Nocturnal increases in the use of near-surface water by pond animals

We assessed diel animal habitat use in three shallow ponds, using unbaited funnel traps, a large column sampler, and sweep net collections in the upper stratum (0–0.3 m) of littoral and open habitats. In all three ponds, more animals were caught at night than during the day, indicating that use of near-surface waters was greatest at night, particularly in the fishless ponds. All methods yielded similar patterns. Our results demonstrate that nocturnal observations of pond animals are necessary to describe their ecology, even in fishless ponds where diel differences in habitat use or behavior might not be anticipated.     

Movement response of Orange-Vaal largemouth yellowfish (<Emphasis Type="Italic">Labeobarbus kimberleyensis</Emphasis>) to water quality and habitat features in the Vaal River,South Africa

Many threatened fish species that utilize riverine habitats are faced with habitat degradation and subsequent deterioration in their ecological surroundings. Habitat degradation is a consequence of water quality parameters associated with anthropogenic activities including mining, industrial, agricultural and urban activities. We examined how the movement behaviour of radio-tracked Orange-Vaal largemouth yellowfish (Labeobarbus kimberleyensis) responded to a suite of water quality chemical parameters and habitat features in the Vaal River, South Africa. We found that the probability of their movement increased with a decrease in water clarity, presence of emergent and overhanging marginal vegetation and fast flowing rapids. High mobility in conditions of low water clarity was probably related to low prey capture success of this piscivorous fish. High movement of largemouth yellowfish in emergent and overhanging marginal vegetation areas and rapid habitat biotopes were attributed to accessibility of prey within these important cover features. When water quality parameters were considered, the probability of largemouth yellowfish movement increased with increasing levels of dissolved chloride (Cl) and silicon (Si), whereas movement decreased with high phosphate (PO₄) concentrations and increased with alkalinity levels in the river. High nutrient levels associated with eutrophication caused reductions in the movement of largemouth yellowfish. The association of increased movement of largemouth yellowfish with increasing Cl and Si is indicative of degraded habitat condition in the Vaal River system. Our study showed the importance of monitoring fish behavioural movement patterns to multiple environmental parameters, as these fish are important ecological indicators when appropriate conservation and management plans of freshwater ecosystems are required.     

THE HABITATS AND HABITAT PREFERENCES OF CLARIAS GARIEPINUS (PISCES: CLARIIDAE) IN A CLEAR COASTAL LAKE (LAKE SIBAYA,SOUTH AFRICA)

SUMMARY

The habitat preferences of the sharptooth catfish, Clarias gariepinus, in a clear coastal lake, Lake Sibaya, are described. Larval and small juvenile catfish inhabit flotsam and the rootstocks of plants in very shallow marginal areas. Larger juveniles inhabit densely-vegetated marginal pools, and may venture into open areas at night. Adult catfish (over 350 mm TL) occupy offshore areas, where their density is highest in terrace and sheltered bay habitats, and least in slope and profundal habitats. Adult catfish undergo a diel migration into shallow water at night and into deeper water during the day which is more marked at low lake levels. The methods used to capture catfish, and the main habitats in Lake Sibaya, are also described.     

Light,temperature and flow regimes of the Vaal River at Balkfontein,South Africa

A detailed investigation into light, temperature and flow regimes of the Vaal River was done for the first time. The Vaal River is ecologically and economically one of the most important rivers in South Africa. In a South African context, the Vaal River, with an average Secchi disk depth (Z _sd) of 0.41 m and an average euphotic zone depth (Z _eu) of 1.3 m, is a moderately turbid system. Red light (approx. 670 nm) penetrated Vaal River water to a greater degree than other components of the light spectrum. Water temperature in the Vaal River (min. = 10, max. = 27 °C at midday), closely followed seasonal changes in average atmospheric temperatures. The water-column shows little temperature stratification. The Vaal River is a highly regulated system with relatively high discharge rates at Balkfontein (average 112 m³ s^–1), but relatively low current velocity (approx. 0.35 m s^–1). Discharge was the most important variable to influence transparency of Vaal River water. Higher discharge resulted in higher total suspended solids (TSS) concentration and higher extinction coefficient (k) values as well as in higher turbidity and thus in lower Z _eu and Z _sd. From the mean TSS (141 mg l^–1) it was calculated that the Vaal River (mean annual run-off of 3532 million m³) transported 498016 tonnes of suspended solids per annum.     

