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Journal: ZooKeys


A new species of Anolis lizard from the Andean slopes of southwestern Colombia and northwestern Ecuador, from between 1187 and 2353 m in elevation, is described. The new species can be distinguished from other Anolis in squamation, cranial osteology, hemipenial morphology, and nuclear and mitochondrial DNA. The new species is sister to Anolisaequatorialis, and it is suggested that previous records of A.aequatorialis in Colombia correspond to the new species described herein.


Exosphaeromaamplicauda (Stimpson, 1857) from the west coast of North America is reviewed and redescribed and revealed to be a group of closely related species. A neotype is designated and the species redescribed based on the neotype and topotypic specimens. Exosphaeromaamplicauda is known only from the coast of California, at Marin, Sonoma and San Mateo Counties. Exosphaeromaaphrodita (Boone, 1923), type locality La Jolla, California and previously considered nomen dubium is taken out of synonymy and re-validated. A further three species: Exosphaeromapaydenae sp. n., Exosphaeromarussellhansoni sp. n., and Exosphaeromapentcheffi sp. n. are described herein. Sphaeromaoctonctum Richardson, 1899 is placed into junior synonymy with Exosphaeromaamplicauda. A key to the Pacific West Coast Exosphaeroma is provided.

Concepts: United States, California, North America, Los Angeles, San Diego, Zoological nomenclature, San Francisco, Nomen dubium


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.


