Patterns of prey use in four sympatric species of Philanthus (Hymenoptera: Sphecidae) with a review of prey selection in the genus

Summary

Observations were made on four sympatric species of sphecid wasps of the genus Philanthus that nest in late summer and prey upon Hymenoptera. All four species nest in a restricted locale, but with little overlap in nesting areas. Comparison of nest contents of females demonstrated limited overlap in prey utilization among the four species, but only when both size and taxonomic category of prey were considered. The ultimate cause of this resource partitioning is unknown. Review of 30 populations of 16 species of North American Philanthus reveals varying degrees of specialization on prey types. Although populations of one species, separated in either time or space, tend to have similar niche breadths, this is not always the case. Intra-and inter-populational comparisons of prey records suggest behavioural plasticity allowing response to variations in prey availability. Some of the divergence in prey use among species is due to differences in predator size.     

Diet of tropical insectivorous birds in lowland Malaysian rainforest

ABSTRACT

Despite the importance of diet in avian ecology, knowledge of bird diet in tropical regions, particularly South-East Asia, is limited. Assessing predator–prey relationships of how energy flows across ecosystems provides insight into niche segregation of sympatric species. We examined induced-regurgitation samples from 15 species of insectivorous birds – 12 babblers and three flycatcher-like species – from the Krau Wildlife Reserve, central Peninsular Malaysia. The main objective of this study was to compare and characterise the diet of similar trophically insectivorous birds so as to examine the occurrence of dietary overlap or segregation. The majority of identified prey items belong to the following arthropod taxa: Coleoptera (53%), Hymenoptera (19%), Blattodea (11%), and Araneae (11%). The diet of all bird species slightly overlapped (p = 0.070) but network analysis yielded a relatively low value (R_o = 0.53), indicating dietary divergence may have occurred among the studied bird species, possibly reflected by the variation in proportion of arthropod groups. A high proportion of Coleoptera and three other arthropod groups (i.e. Hymenoptera, Blattodea and Araneae) in the diet of studied birds could be related to their richness and abundance in aerial leaf-litter, the substrate preferred by most babblers. Dietary differences may further be explained by variations in bill shapes and sizes, likely corresponding to prey size. Unique foraging strategies (utilising different vertical strata, microhabitats, and attack manoeuvres) could also lead to dietary separation among sympatric species. Flycatcher-like species were expected to flycatch different prey items (families or lower taxa) versus the gleaning birds (most babblers). This study successfully discerned more details on the diets of insectivorous birds inhabiting Malaysian forests, providing a foundation for future studies on the ecology and biology of understorey birds.     

Natural prey of the crab spider Thomisus onustus (Araneae: Thomisidae), an extremely powerful predator of insects

The natural prey of the crab spider Thomisus onustus inhabiting Eryngium plants was studied in the Absheron Peninsula, Azerbaijan. The total percentage of specimens of T. onustus found while feeding was low (9.1%). However, solitary late instar and adult females exhibited a very high feeding percentage (29.9%). Other sex–age groups of T. onustus, including small juveniles, adult males, and females guarding egg sacs, were found feeding significantly less frequently. Investigation has shown that T. onustus is a polyphagous predator, with representatives of four arthropod orders found in its diet. The primary food of T. onustus was Diptera and Hymenoptera, which collectively accounted for 94.2% of total prey. Worker ants constituted about one‐fifth of the prey, suggesting that T. onustus is a myrmecophagic spider. The length of prey killed by T. onustus ranged between 1.25 and 16.00 mm (mean 6.21 mm) and constituted from 34.4 to 587.9% (mean 194.9%) of length of their captors. The most frequently captured were large arthropods, exceeding the size of the spiders (83.1%). Small and medium‐sized juveniles captured considerably larger prey (in terms of prey–predator length ratio) than late instar and adult females. Over half of their prey constituted arthropods more than two‐fold larger than the spiders.     

A method of obtaining dietary data for slow worms (Anguis fragilis) by means of non‐harmful cooling and results from a Danish population

Diet composition of slow worms (Anguis fragilis) from a Danish population was recorded from May to September 2006. Slow worms were maintained in cool conditions (at 8°C) for a maximum of 126 h, which made approximately half of the animals regurgitate. The method worked equally well on juveniles and adults. The regurgitations revealed that the slow worms preyed on small snails, slugs, pill millipedes (Glomeris marginata), earthworms and Lepidoptera larvae. There were seasonal changes in taxon composition of the diet but no ontogenetic or sex‐related differences. The food quality of selected prey types was tested on juveniles in a laboratory experiment. Mealworms gave a significantly lower gain in mass and snout–vent length than slugs, earthworms and a mix of all three prey types. A supplementary experiment revealed that slow worms from the present population completely rejected mealworms as prey. Earthworms seemed to be nutritionally inferior to molluscs.     