Diel variation in beaked whale diving behavior

We investigate diel variation in beaked whale diving behavior using data from time–depth recorders deployed on six Blainville's ( Mesoplodon densirostris) (255 h) and two Cuvier's ( Ziphius cavirostris ) (34 h) beaked whales. Deep foraging dives (>800 m) occurred at similar rates during the day and night for Blainville's beaked whales, and there were no significant diel differences in ascent rates, descent rates, or mean or maximum depths or durations for deep dives. Dive to mid-water depths (100–600 m) occurred significantly more often during the day (mean = 1.59 h⁻¹) than at night (mean = 0.26 h⁻¹). Series of progressively shallower "bounce" dives were only documented to follow the deep, long dives made during the day; at night whales spent more time in shallow (<100 m) depths. Significantly slower ascent rates than descent rates were found following deep foraging dives both during the day and night. Similar patterns were found for the Cuvier's beaked whales. Our results suggest that so-called "bounce" dives do not serve a physiological function, although the slow ascents may. This diel variation in behavior suggests that beaked whales may spend less time in surface waters during the day to avoid near-surface, visually oriented predators such as large sharks or killer whales ( Orcinus orca ).     

Diel variation in predator abundance, predation risk and prey distribution in shallow-water estuarine habitats

Predation by visual predators is often affected by light conditions and may therefore exhibit strong diel variation. The dominant predators on grass shrimp, Palaemonetes pugio, are finfish predators that are thought to locate their prey by visual cues. We examined the response of grass shrimp to diel variation in predation risk in the nearshore shallow waters of the Chesapeake Bay. We used diel shoreline seines to assess the relative abundance of predators. We assessed the relative risk of predation with shrimp tethered at refuge (30 cm) and nonrefuge (60 cm) depths. To measure grass shrimp response to predation risk, we used dipnets to monitor habitat use. Four predominantly visual predators dominated the shoreline seine catches, Fundulus heteroclitus, Micropogonias undulatus, Morone americana and Morone saxatilis. Total predator abundance had a diel component, with dramatic nighttime decreases in total abundance, whereas guild composition and relative abundance remained unchanged. Relative predation risk for tethered shrimp exhibited significant time by habitat interaction. During the day, depth negatively affected survivorship of tethered shrimp while at night overall survivorship increased and there was no effect of depth. Shrimp habitats use reflected diel predation risks. Abundances in the near shore were highest during the day with decreased abundances at night. Together, the seine and tethering data highlight the importance for a refuge (e.g., shallow water) from predation during the daytime and a relaxation of predation pressure at night.     

Ground-truthing the impact of invasive species: spatio-temporal overlap between native least chub and introduced western mosquitofish

Western mosquitofish (Gambusia affinis) are one of the most destructive introduced species in the American West. The negative impact of introduced species on native taxa depends on their spatio-temporal overlap, which will determine the availability of refugia for native species. Experiments on the mechanisms underlying the interactions between introduced and native species rarely address habitat use, overlap, and refugia because individuals are confined to enclosures. In a previous study we used cages, microcosms, and aquaria to show that mosquitofish could prey on and out-compete native least chub (Iotichthys phlegethontis). In this study, we examined the spatio-temporal overlap between mosquitofish and least chub under natural conditions. We found periods of overlap and partitioning in the seasonal and diel habitat use of these species. Both species used shallow habitats during the day and night throughout the spring when least chub were spawning. Predation by adult mosquitofish on young least chub during the spring likely explains the reduction in least chub recruitment in the presence of mosquitofish. During the summer least chub avoided mosquitofish by exclusively using cooler habitats, or by occupying deeper, cooler habitats during the day when mosquitofish were active, and shallower, warmer habitats at night when mosquitofish were inactive. A shift to cooler habitats in the presence of mosquitofish may result in decreased growth and fecundity of least chub. However, we suggest that a reduction of warm shallow habitat may have a disproportionately greater negative impact on mosquitofish than species native to temperate regions, such as least chub. Habitat manipulations that reduce or eliminate warm habitats may ameliorate the harmful effects of mosquitofish and promote the long-term persistence of native taxa.     

