The sound of a blood meal: Acoustic ecology of frog‐biting midges (Corethrella) in lowland Pacific Costa Rica

Animal communication systems are often exploited by eavesdropping antagonists, especially predators and ectoparasites. Female frog‐biting midges (Diptera: Corethrellidae) are known to use male anuran advertisement calls to locate their blood hosts, frogs. Here, we use acoustic midge traps broadcasting synthetic and recorded calls to identify those frog call parameters that affect midge attraction. At our study site in Pacific lowland Costa Rica, we found that overall midge attraction was influenced by both spectral and temporal acoustic parameters. Low dominant frequencies (below 1 kHz) and short pulse durations (between 125 and 500 ms) attracted the highest numbers of midges in tests with synthetic sinusoidal pure tones. These preferences partially explained the relative attractiveness of the advertisement calls of ten local frog species. The advertisement calls of the common and widespread Giant Bullfrog, Leptodactylus savagei (Anura: Leptodactylidae), attracted by far the largest number of midges, suggesting that this frog species might be a key resource for frog‐biting midges in Costa Rica. Although we found that calls of different frog species attracted different midge species/morphospecies in statistically different proportions, acoustic niche differentiation among midge species appeared to be moderate, suggesting either a limited degree of host specialization or the use of additional short‐range, that is, chemical, host recognition cues.     

Acoustic Preference of Frog‐Biting Midges (Corethrella spp) Attacking Túngara Frogs in their Natural Habitat

In many animals, males aggregate to produce mating signals that attract conspecific females. These leks, however, also attract eavesdropping predators and parasites lured by the mating signal. This study investigates the acoustic preferences of eavesdroppers attracted to natural choruses in a Neotropical frog, the túngara frog (Engystomops pustulosus). In particular, we examined the responses of frog‐biting midges to natural variation in call properties and signaling rates of males in the chorus. These midges use the mating calls of the frogs to localize them and obtain a blood meal. Although it is known that the midges prefer complex over simple túngara frog calls, it is unclear how these eavesdroppers respond to natural call variation when confronted with multiple males in a chorus. We investigated the acoustic preference of the midges using calling frogs in their natural environment and thus accounted for natural variation in their call properties. We performed field recordings using a sound imaging system to quantify the temporal call properties of males in small choruses. During these recordings, we also collected frog‐biting midges attacking calling males. Our results revealed that, in a given chorus, male frogs calling at higher rates and with higher call complexity attracted a larger number of frog‐biting midges. Call rate was particularly important at increasing the number of midges attracted when males produced calls of lower complexity. Similarly, call complexity increased attractiveness to the midges especially when males produced calls at a low repetition rate. Given that female túngara frogs prefer calls produced at higher repetition rates and higher complexity, this study highlights the challenge faced by signalers when increasing attractiveness of the signal to their intended receivers.     

Does weather affect biting fly abundance in avian nests?

Environmental factors may strongly affect avian‐biting fly interactions in different ways because insects are heterothermic organisms that depend greatly on environmental variables to activate their metabolism and behaviour. We studied the effects of weather on both blackfly (Simuliidae) and biting midge Culicoides (Ceratopogonidae) abundance in nests of three passerine species: blue tits Cyanistes caeruleus, great tits Parus major and pied flycatchers Ficedula hypoleuca, breeding in the same area. We controlled for different host‐related factors (hatching date, brood size and host species). Blackfly abundance was negatively related to minimum temperature. In addition, blackfly and biting midge abundances were negatively affected by wind speed measured at 07.00 h, but blackfly abundance was positively associated to wind speed at 18.00 h. We found higher blackfly and biting midge abundances in nests with larger broods breeding later in the season, and significantly higher biting midge abundance in pied flycatcher nests as compared to tit nests. These results represent, to our knowledge, the first report of both environmental and host‐related effects on haematophagous fly abundance in the nests of wild hole‐nesting birds.     