Genetic diversity of freshwater crabs (Brachyura: Sesarmidae) from central Jamaica with description of a new species

Jamaica is the only island of the Greater Antilles where freshwater streams are not populated by representatives of the old lineage of freshwater crabs, the Pseudothelphusidae. Instead, a very diverse fauna of endemic sesarmid crabs inhabits freshwater and terrestrial habitats throughout the island, thereby showing complete independence from the sea. They have been reported thriving in bromeliad leaf axils, rock rubble, empty snail shells, caves and mountain streams. Otherwise, the Sesarmidae are typical inhabitants of soft‐sediment littoral habitats like marshes and mangroves. For many years, crabs from Jamaican mountains streams had been considered to belong to a single species, Sesarma bidentatum Benedict . However, recent morphological and molecular studies have shown that crabs from mountain streams of different regions of the island belong to different species. Consequently, four new crab species have been described over the last 10 years. In this study, we give evidence that freshwater streams in central Jamaica also host two different species of crabs. In streams draining to the north we exclusively found the species S. windsor Türkay and Diesel, , while streams draining to the south were inhabited by a closely related but undescribed species of crab. The southern species is here described as new and is referred to as Sesarma meridies sp. n. Morphological and molecular (12S and 16S mtDNA) evidence is provided that allows these two species of freshwater crabs from central Jamaica to be distinguished. The species richness of Jamaican endemic sesarmids thereby increases to 10, which makes the island unique in terms of its diversity of land‐dwelling crabs.     

Distribution and postbreeding environmental relationships of northern leopard frogs ( Rana [ Lithobates ] pipiens ) in Washington

Northern leopard frogs ( Rana [ Lithobates ] pipiens ) are considered sensitive, threatened, or endangered in all western states and western Canadian provinces. Historically present in eastern Washington in 6 major river drainages, leopard frogs are now only known to occur at 2 localized areas in the Crab Creek drainage in Grant County. During the summers of 2002–2005, we surveyed both areas to document extent of leopard frog distributions and to describe habitat and vertebrate community characteristics associated with leopard frog site occupancy. At Gloyd Seeps, 2 juvenile leopard frogs were observed in a total of 8.2 person-days of searching along a 5-km stream reach. At Potholes Reservoir, we surveyed 243 wetland sites in 7 management units known to have been occupied by leopard frogs during the 1980s. We confirmed leopard frog presence at only 87 sites (36%) in 4 management units. Site occupancy models for individual ponds indicated that, compared to unoccupied sites, occupied sites had slightly greater pond depths, less tall emergent vegetation, more herbaceous vegetative cover, and fewer neighboring ponds containing nonnative predatory fish. Models developed at the 1-km 2 scale indicated that occupied areas had greater average midsummer pond depths, fewer ponds occupied by bullfrogs ( Rana [ Lithobates ] catesbeiana ) and carp ( Cyprinus carpio ), and more herbaceous vegetation surrounding ponds. The Gloyd Seeps population now appears defunct, and the Potholes Reservoir population is in sharp decline. Unless management actions are taken to reduce nonnative fish and bullfrogs and to enhance wetland vegetation, leopard frogs may soon be extirpated from both sites and possibly, therefore, from Washington.     

Food habits and dietary overlap of nongame insectivorous fishes in Flint Creek, Oklahoma, a western Ozark foothills stream

Insectivorous fishes were sampled from March, 1983 to February 1984, in Flint Creek, Delaware Co., Oklahoma. There was insignificant habitat segregation between Etheostoma spectabile and E. punctulatum and seasonal habitat partitioning between Cottus carolinae and both darters. Mature E. spectabile ate primarily chironomids and mayflies, whereas juveniles fed primarily on microcrustaceans. Mature E. punctulatum consumed fewer Ephemerella and Leptophlebia than E. spectabile , feeding on Stenonema and other crustaceans. Juvenile E. punctulatum fed mainly on amphipods and mayflies, and juvenile E. spectabile ate primarily microcrustaceans. Cottus carolinae elected primarily mayflies in spring - summer and chironomids in January - February. Coefficients of dietary overlap were highest between larger E. spectabile and juvenile E. punctulatum and lowest between immature E. spectabile and mature E. punctulatum . Overlap between the two darters was significantly correlated with differences in mean prey size (p E. spectabile was also significantly correlated to differences in mean prey sizes. Etheostoma spectabile generally preferred smaller prey than E. punctulatum . All three species avoided Stenelmis . Cottus carolinae avoided microcrustaceans. The study showed that resource partitioning among these three insectivorous fishes is affected by complex interactions of habitat and prey electivity, and prey size selectivity.         

