A new,sibling, tree frog from Jerusalem (Amphibia: Anura: Hylidae)∗

A new, sibling, species of tree frog of the Hyla arborea group is described from Jerusalem and the adjacent Judean hills. Hyla heinzsteinitzi sp. n. differs from the sympatric common H. savignyi of the Middle East in head shape, as the head is relatively wider and the snout more truncate; in call structure, as in each segment of its advertisement call the energy peaks abruptly near the segment's onset; and in colouration, as its dark lateral band is highly disrupted. In at least one of the H. heinzsteinitzi populations the frogs' green‐phase colour is turquoise rather than green. The distinction of the two species is demonstrable by Principal Coordinate Analysis. Hyla heinzsteinitzi seems to be endemic to a small area within the range of distribution of H. savignyi; here the two are sympatric and at least sometimes apparently syntopic. The ecological relations of the two are unknown. The new species is probably endangered.     

Genetic differentiation and gene flow of Porcellionides pruinosus (Crustacea: Isopoda: Oniscidea) in Tunisia

Iso-enzymatic polymorphism analysis was carried out on Tunisian populations of Porcellionides pruinosus to clarify their confused taxonomic status, previously revealed by reproductive and morphological approaches. Seventeen allozyme loci were studied, using horizontal starch gel electrophoresis, in 230 adult specimens belonging to nine populations of P. pruinosus located in north, central and south Tunisia, to assess the levels of their intra-genetic and inter-genetic diversity, and to analyse their genetic structure. Our results indicate a low genetic variability with a polymorphism ratio ranging from 29.4% in Tabarka to 47.1% in the Garat Nâam and Raccada populations, and the mean number of alleles per locus was estimated at 1.51 ± 0.15. The direct-count of heterozygosity ranged between 8.1% and 15%. Substantial estimates of demic structuring were found at the total population (F_ST = 0.198), with a generalized and significant heterozygote deficiency detected in every population (with a mean value for all populations of F_IS = 0.372). Genetic distances between the studied populations were low, indicating that they belong to the same species. Nevertheless, the genetic analyses showed the divergence of the population of Tabarka from all the other studied populations, with the lowest mean gene flow, 0.58, and the highest mean genetic distance, (0.182) obtained also by the morphological and reproductive approaches.     

Morphology and larval development of Polydora cf. websteri (Polychaeta: Spionidae) in a tropical region of north-eastern Brazil

Polydora websteri belongs to a group of the Spionidae known as the Polydora complex, members of which are known for building tubes within the shells of mussels and for their ability to infest commercially valuable bivalves, which causes serious losses for malacoculture. P. websteri is a species subjected to increasing research due to the global scale at which these infestations occur. In this paper, we describe the larval development of P.cf. websteri associated with the oyster Crassostrea cf. brasiliana in a tropical region of north-east Brazil. The main structures of all stages of development were analysed and compared to other populations of P. websteri. Ovigerous capsules of P.cf. websteri were cultured in the laboratory and the larval stages were analysed until metamorphosis. Ovigerous capsules were present in all sampling intervals, indicating that this population exhibits continuous reproduction. Up to 21 capsules per tube were found, and the number of eggs per capsule varied from 20 to 40. All observed eggs developed into larvae. Larval development from egg to recruitment stage did not exceed 13 days. Metamorphosis occurred when larvae attained 17 chaetigers, at which stage important diagnostic characteristics were revealed, such as the spines on chaetiger 5. The developmental stages of P.cf. websteri larvae, and the morphological features found at each stage, are in accordance with what has been recorded for temperate populations. In our study, the planktonic stage was found to be of shorter duration than elsewhere, which might be associated with the higher temperatures at which larvae were cultured compared to previous studies.     

