Cuticular hydrocarbons corroborate the distinction between lowland and highland Natal fruit fly (Tephritidae,Ceratitis rosa) populations

The cuticular hydrocarbons (CHs) and morphology of two Ceratitis rosa Karsch (Diptera: Tephritidae) populations, putatively belonging to two cryptic taxa, were analysed. The chemical profiles were characterised by two-dimensional gas chromatography with mass spectrometric detection. CHs of Ceratitis rosa that originated from the lowlands and highlands of Kenya comprised of n-alkanes, monomethylalkanes, dimethylalkanes and unsaturated hydrocarbons in the range of the carbon backbone from C₁₄ to C₃₇. Hydrocarbons containing C₂₉, C₃₁, C₃₃ and C₃₅ carbon atoms predominated in these two populations. 2-Methyltriacontane was the predominant compound in both populations. Quantitative differences in the distribution of hydrocarbons of different chain lengths, mainly the C₂₂, C₃₂, C₃₃ and C₃₄ compounds of these two populations, were observed despite indistinct qualitative differences in these hydrocarbons. Morphological analyses of male legs confirmed that the flies belong to different morphotypes of Ceratitis rosa previously labelled as R1 and R2 for lowland and highland populations, respectively. A statistical analysis of the CH compositions of the putative R1 and R2 species showed distinct interspecific identities, with several CHs specific for each of the lowland and highland populations. This study supports a hypothesis that the taxon Ceratitis rosa consists of at least two biological species.     

An integrative approach to unravel the Ceratitis FAR (Diptera,Tephritidae) cryptic species complex: a review

This paper reviews all information gathered from different disciplines and studies to resolve the species status within the Ceratitis FAR (Ceratitis fasciventris, Ceratitis anonae, Ceratitis rosa) complex, a group of polyphagous fruit fly pest species (Diptera, Tephritidae) from Africa. It includes information on larval and adult morphology, wing morphometrics, cuticular hydrocarbons, pheromones, microsatellites, developmental physiology and geographic distribution. The general consensus is that the FAR complex comprises Ceratitis anonae, two species within Ceratitis rosa (so-called R1 and R2) and two putatitve species under Ceratitis fasciventris. The information regarding the latter is, however, too limited to draw final conclusions on specific status. Evidence for this recognition is discussed with reference to publications providing further details.     

Wing morphometrics as a possible tool for the diagnosis of the Ceratitis fasciventris,C. anonae,C. rosa complex (Diptera,Tephritidae)

Previous attempts to resolve the Ceratitis FAR complex (Ceratitis fasciventris, Ceratitis anonae, Ceratitis rosa, Diptera, Tephritidae) showed contrasting results and revealed the occurrence of five microsatellite genotypic clusters (A, F1, F2, R1, R2). In this paper we explore the potential of wing morphometrics for the diagnosis of FAR morphospecies and genotypic clusters. We considered a set of 227 specimens previously morphologically identified and genotyped at 16 microsatellite loci. Seventeen wing landmarks and 6 wing band areas were used for morphometric analyses. Permutational multivariate analysis of variance detected significant differences both across morphospecies and genotypic clusters (for both males and females). Unconstrained and constrained ordinations did not properly resolve groups corresponding to morphospecies or genotypic clusters. However, posterior group membership probabilities (PGMPs) of the Discriminant Analysis of Principal Components (DAPC) allowed the consistent identification of a relevant proportion of specimens (but with performances differing across morphospecies and genotypic clusters). This study suggests that wing morphometrics and PGMPs might represent a possible tool for the diagnosis of species within the FAR complex. Here, we propose a tentative diagnostic method and provide a first reference library of morphometric measures that might be used for the identification of additional and unidentified FAR specimens.     