The influence of environmental management and animal shelters in vector control of Culicoides (Diptera,Ceratopogonidae) in northeastern Brazil

We investigated whether biting midges in peridomestic environments are affected by environmental management practices and the presence of domestic animals. We used CDC light traps to collect midges in 112 residences across 24 locations along tourism routes of Maranhão, Brazil. The collection areas were characterized as follows: i) peridomestic area with domestic animals and without management (dirty); ii) peridomestic with domestic animals and management (clean); iii) peridomestic without animals and with management (clean); iv) peridomestic without animals and without management (dirty). The first two treatments had higher biting midge species richness and abundance, respectively. Generalized linear models indicated a positive correlation between the presence of domestic animals and midge abundance, with an approximate four‐fold increase in Culicoides (Diptera: Ceratopogonidae) abundance in peridomestic areas with animals. The same model showed that domestic animals have no influence on richness. Environmental management does not appear to influence species richness or abundance of biting midges.     

Cues used in host‐seeking behavior by frog‐biting midges (Corethrella spp. Coquillet)

We investigated the role of carbon dioxide and host temperature in host attraction in frog‐biting midges (Corethrella spp). In these midges, females are known to use frog calls to localize their host, but the role of other host‐emitted cues has yet not been investigated. We hypothesized that carbon dioxide acts as a supplemental cue to frog calls. To test this hypothesis, we determined the responses of the midges to carbon dioxide, frog calls, and both cues. A significantly lower number of midges are attracted to carbon dioxide and silent traps than to traps broadcasting frog calls. Adding carbon dioxide to the calls does not increase the attractiveness to the midges. Instead, carbon dioxide can have deterrent effects on frog‐biting midges. Temperature of calling frogs is not a cue potentially available to the midges. Contrary to our hypothesis, there was no supplemental effect of carbon dioxide when presented in conjunction to calls. Midge host‐seeking behavior strongly depends on the mating calls emitted by their anuran host. Overall, non‐acoustic cues such as host body temperature and carbon dioxide are not important in long‐distance host location by frog‐biting midges.     

Determinants of abundance and effects of blood-sucking flying insects in the nest of a hole-nesting bird

Compared to non-flying nest-dwelling ectoparasites, the biology of most species of flying ectoparasites and its potential impact on avian hosts is poorly known and rarely, if ever, reported. In this study we explore for the first time the factors that may affect biting midge (Diptera: Ceratopogonidae) and black fly (Diptera: Simuliidae) abundances in the nest cavity of a bird, the hole-nesting blue tit Cyanistes caeruleus, and report their effects on adults and nestlings during reproduction. The abundance of biting midges was positively associated with nest mass, parental provisioning effort and abundance of blowflies and black flies, while negatively associated with nestling condition. Furthermore, a medication treatment to reduce blood parasitaemias in adult birds revealed that biting midges were more abundant in nests of females whose blood parasitaemias were experimentally reduced. This finding would be in accordance with these insect vectors attacking preferentially uninfected or less infected hosts to increase their own survival. The abundance of black flies in the population was lower than that of biting midges and increased in nests with later hatching dates. No significant effect of black fly abundance on adult or nestling condition was detected. Blood-sucking flying insects may impose specific, particular selection pressures on their hosts and more research is needed to better understand these host–parasite associations.     

Seasonal variation in the composition of fish assemblages in small Amazonian forest streams: evidence for predictable changes

1. The effects of seasonal inundation on the biology of fishes on floodplains of large Amazonian rivers are well studied. However, the small seasonal changes in headwater streams are generally considered to have little effect on fish assemblages. 2. In this study, we analysed seasonal changes in the species composition and abundance of fish in small Amazonian forest streams. We sampled fish with hand and seine nets in headwater streams in a 10 000 ha terra‐firme forest reserve near Manaus, Brazil. Each stream was surveyed at the end of the 2005 dry season, at the beginning of the 2006 rainy season and at the beginning of the 2006 dry season, by means of a standardized sampling effort. 3. The numbers of individuals and species caught were higher in the dry season, but rarefaction analyses indicated that greater species numbers could have been due simply to the larger number of individuals caught. 4. Between the dry and rainy season, the direction of changes in species composition in multivariate space varied among sites, especially for quantitative (abundance) data. However, the observed variation among sites was the less than expected if the directions of change were random. 5. Fish assemblages in the second dry season were more similar to those in the previous dry season than expected if changes in species composition among seasons were random. This indicates that a general seasonal pattern in fish assemblages can be detected, despite the existence of some erratic site‐specific changes. 6. Most of the species that showed large seasonal variations in density occupy temporary ponds during the rainy season, when much of the valley is inundated and pond networks form adjacent to streams. Short‐duration lateral migrations to these ponds may play an important role in the seasonal fish‐assemblage dynamics in Amazonian headwater streams. 7. Our results contrast with previous studies on small Amazonian streams, which have found little seasonal change in fish assemblages, and highlight the importance of the abundance of common species as an indicator of general fish assemblage structure in biological monitoring programmes.     

