Local modification of benthic flow environments by suspension-feeding stream insects

Larval black flies often exhibit spatially aggregated distributions, and individuals within patches can potentially reduce the supply of suspended food particles to downstream neighbors by modifying local flow characteristics. We used hot-film anemometry to quantify the magnitude and spatial extent of flow modifications downstream from feeding Simulium vittatum larvae in a laboratory flume, and to determine whether temporal patterns of flow variation are related to movements of the larval feeding appendages. Mean velocity 1 mm downstream from feeding larvae was reduced by 75%, and the percent reduction in velocity diminished asymptotically with downstream distance. Reduced velocities were evident as much as 60 mm downstream from, and 3 mm to either side of, larvae. Turbulence intensity (i.e., the SD of the velocity time series) was generally higher in this region relative to control flow conditions. Three results demonstrate the major contribution of the larval feeding appendages (i.e., labral fans) to such flow modification. First, there was a minimal reduction in mean velocity 5 mm downstream from non-feeding larvae (i.e., with closed labral fans), whereas mean velocity at the same location was reduced markedly when larvae were feeding. Second, the power spectrum of the velocity time series exhibited greatest power at frequencies that corresponded to the frequency of labral fan motions. Third, fan flick times accounted for most of the variance in the velocity power spectrum. The large local flow modifications that we documented have potentially important consequences for the feeding performance and growth of individuals located within larval aggregations, and are likely to influence behavioral interactions and spacing patterns.     

Feeding behavior of larval black flies (Diptera: Simuliidae) with and without exposure to Bacillus thuringiensis var. israelensis.

Behavioral time budgets were constructed for larval black flies of the Simulium jenningsi and S. tuberosum groups in a recirculating flume. Larvae spent the majority (54-80%) of time filter feeding. Flick rates of the labral fans, which averaged 0.7-1.9 flicks/sec, did not differ significantly between regimens of 0 mg/l and 10 mg/l of food. Larvae consistently flicked the fan nearest the water surface significantly more often than the lower fan. They also tended to flick the right fan more than the left, although this trend was significant in only one of four treatments. Although handedness cannot be dismissed, the differential flick rate appeared to be related to the position of the fans relative to the water surface, possibly because of different particulate concentrations in the microcurrents received by the upper and lower fans. Cessation of feeding by larvae of the S. jenningsi group from rivers regularly treated with Bacillus thuringiensis var. israelensis was independent of exposure to this biological control agent. Of larvae that ceased to feed, the mean time to cessation of feeding after exposure to Bti was 12-18 s, sufficient time to ingest a lethal dose.     

Stream insects as passive suspension feeders: effects of velocity and food concentration on feeding performance

Benthic suspension feeders are important components of aquatic ecosystems, often dominating the use of space and influencing patterns of material cycling between the water column and benthos. Biomechanical theory predicts that feeding by these consumers is governed by the flux (i.e., product of food concentration and velocity) of particulate material to their feeding appendages. We performed a laboratory flume experiment to test how feeding by larval black flies (Simulium vittatum Zett.) responds to independent manipulations of flow and food concentration. We quantified larval body posture, flick rate of the labral fans, and ingestion rate as a function of two concentrations of a baker's yeast/chalk suspension (0.96 and 4.44 mg l^-1) and five water velocities (20, 30, 45, 60, and 90 cm s^-1). Using analysis of covariance, we found that both flick rate and ingestion rate increased in a decelerating manner with increasing velocity, while fan height decreased linearly with increasing velocity. In contrast, food concentration had no effect on any aspect of feeding behavior. Thus, although both velocity and food concentration contribute to particle flux, our results indicate that the two were not substitutable under the range of conditions tested here.     

Balancing food availability and hydrodynamic constraint: phenotypic plasticity and growth in Simulium noelleri blackfly larvae

Organisms through phenotypic plasticity can cope with multiple changed environmental conditions. Theory predicts that animals in streams and rivers should be able to balance demands of the needs to obtain food efficiently and to adjust response to hydrodynamic variability. This study examined effects of variations in food availability and current velocity on the feeding structure and growth rate of Simulium noelleri blackfly larvae. The larvae developed larger labral fans and more rays under slow current and low food regimes than in fast current and high food conditions. In both fast and slow current regimes, growth rates were higher and development periods to the final-instar stage were shorter in high food treatments. The estimated flux rates of food particles through labral fans under high food treatments for both fast and slow current regimes were higher than those under low food treatments. Although both food and current velocity appeared to have selected for flexibility of feeding structure and growth rate, food availability was a more important factor for phenotypic and developmental plasticity than current velocity. The results indicate a strong link between environmental changes in food availability and current velocity, phenotypic plasticity, and growth rate of S. noelleri. This study suggests that plasticity of ecomorphs with macroevolutionary significance may play a role in the early evolutionary stages of blackfly larvae.     

