Browsing by Author "Means, Jackson C."

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    Apheloria polychroma, a new species of millipede from the Cumberland Mountains (Polydesmida: Xystodesmidae)
    Marek, Paul E.; Means, Jackson C.; Hennen, Derek A. (Zootaxa, 2018-01-25)
    Millipedes of the genus Apheloria Chamberlin, 1921 occur in temperate broadleaf forests throughout eastern North America and west of the Mississippi River in the Ozark and Ouachita Mountains. Chemically defended with toxins made up of cyanide and benzaldehyde, the genus is part of a community of xystodesmid millipedes that compose several Müllerian mimicry rings in the Appalachian Mountains. We describe a model species of these mimicry rings, Apheloria polychroma n. sp., one of the most variable in coloration of all species of Diplopoda with more than six color morphs, each associated with a separate mimicry ring.
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    Biodiversity, conservation and mimicry rings of Appalachian millipedes
    Means, Jackson C. (Virginia Tech, 2019-01-18)
    The Appalachian Mountains contain some of the highest millipede diversity on the planet, including many endemic and range-restricted species. Millipedes have extremely low dispersal capacity, strict environmental requirements for survival and, in the case of the family Xystodesmidae, are completely blind. Coupled with the complex topology of the Appalachians, xystodesmid millipedes have speciated in the eastern United States to a greater degree than in any other region. Many of the taxonomic relationships within the Xystodesmidae remain to be tested using modern molecular techniques, and numerous species await description. The subtribe Nannariina, for example, contains 3 genera and 19 species, yet evidence suggests the tribe may actually contain over 100 species. Traditional taxonomic work has been hindered by a paucity of researchers in the field, and a necessary reliance on morphological characters for species delimitation. The external male genitalia, known as gonopods, are the structures which have most often been used to define species boundaries. However, in many groups, including the Xystodesmidae, the gonopods are extremely simple, making it difficult to discern whether similarity between gonopods is a result of shared ancestry or morphological convergence. Further complicating matters, somatic characters are often uniform between species, and some obvious phenotypes, such as color pattern and hue, are shared between genera in aposematic mimicry rings. Leading to the formation of mimicry rings between distantly related genera, their bright coloration functions to advertise a cyanide-based defense. Some species have been recorded as having six color patterns, each a member of a different mimicry rings. The factors which drive some taxa to participate in mimicry rings, while others have a myriad of hues and patterns, unmatched by other species, are unknown. Here I investigate the phylogenetic relationships between the tribes and genera of the Xystodesmidae, with the aims of (1) understanding the role of homoplasy within the Xystodesmidae, (2) delineating tribal and generic boundaries within the family, (3) describing species in the genus Nannaria for the purposes of revising the subtribe Nannariina, and (4) investigating mimicry and color variability through a detailed color atlas for North American Xystodesmidae. I found that 95% of the morphological characters traditionally used in xystodesmid taxonomy are homoplasious (HI > 0.5), delineated the Apheloriini, Appalachioria and Sigmoria, described 34 new Nannaria species, and created a color and mimicry dataset of 956 Xystodesmidae for use in future studies of color variability.
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    Development of a Precision Mite Management Program for the Control of the Ectoparasite Varroa destructor in Hives of Apis mellifera L.