Seasonality affects the parasitism levels in two fish species in the eastern Amazon region

Fish parasite communities have strong interactions with the environment, the host fish, and the aquatic invertebrate communities. Thus, factors directly involving their different life cycles, such as different host populations and environments, as well as seasonal fluctuations in water levels can cause different responses. The main factors structuring parasite communities may be the hydrodynamic variations, together with seasonal variations in the availability of infectious stages of parasites in the environment. The purpose of this study was therefore to investigate the influence of the rainy and dry seasons on parasite infracommunities in fish species in the Amazon River System of Brazil. Between October 2009 and April 2011, specimens of Colossoma macropomum and Colossoma macropomum × Piaractus brachypomus hybrids (tambatinga) were examined for the presence of parasites. The parasite communities were similar in these two species that both showed aggregate dispersion, although the hybrid specimens were less parasitized. For both hosts, the Brillouin diversity, species richness, evenness and Berger‐Parker dominance indices were similar in the rainy season and dry season, except that the Berger‐Parker dominance index for the C. macropomum × P. brachypomus hybrid was higher during the dry season. Ichthyophthirius multifiliis was the dominant parasite, followed by Piscinoodinium pillulare, in both hosts. However, C. macropomum had a higher prevalence and abundance in the dry season, while in the C. macropomum × P. brachypomus hybrid, only the I. multifiliis prevalence increased. The abundance of monogenean species (Anacanthorus spathulatus, Linguadactyloides brinkmanni, Mymarothecium and Notozothecium janauachensis) was higher during the dry season in the gills of C. macropomum. Despite overdispersion of the lernaeid Perulernaea gamitanae, a seasonal pattern was seen only in the C. macropomum × P. brachypomus hybrid, with higher prevalence in the rainy season. The low prevalence and abundance of the ectoparasites Tetrahymena sp., Trichodina sp. and Braga patagonica, as well as of the endoparasites Procamallalus (Spirocamallanus) inopinatus and Neoechinorhynchus buttnerae, were not affected by seasonality. This information is applicable in farming practice for these economically important fish, as it indicates the best time for prophylactic management and treatment against parasites, in order to prevent economic losses in fish farms.     

Effect of vertebrate feces on the attraction of Culicoides (Diptera: Ceratopogonidae) in an Amazonian mangrove

Biting midges are of interest to public health because they play an important role as vectors of disease‐causing pathogens, as well as being a biting nuisance to humans and domestic animals. Although these insects are common in mangrove areas, they have not yet been studied in this ecosystem in the state of Maranhão (MA), Brazil. The objective of this study was to characterize the Culicoides community structure found in a mangrove swamp and verify if the use of vertebrate feces as bait interferes with their composition, richness, abundance, or seasonality. CDC light traps with vertebrate (chicken, ox, control, donkey, capuchin monkey and pig) feces baits were used to capture biting midges in a mangrove area of the Island of São Luís, MA. A total of 4,087 individuals representing 22 species of Culicoides were captured, the most abundant being C. (Oecacta) furens Poey (23.46%), C. (gr. Fluviatilis) leopoldoi Ortiz (21.58%), C. (Hoffmania) ignacioi Forattini (16.98%), C. (Hoffmania) maruim Lutz (13.85%), C. (Diphaomyia) iriartei Fox (10.57%), C. (Hoffmania) insignis Lutz (7.07%), and C. (gr. Limai) limai Barreto (3.03%). Species richness and abundance were higher when baits of capuchin monkey (15 species; 26.84% of the individuals) and pig (15; 25.3%) feces were used. The least attractive baits were donkey (12 species; 9.3%) and ox (nine species; 11.52%) feces. Biting midges were more abundant in the rainy season (67%), but richness was higher in the dry season (19 species). These results show that vertebrate feces may serve as olfactory cues and increase the attraction of biting midges to traps.     

