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Journal: ZooKeys


The geographic distribution of coyotes (Canis latrans) has dramatically expanded since 1900, spreading across much of North America in a period when most other mammal species have been declining. Although this considerable expansion has been well documented at the state/provincial scale, continent-wide descriptions of coyote spread have portrayed conflicting distributions for coyotes prior to the 1900s, with popularly referenced anecdotal accounts showing them restricted to the great plains, and more obscure, but data-rich accounts suggesting they ranged across the arid west. To provide a scientifically credible map of the coyote’s historical range (10,000-300 BP) and describe their range expansion from 1900 to 2016, we synthesized archaeological and fossil records, museum specimens, peer-reviewed reports, and records from wildlife management agencies. Museum specimens confirm that coyotes have been present in the arid west and California throughout the Holocene, well before European colonization. Their range in the late 1800s was undistinguishable from earlier periods, and matched the distribution of non-forest habitat in the region. Coyote expansion began around 1900 as they moved north into taiga forests, east into deciduous forests, west into costal temperate rain forests, and south into tropical rainforests. Forest fragmentation and the extirpation of larger predators probably enabled these expansions. In addition, hybridization with wolves (C. lupus, C. lycaon, and/or C. rufus) and/or domestic dogs has been documented in the east, and suspected in the south. Our detailed account of the original range of coyotes and their subsequent expansion provides the core description of a large scale ecological experiment that can help us better understand the predator-prey interactions, as well as evolution through hybridization.


Existing models for assigning species, subspecies, or no taxonomic rank to populations which are geographically separated from one another were analyzed. This was done by subjecting over 3,000 pairwise comparisons of vocal or biometric data based on birds to a variety of statistical tests that have been proposed as measures of differentiation. One current model which aims to test diagnosability (Isler et al. 1998) is highly conservative, applying a hard cut-off, which excludes from consideration differentiation below diagnosis. It also includes non-overlap as a requirement, a measure which penalizes increases to sample size. The “species scoring” model of Tobias et al. (2010) involves less drastic cut-offs, but unlike Isler et al. (1998), does not control adequately for sample size and attributes scores in many cases to differentiation which is not statistically significant. Four different models of assessing effect sizes were analyzed: using both pooled and unpooled standard deviations and controlling for sample size using t-distributions or omitting to do so. Pooled standard deviations produced more conservative effect sizes when uncontrolled for sample size but less conservative effect sizes when so controlled. Pooled models require assumptions to be made that are typically elusive or unsupported for taxonomic studies. Modifications to improving these frameworks are proposed, including: (i) introducing statistical significance as a gateway to attributing any weighting to findings of differentiation; (ii) abandoning non-overlap as a test; (iii) recalibrating Tobias et al. (2010) scores based on effect sizes controlled for sample size using t-distributions. A new universal method is proposed for measuring differentiation in taxonomy using continuous variables and a formula is proposed for ranking allopatric populations. This is based first on calculating effect sizes using unpooled standard deviations, controlled for sample size using t-distributions, for a series of different variables. All non-significant results are excluded by scoring them as zero. Distance between any two populations is calculated using Euclidian summation of non-zeroed effect size scores. If the score of an allopatric pair exceeds that of a related sympatric pair, then the allopatric population can be ranked as species and, if not, then at most subspecies rank should be assigned. A spreadsheet has been programmed and is being made available which allows this and other tests of differentiation and rank studied in this paper to be rapidly analyzed.


The root-knot nematode Meloidogyne ulmi is synonymised with Meloidogyne mali based on morphological and morphometric similarities, common hosts, as well as biochemical similarities at both protein and DNA levels. M. mali was first described in Japan on Malus prunifolia Borkh.; and M. ulmi in Italy on Ulmus chenmoui W.C. Cheng. Morphological and morphometric studies of their holo- and paratypes revealed important similarities in the major characters as well as some general variability in a few others. Host test also showed that besides the two species being able to parasitize the type hosts of the other, they share some other common hosts. Our study of the esterase and malate dehydrogenase isozyme phenotypes of some M. ulmi populations gave a perfectly comparable result to that already known for M. mali. Finally, phylogenetic studies of their SSU and LSU rDNA sequence data revealed that the two are not distinguishable at DNA level. All these put together, leave strong evidences to support the fact that M. ulmi is not a valid species, but a junior synonym of M. mali. Brief discussion on the biology and life cycle of M. mali is given. An overview of all known hosts and the possible distribution of M. mali in Europe are also presented.

