of specimen and identification management that establishes a basis for well-vouchered molecular identification. We also present an expanded Bayesian phylogenetic analysis based on concatenated fragments of 4 genetic loci: the large ribosomal subunit (28S), elongation factor-1 alpha (EF-1α), cytochrome oxidase I and II (COI‒II), and the small ribosomal subunit (16S) of the primary endosymbiont, Uzinura diaspidicola (Bacteroidetes: Flavobacteriales). Our sample includes 1,389 individuals, representing 11 outgroup species and at least 311 described and 61 undescribed diaspidid species. The results broadly support Takagi's 2002 classification but indicate that some revisions are needed. We propose a revised classification recognizing 4 subfamilies: Ancepaspidinae Borchsenius, new rank, Furcaspidinae Balachowsky, new rank, Diaspidinae Targioni Tozzetti, and Aspidiotinae Westwood. Within Aspidiotinae, in addition to the existing tribes Aspidiotini Westwood, Parlatoriini Leonardi, Odonaspidini Ferris, Leucaspidini Atkinson, and Smilacicolini Takagi, we recognize as tribes Gymnaspidini Balachowsky, new rank, and Aonidiini Balachowsky, new rank. Within Diaspidinae we recognize the 2 tribes Lepidosaphidini Shimer and Diaspidini Targioni Tozzetti, and within Diaspidini we recognize three subtribes: Diaspidina Targioni Tozzetti, Fioriniina Leonardi, and Chionaspidina Brues Melander. We regard Kuwanaspidina Borchsenius as a junior synonym of Fioriniina, Thysanaspidini Takagi as a junior synonym of Leucaspidini, and Protodiaspidina Takagi and Ulucoccinae Takagi as junior synonyms of Chionaspidina. To clarify the composition of the higher taxa we describe 2 new genera for Australian species heretofore misplaced in the genus Ancepaspis Ferris: Brimblecombia Normark (Aonidiini) and Hendersonaspis Normark (Leucaspidini). We also propose many additional minor modifications to the taxonomy of Diaspididae, including the following new combinations, revived combinations, and replacement names: Aonidia edgerleyi (Mamet), new combination (from Bigymnaspis Balachowsky); Aonidomytilus espinosai Porter, revived combination (from Porterinaspis González); Aspidiotus badius (Brain), new combination (this and the next 5 Aspidiotus species all from Aonidia Targioni Tozzetti); Aspidiotus biafrae (Lindinger), new combination; Aspidiotus chaetachmeae (Brain), new combination; Aspidiotus laticornis (Balachowsky), new combination; Aspidiotus rhusae (Brain), new combination; Aspidiotus sclerosus (Munting), new combination; Brimblecombia asperata (Brimblecombe), new combination (this and the next 5 Brimblecombia species all from Ancepaspis); Brimblecombia longicauda (Brimblecombe), new combination; Brimblecombia magnicauda (Brimblecombe), new combination; Brimblecombia reticulata (Brimblecombe), new combination; Brimblecombia rotundicauda (Brimblecombe), new combination; Brimblecombia striata (Brimblecombe), new combination; Cooleyaspis pseudomorpha (Leonardi), new combination (from Dinaspis Leonardi); Cupidaspis wilkeyi (Howell Tippins), new combination (from Paracupidaspis Howell Tippins); Cupressaspis isfarensis Borchsenius, revived combination (this species, the next 2 species in Cupressaspis Borchsenius, revived genus, and the next 9 species in Diaspidiotus Cockerell all from Aonidia); Cupressaspis mediterranea (Lindinger), revived combination; Cupressaspis relicta (Balachowsky), new combination; Diaspidiotus atlanticus (Ferris), new combination; Diaspidiotus marginalis (Brain), new combination; Diaspidiotus maroccanus (Balachowsky), new combination; Diaspidiotus mesembryanthemae (Brain), new combination; Diaspidiotus opertus (De Lotto), new combination; Diaspidiotus shastae (Coleman), new combination; Diaspidiotus simplex (Leonardi), new combination; Diaspidiotus visci (Hall), new combination; Diaspidiotus yomae (Munting), new combination; Diaspis arundinariae (Tippins Howell), new combination (from Geodiaspis Tippins Howell); Duplachionaspis arecibo (Howell), new combination (this and the next 10 Duplachionaspis MacGillivray species all from Haliaspis Takagi); Duplachionaspis asymmetrica Ferris, revived combination; Duplachionaspis distichlii (Ferris), revived combination; Duplachionaspis litoralis Ferris, revived combination; Duplachionaspis mackenziei McDaniel, revived combination; Duplachionaspis milleri (Howell), new combination; Duplachionaspis nakaharai (Howell), new combination; Duplachionaspis peninsularis (Howell), new combination; Duplachionaspis spartinae (Comstock), revived combination; Duplachionaspis texana (Liu Howell) new combination; Duplachionaspis uniolae (Takagi), new combination; Duplachionaspis mutica (Williams) (from Aloaspis Williams), new combination; Epidiaspis doumtsopi (Schneider), new