垂叶榕榕小蜂群落及种间互作网络季节动态

群落中的物种相互作用构成了复杂的生态网络。有关物种的数量和组成的季节性动态变化已有较多的研究, 但是对于生态网络的动态变化知之甚少。揭示生态网络的动态变化对于理解群落的稳定性以及群落的动态变化过程和机理具有重要意义。本研究以垂叶榕(Ficus benjamina)榕小蜂群落为研究对象, 分别在西双版纳的干季和雨季采集了榕小蜂的种类和数量信息。比较了两个季节榕小蜂群落的动态变化以及共存网络的参数(例如网路直径、连接数、嵌套性和群落矩阵温度)变化。结果显示: 雨季榕果内传粉榕小蜂Eupristina koningsbergeri所占比例高于干季, 传粉榕小蜂的种群数量也高于干季, 而在干季非传粉榕小蜂的种类增加(干季15种小蜂, 雨季14种)。从榕树-传粉榕小蜂互利共生系统的适合度来看, 干季非传粉小蜂的增加对传粉榕小蜂和榕树的适合度是不利的。在干季, 共存网络物种间的连接数(干季0.95, 雨季0.47)多于雨季, 群落矩阵温度(干季23.24, 雨季2.64)也显著高于雨季。表明干季榕小蜂群落组成及种间关系较雨季更为复杂而多样, 高的矩阵温度暗示群落受到的干扰更大。     

Feeding Site Selection by Frog-Biting Midges (Diptera: Corethrellidae) on Anuran Hosts

A critical challenge of obtaining a blood meal resides in selecting a feeding site at the host. We investigate the feeding sites of species of frog-biting midges (Corethrella spp) on túngara frogs, Engystomops pustulosus (Cope), and two species of treefrogs, Dendropsophus ebraccatus (Cope) and Dendropsophus microcephalus (Cope). To investigate the differences in midge feeding sites, we performed field observations, histological sections of the frogs and examined the mouthparts of the midges. Feeding sites are predicted by the vascular properties of the skin in different body areas of the frogs. According to optimal foraging theory, the midges should feed on host body areas that maximize blood intake. Contrary to optimal foraging theory predictions, however, their feeding sites correspond to body areas with high intensity host defensive behavior.     

Seasonal variation in diet breadth of folivorous Lepidoptera in the Brazilian cerrado

Documented patterns of specialization and species interactions often omit plasticity in resource use across space and time, yet such variation is an important part of species interactions. To examine temporal variation in resource use, we compared species‐ and community‐level patterns of host plant use by folivorous caterpillars between the dry and rainy seasons in four preserved areas of cerrado vegetation in the Distrito Federal, Brazil. We sampled plants and caterpillars in 10 m circular plots monthly from March 2010 to March 2011. At the community level, we found a significant increase in the mean diet breadth of dry season caterpillars relative to those collected in the rainy season. Families and species of moths varied in diet breadth, but most exhibited a seasonal expansion. In particular, intraspecific comparisons showed a 30% increase in the number of host plants used in the dry season compared to the rainy season. These results provide a clear example of how temporal variation in resource use is heterogeneous, and more generalized patterns of resource use can emerge from studies at large temporal scales. Thus, seasonal or annual heterogeneity may obscure ecologically relevant specialized interactions that occur at smaller scales.     

Parasite fauna of seabass (Lates calcarifer) under mariculture conditions in Lampung Bay, Indonesia