Diversity, distribution and larval habitats of North Swedish blackflies (Diptera: Simuliidae)

1. Blackfly species richness and community structure were analysed at fifty-six sites in northern Sweden in two seasons. The sites were situated in a wide range of streams and rivers from small springbrooks, bog streams and lake-outlet streams to medium-sized forest rivers and large rivers draining montane regions.
2. Thirty-nine blackfly species were found, with between two and thirteen species per site. Neither species richness nor abundance could be related to the environmental variables measured.
3. An analysis of labral fan size of blackflies indicated a clear trend for the prevalence of larvae with small fans in large rivers and larvae with larger fan size in small streams. Similarly, fan size related to current velocities so that large fans were associated with slow current velocities and small fans with high velocities.
4. A strong relationship existed between species composition and habitat, as seen in ordination by non-metric multidimensional scaling. The relationship found between fan size and habitat size-related variables, such as channel width, depth, velocity and substratum particle size, along with longitude and altitude, in partial least squares regression analysis offered an explanation of the species composition–habitat relationship.
5. In addition to testing that distributions of blackfly larvae reflect morphological traits, we tested two general hypotheses pertaining to distribution patterns: (a) that blackfly communities show bimodal distributions; and (b) that their distributions are nested. Neither of these two hypotheses was supported by our observations. However, widespread blackfly species were locally more abundant than those found at relatively few sites, thus showing a positive abundance–occupancy relationship.     

Drift and settlement of stream insects in a complex hydraulic environment

1. The hydraulic and geomorphic characteristics of stream patches are often associated with distinctive assemblages or densities of stream invertebrates, and it is routinely presumed that these patterns reflect primarily species‐specific habitat requirements. An alternative hypothesis is that such patterns may be influenced by constraints on movement, such as the results of departure and settlement processes. We describe a manipulative experiment that examined how the hydraulic environments created by topographic bedforms influenced the drift behaviour and potential settlement sites for two species of mayfly (Baetis rhodani and Ecdyonurus torrentis). These species are common in the drift and often co‐occur in streams, but differ in their small‐scale distribution patterns, body shape and movement behaviour. 2. Flume experiments were carried out to determine how the hydraulic environments conditioned by a step bedform influence the behaviour of mayflies in the drift (swimming, posturing, tumbling), and the consequences of those behaviours (drift distance and time), compared to drift over a plane bed. The ramped step in the flume mimicked step bedforms that are common in coarse‐grained, high‐gradient streams. In contrast to the plane bed, a zone of recirculating flow was created downstream of the step, above which flow was faster and more turbulent. Uniform flows are used in most flume studies of drift; our approach is novel in recreating a complex hydraulic environment characteristic of stream channels. 3. Both species had some behavioural control over drift, and drift distances and times were shorter for live larvae than for dead larvae over the plane bed. The step had no impact on drift time or distance for live Baetis, but dead larvae were trapped in the flow separation eddy and drift time increased accordingly. Some Ecdyonurus also became trapped in the eddy, but live larvae drifted farther than dead larvae, and farther over the step than the plane bed. 4. Whilst in the drift, larvae altered their behaviour according to the ambient hydraulic environment, but in a species‐specific manner. Over the plane bed, Baetis had occasional swimming bursts, but primarily postured (maintained a stable body orientation), whereas Ecdyonurus spent roughly equal time posturing and swimming. In the more turbulent flows generated by the step, Baetis spent proportionately more time swimming, whereas Ecdyonurus spent more time posturing and often tumbling as body orientation became unstable. 5. In a high‐gradient stream, Baetis was more abundant close to steps than in plane bed patches with less complex flow, whereas the opposite pattern held for Ecdyonurus. Thus, the small‐scale distribution patterns of these species within streams correspond to their drift behaviours and ability to access various hydraulic patch types in our flume. These results are consistent with the hypothesis that constraints on movement and settlement may be important driver of distribution patterns within streams.     