Concepts: DNA, Gene, Parasites, Evolution, Biology, Nematode, Nematodes, Root-knot nematode


Ptilohyale explorator (formerly Parhyale explorator), described by Arresti (1989), can be considered to be a synonym of west-Atlantic Ptilohyale littoralis (Stimpson, 1853), based on morphological observations of paratypes and specimens recently collected in the type locality of Ptilohyale explorator. The first collections of Ptilohyale littoralis, from the eastern Atlantic were from the port of Rotterdam (The Netherlands) in 2009 and later in Wimereux, Opal Coast (France) in 2014; however, the synonymy of Ptilohyale explorator with Ptilohyale littoralis backdates to the first European record of Ptilohyale littoralis in 1985 at La Vigne, Bay of Arcachon (France). This indicates that Ptilohyale littoralis has been established along European Atlantic coast for many years. An assessment of the nominal valid species belonging to the genus Ptilohyale was carried out and a comparison between the Atlantic Ptilohyale littoralis and the very similar Mediterranean hyalid species, Parhyale plumicornis, is presented based on morphological features and distribution. Due to the invasive ability of Ptilohyale littoralis, a comparison between the two species is necessary.


A total of ca 800,000 occurrence records from the Australian Museum (AM), Museums Victoria (MV) and the New Zealand Arthropod Collection (NZAC) were audited for changes in selected Darwin Core fields after processing by the Atlas of Living Australia (ALA; for AM and MV records) and the Global Biodiversity Information Facility (GBIF; for AM, MV and NZAC records). Formal taxon names in the genus- and species-groups were changed in 13-21% of AM and MV records, depending on dataset and aggregator. There was little agreement between the two aggregators on processed names, with names changed in two to three times as many records by one aggregator alone compared to records with names changed by both aggregators. The type status of specimen records did not change with name changes, resulting in confusion as to the name with which a type was associated. Data losses of up to 100% were found after processing in some fields, apparently due to programming errors. The taxonomic usefulness of occurrence records could be improved if aggregators included both original and the processed taxonomic data items for each record. It is recommended that end-users check original and processed records for data loss and name replacements after processing by aggregators.


An improved and expanded nomenclature for genetic sequences is introduced that corresponds with a ranking of the reliability of the taxonomic identification of the source specimens. This nomenclature is an advancement of the “Genetypes” naming system, which some have been reluctant to adopt because of the use of the “type” suffix in the terminology. In the new nomenclature, genetic sequences are labeled “genseq,” followed by a reliability ranking (e.g., 1 if the sequence is from a primary type), followed by the name of the genes from which the sequences were derived (e.g., genseq-1 16S, COI). The numbered suffix provides an indication of the likely reliability of taxonomic identification of the voucher. Included in this ranking system, in descending order of taxonomic reliability, are the following: sequences from primary types - “genseq-1,” secondary types - “genseq-2,” collection-vouchered topotypes - “genseq-3,” collection-vouchered non-types - “genseq-4,” and non-types that lack specimen vouchers but have photo vouchers - “genseq-5.” To demonstrate use of the new nomenclature, we review recently published new-species descriptions in the ichthyological literature that include DNA data and apply the GenSeq nomenclature to sequences referenced in those publications. We encourage authors to adopt the GenSeq nomenclature (note capital “G” and “S” when referring to the nomenclatural program) to provide a searchable tag (e.g., “genseq”; note lowercase “g” and “s” when referring to sequences) for genetic sequences from types and other vouchered specimens. Use of the new nomenclature and ranking system will improve integration of molecular phylogenetics and biological taxonomy and enhance the ability of researchers to assess the reliability of sequence data. We further encourage authors to update sequence information on databases such as GenBank whenever nomenclatural changes are made.

Concepts: DNA, Genetics, Biology, Taxonomy, Noun, Biological classification, Name, Nomenclature


The Bornean hydrocenids have so far been understudied compared to other non-pulmonate snails in this region. In the present study, we review a first group of minute land snail species belonging to the genus Georissa (Gastropoda, Hydrocenidae) from Malaysian Borneo. This group is restricted to the species with conspicuous scale-like sculpture on the shell. Based on materials from recent fieldwork, museums, and personal collections, Malaysian Borneo hydrocenids are more complex and diverse in shell characters than previously anticipated. Here, a molecular, conchological, and biogeographic study of this “scaly group” is presented. We recognise 13 species of which six are new to science, namely Georissa anyiensissp. n., Georissa muluensissp. n., Georissa bauensissp. n., Georissa silaburensissp. n., Georissa kinabatanganensissp. n., and Georissa sepulutensissp. n.


