Environmental influences on the replenishment of lizardfish (family Synodontidae) in Caribbean Panama

Lizardfish (family Synodontidae) are little studied despite their potentially important predatory role in epibenthic coral reef communities. The present study documented the temporal and spatial larval supply patterns of five Caribbean lizardfish species together with environmental variables (solar radiation, rainfall, water temperature, onshore-offshore winds, alongshore winds and wind-induced turbulence) to examine: (1) whether species within the same family respond differently to their developmental environment and (2) if larval supply in year-round warm waters is influenced by climatic changes. To address these questions, late-stage larvae of Synodus foetens, Synodus intermedius, Synodus poeyi, Saurida suspicio and Saurida brasiliensis were collected in replicate light traps in three different reef habitats (back-reef, lagoon and exposed) in the San Blas Archipelago, Panama, over 18 consecutive lunar months. Although replenishment of lizardfish occurred year-round, the temporal and spatial supply patterns were species-specific: S. foetens, S. intermedius and S. poeyi were most abundant during the dry season while S. suspicio and S. brasiliensis were most prevalent during the wet season. When seasons were analysed separately, water temperature explained 39 and 26% of the variance in light trap catches of S. foetens and S. intermedius, respectively, in the dry season while wind-induced turbulence accounted for 25% of the variability in S. suspicio and S. brasiliensis catches during the wet season. These findings stress the importance of analysing larval supply in conjunction with environmental data at a high taxonomic resolution to better understand the mechanisms that drive replenishment in reef systems at low latitudes.     

A light trap design for stratum-specific sampling of reef fish larvae

The vertical distribution of late stage reef fish larvae may potentially influence their dispersal, recruitment success and energetic expenditure during the recruitment process. To date, methods of examining the vertical distribution of reef fish larvae either under-sample late stage individuals, or are incapable of discretely sampling the water column. The aim of this study was to develop a light trap able to sample a narrow depth range enabling fine scale patterns of vertical distribution to be examined. The experimental traps radiated light in a relatively narrow beam with a maximum vertical angle of radiation of 7.5°, indicating that the traps could be placed 4.8 m apart and still sample discrete depth strata. Their catch efficiency was similar to conventional light traps, indicating that they are adequate sampling units. Preliminary data showed that most families are more abundant near the surface, although many have significant numbers of individuals lower in the water column. Some families (inc. Apogonidae) occurred in higher abundance at greater depths. Our experimental light traps permit increased resolution of the vertical distribution of late stage larval reef fishes in the field.     

Identifying the ichthyoplankton of a coral reef using DNA barcodes

Marine fishes exhibit spectacular phenotypic changes during their ontogeny, and the identification of their early stages is challenging due to the paucity of diagnostic morphological characters at the species level. Meanwhile, the importance of early life stages in dispersal and connectivity has recently experienced an increasing interest in conservation programmes for coral reef fishes. This study aims at assessing the effectiveness of DNA barcoding for the automated identification of coral reef fish larvae through large‐scale ecosystemic sampling. Fish larvae were mainly collected using bongo nets and light traps around Moorea between September 2008 and August 2010 in 10 sites distributed in open waters. Fish larvae ranged from 2 to 100 mm of total length, with the most abundant individuals being <5 mm. Among the 505 individuals DNA barcoded, 373 larvae (i.e. 75%) were identified to the species level. A total of 106 species were detected, among which 11 corresponded to pelagic and bathypelagic species, while 95 corresponded to species observed at the adult stage on neighbouring reefs. This study highlights the benefits and pitfalls of using standardized molecular systems for species identification and illustrates the new possibilities enabled by DNA barcoding for future work on coral reef fish larval ecology.     

High species richness of early stages of fish in a locality of the Mesoamerican Barrier Reef System: a small-scale survey using different sampling gears