Determinants of ingestion rates in filter-feeding larval blackflies (Diptera: Simuliidae)

SUMMARY 1. The functional response of the filter-feeding blackfly larva Frosimulium mixtum/fuscum was examined in laboratory experiments. We focused on the relationship between a component of the particle handling process (the cephalic fan flick rate) and ingestion rate, to determine whether ingestion rate is limited by handling-time. The influence of food deprivation on ingestion rate and feeding efficiency also was assessed. 2. Ingestion rate rose asymptotically with increasing food availability, leveling off at a concentration of about 100 mg1^?1. Satiation did not account for this plateau. Flick rate (the frequency with which larvae open and close their cephalic fans) increased in a similar fashion, though flick rate only accounted for about two-thirds of the variation in ingestion rate. 3. A simple equation describing the components of the feeding process in blackfly larvae was developed to investigate alternate flick rate ‘decision rules’ capable of predicting relationships between food concentration, flick rate and ingestion rate. The flick rate did not follow a fixed time rule, in which larvae allow particles to accumulate in the fan for a fixed amount of time prior to retraction. For most food densities, flick rate patterns more closely approximated a fixed number rule, in which fans are retracted after a fixed number of particles accumulate. 4. Flick rate rose with increasing food concentration even after the ingestion rate had levelled off, whereas several optimal filter-feeding models predict that filtering rate should decline as food concentration increases above this level. This predicted behavior was not observed because the number of particles ingested per flick failed to increase with increasing food concentration. 5. The efficiency of particle ingestion (number ingested × number available^?1× 100) was very low (<0.1%), and declined with increasing food concentration. 6. An increase in food deprivation time prior to a feeding trial produced a significant increase in flick rate, ingestion rate and feeding efficiency. This adjustment of feeding behaviour allowed animals to replace their gut contents more quickly following longer periods of food deprivation, which agrees with the qualitative predictions of optimal digestion-time models.     

Live blind snakes (Leptotyphlops dulcis) in eastern screech owl (Otus asio) nests: a novel commensalism

Summary Eastern screech owls bring live blind snakes to their nestlings, whereas all other prey are delivered dead. Some of the snakes are eaten but most live in nest debris, where they eat soft-bodied insect larvae from the decomposer community in fecal matter, pellets, and uneaten prey. Consumption of larvae may reduce larval parasitism on owl nestlings or larval competition with nestlings for food stored in the nest, because nestlings with live-in blind snakes grow faster and experience lower mortality than same-season broods lacking snakes. We propose a commensalistic association in which the screech owl benefits reproductively and the live-in blind snake is not affected.     

Distribution of Simulium buissoni Roubaud, 1906 and S. sechani Craig & Fossati, 1995 (Diptera: Simuliidae) in Nuku-Hiva and Eiao (Marquesas Archipelago, French Polynesia)

Nuku-Hiva and Eiao are the only islands of the Marquesas archipelago where blackflies bite humans. Knowledge of the geographic distribution of species is necessary to optimize control operations. Looking at frequency distributions of labral fan rays numbers from a large sample of larvae allowed redefinition of species. Simulium sechani Craig & Fossati, 1995, become the only species represented in Eiao. This species live in Nuku-Hiva too, but is limited to the external part of the main caldera. Presence of S. buissoni Roubaud, 1906, in all part of the island is confirmed, as is presence of S. hukaense Séchan, 1983 but in reduced number. Large samples of S. sechani extends its labral fan rays numbers. Stability of the described repartition over several decades is shown.     

Feeding behaviour of Japanese flounder larvae under laboratory conditions

Tank–reared Japanese flounder larvae, Paralichthys olivaceus , had a major feeding peak in the morning and a secondary peak in the afternoon throughout the larval development, with light being the primary factor regulating their feeding activity. The larvae consumed rotifers in preference to Artemia for up to 10 days, after which the food preference shifted to Artemia . Feeding rates of the larvae prior to 10 days post–hatch depended on prey density, but in the old larvae, feeding rates were independent of prey density. Maximum feeding rate occurred at 19° C. The occurrence of the attack posture, after its onset at first feeding (2 days post–hatch), increased up to 25 days, began to decrease when the larvae prepared to settle down, then disappeared after settlement. The occurrence frequency of the attack posture was positively related to fish density, but inversely related to starvation duration, and occurred most frequently at 19° C. This posture depended on prey density in larvae prior to 10 days post–hatch, but became independent of prey density as the larvae developed. It was obvious that, for flounder larvae, attack posture was a behavioural character closely related to feeding and subject to larval development and environmental factors.     