Analyses of volatiles produced by the African fruit fly species complex (Diptera,Tephritidae)

Ceratitis fasciventris, Ceratitis anonae and Ceratitis rosa are polyphagous agricultural pests originating from the African continent. The taxonomy of this group (the so-called Ceratitis FAR complex) is unclear. To clarify the taxonomic relationships, male and female-produced volatiles presumably involved in pre-mating communication were studied using comprehensive two-dimensional gas chromatography with time-of-flight mass spectrometry (GC×GC-TOFMS) followed by multivariate analysis, and gas chromatography combined with electroantennographic detection (GC-EAD). GC×GC-TOFMS analyses revealed sex specific differences in produced volatiles. Male volatiles are complex mixtures that differ both qualitatively and quantitatively but share some common compounds. GC-EAD analyses of male volatiles revealed that the antennal sensitivities of females significantly differ in the studied species. No female volatiles elicited antennal responses in males. The results show clear species-specific differences in volatile production and provide complementary information for the distinct delimitation of the putative species by chemotaxonomic markers.     

Three new species of the genus Aporcelaimoides Heyns, 1965 from Vietnam (Nematoda,Dorylaimida, Aporcelaimidae), with an updated taxonomy of the genus

Three new species of Aporcelaimoides from natural habitats in Vietnam are studied, described and illustrated, including line drawings, LM and/or SEM pictures. Aporcelaimoides brevistylum sp. n. is characterized by its body 1.95–2.90 mm long, lip region offset by deep constriction and 17–18 µm broad, ventral side of mural odontostyle 11–14 µm long with aperture occupying 62–71% of its length, neck 663–767 µm long, pharyngeal expansion occupying 58–66% of total neck length, uterus a simple tube 85–182 µm long, pars refringens vaginae absent, V = 55–63, tail short and rounded (34–46 µm, c = 49–76, c’ = 0.6–0.8), spicules 67–86 µm long, and one ventromedian supplement out the range of spicules. Aporcelaimoides minor sp. n. is distinguished in having body 2.09–2.61 mm long, lip region offset by deep constriction and 19–20 µm broad, mural odontostyle 14–16 µm long at its ventral side with aperture occupying 73–84% of its length, neck 579–649 µm long, pharyngeal expansion occupying 57–66% of total neck length, uterus a simple tube 44–69 µm long, pars refringens vaginae well developed, V = 48–56, female tail very short, rounded conoid or truncate (14–26 µm, c = 90–146, c’ = 0.3–0.6), and male unknown. Aporcelaimoides silvaticum sp. n. is characterized by its body 2.09–2.60 mm long, lip region offset by depression and 17–18 µm broad, mural odontostyle 11–12 µm long at its ventral side with aperture occupying 60–66% of its length, neck 597–720 µm long, pharyngeal expansion occupying 58–64% of total neck length, uterus a simple tube 128–243 µm long, pars refringens vaginae well developed, V = 58–60, tail short and rounded (27–37 µm, c = 67–94, c’ = 0.6–0.7), spicules 64–75 µm long, and two or three widely spaced ventromedian supplements bearing hiatus. The genus Aporcelaimoides is restored, its diagnosis emended, and three species of Sectonema, namely Sectonema amazonicum, Sectonema haguei and Sectonema moderatum, transferred to it. An updated list of its species, a key to their identification and a tabular compendium with the most important morphometric features are also presented.     

Distribution of pelagic squids Abraliopsis Joubin, 1896 (Enoploteuthidae) and Pterygioteuthis P. Fischer, 1896 (Pyroteuthidae) (Cephalopoda,Decapodiformes, Oegopsida) in the Mexican Pacific

The oegopsid squids Abraliopsis and Pterygioteuthis are abundant and diverse genera with taxonomic and distributional problems. Identification and distribution of species in the Mexican Pacific has been somewhat controversial. Here are provided a large series of new records for Abraliopsis affinis, Abraliopsis falco, Pterygioteuthis gemmata, Pterygioteuthis giardi and Pterygioteuthis hoylei from the Gulf of California and off the SW coast of Mexico. All five species were collected in the central or the southern Gulf of California, or in both. Abraliopsis affinis was found in seven samples with a total of 48 specimens, from 21°59'' to 24°53''12"N. Abraliopsis falco was much less represented in the samples (14 specimens) but it was found in 10 localities, four of which correspond to the central-southern Gulf of California (north to 27°44''53"N) and six to SW Mexico (south to 16°49''18"N). In the case of Pterygioteuthis gemmata, only two records (three specimens) were obtained, both in the SW Gulf of California, while Pterygioteuthis giardi (nine specimens) records were all from the central Gulf of California (27°44''53” to 25°39''59"N). In the case of Pterygioteuthis hoylei (nine specimens), material was obtained in six localities, also in a restricted latitudinal range (24°23''48” to 25°56''56"N).     