Revisiting the saproxylic beetle ‘Propomacrus cypriacus Alexis & Makris, 2002’ (Coleoptera: Euchiridae) using molecular,morphological and ecological data

Propomacrus cypriacus a saproxylic beetle endemic to Cyprus, was described as a species distinct from P. bimucronatus on the basis of a few morphological characters and has been evaluated as Critically Endangered by IUCN, as it depends on veteran trees and has a restricted distribution. Herein we explored the distribution, ecology, morphology and genetic variation of the populations from Cyprus, and we attempted a phylogenetic analysis including related species based on two mitochondrial and one nuclear DNA markers. We also used published mtDNA divergence rates to date cladogenetic events. Despite extensive sampling, we found only a small number of individuals, adding a few additional localities to the known distribution of the species, and we identified two more host species to those previously reported. Morphological examination showed that only the form of male protibiae differs consistently in Cyprus populations compared to those of P. bimucronatus from Turkey and the Lebanon. Nevertheless, genetic analyses do not support a distinct species status for populations from Cyprus, which should instead be considered as a subspecies of P. bimucronatus. Morphological and ecological data further support this conclusion. Cyprus populations have diverged from nearby continental ones in mid to late Pleistocene.     

Do male tree frogs feed during the breeding season? Stomach flushing of five syntopic hylid species in Rio Grande do Sul,Brazil1

The diet of males of five syntopic hylid species from the Araucaria plateau of Rio Grande do Sul was studied by stomach flushing. Because the duration of their breeding seasons was rather different, the levels of food intake at the respective breeding sites were analysed. We found a positive correlation between the length of the male calling period and the ingestion of prey. The longer the species‐specific reproductive activity of the males, the more individuals fed during that time. The results are discussed under congruence aspects of feeding and mating microhabitats in these tree frogs. In addition, differences in niche breadth may influence the temporal and spatial partitioning between the species studied and result in quantitative as well as qualitative patterns of prey consumption, thereby ensuring a maximum of fitness gain.     

Non-monogamous mating system and evidence of lekking behaviour in the Helmeted Manakin (Aves: Pipridae)

ABSTRACT

Manakins (Pipridae) are a group of Neotropical birds well known for their spectacular lekking displays and non-monogamous mating system. Nevertheless, the two species of Antilophia have been traditionally considered monogamous and, therefore, an exception to this rule. In this paper, we studied the home range and the mating system of a colour-ringed population of the Helmeted Manakin Antilophia galeata in southeastern Brazil. We propose that it is a non-monogamous species that ranges widely during the breeding season, presenting a lekking behaviour in the form of unspectacular but aggressive chasing courtship displays.     

A morphological study of Phyllotreta tetrastigma (Coleoptera: Chrysomelidae: Alticinae) including two colour forms

P. tetrastigma is a chrysomelid beetle with two colour forms, and has populations feeding on two different host plants. Morphological variation was studied within both males and females of six populations of P. tetrastigma. Morphological difference was detected between the sexes and between populations from different localities but not between the colour forms. It is not clear whether host plant influences morphology.     

Mating behaviour in the bees,Dieunomia heteropoda and Nomia tetrazonata,with a review of courtship in Nomiinae (Hymenoptera: Halictidae)

Males of many bees in the subfamily Nomiinae (Halictidae) have numerous secondary sexual characters, which provide much systematic information. Similarly, the use of these structures by males during sexual behaviour may provide useful characters, but the courtship and copulatory behaviour of most of these bees is not known. Structures and behaviour used for courtship and mating are described for two species of nomiine bees, Dieunomia heteropoda and Nomia tetrazonata, along with brief observations of mating in a parasitic bee, Triepeolus verbesinae (Apidae: Nomadinae). A review of mating behaviour within the Nomiinae shows no obvious phyletic patterns, based upon present limited knowledge. The species-specific nature of the male secondary sexual characters suggests they may have evolved in the context of sexual selection, but behavioural data from populations of additional species are needed to evaluate the hypothesis adequately.     