Flow over a ski jumper in flight: Prediction of the aerodynamic force and flight posture with higher lift-to-drag ratio

Large eddy simulations (LESs) are performed to study the flow characteristics around two flight posture models of ski jumping. These models are constructed by three-dimensionally scanning two national-team ski jumpers taking flight postures. The drag and lift forces on each component of a ski jumper and skis (head with helmet and goggle, body, arms, legs and skis) and their lift-to-drag ratios are obtained. For the two posture models, the drag forces on the body, legs and skis are larger than those on the arms and head with helmet and goggle, but the lift forces on the body and skis are larger than their drag forces, resulting in high lift-to-drag ratios on the body and skis and low lift-to-drag ratio on the legs. We construct simple geometric models, such as the circular cylinder, sphere and thin rectangular plate, predicting the drag and lift forces on each component of a ski jumper and skis, and validate them with those obtained from LES. Using these geometric models, we perform a parametric study on the position angles of flight posture for higher total lift-to-drag ratio. The flight postures obtained increase the total lift-to-drag ratios by 35% and 21% from those of two base postures, respectively. Finally, LESs are performed for the postures obtained and show the increases in the total lift-to-drag ratios by 21% and 16%, respectively, indicating the adequacy of using the simple geometric models for finding a flight posture of ski jumping having a higher lift-to-drag ratio at low cost.     

Dietary ascorbic acid requirements during the hatchery production of turbot larvae

The effect of high ascorbic acid (AA) levels transferred through enriched live food was evaluated for turbot Scophthalmus maximus larvae in two consecutive feeding experiments. The same feeding strategy was applied to all treatments, except for the AA content of the live food which was manipulated through bioencapsulation with ascorbyl palmitate. This resulted finally in a low, medium and high-AA treatment. The AA incorporation levels in the turbot larvae (up to 1400 μg AA g DW⁻¹) were correlated with the AA content of the live food administered. However, feeding the high AA concentration resulted in the same values as for the medium treatment, indicating a saturation of the body AA reserves. Under standard culture conditions, no differences in growth nor overall survival could be detected among the different groups, illustrating that the dietary AA requirements of larval turbot are met by non-enriched live food containing already 500 μg AA g DW⁻¹. The larvae of the high-AA treatment, however, showed a better pigmentation rate (47 and 32% for experiments 1 and 2, respectively) compared to the other groups (35 and 25%, respectively). Evaluation of the physiological condition applying a salinity stress test revealed an improvement by feeding extra AA, significantly in the medium-AA treatment. Though not significantly different, cumulative mortalities after challenge with Vibrio anguillarum amounted to 50% for the control v. 40% for the fish fed medium and high-AA diets, respectively. Moreover, the onset of mortalities in this study was slower (not significantly) for the fish fed the extra AA.     

Relationships between labral fan morphplogy, body size and habitat in North Swedish blackfly larvae (Diptera: Simuliidae)

A comparison was made of the labral fan morphology of 29 populations belonging to at least 21 different simuliid species present as mature larvae in spring-early summer in North Swedish streams and rivers. The study revealed that there are both adaptive, morphological and phylogenetic traits evident in the material. Two basically different particle capture techniques can be deduced from morphology, where two prosimuliids, Cnephia pallipes and Metacnephia trigonia , represent species which mainly rely on a sieving technique, whereas all other species investigated capture particles by some other process, such as diffusional deposition. Within tribe Simuliini there are significant relationships between several fan traits and habitat. Thus, fast rivers are inhabited by species with small fans, having short and stout rays, whereas small, slow streams harbour species with large fans consisting of elongated, delicate rays. The finer morphology of the fan rays in terms of secondary structures, the microtrichia, appears to be more related to phylogeny than" habitat. Since a phylogeny of blackfly species still is lacking, this conclusion cannot be tested. In two subgenera, however, which are particularly well represented in the present material ( Simulium s. str. and Nevermannia ), and whose taxonomy is based on criteria other than fan morphology, there is consistent support for the conclusions about habitat-morphology relations.     