The Mesoamerican Barrier Reef System (MBRS) contains a diverse array of coastal habitats that are critical for the survival of the early stages of reef fish; however, the knowledge on the abundance and distribution of the early stages of coastal fishes is still limited in this region. This study investigated the species richness of larval and juvenile fishes using a combination of a sled net, standard plankton net, and a nightlight lift-net; these were deployed simultaneously at Bacalar Chico, a site on the MBRS within the protected “Parque Nacional Arrecifes de Xcalak” (PNAX). We collected 53 families and 118 species of larval and juvenile fishes in a small area of about 3 km². This species diversity of early life-history stages is greater than previously found in surveys on the MBRS. Each gear caught a number of species exclusively, so combined sampling with the three methods provided a much fuller picture of the local larval and juvenile fish assemblage. A species-accumulation model estimated that the samples likely represented 84% of the total assemblage. Many species caught were represented predominantly by newly settled juveniles, underscoring the importance of this coastal habitat for settlement of many ecologically and economically important fish species. This information is expected to improve the conservation and management strategies in the fragile PNAX coastal zones by providing additional information based on original field data to raise awareness among managers about the ecological relevance of these coastal habitats. This study provides encouraging evidence that the PNAX is a well-suited natural protected area to preserve a critical fish habitat in a hotspot of marine biodiversity.     

Jetski-based plankton towing as a new method of sampling larval fishes in shallow marine habitats

I tested jetski-based plankton towing as a new method of sampling larval fishes in a shallow marine environment in warm temperate South Africa. Larval fishes were collected by towing a stainless-steel ring net behind a three-man jetski. The sampling technique was designed and tested to overcome problematic sampling of shallow sandy beach surf zones and the adjacent, shallow nearshore immediately seaward of the surf zone (behind the breaker line) with the same type of equipment. The composition, density, stage of development, estuary association, and length frequency of larval fish catches obtained by the new sampling method in the surf zone and in the adjacent shallow nearshore were compared. The success of the technique was evaluated by determining the extent of replication of known trends observed in other surf studies in the same climatic region of South Africa using either a seine or pushnet. Preflexion and postflexion larval fishes were collected. Larvae of the family Sparidae dominated the catches. Trends in catch composition, estuary association, and length frequency of the larvae were similar to those found in other studies of warm temperate South African surf zones. Density of larvae varied depending on the type of equipment used, although these results are tentative given the subjectivity of calculation of the water volume sampled and the extent of nursery use by larvae in the habitats sampled in each study. Increased towing times are recommended. Jetski-based plankton towing is a useful alternative technique for sampling larval fishes in shallow habitats and can easily be applied in surf zones, shallow nearshore areas, and estuaries. The technique greatly increases sampling agility in previously problematic sampling areas.     

Occupancy Models for Monitoring Marine Fish: A Bayesian Hierarchical Approach to Model Imperfect Detection with a Novel Gear Combination

Occupancy models using incidence data collected repeatedly at sites across the range of a population are increasingly employed to infer patterns and processes influencing population distribution and dynamics. While such work is common in terrestrial systems, fewer examples exist in marine applications. This disparity likely exists because the replicate samples required by these models to account for imperfect detection are often impractical to obtain when surveying aquatic organisms, particularly fishes. We employ simultaneous sampling using fish traps and novel underwater camera observations to generate the requisite replicate samples for occupancy models of red snapper, a reef fish species. Since the replicate samples are collected simultaneously by multiple sampling devices, many typical problems encountered when obtaining replicate observations are avoided. Our results suggest that augmenting traditional fish trap sampling with camera observations not only doubled the probability of detecting red snapper in reef habitats off the Southeast coast of the United States, but supplied the necessary observations to infer factors influencing population distribution and abundance while accounting for imperfect detection. We found that detection probabilities tended to be higher for camera traps than traditional fish traps. Furthermore, camera trap detections were influenced by the current direction and turbidity of the water, indicating that collecting data on these variables is important for future monitoring. These models indicate that the distribution and abundance of this species is more heavily influenced by latitude and depth than by micro-scale reef characteristics lending credence to previous characterizations of red snapper as a reef habitat generalist. This study demonstrates the utility of simultaneous sampling devices, including camera traps, in aquatic environments to inform occupancy models and account for imperfect detection when describing factors influencing fish population distribution and dynamics.     