Reproduction,phenology and mortality sources of a species of Physalaemus (Anura: Leptodactylidae)

Physalaemus cf. fuscomaculatus is a foam-nesting frog of seasonally dry environments of south-eastern Brazil. We report on the reproduction and other natural history features of this species. Population parameters were based on trapped specimens. The advertisement call is described. The calling and spawning season is coincident with the rainy period. Calling and spawning were performed in ponds. Most of the adults were trapped in October and the juveniles during wetter months and the beginning of the dry season. At the end of the reproductive season, the median size of adults lowered. Egg clutches averaged 2350 eggs, each egg being about 1.4?mm in diameter. Males were more abundant than females in traps. Communal foam nests were frequent, involving up to 40 individual nests. Tadpoles entered the water within about 40?h. Maggots (Diptera) infested egg clutches; water bugs and mammals preyed upon adults. In this region, the dry/cold season may be so difficult for survival that early maturation at small size would be advantageous. The male-biased sample may be related to the greater motility of males. The rapid abandonment of the foam by the tadpoles and communal nesting may help to reduce predation by maggots. Skin toxins were not effective in deterring predation upon adults. The call of P. cf. fuscomaculatus differs from that of other species in the biligonigerus group and can help to define the specific status of the studied population.     

The systematics of Pyrenean and Cantabrian Cochlostoma (Gastropoda,Cyclophoroidea) revisited

The systematics of Pyrenean and Cantabrian species of the terrestrial prosobranch genus Cochlostoma Jan, 1830 is readdressed using morphology of shell and soft parts, anatomy and allozyme electrophoresis. Ten biological species are recognized: C. obscurum (Draparnaud, 1805), C. crassilabrum (Dupuy, 1849), C. partioti (Moquin-Tandon, 1848), C. martorelli (Bourguignat, 1880), C. gigas Gofas and Backeljau, 1994, C. nouleti (Dupuy, 1851), C. hidalgoi (Crosse, 1864), C. bicostulatum Gofas, 1989, C. oscitans Gofas, 1989 and C. asturicum (Raven, 1990); forty other available species-group names are synonymized accordingly. The populations are very isolated one from another in the southern Pyrenees, with a high incidence of fixed alleles. Cochlostoma gigas is given species status but may be interpreted as a peripheral isolate from C. martorelli. The amount of divergence found between populations of C. martorelli is high, and the intrapopulation polymorphism is very low, a setting which is very prone to further breakup. In the Cantabrian range, C. asturicum has hardly diverged from C. hidalgoi, but fixed or nearly fixed alleles for three enzymatic loci are evidence for reproductive isolation in fully sympatric populations. Cochlostoma asturicum is nevertheless genetically closer to the local C. hidalgoi than the latter is to distant conspecific populations. This points to a sympatric speciation for C. asturicum. It is suggested that the montane habitat of several species is the consequence of an adaptation developed when a colder climate forced the species into coastal refuges. The onset of temperate climate could then result in a disruptive selection, with alternate pathways of a retreat to higher altitude, or colonisation of newly available space in the lowlands; this process is considered as a possible explanation for the lowland/montane divergence between C. obscurum and C. crassilabrum.     

Expulsion and consumption of male ejaculates by promiscous female Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae)

While expulsion of male ejaculates by the females after copulation has been reported for various animal groups, expulsion followed by consumption of the expelled ejaculate is a rare behaviour outside spermatophylax-producing orthopterans. Among Diptera, this behaviour has been reported for a few species of Piophilidae, Empididae and Ulidiidae. Here we report on its occurrence among Euxesta eluta and Euxesta mazorca (Diptera: Ulidiidae). We also attempt to characterize the mating system of E. eluta in order to facilitate future hypothesis testing to understand the behavioural factors leading to the evolution of this peculiar behaviour. For this, courtship sequences, copulation duration, frequency of ejaculate expulsion and subsequent consumption, and latency to ejaculate expulsion for both E. eluta and E. mazorca were recorded. The time of sexual maturation, the time window of sexual receptivity during the day, and the mating frequency and variance in mating success for males and females (degree of polygamy) were determined for E. eluta. Both E. eluta and E. mazorca males engaged in elaborate courtship sequences involving visual and tactile displays before copulation. Females of both species almost invariably expelled and consumed ejaculates after copulation. Female E. eluta, required a 6- to 9-day period feeding on protein and sugar before becoming sexually receptive. Reproductive activity occurred continuously over the day with an early morning and late afternoon peak. Both males and females could mate multiply, with multiple partners over a 2-h observational period. Sexual network analysis revealed that some males and females had greater mating success than others. Ejaculate consumption appears to be widespread in the genus Euxesta. It is possible that females obtain nutrients from this behaviour while exerting some control over egg fertilization.     