The genus Pterostichus in China II: the subgenus Circinatus Sciaky,a species revision and phylogeny (Carabidae,Pterostichini)

All of the known species of the Chinese endemic subgenus Pterostichus (Circinatus) are revised, keyed, and illustrated. Eleven new species and one new subspecies are described: Pterostichus adelphus sp. n. (Sichuan: Meigu, N28.66°, E103.06°); Pterostichus ailaoicus sp. n. (Yunnan: Xinping, N23.94°, E101.50°); Pterostichus camelus sp. n. (Sichuan: Mianning, N28.97°, E102.16°); Pterostichus dimorphus sp. n. (Yunnan: Dayao, N26.08°, E101.03°); Pterostichus maitreya sp. n. (Guizhou: Fanjingshan, N27.90°, E108.70°); Pterostichus miao sp. n. (Guangxi: Maoershan, N25.87°, E110.41°); Pterostichus tumulus sp. n. (Guizhou: Fanjingshan, N27.90°, E108.70°); Pterostichus wangjiani sp. n. (Yunnan: Dongchuan, N26.08°, E102.87°); Pterostichus yan sp. n. (Hubei: Shennongjia, N31.47°, E110.39°); Pterostichus yuxiaodongi sp. n. (Sichuan: Wolong, N30.99°, E103.15°); Pterostichus zhygealu sp. n. (Sichuan: Meigu, N28.67°, E103.05°); and Pterostichus cavazzutianus mianningensis subsp. n. (Sichuan: Mianning, N28.97°, E102.16°). Pterostichus cavazzutianus is proposed as a replacement name for Pterostichus cavazuttii Allegro and Sciaky 2010, preoccupied by Pterostichus (Sinosteropus) barbarae cavazuttii Sciaky and Facchini 2003. A lectotype is designated for Pterostichus baenningeri Jedlička 1931. Two species, Pterostichus schuelkei Sciaky & Wrase and Pterostichus wenxianensis Allegro & Sciaky, are moved from the subgenus Circinatus to Morphohaptoderus. An infra-subgeneric taxonomy is proposed for the subgenus Circinatus with four species groups. The male endophallus characters for most species of Circinatus were well studied, with three types of endophallus defined. A phylogenetic analysis based on adult morphological characters confirmed the infra-subgeneric classification and clarified some of the relationships among species. Two main lineages within Circinatus were identified from the phylogenetic analyses. Three of the four species groups were monophyletic, whereas the fourth group was paraphyletic.     

Does temperature and oxygen affect duration of intramarsupial development and juvenile growth in the terrestrial isopod Porcellio scaber (Crustacea,Malacostraca)?

According to the temperature-size rule (TSR), ectotherms developing under cold conditions experience slower growth as juveniles but reach a larger size at maturity. Whether temperature alone causes this phenomenon is unknown, but oxygen limitation can play a role in the temperature-size relationship. Oxygen may become limited under warm conditions when the resulting higher metabolism creates a greater demand for oxygen, especially in larger individuals. We examined the independent effects of oxygen concentration (10% and 22% O₂) and temperature (15 °C and 22 °C) on duration of ontogenic development, which takes place within the maternal brood pouch (marsupium), and juvenile growth in the terrestrial isopod common rough woodlouse (Porcellio scaber). Individuals inside the marsupium undergo the change from the aqueous to the gaseous environment. Under hypoxia, woodlice hatched from the marsupium sooner, but their subsequent growth was not affected by the level of oxygen. Marsupial development and juvenile growth were almost three times slower at low temperature, and marsupial development was longer in larger females but only in the cold treatment. These results show that temperature and oxygen are important ecological factors affecting developmental time and that the strength of the effect likely depends on the availability of oxygen in the environment.     