Marine depth use of sea trout Salmo trutta in fjord areas of central Norway

The vertical behaviour of 44 veteran sea trout Salmo trutta (275–580 mm) in different marine fjord habitats (estuary, pelagic, near shore with and without steep cliffs) was documented during May–February by acoustic telemetry. The swimming depth of S. trutta was influenced by habitat, time of day (day v. night), season, seawater temperature and the body length at the time of tagging. Mean swimming depth during May–September was 1·7 m (individual means ranged from 0·4 to 6·4 m). Hence, S. trutta were generally surface oriented, but performed dives down to 24 m. Mean swimming depth in May–September was deeper in the near‐shore habitats with or without steep cliffs (2·0 m and 2·5 m, respectively) than in the pelagic areas (1·2 m). May–September mean swimming depth in all habitats was slightly deeper during day (1·9 m) than at night (1·2 m), confirming that S. trutta conducted small‐scale diel vertical movements. During summer, S. trutta residing in near‐shore habitat progressively moved deeper over the period May (mean 1·1 m) to August (mean 4·0 m) and then reoccupied shallower areas (mean 2·3 m) during September. In winter (November and February), individuals residing in the innermost part of the fjords were found at similar average depths as they occupied during the summer (mean 1·3 m). The swimming depths of S. trutta coincide with the previously known surface orientation of salmon lice Lepeophtheirus salmonis. Combined with previous studies on horizontal use of S. trutta, this study illustrates how S. trutta utilize marine water bodies commonly influenced by anthropogenic factors such as aquaculture, harbours and marine constructions, marine renewable energy production or other human activity. This suggests that the marine behaviour of S. trutta and its susceptibility to coastal anthropogenic factors should be considered in marine planning processes.     

Distribution,movements and diet of nocturnal fishes on temperate reefs

We counted nocturnal fishes both day and night, and monitored the position of tagged individuals on temperate reefs in New South Wales, Australia. Pempheris affinis and P. multiradiata were the most abundant nocturnal planktivores on Sydneys rocky reefs and showed great differences in diel migration behaviour. Both species were observed in deep shelter sites during the day (5–10m), and most emerged into the water column at night. P. multiradiata was found to undergo extensive vertical and horizontal migrations. In contrast, P. affinis remained within daytime depth strata, with tagged individuals often moving less than 20m at night. Tagged adult P. affinis returned to tagging sites for up to 7weeks, indicating high site fidelity. Dietary analysis demonstrated that small and large pempherids differed in diet and the timing of foraging, suggesting a size-based transition from diurnal to nocturnal foraging. Stratified sampling of planktonic assemblages at different depths during the day and night showed spatial variation in the availability of prey items at different times of the day. Amphipods, the main prey of large fish, were only available during the night, and concentrated in shallow water, whereas decapod larvae, consumed mainly by small fish, were abundant day and night. Large P. affinis also fed on polychaetes, which were never found in the stomachs of P. multiradiata, suggesting that these species may have different prey requirements, or that these polychaetes are only found in deep water where foraging P. affinis were abundant. We found no general model for the Pempheridae. The movements and behaviour of nocturnal fishes varied greatly by species, and this may be due to differences in body size, and/or physiological (e.g. visual ability) and ecological constraints.     

马鞍列岛多种生境中鱼类群聚的昼夜变化

为了解岛礁水域鱼类群聚的昼夜变化特征,以便更全面地设计采样方法和掌握采样的时间尺度,于2009年9月对马鞍列岛7种生境进行了共计24网次的刺网昼夜采样,结合排序和聚类方法,从种类组成、相对生物量和丰度、种类丰富度、多样性和相似性等方面对研究海域鱼类群聚特征的昼夜变化作了探讨.在采获的55种鱼类中,昼夜出现的分别为41和46种,数量差别不大,但其昼夜组成却随栖息水层的变化而不同,底层鱼类更趋向于夜间在硬相生境集群活动;近底层鱼类的昼夜集群随生境变化而变化,在同一生境中既有偏向白天也有趋向夜间的;中上层鱼类更多地出现在白天的人工生境(AH).AH白天的丰度渔获率显著大于晚上,而天然生境(NH)昼夜差别不大;生物量渔获率无论NH还是AH皆无显著昼夜差异.具体到种类,仅有小黄鱼Larimichthys polyactis和赤鼻棱鳗Thryssa kammalensis等少数种类的数量在AH有显著的昼夜差别,其他多数种类虽然昼夜的出现率大多有别,但渔获率昼夜差异皆不明显.多样性差异更多的表现在不同生境之间,而同一生境的昼夜差异往往不甚显著.各个生境中鱼类的昼夜种类交替现象非常明显,形成了以褐菖(鲐)Sebastiscus marmoratus和鳗鲇Plotosus anguillaris为代表的夜间优势类群为主的硬相生境群聚格局、以丝背细鳞鲀Stephanolepis cirrhifer和细刺鱼Microcanthus strigatus为代表的白天优势类群为主的硬相生境群聚格局以及缺乏底层优势类群、以石首鱼科鱼类为代表的近底层鱼类为绝对优势类群的软相生境群聚格局.因此,采用被动性渔具在近岸典型生境进行鱼类等相关生物调查时,应使采样时间覆盖昼夜两个时段,且至少保证24h.     