We present a pinned insect manipulator (IMp) constructed of LEGO® building bricks with two axes of movement and two axes of rotation. In addition we present three variants of the IMp to emphasise the modular design, which facilitates resizing to meet the full range of pinned insect specimens, is fully customizable, collapsible, affordable and does not require specialist tools or knowledge to assemble.

Concepts: Design, Rotation, IMP


The aganippine shield-backed trapdoor spiders of the monophyletic nigrum-group of Idiosoma Ausserer s. l. are revised, and 15 new species are described from Western Australia and the Eyre Peninsula of South Australia: I. arenaceum Rix & Harvey, sp. n., I. corrugatum Rix & Harvey, sp. n., I. clypeatum Rix & Harvey, sp. n., I. dandaragan Rix & Harvey, sp. n., I. formosum Rix & Harvey, sp. n., I. gardneri Rix & Harvey, sp. n., I. gutharuka Rix & Harvey, sp. n., I. incomptum Rix & Harvey, sp. n., I. intermedium Rix & Harvey, sp. n., I. jarrah Rix & Harvey, sp. n., I. kopejtkaorum Rix & Harvey, sp. n., I. kwongan Rix & Harvey, sp. n., I. mcclementsorum Rix & Harvey, sp. n., I. mcnamarai Rix & Harvey, sp. n., and I. schoknechtorum Rix & Harvey, sp. n. Two previously described species from south-western Western Australia, I. nigrum Main, 1952 and I. sigillatum (O. P.-Cambridge, 1870), are re-illustrated and re-diagnosed, and complementary molecular data for 14 species and seven genes are analysed with Bayesian methods. Members of the nigrum-group are of long-standing conservation significance, and I. nigrum is the only spider in Australia to be afforded threatened species status under both State and Commonwealth legislation. Two other species, I. formosum Rix & Harvey, sp. n. and I. kopejtkaorum Rix & Harvey, sp. n., are also formally listed as Endangered under Western Australian State legislation. Here we significantly relimit I. nigrum to include only those populations from the central and central-western Wheatbelt bioregion, and further document the known diversity and conservation status of all known species.


Fossil elytra of a small trechine carabid are reported from the Oliver Bluffs on the Beardmore Glacier at lat. 85°S. They were compared with counterparts from the extant genera Trechisibus, Tasmanorites, Oxytrechus and Pseudocnides. The fossils share some characters but are sufficiently different to be described as a new genus and species. We named the new species Antarctotrechus balli in honour of George E. Ball who made major contributions to the study of carabids through his own research and the training of students while at the University of Alberta, Edmonton, Alberta, Canada. The closest extant relatives to the extinct Antarctotrechus balli are species of Trechisibus, which inhabit South America, the Falkland Islands and South Georgia, and Tasmanorites, which inhabit Tasmania, Australia. Plant fossils associated with Antarctotrechus balli included Nothofagus (southern beech), Ranunculus (buttercup), moss mats and cushion plants that were part of a tundra biome. Collectively, the stratigraphic relationships and the growth characteristics of the fossil plants indicate that Antarctotrechus balli inhabited the sparsely-vegetated banks of a stream that was part of an outwash plain at the head of a fjord in the Transantarctic Mountains. Other insects represented by fossils in the tundra biome include a listroderine weevil and a cyclorrhaphan fly. The age of the fossils, based on comparison of associated pollen with (40)Ar/(39)Ar dated pollen assemblages from the McMurdo Dry Valleys, is probably Early to Mid-Miocene in the range 14-20 Ma. The tundra biome, including Antarctotrechus balli, became extinct in the interior of Antarctica about 14 Ma and on the margins of the continent by 10-13 Ma. Antarctotrechus balli confirms that trechines were once widely distributed in Gondwana. For Antarctotrechus balli and other elements of the tundra biome it appears they continued to inhabit a warmer Antarctica for many millions of years after rifting of Tasmania (45 Ma) and southern South America (31 Ma).

Concepts: Plant, Beetle, South America, Antarctica, Ground beetle, Victoria Land, Transantarctic Mountains, Beardmore Glacier