New damselfly hosts and species identification of an aquatic parasitoid Hydrophylita emporos (Hymenoptera: Trichogrammatidae) in Taiwan

ABSTRACT

The host–parasitoid relationship and species identity of aquatic parasitoids of two damselflies Coeliccia cyanomelas (Platycnemididae) and Psolodesmus mandarinus dorothea (Calopterygidae) from Fushan and Lienhuachih in Taiwan were studied using morphological characters and DNA barcoding sequences. The parasitoids reared from the damselflies’ eggs, and the field-collected parasitoids, were morphologically identified as Hydrophylita emporos (Trichogrammatidae), a recently described parasitoid of the damselfly P. m. mandarinus from Northern Taiwan. The CO1 (cytochrome c oxidase I) gene tree supported the identification as H. emporos, as well as all parasitoid samples from C. cyanomelas, P. m. dorothea and P. m. mandarinus. The sampled H. emporos populations did not differ genetically despite their different host associations. However, some genetic differences were found between H. emporos populations from Northern and Central Taiwan, indicating that the dispersal of H. emporos may be limited by geographical distances. Our results suggest that H. emporos can parasitise not only closely related sister subspecies, P. m. mandarinus and P. m. dorothea, but also phylogenetically distant species of another damselfly family, C. cyanomelas. This is the first record of multiple damselfly hosts for the aquatic parasitoid genus Hydrophylita. This finding implies that the host range of H. emporos and congeneric species may be broader than previously thought.     

Asymmetrical male mandibular horns and mating behavior in Agathidium Panzer (Coleoptera: Leiodidae)

Males of some species of Agathidium Panzer have a prominent horn on the dorsal surface of the left mandible. The horn is unique in that it is highly asymmetrical. One species, A. marae Miller and Wheeler, has an additional horn on the right side of the frons. The horns are categorized into four general morphological types. Scaling relationships are investigated for two species, A. angulare Mannerheim and A. pulchrum LeConte. These species exhibit a scaling relationship that is strongly discontinuous (sigmoid) with a prominent “switch point”. Measurement data (pronotal width versus mandible height) are fit to a nonlinear regression model to determine the switch points and slopes at the switch points for the populations of each species. Behaviors associated with mating and other behaviors are described for A. pulchrum. Males use a head‐thrusting behavior to dislodge rival males from the substrate. Horned males appear better able to dislodge opponents. Head thrusting is also used less vigorously on females before and after mating.     

Reproductive ecology of the glass frog Espadarana prosoblepon (Anura: Centrolenidae) in an urban forest of the Central Andes of Colombia

Population and reproductive behaviour aspects of the glass frog Espadarana prosoblepon are well known for populations from Central America, but similar information is lacking for populations from South America. We recorded the reproductive ecology of a population of the glass frog E. prosoblepon in the city of Armenia, Central Andes of Colombia. With weekly surveys with mark-recapture between December 2013 and November 2015, we monitored activity patterns and evaluated if the probability of males mating is associated to their body size or to chorus tenure (i.e. the number of nights a same individual is calling for attracting a mate). In addition, upon observing an amplexus, we recorded the behaviour of the individuals until oviposition and noted characteristics of egg clutch. We recorded 47 males and 32 females, nine amplexus events, eight ovipositions, and 21 egg clutches (35.4 ± 4.79 eggs per clutch, N = 29). Activity of males and females and the number of egg clutches were positively correlated to rainfall. Mating success of males was not associated to their body size, but it was positively associated to longer chorus tenure; clutches from large females had a higher number of eggs than clutches from small females. Oviposition occurred on average 3.76 ± 1.74 hours after we first observed the amplexus, occurring 93.1% of the time in fronds of Selaginella geniculata at an average height of 1.58 ± 0.44 m. After the oviposition, the male left the site, while the female remained between 30 and 45 min, partially covering the eggs. The reproductive behaviour of E. prosoblepon did not vary widely between a population in Costa Rica and our population in time to oviposition, and mating success of males associated to chorus tenure; however, in our study population we recorded a larger clutch size and the preference for S. geniculata fronds as oviposition substrate.     