Comparison between anopheline mosquitoes (Diptera: Culicidae) caught using different methods in a malaria endemic area of Papua New Guinea

The mosquito sampling efficiency of CDC (Centers for Disease Control) miniature light traps hung adjacent to mosquito nets, was compared with that of both indoor and outdoor human-bait collections in ten villages in the Wosera area of Papua New Guinea. The most frequently collected anopheline in the matched indoor and light trap samples was Anopheles koliensis Owen, followed by A. punctulatus D?nitz, A. karwari (James), A. farauti Laveran (sensu lato), A. longirostris Brug and A. bancroftii Giles. All species were much less frequent in the light traps than in landing catches. The hypothesis that the numbers of mosquitoes in light traps are proportional to human landing catches was examined using regression models that allowed for sampling error in both entomological measurements. Light traps under-sampled A. punctulatus and A. farauti s.l. at high densities. The models indicated that the ratio of light trap to landing catch females of A. koliensis and A. karwari increased with increasing mosquito density. Light trap catches of A. longirostris were proportional to indoor landing rates but when outdoor landing rates were high this species was under-sampled by light traps. Numbers of A. bancroftii in light traps were found to be proportional to those in outdoor landing catches, but were negatively related to those attempting to bite indoors. Circumsporozoite positivity rates for both Plasmodium falciparum Welch and P. vivax (Grassi & Feletti) in A. punctulatus and A. farauti s.l. were significantly higher in light trap collections than in either indoor or outdoor landing catches, suggesting that light traps may selectively sample older mosquitoes of these species.     

Ecology of the introduced snapper Lutjanus kasmiva (Forsskal) in the reef fish assemblage of a Hawaiian bay

Since its first introduction in 1955, the snapper Lutjanus kasmira (taape) has developed large populations in shallow coastal waters of Hawaii. Visual abundance estimates and trap catches of taape were different between the main habitat types of the reef. Overall, taape was the second most abundant species by numbers and biomass over a hard substratum and was patchily distributed, even within habitat type. The largest individuals occurred singly or in small groups on shallow reef slopes; on the deep slopes and in spur-and-groove habitat, taape occurred in larger groups of smaller sized fish. Catches in fish traps showed a negative relationship with the complexity of the adjacent natural substratum, suggesting that traps are more productive where they offer additional cover in areas with less natural cover. In all areas in which traps were used, from fringing reef to open sand, the catch was dominated by taape. Tagging confirmed the theory that taape tend to maintain a limited range over long periods. Tagging results also indicated occasional long-range movements among major habitats and provided data confirming patterns of movement between reef habitat by day and sand habitat by night. Taape collected over open sand substratum by trap and by hook-and-line were smaller than those collected by the same means at the reef. Observations and collections suggested an ontogenetic trend in habitat use by taape from initial settlement in fringe areas toward ultimate residence on the main reef. Other common fish species in the reef assemblages were tested for statistical associations with taape. Although several positive species associations were found, multivariate analysis did not reveal patterns that indicated strong ecological relationships.     

Diversity of settlement-stage reef fishes captured by light-trap in a tropical south-west Atlantic Ocean coastal reef system

This study reports the results of 5 years of monitoring reef fish post-larvae using light traps in the Bay of Tamandaré, north-east Brazil. An annotated checklist of pre-settlement fish species, their frequency of occurrence and taxonomic characteristics are provided. In total, 4,422 post-larval fishes belonging to 36 families, 56 genera and 76 species were captured. The most species-rich families were Carangidae (7), Lutjanidae (6) and Pomacentridae (4), while the families Gerreidae (30.47%), Holocentridae (16.54%), Blenniidae (12.01%), Labrisomidae (8.36%), Lutjanidae (8.29%) and Acanthuridae (5.95%) were the most abundant. This is the first study of the taxonomic diversity and assemblage structure of settlement-stage reef fishes in the tropical south-west Atlantic Ocean. Although a few common species were not captured due to selectivity of light traps, the composition and taxonomic diversity of this first collection suggests that light traps are useful for studies of the early life history of a wide range of pre-settlement reef fishes.     

Coral-reef sounds enable nocturnal navigation by some reef-fish larvae in some places and at some times