Not so fast: Speed effects on forelimb kinematics in cercopithecine monkeys and implications for digitigrade postures in primates

Terrestrial mammals are characterized by their digitigrade limb postures, which are proposed to increase effective limb length (ELL) to achieve preferred or higher locomotor speeds more efficiently. Accordingly, digitigrade postures are associated with cursorial locomotion. Unlike most medium‐ to large‐sized terrestrial mammals, terrestrial cercopithecine monkeys lack most cursorial adaptations, but still adopt digitigrade hand postures. This study investigates when and why terrestrial cercopithecine monkeys adopt digitigrade hand postures during quadrupedal locomotion. Three cercopithecine species (Papio anubis, Macaca mulatta, Erythrocebus patas) were videotaped moving unrestrained along a horizontal runway at a range of speeds (0.4–3.4 m/s). Three‐dimensional forelimb kinematic data were recorded during forelimb support. Hand posture was measured as the angle between the metacarpal segments and the ground (MGA). As predicted, a larger MGA was correlated with a longer ELL. At slower speeds, subjects used digitigrade postures (larger MGA), however, contrary to expectations, all subjects used more palmigrade hand postures (smaller MGA) at faster speeds. Digitigrade postures at slower speeds may lower cost of transport by increasing ELL and step lengths. At higher speeds, palmigrade postures may be better suited to spread out high ground reaction forces across a larger portion of the hand thereby potentially decreasing stresses in hand bones. It is concluded that a digitigrade forelimb posture in primates is not an adaptation for high speed locomotion. Accordingly, digitigrady may have evolved for different reasons in primates compared to other mammalian lineages. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Role of silk threads in the dispersal of larvae through stream pools

The slow flow found in stream pools should lead to increased larval deposition in habitat unsuitable to many benthic invertebrates such as the larvae of the black fly Simulium tribulatum . To increase their chances of successfully transiting a pool, these larvae may use silk threads previously thought to facilitate settlement in fast-flowing habitat. In this study, we tested two ways in which these threads can affect a larva's transport in slow flow. First, a significant decrease in the fall velocity, which should lead to increased drift distance, was found between live (with silk) and dead (without silk) larvae falling in a laboratory chamber. The reduction due to the presence of silk varied over the natural range of larval sizes from a maximum of a 36% reduction for neonates to only 14% for the largest late-instar larvae. A second possible role of silk, that of increasing the chance of resuspension in slow flow due to increased drag forces, was tested using dynamically scaled models of neonate and late-instar larvae. The coefficient of drag ( C _d) was determined for both model types, with and without scaled silk threads, under conditions recreating a range of naturally occurring Reynolds numbers ( Re ). A significant increase in C _d was found due to the presence of silk for both life stages across relevant ranges in Re . The results of this study show that the presence of silk threads should significantly increase the distance larvae travel and their chances of being resuspended should they be deposited on the bed of a pool. Further, the same threads may act to both limit transport in regions of preferred fast flow, while maximizing transport through regions of unsuitably slow flow.     

Gut evacuation of walleye pollock larvae in response to feeding conditions

Gut residence times of first-feeding walleye pollock larvae were measured at 6°C in continuous and discontinuous feeding regimes. Larvae fed tagged copepod prey evacuated their guts more quickly in continuous feeding as compared to the discontinuous feeding treatment. The mean gut residence time was estimated to be 5.0 h for larvae feeding continuously. Gut evacuation by larvae fed tagged prey and then isolated without food (discontinuous treatment) was slower and more variable, with an estimated gut residence time exceeding 8.0 h. Field and laboratory observations suggest that the larval fish gut may be modeled as an intermittent plug-flow reactor (PFR) in response to diel feeding patterns. The cyclical nature of gut dynamics has implications for gut content analyses and the estimation of daily food rations.     

Carnivorous and detritivorous feeding of Mesocyclops leuckarti Claus (Cyclopoida Copepoda)

Food preference of M. leuckarti was experimentally analysed in relation to different species of live and dead crustaceans. Also their possibility of using detritus from bottom sediments as a source of food was tested.It has been found that M. leuckarti readily feed on dead organisms, and detritus from bottom sediments is sufficient for their survival.On the basis of experimental results the feeding strategy of M. leuckarti in lakes is discussed. It seems that the availability of different species of live prey depends on the mode of their swimming and on their body cover, while the availability of dead prey depends only on the body cover. Detritus may be an important source of food, particularly when the live prey are scarce in the pelagic zone, or not easily available.     