Fish parasites have been repeatedly reported to be a major threat to the developing industry of finfish mariculture in Indonesia, due to severe parasite and disease outbreaks. The aim of this study was to identify the metazoan parasite fauna and trichodinid ciliates that infect Lates calcarifer in a representative mariculture farm in Indonesia. Examined were 105 L. calcarifer (seabass) for the metazoan parasite fauna and trichodinid ciliates. Thirty‐five specimens each from the net cages of the National Sea Farming Development Centre (Balai Budidaya Laut, BBL) in Lampung Bay, South Sumatra, Indonesia were investigated in three consecutive seasons (two dry and one rainy season from 2002 to 2003). Nineteen parasite species were identified; all fish specimens were infected with two to 10 parasite species, demonstrating a species‐rich parasite fauna. Protozoans (1 species), myxozoans (1), digeneans (3), monogeneans (5), cestodes (3), nematodes (5) and acanthocephalans (1) were found, including 11 new host records in cultured L. calcarifer from Indonesia. Larval and adult parasite stages were isolated, demonstrating that this fish species, although kept inside the net cages, still functions as an intermediate and final host for marine fish parasites. During all seasons, the six detected monoxenous (single host life cycle) parasite species showed a higher prevalence than the 13 heteroxenous (multiple hosts) species. Most abundant were the fish pathogenic monogeneans Pseudorhabdosynochus epinepheli, Pseudorhabdosynochus lantauensis, Benedenia epinepheli and Neobenedenia melleni with a high prevalence. Most heteroxenous parasites (Digenea, Cestoda, Nematoda and Acanthocephala) occurred with a low prevalence below 26%, caused by the specific culture conditions. Diversity of the heteroxenous parasites was higher in the dry seasons than in the rainy season. Though some seasonality could be observed for the fish pathogenic monogeneans, severe disease outbreaks of these ectoparasites cannot be excluded in either the dry or rainy season.     

Species assemblages of cyclopoid Copepoda from Mantecal,Venezuela

A 14 year monitoring of species composition and abundance of cyclopoid copepods throughout the rainy and dry seasons in a flooded savanna at Mantecal, Apure State, Venezuela, is presented. Two characteristic habitats within these water bodies were selected: the open center of small ponds free of emergent vegetation and the surrounding flooded grassland. In total, 13 species were registered, all of which appeared in the flooded grassland and 11 of which occurred in the open water. Ten species were common for the two habitats during the dry season. The highest abundance was found in flooded grassland during the dry season. Eight species showed abundances greater than 1.0 individual per liter in flooded grassland during both seasons; 3 and 7 species were this abundant in the open water during rainy and dry seasons, respectively. Mesocyclops meridianus and Microcyclops varicans were the most abundant species in both habitats and seasons, while Thermocyclops decipiens was the dominant species during rainy season in open water. Principal component analysis indicates that the important species were positively correlated between them (sharing habitat and season).     

Phytoplankton community structure and environment in the Kenyan waters of Lake Victoria

• 1 Phytoplankton species composition, numerical abundance, spatial distribution and total biomass measured as chlorophyll a concentration were studied in relation to environmental factors in September 1994 (dry season) and March 1995 (rainy season), respectively, in the Kenyan waters of Lake Victoria; 103 species were recorded.
• 2 Blue‐green algae (Cyanophyceae) were most diverse, followed by diatoms (Bacillariophyceae), green algae (Chlorophyceae) and dinoflagellates (Dinophyceae).
• 3 Twinspan separated the phytoplankton communities in the Nyanza Gulf and those in the open lake during both seasons. During the dry season, the Nyanza Gulf was strongly dominated by blue‐greens, while diatoms dominated in the open lake. During the rainy season, blue‐greens remained dominant in the Nyanza Gulf although the number of species found was lower than during the dry season; in the open lake, blue‐greens replaced diatoms as the dominant group and there were more species than in the dry season.
• 4 Canonical correspondence analysis indicated that the phytoplankton species distribution was significantly correlated with turbidity during the dry season and with SiO₂ during the rainy season. Chlorophyll a concentrations ranging from 2.0 to 71.5 mg m^‐3 in the dry season and 2.0–17.2 mg m^‐3 in the rainy season confirm earlier reports of increasing phytoplankton biomass in Lake Victoria since the 1960s.

     

Diversity of parasites in Cichlasoma amazonarum Kullander, 1983 during rainy and dry seasons in eastern Amazon (Brazil)