At Lizard Island, Great Barrier Reef, catches of fish larvae by light traps that broadcast nocturnal reef sounds (noisy traps) were compared with catches by quiet traps over two 2·5 week new-moon periods in November (XI) 2000 and January (I) 2001. Three areas were sampled: near-reef (NR, 500 m from the shore) in I, middle (M, 650 m) in I and XI and offshore (O, >1000 m) in XI. The most abundant taxa captured were Apogonidae, Blenniidae, Chaetodontidae, Lethrinidae, Mullidae and Pomacentridae. Significant differences in catch were found between areas, and a position effect was found at the O and M areas. At the NR and M areas, no taxa had significantly greater catches in quiet traps, but larvae of five taxa had significantly greater catches in noisy traps. These were (areas and times of greater catches): Apogonidae (NR; M XI), Mullidae (M I & XI), Pomacentridae (NR; M I & XI), Serranidae (M I) and Sphyraenidae (NR). At the offshore area, five taxa (Apogonidae, Blenniidae, Chaetodontidae, Mullidae and Pomacentridae) had significantly greater catches in quiet traps and only Lethrinidae had significantly greater catches in noisy traps. Thus some taxa (particularly apogonids and pomacentrids which had catches up to 155% greater in noisy traps, but also lethrinids and mullids, and perhaps others), were attracted to reef sounds at night, but this apparently varied with location and time. The sound-enhanced catches imply a radius of attraction of the sound 1·02–1·6 times that of the light. More than 65 m from the speaker,the broadcast sound levels at frequencies typical of fish hearing were equivalent to background levels, providing a maximum radius of sound attraction in this experiment.     

Spatio-temporal variability in fish recruitment to a coral reef (Moorea,French Polynesia)

Spatial and temporal patterns of recruitment of juvenile coral reef fishes were studied on the reefs of the high island of Moorea (Society Archipelago, French Polynesia) during the wet season (October 1988 to April 1989). Two size-classes of fishes (new-recruits and juveniles) were censused by visual counts within 24 quadrats along a transect across the lagoon. Correspondence analysis was used to calculate the hypothetical movements of the two size classes. Spatial variability was far more important than temporal variability; six main spatial communities were revealed, which could result from differences among habitats. Temporal variability in recuitment occurred only at nearshore stations; stations on the outer fringing reef and inner barrier reef displayed stable recruitment patterns.     

Early life history and settlement of the slender filefish, <Emphasis Type="Italic">Monacanthus tuckeri</Emphasis> (Monacanthidae), at Calabash Caye,Turneffe Atoll,Belize

Synopsis We examined early life history traits and patterns of settlement of the slender filefish, Monacanthus tuckeri, at Calabash Caye, Turneffe Atoll, Belize. A settlement peak was evident at the new moon, and no settlement occurred at the full moon. However, settlement rates at the quarter moons could not be estimated due to sampling gaps. Many reef fishes show new moon settlement peaks, so M. tuckeri shares some characteristics with the primarily perciform species on coral reefs. Pelagic larval duration was long (mean = 42 days) and variable, suggesting that dispersal patterns might be diverse. Size at settlement was large (mean = 32 mm total length) and also variable. Larval duration and size at settlement were outside of the average values exhibited by reef fishes, but are not beyond the extreme end of the range, and might be explained by association with pelagic debris prior to settlement. There were no differences in overall settlement rates on reef and seagrass habitats, and fish settling to either habitat did not differ in larval duration, size at settlement, or larval growth rate. This suggests that settlement to alternative habitats may be random, or driven by availability of suitable microhabitat, rather than habitat quality or individual traits.     

Eukaryotic plankton communities across reef environments in Bocas del Toro Archipelago,Panamá

Variation in light and temperature can influence the genetic diversity and structure of marine plankton communities. While open-ocean plankton communities receive much scientific attention, little is known about how environmental variation affects plankton communities on tropical coral reefs. Here, we characterize eukaryotic plankton communities on coral reefs across the Bocas del Toro Archipelago, Panamá. Temperature loggers were deployed, and midday light levels were measured to quantify environmental differences across reefs at four inshore and four offshore sites (Inshore = Punta Donato, Smithsonian Tropical Research Institute (STRI) Point, Cristobal, Punta Laurel and Offshore = Drago Mar, Bastimentos North, Bastimentos South, and Cayo de Agua). Triplicate vertical plankton tows were collected midday, and high-throughput 18S ribosomal DNA metabarcoding was leveraged to investigate the relationship between eukaryotic plankton community structure and inshore/offshore reef environments. Plankton communities from STRI Point were additionally characterized in the morning (~ 08:00), midday (~ 12:00), and late-day (~ 16:00) to quantify temporal variation within a single site. We found that inshore reefs experienced higher average seawater temperatures, while offshore sites offered higher light levels, presumably associated with reduced water turbidity on reefs further from shore. These significant environmental differences between inshore and offshore reefs corresponded with overall plankton community differences. We also found that temporal variation played a structuring role within these plankton communities, and conclude that time of community sampling is an important consideration for future studies. Follow-up studies focusing on more intensive sampling efforts across space and time, coupled with techniques that can detect more subtle genetic differences between and within communities will more fully capture plankton dynamics in this region and beyond.