A comprehensive taxonomic study is presented for the four genera and 286 species of the doryctine tribe Heterospilini occurring in Costa Rica. The tribe is represented almost entirely by the 280 species of the genus Heterospilus Haliday. Keys for identification of the genera and species are provided and the genera and species are described and illustrated. An interactive key to the species of Heterospilus also was prepared using Lucid Builder. The following new genus and species are described from Costa Rica: Paraheterospilus gen. n., P. ceciliaensis sp. n., P. eumekus sp. n., P. wilbotgardus sp. n., Heterospilus achi sp. n., H. achterbergi sp. n., H. aesculapius sp. n., H. agujas sp. n., H. agujasensis sp. n., H. alajuelus sp. n., H. albocoxalis sp. n., H. alejandroi sp. n., H. amuzgo sp. n., H. angelicae sp. n., H. angustus sp. n., H. aphrodite sp. n., H. apollo sp. n., H. arawak sp. n., H. areolatus sp. n., H. artemis sp. n., H. athena sp. n., H. attraholucus sp. n., H. aubreyae sp. n., H. austini sp. n., H. azofeifai sp. n., H. bacchus sp. n., H. barbalhoae sp. n., H. bennetti sp. n., H. bicolor sp. n., H. boharti sp. n., H. borucas sp. n., H. braeti sp. n., H. brethesi sp. n., H. breviarius sp. n., H. brevicornus sp. n., H. bribri sp. n., H. brullei sp. n., H. bruesi sp. n., H. cabecares sp. n., H. cacaoensis sp. n., H. cachiensis sp. n., H. cameroni sp. n., H. cangrejaensis sp. n., H. careonotaulus sp. n., H. caritus sp. n., H. carolinae sp. n., H. cartagoensis sp. n., H. catiensis sp. n., H. catorce sp. n., H. cero sp. n., H. chaoi sp. n., H. chilamatensis sp. n., H. chocho sp. n., H. chorotegus sp. n., H. chorti sp. n., H. cinco sp. n., H. cocopa sp. n., H. colliletus sp. n., H. colonensis sp. n., H. complanatus sp. n., H. conservatus sp. n., H. cora sp. n., H. corcovado sp. n., H. corrugatus sp. n., H. costaricensis sp. n., H. cressoni sp. n., H. cuatro sp. n., H. curtisi sp. n., H. cushmani sp. n., H. dani sp. n., H. demeter sp. n., H. dianae sp. n., H. diecinueve sp. n., H. dieciocho sp. n., H. dieciseis sp. n., H. diecisiete sp. n., H. diez sp. n., H. doce sp. n., H. dos sp. n., H. dulcus sp. n., H. eberhardi sp. n., H. ektorincon sp. n., H. emilius sp. n., H. empalmensis sp. n., H. enderleini sp. n., H. escazuensis sp. n., H. fahringeri sp. n., H. fischeri sp. n., H. flavidus sp. n., H. flavisoma sp. n., H. flavostigmus sp. n., H. foersteri sp. n., H. fonsecai sp. n., H. fournieri sp. n., H. gahani sp. n., H. garifuna sp. n., H. gauldi sp. n., H. golfodulcensis sp. n., H. gouleti sp. n., H. granulatus sp. n., H. grisselli sp. n., H. guanacastensis sp. n., H. guapilensis sp. n., H. hachaensis sp. n., H. halidayi sp. n., H. hansoni sp. n., H. hansonorum sp. n., H. haplocarinus sp. n., H. hedqvisti sp. n., H. hera sp. n., H. heredius sp. n., H. hespenheidei sp. n., H. holleyae sp. n., H. huddlestoni sp. n., H. huetares sp. n., H. hypermekus sp. n., H. itza sp. n., H. ixcatec sp. n., H. ixil sp. n., H. jabillosensis sp. n., H. jakaltek sp. n., H. janzeni sp. n., H. jennieae sp. n., H. jonmarshi sp. n., H. jupiter sp. n., H. kellieae sp. n., H. kiefferi sp. n., H. kikapu sp. n., H. kulai sp. n., H. kuna sp. n., H. lapierrei sp. n., H. lasalturus sp. n., H. laselvus sp. n., H. leenderti sp. n., H. leioenopus sp. n., H. leiponotaulus sp. n., H. lenca sp. n., H. levis sp. n., H. leviscutum sp. n., H. levitergum sp. n., H. limonensis sp. n., H. longinoi sp. n., H. longisulcus sp. n., H. longius sp. n., H. luteogaster sp. n., H. luteoscutum sp. n., H. luteus sp. n., H. macrocarinus sp. n., H. macrocaudatus sp. n., H. magnus sp. n., H. malaisei sp. n., H. mam sp. n., H. maritzaensis sp. n., H. mars sp. n., H. masneri sp. n., H. masoni sp. n., H. mellosus sp. n., H. menkei sp. n., H. mercury sp. n., H. milleri sp. n., H. miskito sp. n., H. mixtec sp. n., H. monteverde sp. n., H. mopanmaya sp. n., H. muertensis sp. n., H. muesebecki sp. n., H. nahua sp. n., H. neesi sp. n., H. nemestrinus sp. n., H. nephilim sp. n., H. nephus sp. n., H. nigracapitus sp. n., H. nigragonatus sp. n., H. nigricoxus sp. n., H. nixoni sp. n., H. noyesi sp. n., H. nueve sp. n., H. nunesi sp. n., H. once sp. n., H. orbitus sp. n., H. orosi sp. n., H. paloverde sp. n., H. pappi sp. n., H. parkeri sp. n., H. parvus sp. n., H. pech sp. n., H. penosa sp. n., H. petiolatus sp. n., H. petralbus sp. n., H. phaeocoxus sp. n., H. phaeoskelus sp. n., H. pharkidodus sp. n., H. phytorius sp. n., H. pitillaensis sp. n., H. poqomchi sp. n., H. poqomom sp. n., H. puertoviejoensis sp. n., H. puntarensis sp. n., H. qanjobal sp. n., H. quickei sp. n., H. quitirrisi sp. n., H. racostica sp. n., H. rama sp. n., H. ramirezi sp. n., H. ratzeburgi sp. n., H. reagani sp. n., H. reinhardi sp. n., H. retheospilus sp. n., H. rhabdotus sp. n., H. ricacosta sp. n., H. rinconensis sp. n., H. robbieae sp. n., H. rohweri sp. n., H. rojasi sp. n., H. romani sp. n., H. rugosus sp. n., H. sabrinae sp. n., H. saminae sp. n., H. sanjosensis sp. n., H. santarosensis sp. n., H. sanvitoensis sp. n., H. saturn sp. n., H. seis sp. n., H. sergeyi sp. n., H. sharkeyi sp. n., H. shawi sp. n., H. shenefelti sp. n., H. shonan sp. n., H. siete sp. n., H. similis sp. n., H. sinuatus sp. n., H. smithi sp. n., H. spiloheterus sp. n., H. staryi sp. n., H. stelfoxi sp. n., H. strazanaci sp. n., H. sumo sp. n., H. szepligeti sp. n., H. terrabas sp. n., H. thereospilus sp. n., H. tobiasi sp. n., H. tolupan sp. n., H. townesi sp. n., H. trece sp. n., H. tres sp. n., H. tricolor sp. n., H. trienta sp. n., H. tuberculatus sp. n., H. turrialbaensis sp. n., H. tzutujil sp. n., H. ugaldei sp. n., H. uno sp. n., H. variabilis sp. n., H. veinte sp. n., H. veintidos sp. n., H. veintitres sp. n., H. veintiuno sp. n., H. vierecki sp. n., H. villegasi sp. n., H. vittatus sp. n., H. vulcanus sp. n., H. wahli sp. n., H. warreni sp. n., H. washingtoni sp. n., H. wesmaeli sp. n., H. whartoni sp. n., H. whitfieldi sp. n., H. wildi sp. n., H. wilkinsoni sp. n., H. wrightae sp. n., H. xanthus sp. n., H. xerxes sp. n., H. xinca sp. n., H. yaqui sp. n., H. ypsilon sp. n., H. zapotec sp. n., H. zeus sp. n., H. zitaniae sp. n., H. zoque sp. n., H. zunigai sp. n., H. zurquiensis sp. n. One new combination is proposed, Pioscelus costaricensis (Marsh) comb. n.

Concepts: Costa Rica, Athena


The genus Admetovis Grote is revised. Admetovisicarussp. n. is described from the mountains of western North America. A lectotype of Admetovisoxymorus Grote is designated. Illustrations of the adults, male and female genitalia, and distribution maps are presented, together with an identification key. The classification of the genus is reviewed resulting in its reassignment to the tribe Hadenini from Orthosiini.


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