Checklist of the helminth parasites of the genus Profundulus Hubbs, 1924 (Cyprinodontiformes,Profundulidae), an endemic family of freshwater fishes in Middle-America

From December 2012 to November 2014, 267 fish belonging to the family Profundulidae (representing nine of the 11 species of the genus Profundulus) were collected in 26 localities of Middle-America, across southern Mexico, Guatemala, and Honduras, comprising the distribution range of the genus, and analyzed for helminth parasites. Additionally, a database with all ten available published accounts of the helminth parasite fauna of this genus (the only genus within the family) was assembled. Based on both sources of information, a checklist containing all the records was compiled as a tool to address future questions in the areas of evolutionary biology, biogeography, ecology and phylogeography of this host-parasite association. The helminth parasite fauna of this fish group consists of 20 nominal species, classified in 17 genera and 14 families. It includes six species of adult digeneans, five metacercariae, two monogeneans, one adult cestode, three adult nematodes and three larval nematodes. The profundulid fishes are parasitized by a specialized group of helminth species (e.g. Paracreptotrema blancoi sensu Salgado-Maldonado et al. (2011b), Paracreptotrema profundulusi Salgado-Maldonado, Caspeta-Mandujano & Martínez Ramírez, 2011, Phyllodistomum spinopapillatum Pérez-Ponce de León, Pinacho-Pinacho, Mendoza-Garfias & García-Varela, 2015, Spinitectus humbertoi Mandujano-Caspeta & Moravec, 2000, Spinitectus mariaisabelae Caspeta-Mandujano Cabañas-Carranza & Salgado-Maldonado, 2007 and Rhabdochona salgadoi Mandujano-Caspeta & Moravec, 2000), representing the core helminth fauna that are not shared with other Middle-American fish species.     

First record of harpacticoid copepods from Lake Tahoe,United States: two new species of Attheyella (Harpacticoida,Canthocamptidae)

Benthic harpacticoids were collected for the first time at Lake Tahoe, California-Nevada, United States. Two species were identified as members of the genus Attheyella Brady, 1880. The genus Attheyella comprises about 150 species within six subgenera, but only twelve species have previously been reported from North American freshwater habitats. The two new species of Attheyella described here have a 3-segmented endopod on P1 and 2-segmented P2–P4 endopods, the distal segment of exopod of P2–P4 has three outer spines, and the P5 has five setae on the exopod and six setae on the baseoendopod. Attheyella (Attheyella) tahoensis sp. n. most closely resembles Attheyella (Attheyella) idahoensis (Marsh, 1903) from Idaho, Montana, and Alaska (United States) and Attheyella (Attheyella) namkungi Kim, Soh & Lee, 2005 from Gosu Cave in South Korea. They differ mainly by the number of setae on the distal endopodal segment of P2–P4. In addition, intraspecific variation has been observed on the caudal rami. Attheyella (Neomrazekiella) tessiae sp. n. is characterized by the extension of P5 baseoendopod, 2-segmented endopod of female P2–P3, and naked third seta of male P5 exopod. The two new species are likely endemic to Lake Tahoe, an isolated alpine lake within the Great Basin watershed in the western United States.     

Population structure and cryptic genetic variation in the mango fruit fly,Ceratitis cosyra (Diptera,Tephritidae)