Microdistribution of juveniles and adults of the mud crab Panopeus americanus (Brachyura,Panopeidae) in a remnant mangrove area in the southwest Atlantic

The distribution and density of a population of the narrowback mud crab Panopeus americanus were evaluated, with emphasis on the different developmental stages (juveniles and adults) and sexes. Specimens were collected in a remnant human‐impacted mangrove at Araçá on the northern coast of São Paulo, Brazil, at two‐month intervals from January through July 2006, using 1 m×1 m sample plots randomly distributed in the intertidal zone. The population was abundant, with a mean density of 14.3±9.3 individuals/m². Larger individuals were more abundant in the upper and middle intertidal zones, whereas smaller crabs were predominant in the lower intertidal and thus remained submerged longer. Males and juveniles were homogeneously distributed. Females preferred regions farther from the waterline. A clustered distribution pattern suggested gregarious behaviour and settling of juveniles in the vicinity of the adults' habitats, in an adaptive process as a function of tidal rhythms.     

Notes on Simulium ovazzae Grenier and Mouchet (Diptera: Simuliidae) and river crabs (Malacostraca: Potamidae) and their association

Summary

Collections of crabs in the forest zone of West Cameroon produced three species—Potamonautes africanus, P. orthostylis and P. pobeguini. The P. africanus carried populations of immature Simulium ovazzae.

Analysis of crabs collected by hand and those collected in traps indicates very little overlap in the sizes of the crabs obtained by the two techniques. This problem is discussed.

The infestation rates of S. ovazzae on P. africanus were found to be highest on medium-sized crabs. The attachment sites of the S. ovazzae on the crabs were found to vary with both the size of the crab and the size of the Simulium larva.

The size distributions of S. ovazzae larvae on small crabs were found to be different from those on medium and large crabs. The characteristic deficiency of smaller larvae on medium and large crabs reported for other crab-phoretic species was also found with S. ovazzae but the smaller crabs showed a deficiency of larger larvae. A hypothesis is put forward to explain the dearth of larger larvae on the smaller crabs and the higher infestation rate on medium crabs.

S. ovazzae are shown to arrive on their hosts as single larvae carried by the current.

Six adult S. ovazzae were reared from pupae in the laboratory. Small numbers of presumed S. ovazzae biting man in Cameroon are reported.     

Dietary overlap in giant salamanders (Dicamptodon): applying null models to resource partitioning

We examined stomach contents of preserved specimens of larval Pacific giant salamander ( Dicamptodon tenebrosus ) and Cope's giant salamander ( D. copei ) collected from sympatric and allopatric stream populations. The dietary components of these specimens were used to calculate dietary overlap between the 2 species and to determine if changes in overlap existed between sympatric and allopatric populations. To statistically test overlap values, a randomization algorithm was used to construct a simulated data matrix (i.e., null model) in order to compare observed values of dietary overlap to a distribution of overlap values from the null model. Significant levels ( P < 0.05) of dietary overlap occurred in all cases of sympatry as well as allopatry. Average dietary overlap in sympatry was significantly lower than in allopatry, suggesting a dietary shift when in sympatry to reduce competition. Diet composition also differed between sympatric and allopatric populations of each species, further suggesting a partitioning of food resources by one or both species when in the presence of its congener.     

On the natural history of Leptodactylus syphax with comments on the evolution of reproductive features in the L. pentadactylus species group (Anura,Leptodactylidae)

We describe the reproduction of Leptodactylus syphax from Brazil and attempt to trace the evolution of reproductive traits within the L. pentadactylus species group. We quantified the proportion of eggs/embryos from fresh nests. We test whether L. labyrinthicus and L. syphax tadpoles can prey on eggs from heterospecific foam nests. Egg clutches were found outside the water at the edges of rocky/seasonal streams, usually within excavated basins. Males beat the foam with their legs moving from side to side. Clutches do not have trophic eggs, since most of their eggs develop into tadpoles. Tadpoles cling on rocks in current/shallow waters. Only L. labyrinthicus tadpoles could prey on eggs from heterospecific foam nests. If the L. pentadactylus group is monophyletic, excavated basins and foam‐beating in lateral motions may be possible synapomorphies of this group. However, large adults, carnivorous tadpoles and trophic eggs appear to be associated synapomorphies of a less inclusive group.     