    Means, Jackson C. (Virginia Tech, 2014-06-03)
    The European honey bee, Apis mellifera, is an important pollinator of horticultural and agricultural field crops, providing ≈ 90% of all commercial pollination services (Genersch et al. 2010). The recent rise in colony loss due to Colony Collapse Disorder (CCD) has been a source of concern for both beekeepers and the apiculture industry. One of the factors implicated in CCD is infestation by the ectoparasitic mite, Varroa destructor. Initial efforts to control the mite relied heavily on regular application of miticides without regard to actual mite infestation levels. This approach has led to problems of resistance in the mite and contamination of the hive and hive-products. Because it is unlikely that miticides will be removed as an option for mite management, a precision mite management (PMM) approach using information on the spatiotemporal distribution of the mite to improve sampling and treatments is seen as a viable option, particularly with respect to treatment costs and impacts on the environment. The primary objective of this study was to develop an understanding of the spatiotemporal distribution of the Varroa mite and bee brood within hives for the purpose of developing a PMM approach for the mite. Varroa mite populations were sampled from May to June, 2012 and February to October, 2013. Sampling was conducted with three commonly used sampling methods: soapy water roll (SWR), brood uncapping, and a modified sticky board; brood uncapping, however, was discontinued during the study due to hive the labor cost and harmful effects of this method to the hives. Similar trends in mite population levels were observed using the soapy water roll and sticky board sampling methods. Spearman's nonparametric analysis showed that there was a significant correlation (ρ = 0.47, P<0.001) in mite population levels for the soapy water roll and sticky board methods for sampling conducted from February to September, 2013 (the SWR method was not used in October). This was despite the fact that there was no significant correlation (ρ= -0.03, P = 0.8548) between the two sampling methods during the spring sampling period from February to April, 2013. The observed lack of correlation between the two sampling methods in early spring was likely due to the low population of brood in the hive, which caused the majority of the mites to remain on adult bees. Mites per 100 adult bees, therefore, appear to reflect mite population levels within the hive more closely than mite fall on sticky borad during the February to April sampling period. This suggests that the soapy water roll method is a better method for estimating mite population levels within the hive in the early spring compared with the sticky board method. Geospatial analyses of the distributions of mite fall on the sticky boards were conducted using geostatistics and Spatial Analysis by Distance IndicEs (SADIE). Both analyses showed that mite fall on the sticky board was generally aggregated and the aggregation increased with mite population levels. The average range of the variogram from geostatistical analysis was estimated at 4 sticky board cells; this range value was increased to 5 cells and was used to develop a systematic outside-range sampling protocol for mites on a sticky board. The results showed that the accuracy of the systematic outside-range sampling compared well with that of the traditional sticky board counting method in estimating total mite fall, but required only 60% of the effort (i.e., counting 63 instead of 105 cells). SADIE analysis showed that there is an overall association between the distribution of mite fall on a sticky board and the distribution of brood within a hive. A greater degree of correspondence was also observed in the association of drone and mite distributions during May to June; greater correspondence in worker brood and mite associations was observed in August and September. These differences may be due to relative amounts of the two types of brood present within the hive. A test of the efficacy of precision application of Varroa mite treatment based on the association between drone brood and mite fall resulted in a significantly greater reduction in mite levels on the sticky board using a traditional miticide treatment method compared with the control and precision treatments (𝜒2 =362.571; df = 2; P <0.0001); mite population levels with the precision method, however, were significantly reduced compared with the control.
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    The first true millipede—1306 legs long
    Marek, Paul E.; Buzatto, Bruno A.; Shear, William A.; Means, Jackson C.; Black, Dennis G.; Harvey, Mark S.; Rodriguez, Juanita (Springer, 2021-12-01)
    The name “millipede” translates to a thousand feet (from mille “thousand” and pes “foot”). However, no millipede has ever been described with more than 750 legs. We discovered a new record-setting species of millipede with 1,306 legs, Eumillipes persephone, from Western Australia. This diminutive animal (0.95 mm wide, 95.7 mm long) has 330 segments, a cone-shaped head with enormous antennae, and a beak for feeding. A distant relative of the previous record holder, Illacme plenipes from California, it belongs to a different order, the Polyzoniida. Discovered 60 m below ground in a drill hole created for mineral exploration, E. persephone possesses troglomorphic features; it lacks eyes and pigmentation, and it has a greatly elongated body—features that stand in stark contrast to its closest surface-dwelling relatives in Australia and all other members of its order. Using phylogenomics, we found that super-elongation (> 180 segments) evolved repeatedly in the millipede class Diplopoda. The striking morphological similarity between E. persephone and I. plenipes is a result of convergent evolution, probably for locomotion in similar soil habitats. Discovered in the resource-rich Goldfields-Esperance region and threatened by encroaching surface mining, documentation of this species and conservation of its habitat are of critical importance.
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    Is geography an accurate predictor of evolutionary history in the millipede family Xystodesmidae?