     

The biogeography of tropical reef fishes: endemism and provinciality through time

The largest marine biodiversity hotspot straddles the Indian and Pacific Oceans, driven by taxa associated with tropical coral reefs. Centred on the Indo‐Australian Archipelago (IAA), this biodiversity hotspot forms the ‘bullseye’ of a steep gradient in species richness from this centre to the periphery of the vast Indo‐Pacific region. Complex patterns of endemism, wide‐ranging species and assemblage differences have obscured our understanding of the genesis of this biodiversity pattern and its maintenance across two‐thirds of the world's oceans. But time‐calibrated molecular phylogenies coupled with ancestral biogeographic estimates have provided a valuable framework in which to examine the origins of coral reef fish biodiversity across the tropics. Herein, we examine phylogenetic and biogeographic data for coral reef fishes to highlight temporal patterns of marine endemism and tropical provinciality. The ages and distribution of endemic lineages have often been used to identify areas of species creation and demise in the marine tropics and discriminate among multiple hypotheses regarding the origins of biodiversity in the IAA. Despite a general under‐sampling of endemic fishes in phylogenetic studies, the majority of locations today contain a mixture of potential paleo‐ and neo‐endemic fishes, pointing to multiple historical processes involved in the origin and maintenance of the IAA biodiversity hotspot. Increased precision and sampling of geographic ranges for reef fishes has permitted the division of discrete realms, regions and provinces across the tropics. Yet, such metrics are only beginning to integrate phylogenetic relatedness and ancestral biogeography. Here, we integrate phylogenetic diversity with ancestral biogeographic estimation of lineages to show how assemblage structure and tropical provinciality has changed through time.     

Spatial relationships between two Agriotes click-beetle species and wireworms in agricultural fields

1 The production of new insect pheromones for pest monitoring proceeds at a greater rate than their evaluation, with the consequential possibility of premature introduction. Fundamental to their successful deployment is the determination of a consistent relationship between adult male pheromone trap catches and pest damage. In the present study, adult pheromone traps and larval bait traps were used to examine spatial relationships between two species of Agriotes beetle and wireworms at the field scale. 2 The spatial distributions of adult male Agriotes lineatus and Agriotes obscurus in two fields were determined and compared with the distribution of their larvae. Data were assembled as spatially referenced trap counts, and analysed for evidence of aggregation and clustering using Spatial Analysis by Distance IndicEs (SADIE) methodology. Spatial stabilities of adult populations between sampling dates were tested using association tests. Spatial and quantitative linkages between adult and larval trap catches were also tested. Moreover, a new way of adapting SADIE methodologies is presented for situations where two datasets within an area do not share the same sampling points. 3 There was no significant difference in variance : mean relationships for the two species but there were differences in their spatial distributions, and this is a definitive example of the general argument stating that it is important to consider spatial as well as count data in ecological studies. The spatial distribution of A. lineatus varied between sampling occasions at both sites whereas A. obscurus had consistently significant SADIE indices over time at one site, and adult catches could also be linked to larval distributions and counts. It is proposed that observed differences between the two species can be explained by interference between traps and dissimilar movement rates. There was some evidence of an edge effect at the field boundaries. 4 The distance between pheromone traps is related to the time that elapses before adjacent traps interfere with trap captures and this limits the detection of statistically significant spatial patterns. It is shown that the current practice of adding trap counts for different Agriotes species and treating them as numerically equivalent is insufficiently robust to be recommended at this stage. 5 The implications for the use of sex pheromone traps in wireworm pest management are considered. It is concluded that pheromone traps, as currently used, will not reliably indicate where wireworms occur in a field, and that the complexity of interpreting adult male trap counts limits quantitative predictions of population size.     

Can differences in the structure of larval,juvenile and adult coral‐reef fish assemblages be detected at the family level?