The fruit fly Ceratitis cosyra is an important agricultural pest negatively affecting the mango crop production throughout Africa and also feeding on a variety of other wild and cultivated hosts. The occurrence of deeply divergent haplotypes, as well as extensive morphological variability, previously suggested possible cryptic speciation within Ceratitis cosyra. Here we provide the first large-scale characterisation of the population structure of Ceratitis cosyra with the main objective of verifying cryptic genetic variation. A total of 348 specimens from 13 populations were genotyped at 16 polymorphic microsatellite loci. Hardy-Weinberg equilibrium (HWE) deviations were observed in 40.4% of locus-population combinations and suggested the occurrence of genetic substructuring within populations. Discriminant Analysis of Principal Components (DAPC) showed genetic divergence between the vast majority of vouchers from Burundi and Tanzania (plus a few outliers from other African countries) and all other specimens sampled. Individual Bayesian assignments confirmed the existence of two main genotypic groups also occurring in sympatry. These data provided further support to the hypothesis that Ceratitis cosyra might include cryptic species. However, additional integrative taxonomy, possibly combining morphological, ecological and physiological approaches, is required to provide the necessary experimental support to this model.     

Seven new hypselostomatid species from China,including some of the world’s smallest land snails (Gastropoda,Pulmonata, Orthurethra)

Seven new species of Hypselostomatidae are described from the Chinese province Guangxi: Angustopila dominikae Páll-Gergely & Hunyadi, sp. n., Angustopila fabella Páll-Gergely & Hunyadi, sp. n., Angustopila subelevata Páll-Gergely & Hunyadi, sp. n., Angustopila szekeresi Páll-Gergely & Hunyadi, sp. n., Hypselostoma socialis Páll-Gergely & Hunyadi, sp. n., Hypselostoma lacrima Páll-Gergely & Hunyadi, sp. n. and Krobylos sinensis Páll-Gergely & Hunyadi, sp. n. The latter species is reported from three localities. All other new species are known only from the type locality. Specimens nearly identical to the type specimens of Angustopila huoyani Jochum, Slapnik & Páll-Gergely, 2014 were found in a cave in northern Guangxi, 500 km from the type locality. Adult individuals of Angustopila subelevata sp. n. (shell height = 0.83–0.91 mm, mean = 0.87 mm) and Angustopila dominikae sp. n. (shell height of the holotype = 0.86 mm) represent the smallest known members of the Hypselostomatidae, and thus are amongst the smallest land snails ever reported. We note that Pyramidula laosensis Saurin, 1953 might also belong to Krobylos. Paraboysidia neglecta van Benthem Jutting, 1961, which was previously included in Angustopila, is classified in Hypselostoma.     

Systematics of the Rhinella margaritifera complex (Anura,Bufonidae) from western Ecuador and Panama with insights in the biogeography of Rhinella alata

The Rhinella margaritifera species group consists of 17 species of toads distributed in tropical and subtropical South America and eastern Central America. The identity of some of its species is poorly understood and there are numerous undescribed cryptic species. Among them, the status of Rhinella margaritifera is one of the most problematic. Its range includes lowland rainforests separated by the Andes, the Chocoan rainforest to the west and the Amazonian rainforest to the east. This distribution is puzzling because the Andes are an old and formidable barrier to gene flow and therefore should generate vicariant speciation between disjunct lowland populations. Herein we clarify the taxonomy of populations of the Rhinella margaritifera complex from Central America and the Chocó region of South America. The morphological and genetic variation of Rhinella margaritifera was examined from 39 populations from Chocó, 24 from the upper Amazon region of Ecuador, and 37 from Panama, including the holotype of the Panamanian Rhinella alata. Phylogenetic analyses were performed based on mitochondrial genes 12S rRNA, 16S rRNA, and cytochrome c oxidase I (COI) and the nuclear gene Tyrosinase (Tyr). The genetic and morphological data show that Panamanian and Chocoan populations are conspecific. In the phylogeny, populations from Chocó and Panama form a well-supported clade. The morphology of the holotype of Rhinella alata falls within the variation range of Panamanian and Chocoan populations. Based on all this evidence, we assign the populations from western Ecuador and Panama to Rhinella alata and demonstrate that the unusual distribution pattern of “Rhinella margaritifera” on both sides of the Andes was an artifact of incorrectly defined species boundaries.     