Feeding habits of the endemic tropical parthenogenetic lizard Cnemidophorus nativo (Teiidae) in a restinga area of northeastern Brazil

The population of Cnemidophorus nativo in the restinga of Guaratiba, Bahia State, Brazil, is carnivorous with a diet consisting predominantly of insect larvae and termites, which is similar to that of other cnemidophorines. The absence of seasonal variation in diet may be a consequence of the availability of termites throughout the year. The small size of the termites and the ease with which both juveniles and adults eat larvae independent of their length were probably the cause of the poor relationship between lizard body measurements and mean number and volume of prey consumed.     

The prey and predators of Homalopsine snakes

The diets of all South-East Asian water snakes of the subfamily Homalopsinae are reviewed. New diet observations for 10 species of homalopsine snakes are presented. The data on four of these, Cantoria violacea, Enhydris doriea, E. longicauda and Gerarda prevostina represent the first published diet records for these species. The observations for Fordonia leucobalia, Myron richardsonii andEnhydris enhydris represent the first taxonomically detailed diet records for these three species. The records provided for Cerberus rynchops, Enhydris plumbea and Homalopsis buccata add new taxonomic categories to previous records. The records provided for Cerberus rynchops, E. plumbea and Homalopsis buccata add new taxonomic categories to previous records. Four generalities emerge from our findings: (1) freshwater homalopsines feed primarily on fish and the adults and larvae of amphibians; (2) at least four of the eight estuarial species specialize on crustaceans while two primarily prey on fish; (3) feeding on crustaceans appears to have evolved independently at least twice; (4) prey are usually less than 10% of the predator's mass. The known predators of homalopsines are summarized and include both invertebrates and vertebrates. The microhabitats of the estuarial species are discussed and related to prey and habitat preferences.     

Two new species of freshwater crabs from Mount Elgon,East Africa (Brachyura: Potamoidea: Potamonautidae) and a re-diagnosis of Potamonautes niloticus (H. Milne Edwards, 1837) with notes on their natural history and onchocerciasis

Two new species of potamonautid freshwater crabs from eastern Uganda, Africa are described, and Potamonautes niloticus (H. Milne Edwards, 1837) is re-diagnosed with key characters illustrated photographically. A total of four species of crabs are now known to occur on Mount Elgon, which borders both Kenya and Uganda. Two of these species are endemic to the mountain and the distribution patterns of all four species indicate a distinct altitudinal preference. All taxa are compared with similar species and their distributions and natural history are reviewed. Furthermore, three of the crabs recorded from Mount Elgon have been implicated in the transmission of onchocerciasis in East Africa.     

Diet comparison between rainforest and cave populations of Craugastor alfredi (Anura: Craugastoridae): does diet vary in contrasting habitats?

Information about interpopulational variation of diet is important for thoroughly understanding the ecological niche of a species, the possible attributes that have allowed it to live in different habitats and ecosystems (if it is the case) and the degree of dietary specialization. This study compared dietary composition and diversity, size of prey consumed and feeding intensity between two geographically proximate populations of Craugastor alfredi, inhabiting contrasting habitats: one located in a remnant tropical rainforest and the other population located within a cave entrance, in eastern Mexico. A total of 118 frogs (50 from rainforest and 68 from cave) were stomach-flushed and, from those, 86 had at least one prey item (42 from rainforest and 44 from cave). Consumption proportions of orthopterans differed between habitats, with frogs from the cave having higher importance values of this prey than frogs from the rainforest. Additionally, both dietary diversity and feeding intensity were higher in the rainforest population. It was concluded that the generalized diet of C. alfredi, besides the high consumption of orthopterans by the cave population, may have contributed to its success in cave entrances. Nonetheless, other non-feeding attributes like small body size, direct development and semiarboreality may also facilitate the use of rock crevices in caves for foraging and reproduction.     

Ecological notes of the alien species Godiva quadricolor (Gastropoda: Nudibranchia) occurring in Faro Lake (Italy)

The first record from Sicily of the introduced facelinid nudibranch Godiva quadricolor allowed the detection of trophic relationships with the polycerid Polycera hedgpethi, another non-native nudibranch, and with two bryozoan species, namely the naturalized Cheilostomatida Bugula neritina and the cryptogenic Ctenostomatida Amathia verticillata. The settlement of both nudibranchs was presumably promoted by a trophic shift of P. hedgpethi from the natural prey B. neritina towards the largely available and not exploited A. verticillata. This short food web, without evident links with native fauna and having G. quadricolor as the top predator, is described. A DNA barcoding approach was used to confirm the identity of this facelinid species and to explore the possible genetic divergence occurring among the samples analysed.     