Consumer movement dynamics as hidden drivers of stream habitat structure: suckers as ecosystem engineers on the night shift

Ecosystem engineering can control the spatial and temporal distribution of resources and movement by engineering organisms within an ecosystem can mobilize resources across boundaries and distribute engineering effects. Movement patterns of fishes can cause physical changes to aquatic habitats though nesting or feeding, both of which often vary in space and time. Here we present evidence of ecosystem engineering by the Sonora sucker Catostomus insignis, a dominant fish in streams of the southwestern United States, and show how cryptic nocturnal movement patterns and bioturbation activities control heterogeneity in benthic substrates, and in sediment and carbon export. Sonora suckers exhibit distinct diel movement patterns, spending daylight hours in refuge habitats (typically deep pools) while moving into shallow habitats at night to feed. Feeding by suckers creates substantial disturbance in soft sediments that are patchy in space and time. These disturbances moved up to 2.4 × 10⁴ cm³ of sediment per square meter per week in locations that are up to hundreds of meters away from sucker daytime refuges. The diel cycles in feeding activity (i.e. nocturnal digging in benthic substrates) caused nighttime pulses in suspended sediment that comprised up to 32% of the daily suspended load and organic matter transport of a stream reach. During the daytime, this particulate transport settles in habitats beyond the location of the initial disturbance, thus redistributing both sediment and organic matter. Our data indicate that cryptic movement by ecosystem engineers can distribute their effects in space and time generating heterogeneity in resources and suggest that habitat modifications restricting consumer movement may alter the impact of engineering activities.     

Diel differences in fish assemblages in nearshore eelgrass and kelp habitats in Prince William Sound,Alaska

The importance of a particular habitat to nearshore fishes can be best assessed by both diurnal and nocturnal sampling. To determine diel differences in fish assemblages in nearshore eelgrass and understory kelp habitats, fishes were sampled diurnally and nocturnally at six locations in western Prince William Sound, Alaska, in summer 2007. Abundance of fish between day and night were similar, but species composition and mean size of some fish changed. Species richness and species diversity were similar in eelgrass during the day and night, whereas in kelp, species richness and species diversity were greater at night than during the day. In eelgrass, saffron cod (Eleginus gracilis) was the most abundant species during the day and night. In kelp, the most abundant species were Pacific herring (Clupea pallasii) during the day and saffron cod at night. Diel differences in fish size varied by species and habitat. Mean length of saffron cod was similar between day and night in eelgrass but was greatest during the day in kelp. Pacific herring were larger at night than during the day in kelp. Diel sampling is important to identity nearshore habitats essential to fish and help manage fish stocks at risk.     

Diel Habitat Partitioning by Bull Charr and Cutthroat Trout During Fall and Winter in Rocky Mountain Streams

We used underwater observation to determine diel habitat partitioning between bull charr, Salvelinus confluentus, and cutthroat trout, Oncorhynchus clarki, during fall and winter (0.1–8.3°C) in two Rocky Mountain streams that differed in habitat availability. The majority (>70%) of both species emerged from concealment cover at night, though bull charr exhibited a greater tendency for nocturnal behavior than cutthroat trout. Differences in day and night counts were most pronounced at temperatures <3°C, when very few fish of either species were observed in the water column during the day, but both species were common at night. Both species used concealment cover of large woody debris and boulder substrate crevices in deep pools during the day. At night, fish emerged from cover and habitat use shifted to shallow water with low cover. Microhabitat partitioning among species and size classes occurred at night, cutthroat trout moving into shallower, faster water that was farther from cover compared to bull charr. Smaller fish of both species occupied focal positions in slower, shallower water closer to the substrate than larger fish. Large, mixed-species aggregations also were common in beaver ponds both day and night. High variation in diel and site-specific winter habitat use suggests the need for caution in developing habitat suitability criteria for salmonids based solely on daytime observations or on observations from a few sites. Our results support the need to incorporate nocturnal habitat use and partitioning in studies of salmonid ecology.     