combination (from Diaspis Costa); Fiorinia ficicola (Takahashi), new combination (from Ichthyaspis Takagi); Fiorinia macroprocta (Leonardi), revived combination (this and the next 2 species of Fiorinia Targioni Tozzetti all from Trullifiorinia Leonardi); Fiorinia rubrolineata Leonardi, revived combination; Fiorinia scrobicularum Green, revived combination; Genaparlatoria pseudaspidiotus (Lindinger), revived combination (from Parlatoria); Greeniella acaciae (Froggatt), new combination (this and the next 4 Greeniella Cockerell species all from Gymnaspis Newstead); Greeniella cassida (Hall Williams), new combination; Greeniella grandis (Green), new combination; Greeniella perpusilla (Maskell), new combination; Greeniella serrata (Froggatt), new combination; Hendersonaspis anomala (Green), new combination (from Ancepaspis); Hulaspis bulba (Munting), new combination (this and the next Hulaspis Hall species both from Andaspis MacGillivray); Hulaspis formicarum (Ben-Dov), new combination; Lepidosaphes antidesmae (Rao in Rao Ferris), new combination (this and the next 19 species all from Andaspis); Lepidosaphes arcana (Matile-Ferrero), new combination; Lepidosaphes betulae (Borchsenius), new combination; Lepidosaphes citricola (Young Hu), new combination; Lepidosaphes conocarpi (Takagi), new combination; Lepidosaphes crawi (Cockerell), revived combination; Lepidosaphes erythrinae Rutherford, revived combination; Lepidosaphes incisor Green, revived combination; Lepidosaphes indica (Borchsenius), new combination; Lepidosaphes kashicola Takahashi, revived combination; Lepidosaphes kazimiae (Williams), new combination; Lepidosaphes laurentina (Almeida), new combination; Lepidosaphes maai (Williams Watson), new combination; Lepidosaphes mackieana McKenzie, revived combination; Lepidosaphes micropori (Borchsenius), new combination; Lepidosaphes punicae Laing, revived combination; Lepidosaphes quercicola (Borchsenius), new combination; Lepidosaphes recurrens (Takagi Kawai), new combination; Lepidosaphes viticis (Takagi), new combination; Lepidosaphes xishuanbannae (Young Hu), new combination; Lepidosaphes giffardi (Adachi Fullaway), new combination (from Carulaspis MacGillivray); Lepidosaphes garciniae (Young Hu), new combination (this and the next 2 species all from Ductofrontaspis Young Hu); Lepidosaphes huangyangensis (Young Hu), new combination; Lepidosaphes jingdongensis (Young Hu), new combination; Lepidosaphes recurvata (Froggatt), revived combination (from Metandaspis Williams); Lepidosaphes ficicola Takahashi, revived combination (this and the next 2 species all from Ungulaspis MacGillivray); Lepidosaphes pinicolous Chen, revived combination; Lepidosaphes ungulata Green, revived combination; Lepidosaphes serrulata (Ganguli), new combination (from Velataspis Ferris); Lepidosaphes huyoung Normark, replacement name for Andaspis ficicola Young Hu; Lepidosaphes tangi Normark, replacement name for Andaspis schimae Tang; Lepidosaphes yuanfeng Normark, replacement name for Andaspis keteleeriae Yuan Feng; Leucaspis ilicitana (Gómez-Menor), new combination (from Aonidia); Lopholeucaspis spinomarginata (Green), new combination (from Gymnaspis); Melanaspis campylanthi (Lindinger), new combination (from Aonidia); Mohelnaspis bidens (Green), new combination (from Fiorinia); Parlatoria affinis (Ramakrishna Ayyar), new combination (this and the next 4 Parlatoria species all from Gymnaspis); Parlatoria ficus (Ramakrishna Ayyar), new combination; Parlatoria mangiferae (Ramakrishna Ayyar), new combination; Parlatoria ramakrishnai (Green), new combination; Parlatoria sclerosa (Munting), new combination; Parlatoria bullata (Green), new combination (from Bigymnaspis); Parlatoria leucaspis (Lindinger), new combination (this and the next species both from Cryptoparlatorea Lindinger); Parlatoria pini (Takahashi), new combination; Parlatoria tangi Normark, replacement name for Parlatoria pini Tang; Pseudoparlatoria bennetti (Williams), new combination (from Parlagena McKenzie); Pseudoparlatoria chinchonae (McKenzie), new combination (from Protodiaspis Cockerell); Pseudoparlatoria larreae (Leonardi), revived combination (from Protargionia Leonardi); Quernaspis lepineyi (Balachowsky), new combination (from Chionaspis); Rhizaspidiotus nullispinus (Munting), new combination (from Aonidia); Rolaspis marginalis (Leonardi), new combination (from Lepidosaphes); Salicicola lepelleyi (De Lotto), new combination (from Anotaspis Ferris); Tecaspis giffardi (Leonardi), new combination (from Dinaspis); Trullifiorinia geijeriae (Froggatt), new combination (from Fiorinia); Trullifiorinia nigra (Lindinger), new combination (from Crypthemichionaspis Lindinger); and Voraspis olivina (Leonardi), new combination (from Lepidosaphes).