The juvenile stages of Parastacus brasiliensis (von Martens, 1869) (Crustacea,Decapoda, Parastacidae)

The family Parastacidae consists of the freshwater crustaceans popularly known as crayfishes or crawfishes. Only the genus Parastacus occurs in Brazil, with P. brasiliensis (von Martens, 1869) endemic to the state of Rio Grande do Sul in southern Brazil. Parastacids have direct development, ecloding as juveniles. In laboratory‐reared specimens, juvenile I stage is similar in morphology to the adult, with no setae on most appendages, absence of uropods and presence of a hook on the dactyl of pereiopods 4 and 5. The juvenile II retains juvenile I characteristics but has more setae, which are more varied in morphology. The juvenile III possesses setae as the adults, fully formed uropods and the hooks on pereiopods 4 and 5 are substituted by an apical rectilinear dactylus.     

Food spectrum analysis of the Asian toad,Duttaphrynus melanostictus (Schneider, 1799) (Anura: Bufonidae), from Timor Island,Wallacea

The Asian toad, Duttaphrynus melanostictus (Schneider, 1799), is widespread throughout tropical Asia and very abundant where it occurs. It was relatively recently introduced to Timor, the second largest island in the biogeographic region called Wallacea. Timor Island shows an exceptionally high level of endemism in a wide range of faunal groups and there are concerns that D. melanostictus may have a negative impact on this diversity, including vertebrates, through direct predation. To evaluate the impact the diet of D. melanostictus might have on the local fauna, gut contents of 83 preserved toad specimens from five habitat types in Timor-Leste, a country occupying the eastern half of Timor Island, were examined. We identified 5581 prey items, comprising the following animal groups: annelids; snails and slugs; spiders and harvestmen; woodlice; millipedes and centipedes; grasshoppers, crickets and earwigs; termites; thrips and true bugs; beetles; ants; hymenopterans other than ants; true flies; butterflies; unidentified insects; and insect larvae. Small eusocial insects (ants and termites) constituted the major part of the diet (61.6% and 23.4%, respectively). No vertebrate prey was recorded. Prey item composition did not differ between habitats. The wide prey spectrum well indicates that D. melanostictus is a generalist invertebrate feeder, as other studies, from regions where this species occurs naturally, have already shown. Although the Asian toad seems to not generally prey on vertebrates, vertebrate species that are morphologically similar to invertebrates in their overall appearance may be consumed. Hence, a negative effect on some taxa (e.g. blindsnakes) may be possible. We also present some limited data on intestinal parasites occurring in D. melanostictus.     

Diet of Enyalius bilineatus (Leiosauridae: Squamata) at a site in southeastern Brazil: effects of phylogeny and prey availability

We studied the diet of Enyalius bilineatus (Leiosauridae: Squamata) at the Inhotim Institute, southeastern Brazil, through induced regurgitation. We obtained 27 individuals using pitfall traps with drift-fence, active search, opportunistic encounters and capture by persons not directly involved in the study. We quantified prey availability at the site using pitfall traps and used electivity indices to identify preferred prey items based on ingested and available prey. Preferred prey included Lepidoptera larvae considering number of prey ingested and Orthoptera considering volume ingested. We obtained the available data on diet of Enyalius species from the literature and compared a phylogenetic distance matrix with a diet dissimilarity matrix, showing that phylogenetically closer species tended to have less dissimilar diets. We used independent contrasts to show that environmental impact did not reduce Enyalius population trophic niches, excluding the effects of phylogeny on niche breadth. Species of Enyalius may adapt to some disturbance in their habitats.     

Microhabitat use and feeding habits of Crossodactylus bokermanni Caramaschi and Sazima, 1985 (Anura,Hylodidae) at a site in south‐eastern Brazil

We studied a population of Crossodactylus bokermanni in a stream at the Reserva Particular do Patrimônio Natural, Santuário do Caraça, south‐eastern Brazil, with respect to microhabitat use and diet. We characterized microhabitats based on substrate, distance to water and exposure level of the occupying individual (exposed or sheltered), and adjacent aquatic microhabitats (evaluated as potential nesting sites based on observed nests), according to water current (absent, slow or fast), substrate and depth. Calling males, silent males, ovigerous females, non‐ovigerous females and juveniles showed spatial niche overlap higher than that expected by chance, with calling males showing the strongest preference for microhabitats close to potential nesting sites (with intermediate depths, fast current and sandy bottom). Only 14 males out of 48 individuals examined for stomach contents had prey in their stomachs. The food items with the highest electivity values were Coleoptera and Diptera, but preys with both high and low mobility were exploited.