    Means, Jackson C.; Marek, Paul E. (PeerJ, 2017-10-12)
    For the past several centuries, millipede taxonomists have used the morphology of male copulatory structures (modified legs called gonopods), which are strongly variable and suggestive of species-level differences, as a source to understand taxon relationships. Millipedes in the family Xystodesmidae are blind, dispersal-limited and have narrow habitat requirements. Therefore, geographical proximity may instead be a better predictor of evolutionary relationship than morphology, especially since gonopodal anatomy is extremely divergent and similarities may be masked by evolutionary convergence. Here we provide a phylogenetics-based test of the power of morphological versus geographical character sets for resolving phylogenetic relationships in xystodesmid millipedes. Molecular data from 90 species-group taxa in the family were included in a six-gene phylogenetic analysis to provide the basis for comparing trees generated from these alternative character sets. The molecular phylogeny was compared to topologies representing three hypotheses: (1) a prior classification formulated using morphological and geographical data, (2) hierarchical groupings derived from Euclidean geographical distance, and (3) one based solely on morphological data. Euclidean geographical distance was not found to be a better predictor of evolutionary relationship than the prior classification, the latter of which was the most similar to the molecular topology. However, all three of the alternative topologies were highly divergent (Bayes factor >10) from the molecular topology, with the tree inferred exclusively from morphology being the most divergent. The results of this analysis show that a high degree of morphological convergence from substantial gonopod shape divergence generated spurious phylogenetic relationships. These results indicate the impact that a high degree of morphological homoplasy may have had on prior treatments of the family. Using the results of our phylogenetic analysis, we make several changes to the classification of the family, including transferring the rare state-threatened species Sigmoria whiteheadi Shelley, 1986 to the genus Apheloria Chamberlin, 1921—a relationship not readily apparent based on morphology alone. We show that while gonopod differences are a premier source of taxonomic characters to diagnose species pairwise, the traits should be viewed critically as taxonomic features uniting higher levels.
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    Phylogenetic Systematics of the Millipede Family Xystodesmidae
    Means, Jackson C.; Hennen, Derek A.; Tanabe, Tsutomu; Marek, Paul E. (Oxford University Press, 2021-01-03)
    The millipede family Xystodesmidae includes 486 species distributed primarily in temperate deciduous forests in North America and East Asia. Species diversity of the family is greatest in the Appalachian Mountains of the eastern United States, with 188 species. Although the group includes notable taxa such as those that are bioluminescent and others that display Müllerian mimicry, producing up to 600 mg of cyanide, basic alpha-taxonomy of the group is woefully incomplete and more than 50 species remain undescribed in the Appalachian Mountains alone. In order to establish a robust phylogenetic foundation for addressing compelling evolutionary questions and describing species diversity, we assembled the largest species phylogeny (in terms of species sampling) to date in the Diplopoda. We sampled 49 genera (out of 57) and 247 of the species in the family Xystodesmidae, recollecting fresh material from historical type localities and discovering new species in unexplored regions. Here, we present a phylogeny of the family using six genes (four mitochondrial and two nuclear) and include pivotal taxa omitted from previous studies including Nannaria, Erdelyia, taxa from East Asia, and 10 new species. We show that 6 of the 11 tribes are monophyletic, and that the family is paraphyletic with respect to the Euryuridae and Eurymerodesmidae. Prior supraspecific classification is in part inconsistent with the phylogeny and convergent evolution has caused artificial genera to be proposed. Subspecific classification is likewise incongruent with phylogeny and subspecies are consistently not sister to conspecifics. The phylogeny is used as a basis to update the classification of the family, diagnose monophyletic groups, and to inform species hypotheses.
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    A revision of the wilsoni species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae)
    Hennen, Derek A.; Means, Jackson C.; Marek, Paul E. (Pensoft Publishers, 2022-04-15)
    Although many new species of the millipede genus Nannaria Chamberlin, 1918 have been known from museum collections for over half a century, a systematic revision has not been undertaken until recently. There are two species groups in the genus: the minor species group and the wilsoni species group. In this study, the wilsoni species group was investigated. Specimens were collected from throughout its distribution in the Appalachian Mountains of the eastern United States and used for a multi-gene molecular phylogeny. The phylogenetic tree recovered Nannaria and the two species groups as monophyletic, with Oenomaea pulchella as its sister group. Seventeen new species were described, bringing the composition of the wilsoni species group to 24 species, more than tripling its known diversity, and increasing the total number of described Nannaria species to 78. The genus now has the greatest number of species in the family Xystodesmidae. Museum holdings of Nannaria were catalogued, and a total of 1,835 records used to produce a distribution map of the species group. Live photographs, illustrations of diagnostic characters, ecological notes, and conservation statuses are given. The wilsoni species group is restricted to the Appalachian region, unlike the widely-distributed minor species group (known throughout eastern North America), and has a distinct gap in its distribution in northeastern Tennessee and adjacent northwestern North Carolina. The wilsoni species group seems to be adapted to mesic microhabitats in middle to high elevation forests in eastern North America. New species are expected to be discovered in the southern Appalachian Mountains.