Processes occurring at the end of the larval stage are of major importance in shaping spatial structure of fish assemblages in coral reefs. However, because of the difficulty in identifying larvae to species, many studies dealing with these stages are limited to the family level. It remains unknown if variation in the spatial structure of coral‐reef fish assemblages across life stages can be detected at such a coarse taxonomic level. Two different surveys conducted in a similar area of New Caledonia, Southwest Pacific, provided the opportunity to compare the structure of coral‐reef fish assemblages collected as pre‐settlement larvae, juveniles and adults along a coast to barrier reef gradient. Adult and juvenile fish were sampled using underwater visual counts (UVC) during the warm seasons of 2004 and 2005. Pre‐settlement larvae were sampled with light‐traps during the same seasons. In order to standardize data between sampling methods, analyses were conducted on the relative abundance (for larvae) and density (for juveniles and adults) of 21 families commonly collected with both methods. Relative abundances/densities of families were analysed as a function of life stage (larvae, juveniles or adults), large‐scale spatial location (coast, lagoon or barrier) and years (2004, 2005) using non‐parametric multidimensional scaling (nMDS) and permutational multivariate analysis of variance (permanova ). Kruskal–Wallis tests were then used to examine differences among life stages and locations for individual families. Different levels of spatial and temporal variability characterized fish assemblages from different life stages, and differences among life stages were detected at all locations and years. Differences among life stages were also significant at the level of individual families. Overall results indicate that studies conducted at the family level may efficiently reveal changes in coral‐reef fish spatial structure among successive life stages when large spatial scales are considered.     

浙北近岸典型岛礁瓦氏马尾藻空间分布格局分析

为探明浙北近岸典型岛礁瓦氏马尾藻(Sargassum vachellianum)的空间分布格局, 于2016年5月底至6月初, 分别采用走航观察和水下样方法, 对按离岸距离远近的3条岛礁带上12个岛礁瓦氏马尾藻成藻时期的分布特征进行了调查, 并从不同空间尺度比较分析了3条岛礁带上瓦氏马尾藻在水平和垂直分布上的差异。结果表明: (1)在地区尺度上, 高浊度和高波浪能的水域环境限制了瓦氏马尾藻的分布与生长, 导致浙北近岸的瓦氏马尾藻仅分布在第二条岛礁带这一狭窄的分布带上。依据瓦氏马尾藻最低适宜生长水温要高于10 ℃的特性, 我们可以推断出舟山群岛的绿华岛可能是我国特有种瓦氏马尾藻分布的最北部端线。(2)在站点尺度上, 岛礁东南向瓦氏马尾藻定生密度明显低于西北向, 这与调查站位所受风浪影响的方位和强度相一致。在第二岛礁带上的4个岛礁北向瓦氏马尾藻的平均株高仅为26.3 cm, 说明瓦氏马尾藻不适宜在高波浪能生境中生存。(3)在站点尺度内, 第二岛礁带上浊度最低的北渔山岛瓦氏马尾藻有较大的垂直分布范围, 且定生深度达到了6.4 m, 而浊度高的近岸岛礁瓦氏马尾藻垂直分布范围较小。瓦氏马尾藻株高随着水深的增加而逐渐降低, 由此推测, 瓦氏马尾藻虽不能耐受强光, 但光照对瓦氏马尾藻的分布生长起重要作用。与同海区铜藻的垂直分布格局相比, 瓦氏马尾藻具有一定适应高浊度、高沉积物环境的能力。因此, 在浙北近岸海域瓦氏马尾藻是较适合进行生态移植修复的种类。     

Breeding places of Phlebotomus ariasi in the cevennes focus of leishmaniasis in the south of France

In a search for sandfly breeding sites in the Cévennes, France, six larvae of Phlebotomus ariasi were found in rotted manure in a cellar housing goats, and six adult flies of the same species were collected from emergence traps, one in the same cellar and five on a disused refuse dump. No sandflies were found emerging from holes of long-tailed field mice (Apodemus spp.) and no sandfly larvae were extracted from 77 kg (dry weight) and 130 kg (undried weight) of samples of soil or litter from different places. Total catches and the proportions of male flies in CDC miniature light traps were much higher in peridomestic than silvatic stations. About 15% of males caught in 3 peridomestic stations had unrotated genitalia. It is concluded that the larval breeding sites of P. ariasi are probably peridomestic and have a high content of organic matter. Observations on colonized P. ariasi show that, although immature stages of this fly need moisture, their requirements are comparatively low. It is suggested that female P. ariasi choose oviposition sites by sensing the presence of immature stages already in an optimal place.