Vertical, lateral and longitudinal movement of zooplankton in a large river

1. The spatial distribution and movement patterns of zooplankton in large rivers are little known compared with those in lake environments. We conducted a series of studies in the Ohio River (U.S.A.) during the low flow period to assess diel vertical (DVM), longitudinal and lateral movement of crustacean zooplankton. 2. The dominant large zooplankter, the copepod Eurytemora affinis, showed a consistent vertical migration pattern of daytime ascent and night‐time descent during all sampling periods – the reverse of the most common migratory pattern of zooplankton in lakes. The cladoceran Bosmina migrated in a similar way in two of the three sampling periods. Surveys taken longitudinally in the river showed similar trends for both taxa. 3. During the lateral surveys, E. affinis was significantly more abundant in the shallow littoral zone during the night than in the daytime. The combination of vertical and lateral movement patterns along with the diel distribution of zooplanktivorous fish suggest that these movements are a predator‐avoidance mechanism. 4. Sampling programmes in large rivers should consider that larger zooplankton such as E. affinis may not be randomly distributed in the river channel and behaviours such as diel vertical migration may be just as evident in river habitats as in lakes.     

Comparison of Day and Night Fyke Netting, Electrofishing and Snorkelling for Monitoring a Population of the Threatened Golden Galaxias (Galaxias auratus)

The littoral zone of small off-stream water storage containing a translocated population of Galaxias auratus was sampled fortnightly at day and night with fyke nets, electrofishing and snorkelling over 3 months. Variation in population data provided by each method, including relative abundance indices, size structure, and habitat preferences, were examined. Aspects of behaviour and activity patterns were also investigated. Night sampling using all methods consistently yielded larger catches than day sampling. The size structure of catches varied, with electrofishing at night and fyke netting during the day having higher proportions of juveniles, whilst snorkelling at night and electrofishing during the day had higher proportions of adults. Fyke netting at night yielded by far the largest catches (~3-fold more than other methods) and also captured balanced proportions of juveniles and adults. Galaxias auratus had a strong diel activity pattern and were most active at night. The majority of the population migrated into the littoral zone during the night and back into deeper water during the day. A small number of juveniles remained in the littoral zone and some adults sheltered in the dense cover of species-rich littoral vegetation during the day. Shores with shallow depth profiles appeared to be preferred due to higher catches in these areas using all methods. Based on the results of this study, fyke netting at night in littoral habitats is recommended for monitoring populations of G. auratus. Fyke netting is likely to be an effective method for monitoring other lacustrine galaxiid species; however, further work is required to investigate the effects of habitat variables and fish community structure on activity patterns of galaxiids, and hence their catchability with various methods, in more extensive lentic environments.     

Comparison of nine different methods to assess fish communities in lentic flood‐plain habitats

This study compares the effectiveness and representativeness of electrofishing, snorkelling, seining, baited lift netting, multi‐mesh gillnetting, baited fish traps, fyke netting, angling and longline fishing, considering three typical lentic flood‐plain habitats at different times of day. Electrofishing was by far the most effective method yielding highest species richness, species trait representation and catch per unit of effort (CPUE), followed by seining. For single species like dace Leuciscus leuciscus, European ruffe Gymnocephalus cernua, common bream Abramis brama and silver bream Blicca bjoerkna, seining was more effective than electrofishing. With both methods, some species were more consistently caught during night, dusk or dawn than during daylight. All other methods tested cannot be generally recommended for fish community assessments in shallow backwaters due to their low representativeness of species inventory and generally low CPUE. Based on these results, electrofishing of 30 m transect replicates at different times of day for monitoring the fish community in shallow backwaters, can be recommended, enabling the maximum possible comparability to adjacent river habitats. Seining should be considered as an alternative if accessibility of habitats is restricted or electrofishing is prohibited. The 25 species detected in the backwaters also suggest that these habitats contribute a large proportion of fish diversity and should be included in standard assessments of river ecological status.     