A potential link between lateral semicircular canal orientation,head posture,and dietary habits in extant rhinos (Perissodactyla,Rhinocerotidae)

Extant rhinoceroses share the characteristic nasal horn, although the number and size of horns varies among the five species. Although all species are herbivores, their dietary preferences, occipital shapes, and common head postures vary. Traditionally, to predict the “usual” head posture (the most used head posture of animals during normal unstressed activities, i.e., standing) of rhinos, the occipital shape was used. While a backward inclined occiput implies a downward hanging head (often found in grazers), a forward inclined occiput is related to the horizontal head posture in browsing rhinos. In this study, the lateral semicircular canal (LSC) of the bony labyrinth was virtually reconstructed from µCT‐images in order to investigate a possible link between LSC orientation and head posture in extant rhinoceroses. The usual head posture was formerly reconstructed for several non‐rhinoceros taxa with the assumption that the LSC of the inner ear is held horizontal (parallel to the ground) during normal activity of the living animal. The current analysis of the LSC orientation resulted in a downward inclined usual head posture for the grazing white rhinoceros and a nearly horizontal head posture in the browsing Javan rhinoceros. The other three browsing or mixed feeding species show subhorizontal (closer to horizontal than a downgrade inclination) head postures. The results show that anatomical and behavioral aspects, like occipital shape, presence and size of horns/tusk‐like lower incisors, as well as feeding and feeding height preferences influence the usual head posture. Because quantitative behavioral data are lacking for the usual head postures of the extant rhinos, the here described relationship between the LSC orientation and the resulting head posture linked to feeding preferences gives new insights. The results show, that the inner ear provides additional information to interpret usual head postures linked to feeding preferences that can easily be adapted to fossil rhinoceroses.     

弯曲姿态蛾类幼虫的自动识别方法研究

【目的】为害态幼虫现场识别时, 幼虫常出现姿态弯曲情况, 使提取的特征向量失真, 影响幼虫的匹配识别结果。本文提出了一种基于扇形变换的姿态不变胡氏矩特征向量提取方法, 提取的病害幼虫特征向量具有平移、 比例、 旋转和姿态不变性, 可以实现粗短弯曲姿态幼虫的自动识别。【方法】首先在幼虫图像细化的基础上采用最优一致逼近法确定了幼虫的弯曲区域和非弯曲区域。然后, 幼虫的弯曲区域采用扇形变换实现校正变直, 非弯曲区域经旋转和平移与扇形变换后的区域拼接组成完整虫体; 采用八邻域均值法填充变换后虫体区域中的空白点, 实现幼虫像的弯曲自动校正; 在此基础上提取胡氏不变矩具有姿态不变性, 采用最小距离分类器实现了多姿态幼虫的自动识别。最后, 以多种弯曲姿态的斜纹夜蛾Prodenia litura、 棉铃虫Heliocoverpa armigera、 甜菜夜蛾Spodoptera exigua、 玉米螟Ostrinia nubilalis等病害蛾类幼虫为识别对象进行了识别验证。【结果】对于24种不同姿态的幼虫图像, 在80%的识别阈值条件下, 基于经典胡氏不变矩的幼虫识别率为25%, 基于姿态不变胡氏矩的识别率为100%。【结论】实验结果表明该方法对多种弯曲姿态的粗短幼虫具有较高的识别率。     

Evolution of larval head-fans in Simuliidae (Diptera) as inferred from the structure and biology of Crozetia crozetensis (Womersley) compared with other genera

The life stages of C. crozetensis whose larvae have raking head-fans are described. Functional anatomical study of the fans compared with the filtering fans of normal Simuliidae together with SEM study of first instar larvae of Gymnopais, Twinnia and Prosimulium enable the evolution of cephilic fans to be inferred as follows: a group of lateral labral bristles evolved in two directions:

• 1. 

  To complete loss in Gymnopais and Twinnia whose ancestors therefore never possessed fully developed fans.
• 2. 

  Elaboration of the group of bristles first into raking organs, as in Crozetia , and later into even more complex filtering organs.

It is shown that unflexing of Simuliid fans depends on elastic energy stored in resilin-containing components deformed during flexing.
Adults of Crozetia mate on stones in streams on lie de la Possession, Ties Crozet. The females are non-biting and rest near streams while their eggs mature, thus forming a sub-Antarctic parallel to certain sub-Arctic Simuliidae.