A new species of Paracreptotrema (Digenea,Plagiorchiformes, Allocreadiidae) infecting two species of poeciliids in Río Malila of the Río Pánuco basin,Hidalgo, México,with a key to the species of the genus

Paracreptotrema rosenthali sp. n. was discovered in the intestine of Xiphophorus malinche and Pseudoxiphophorus jonesii, collected from the headwaters of Río Malila, tributary of Río Conzintla, in the Río Pánuco basin, Hidalgo, México, during 2008–2009. The new species differs from the five known species of Paracreptotrema Choudhury, Pérez-Ponce de León, Brooks & Daverdin, 2006 by having vitelline follicles that extend from a level anterior to the pharynx to mid-testes, the seminal vesicle which is more extensively folded, and a wider cirrus sac. The new species resembles Paracreptotrema heterandriae in the length of its ceca, which surpasses the posterior margin of the ovary but do not reach the testes. A key to the species of Paracreptotrema is provided.     

The alien ascidian Styela clava now invading the Sea of Marmara (Tunicata: Ascidiacea)

During the implementation of a large project aimed to investigate the benthic community structures of the Sea of Marmara, specimens of the invasive ascidian species Styela clava were collected on natural substrata (rocks) at 10 m depth at one locality (Karamürsel) in İzmit Bay. The specimens were mature, containing gametes, indicating that the species had become established in the area. The Sea of Marmara seems to provide suitable conditions for this species to survive and form proliferating populations.     

A snail in the long tail: a new Plekocheilus species collected by the ‘Comisión Científica del Pacífico’ (Mollusca,Gastropoda, Amphibulimidae)

Among the historical collection gathered by the ‘Comisión Científica del Pacífico’ during 1862–1865, type material was found of one of the species described on the basis of the material collected shortly afterwards. Inspection of the types revealed that only one specimen may be considered as type material of Bulimus aristaceus Crosse, 1869; this specimen is now designated as the lectotype. The other specimens are described as a new species, Plekocheilus (Plekocheilus) cecepeus.     

Nuclear ribosomal internal transcribed spacer 1 (ITS1) variation in the Anastrepha fraterculus cryptic species complex (Diptera,Tephritidae) of the Andean region

The nuclear ribosomal internal transcribed spacer 1 (ITS1) was sequenced for Anastrepha fraterculus (Wiedemann, 1830) originating from 85 collections from the northern and central Andean countries of South America including Argentina (Tucumán), Bolivia, Perú, Ecuador, Colombia, and Venezuela. The ITS1 regions of additional specimens (17 collections) from Central America (México, Guatemala, Costa Rica, and Panamá), Brazil, Caribbean Colombia, and coastal Venezuela were sequenced and together with published sequences (Paraguay) provided context for interpretation. A total of six ITS1 sequence variants were recognized in the Andean region comprising four groups. Type I predominates in the southernmost range of Anastrepha fraterculus. Type II predominates in its northernmost range. In the central and northern Andes, the geographic distributions overlap and interdigitate with a strong elevational effect. A discussion of relationships between observed ITS1 types and morphometric types is included.     

Alpha taxonomy of the genus Kessleria Nowicki, 1864, revisited in light of DNA-barcoding (Lepidoptera,Yponomeutidae)

The taxonomy of Kessleria, a highly specialized montane genus of Yponomeutidae with larval host restriction to Saxifragaceae and Celastraceae (Saxifraga spp. – subgenus Kessleria; Saxifraga spp. and Parnassia spp. – subgenus Hofmannia), is revised based on external morphology, genitalia and DNA barcodes. An integrative taxonomic approach supports the existence of 29 species in Europe (the two known species from Asia and North America are not treated herein). A full 658 bp fragment of COI was obtained from 135 specimens representing 24 species, a further seven sequences are >560 bp. Five new species are described: Kessleria cottiensis sp. n. (Prov. Torino, Italy; Dep. Hautes Alpes, France), Kessleria dimorpha sp. n. (Dep. Alpes-de-Haute-Provence, France), Kessleria alpmaritimae sp. n. (Dep. Alpes-Maritimes, France), Kessleria apenninica sp. n. (Prov. Rieti, Prov. L´Aquila, Italy), and Kessleria orobiae sp. n. (Prov. Bergamo, Italy).