Genetic diversity in the Carabodes marginatus species group (Acarida,Oribatida, Carabodidae) as inferred from allozymes

Ten populations representing four species of the ‘Carabodes marginatus’ group, as C. marginatus, C. affinis, C. quadrangulus and C. montanus from central Italy have been studied for genetic variation at 16 enzymatic loci by electrophoresis. Morphotypes close to C. marginatus but easily distinguishable by the shorter marginal notogastral setae were also included in the analysis to evaluate their systematic status. Carabodes arduinii, a very different species from the ‘coriaceus’ group, was employed as an out-group. Phylogenetic relationships among different species were studied and inferred data demonstrated that cladogenetic events occurred, even though rates of molecular and morphological evolution do not correlate. Morphotypes related to C. marginatus were genetically very close to the type specimens, but a different pattern of allele frequency was observed in sympatric populations of the two setal forms. Possible explanations for this finding are suggested in relation to historical processes. The rate of gene flow among populations of the two morphs was estimated by both Wright's F _st and Slatkin's Nm methods. These two different procedures yielded contradictory results.     

The advertisement calls of three South American poison frogs (Amphibia: Anura: Dendrobatidae), with comments on their taxonomy and distribution

The advertisement calls of three species of dendrobatid frogs from Bolivia and Peru are described. For each species we present a characteristic audiospectogram and oscillogram. We also provide numerical information about the spectral and temporal features of the calls. Our recordings of these species are compared with previously described calls of the same species from other locations. Comparisons between the calls of three different populations of Colostethus from Peru and Bolivia support their conspecificity. Characteristics of the call of a Peruvian population of Epipedobates differ from those of E. hahneli, suggesting that further taxonomic research is needed. The distribution of the genus Colostethus and Epipedobates in Bolivia is reassessed.     

On some African and Oriental Wolf Spiders (Araneae,Lycosidae): redescription of Pardosa oncka Lawrence from Africa,with notes on its generic position

Pardosa oncka Lawrence (with P. irrasa Roewer, P. kisokwea Roewer, P. resoluta Roewer, P. versuta Roewer and P. wittei Roewer here placed as junior synonyms) is described and illustrated. The male is recognized by its bifurcate embolus. The female has conspicuous mating plugs more or less concealing two pits in the epigyne. The species is widely distributed in Central and South Africa, and may be grouped in a separate genus (here not formally recognized) together with P. venatrix auctt. and some other species listed.     

Reflection of the Neogene–Quaternary phylogeography in the recent distribution limiting climatic factors of eight Mediterranean Phlebotomus species (Diptera: Psychodidae)

The diversification of Mediterranean Phlebotomus species occurred during the Neogene and Quaternary periods. The climatic and geographical history of the Mediterranean area plausibly influenced the climatic requirements of recent sand fly species. Our aim was to investigate the European zoogeography and the climate-based ecological groups of eight Phlebotomus species considering the phylogeography of the studied taxa. Jaccard index calculation, hierarchical cluster and principal component analyses were performed. The results of Jaccard-coefficient calculations may indirectly refer to the former existence of three possible glacial refuges of sand fly populations in Europe: an Iberian, an Apennine and a Balkan Peninsula–Asia Minor refuge. Three main climatic groups of the studied sand fly species were distinguished, namely the so-called ‘Trans-Mediterranean’, ‘East Mediterranean’ and ‘West Mediterranean’ groups. Our results suggest that the ancestor of Phlebotomus similis and Phlebotomus sergenti was adapted to the hot, dry summer Mediterranean climate during the late Neogene. The third member of this climate group, Phlebotomus papatasi because of co-adaptation shows similar climatic requirements as P. similis and P. sergenti. The five Larroussius species studied should have evolved under moister climatic conditions. Species of the ‘West Mediterranean’ group were adapted to the milder and wetter oceanic climate of Atlantic coasts. The species of the ‘East Mediterranean’ group diversified under the less balanced climate of the East Mediterranean. The recent climatic requirements and the European distribution of the studied species is the consequence of the complex Quaternary–Neogene geographical and climatic history of their former habitats in the Old continent. The adaptation of the ancestors of the studied sand fly species to the changing geographical and climatic factors of Europe are discussed.