Determinants of the distribution of juvenile fish in the littoral area of a shallow lake

1. Research has often focused on the pelagic areas of lakes; the littoral zone has received less attention. The few studies concerning fish distribution in littoral habitats have concentrated on stands of submersed macrophytes, whereas other littoral habitat types have seldom been investigated. 2. This study aimed to predict the occurrence of juvenile fish in several littoral habitats of a shallow lake as a function of food availability, complexity of habitat structure, water depth and substrate. Habitats comprising reed, woody structures, and two open water areas differing in depth were sampled for fish and invertebrate biomasses on two shores, over 6 months and during both daylight and at night. 3. The juvenile fish community consisted almost exclusively of 0+ and 1+ roach and perch. There was a strong diel component in habitat use, with a predominant occurrence of fish in complex habitats (mainly woody structures) during the day, and a partial migration towards the open habitats at night, more strongly expressed in roach than in perch. 4. The diet of all fish groups was relatively constant over the seasonal cycle, and was independent of habitat. There was a higher degree of planktivory in roach than in perch, but both species fed on benthic macroinvertebrates to a substantial extent. 5. According to a logistic regression model, the biomass of potential food organisms in the different habitats had little predictive effect on the spatial distribution of the fish, whereas the structural complexity of the habitats combined with the diel cycle explained about 28% of the occurrence patterns in 0+ and 1+ perch and 1+ roach.     

Diel shifts in the structure and function of nearshore estuarine fish communities

Day–night shifts in the nearshore fish fauna of a temperate microtidal estuary were assessed using a holistic suite of structural and functional community attributes. Mean fish species richness and diversity (taxonomic distinctness) were higher at night across all regions of the estuary and seasons, concurring with the findings of numerous comparable studies reviewed worldwide, while the diel period in which mean abundance was higher varied among seasons. Likewise, species and functional guild compositions (the latter based on feeding modes and habitat use) both differed significantly between day and night, with the extent of the diel shift again varying seasonally. Daytime fish communities were characterized by higher abundances of Atherinidae, Sillaginidae and Mugilidae, while Gobiidae were far more abundant at night. Marked shifts in size composition were also evident, with smaller fishes (<100 mm total length, L_T) being more prevalent during the day and larger fishes (≥200 mm L_T) proportionally more abundant at night. The above diel shifts were feasibly related to a range of predator–prey interactions and feeding‐related movements, namely a nocturnal decrease in top‐order avian piscivory coupled with an increase in invertebrate prey availability, resulting in changes in the presence and catchability of certain fish species in shallow estuarine waters.     

DIEL VARIATION IN MIGRATION RATES OF EASTERN PACIFIC GRAY WHALES MEASURED WITH THERMAL IMAGING SENSORS

We recorded the blows of gray whales during their southbound migration past central California in January 1994, 1995, and 1996, using thermal imaging sensors. For our sampling purposes, we defined day (0730–1630) and night (1630–0730) to coincide with the on/off effort periods of the visual counts being conducted concurrently. We pooled data across the three years of sampling and tested for diel variation in surfacing interval, pod size, offshore distance, migration rate, and swimming speed by comparing paired day/night means for samples collected within the respective 24-h period. We performed these tests using data from the entire migration period and then repeated the tests for samples collected prior to and after the approximate median migration date (15 January). Over the entire migration period we observed larger diurnal pod sizes (x?_day= 1.75 ± 0.280, x?_night= 1.63 ± 0.232) and greater diurnal offshore distances (x?_day= 2.30 ± 0.328 km, x?_night= 2.03 ± 0.356 km) but found no diel variation in surfacing interval. For the entire migration period, the nocturnal migration rate (average number of whales passing per hour) was higher than the diurnal rate. During the first half of the migration we detected no diel variation in pod size or surfacing interval, but diurnal offshore distances were larger than at night (x?_day= 2.28 ± 0.273 km, x?_night= 1.96 ± 0.318 km). Diurnal and nocturnal migration rates prior to 15 January were not different. During the second half of the migration, there was no diel variation in surfacing interval, pod size, or distance offshore, but the nocturnal migration rate was higher (28%, SE = 11.6%) than the diurnal rate. We found no diel variation in swimming speed in any comparison. We propose that later migrants socialize more during the day, which effectively slows their diurnal rate of migration relative to nocturnal rates.