A key to the flat grass scale genus Nipponaclerda (Hemiptera, Coccomorpha, Aclerdidae).

The flat grass scale genus Nipponaclerda comprises four species, native to Central and East Asia. Nipponaclerdabiwakoensis has been introduced to the United States and is considered a serious pest of Phragmitesaustralis, the common reed. Heavy infestations of N.biwakoensis in coastal marshes of Louisiana have coincided with extensive die-off of reeds. In this article, dichotomous identification keys to the genera of Aclerdidae and to the species of Nipponaclerda are provided, allowing for accurate identification of species found in the native and invasive range.

An online interactive identification key to common pest species of Aspidiotini (Hemiptera, Coccomorpha, Diaspididae), version 1.0.

Aspidiotini is a species-rich tribe of armored scale insects that includes several polyphagous and specialist pests that are commonly encountered at ports-of-entry to the United States and many other countries. This article describes a newly available online interactive tool that can be used to identify 155 species of Aspidiotini that are recognized as minor to major pests or that are potentially emergent pests. This article lists the species and features included with a description of the development and structure of the key. The interactive key is free to access at https://idtools.org/id/scales/aspidiotini/about_index.php.

Molecular phylogenetics of Aspidiotini armored scale insects (Hemiptera: Diaspididae) reveals rampant paraphyly, curious species radiations, and multiple origins of association with Melissotarsus ants (Hymenoptera: Formicidae).

The armored scale insect tribe Aspidiotini comprises many pest species that are globally invasive and economically damaging. The taxonomy of scale insects is based almost solely upon morphological characters of adult females, and little prior work has been done to test the classification of aspidiotines against molecular evidence. To address these concerns, we reconstruct a molecular phylogeny for aspidiotine armored scales that expands greatly upon taxonomic and character representations from previous studies. Our dataset includes 127 species (356 terminal taxa) and four gene regions: 28S, EF-1α, COI-COII, and CAD. Nearly 50% of the species treated are identified as pests and several more may represent emerging pests. Phylogenetic data were analyzed in a Bayesian framework using MC3 iterations. The majority of sampled aspidiotine genera are not monophyletic as currently defined. Monophyly constraints for 'worst offenders' were imposed on the phylogeny and stepping-stone MCMC was performed to calculate marginal likelihood scores. Comparisons of marginal likelihoods from runs with constrained vs. informative priors support the interpretation that pest-rich genera are not monophyletic. We use character mapping to illustrate signal and convergence for selected traits that have been used to define or recognize genera and evaluate consistency and retention indices for these traits. The phylogeny illustrates a pervasive pattern in which extremely polyphagous pests - typically having large populations and wide geographical distributions - are frequently intertwined with range-limited specialists on the phylogeny. Finally, the phylogeny recovers three origins of ant association among the Aspidiotini. The history of ant/diaspidid symbioses involves periods of sustained partner fidelity, spanning multiple speciation events, which have been punctuated by opportunistic switches to novel partners.