This study investigated the influence of the rainy and dry seasons on the parasite communities of Cichlasoma amazonarum (Kullander, 1983) in a tributary of the Amazon River system, northern Brazil. Of 112 fish examined, 95.5% were parasitized by Ichthyophthirius multifiliis, Piscinoodinium pillulare, Gussevia disparoides, Posthodiplostomum sp., Procamallanus (Spirocamallanus) inopinatus, and Echinorhynchus paranensis, as well as by the Glossiphoniidae leech. Ichthyophthirius multifiliis and P. pillulare were the dominant parasite species, with I. multifiliis the most prevalent and abundant; the leech (Glossiphoniidae) was the least prevalent and least abundant. Parasites presented an aggregate dispersion pattern and seasonal variations in infestation levels, influenced by the environmental conditions in the rainy season. Variation in the parasite dynamics created overall changes in the parasite communities, characterized by greater diversity, species richness and evenness during the rainy season. However, there was a high similarity (99.8%) in the parasite community structure between the rainy and dry seasons. During the dry season the hosts had predominantly 1–3 parasite species compared to 3–4 parasites in the rainy season. Some parasites in the eastern Amazon undergo population changes relating to seasonality. These results thus indicate the correct season to apply adequate prophylactic measures to reduce negative impacts of parasites in this wild ornamental fish when captured.     

Molecular identification of bloodmeals and species composition in Culicoides biting midges

Investigations of host preferences in haematophagous insects, including Culicoides biting midges (Diptera: Ceratopogonidae), are critical in order to assess transmission routes of vector‐borne diseases. In this study, we collected and morphologically identified 164 blood‐engorged Culicoides females caught in both light traps and permanent 12‐m high suction traps during 2008–2010 in Sweden. Molecular analysis of the mitochondrial cytochrome c oxidase subunit I (COI) gene in the biting midges was performed to verify species classification, discern phylogenetic relationships and uncover possible cryptic species. Bloodmeal analysis using universal vertebrate cytochrome b primers revealed a clear distinction in host selection between mammalophilic and ornithophilic Culicoides species. Host sequences found matches in horse (n = 59), sheep (n = 39), cattle (n = 26), Eurasian elk (n = 1) and 10 different bird species (n = 18). We identified 15 Culicoides species previously recorded in Scandinavia and four additional species haplotypes that were distinctly different from the described species. All ornithophilic individuals (n = 23) were caught exclusively in the suction traps, as were, interestingly, almost all mammalophilic species (n = 41), indicating that many biting midge species may be able to cover long distances after completing a bloodmeal. These results add new information on the composition of Culicoides species and their host preferences and their potential long‐distance dispersal while blood‐engorged.     

Temporal shifts and niche overlapping in <Emphasis Type="Italic">Copestylum</Emphasis> (Diptera,Syrphidae) communities reared in cactus species in a central Mexican scrubland

Semiarid scrubland communities are highly dynamic in terms of their species composition, abundance, and functioning, given the drastic changes in climate among seasons. Spatiotemporal patterns of saprophagous Copestylum (Diptera: Syrphidae) communities in different cactus species richness have not yet been studied, although seasonal changes and plant species richness have been shown to strongly impact the diversity and distribution of many insect communities in scrublands. We analyzed the impact of seasonality and of habitat type (disturbed and undisturbed) on Copestylum communities reared from cactus species at the Barranca de Metztitlán Biosphere Reserve, in central Mexico, by comparing their community structure between seasons and habitats, and assessing the contribution of diversity components for the total diversity of this genus. We also measured patterns of temporal niche overlap among hoverfly species considering their breeding medium. Seasonal variation influenced Copestylum community composition most significantly. Species richness and abundance of Copestylum were higher in the rainy season. Additive partitioning of diversity showed that the main component for species richness is beta diversity between seasons. We detected high niche overlap during the dry season and low overlap during the rainy season. This study provides evidence of temporal shifts in xeric hoverfly communities and suggests that the Copestylum species partition resources over time.     

Haemonchosis in sheep and goats in Sierra Leone

The incidence of haemonchosis in sheep and goats in Sierra Leone showed a seasonal variation with a high peak in the dry season (October to January) and a low one from March to May. Mean relative densities were significantly higher in young hosts and showed two peaks, a high one from August to December and a low one from April to June while the mean relative densities of old hosts were low and exhibited an irregular seasonal pattern with no defined peaks. The peak seasons in young hosts coincided approximately with the dry and rainy seasons. Male hosts showed an overall higher but not a significant mean relative density than females but for most of the months mean relative densities of infection of female hosts were not significantly higher than those of males. The roles of climatic conditions and arrested development (hypobiosis) and self-cure and host age immunity on Haemonchus contortus worm burdens are discussed.