Hitchhiking behaviour in the obligatory upstream migration of amphidromous snails

Migratory animals endure high stress during long-distance travel in order to benefit from spatio-temporally fluctuating resources, including food and shelter or from colonization of unoccupied habitats. Along with some fishes and shrimps, nerite snails in tropical to temperate freshwater systems are examples of amphidromous animals that migrate upstream for growth and reproduction after a marine larval phase. Here I report, to my knowledge, the first example of ‘hitchhiking’ behaviour in the obligatory migration of animals: the nerite snail Neritina asperulata appears to travel several kilometres as minute juveniles by firmly attaching to the shells of congeneric, subadult snails in streams of Melanesian Islands, presumably to increase the success rate of migration.     

Two modified Breder traps for quantitative studies of tropical amphidromous migration

Amphidromy is a life cycle of fauna found throughout the tropics and subtropics, including representatives from three phyla. Amphidromy involves a life cycle of oceanic larval development, with postlarval migration into streams where growth and reproduction take place. With increasing industrial and urban development in tropical regions, demands for freshwater are continuously growing, resulting in the construction of dams and diversions that break the freshwater continuum necessary for the amphidromous life cycle; thus, jeopardizing many populations of native and endemic tropical stream fauna. Because of this, more quantitative studies are needed to better understand this unique life cycle, which heretofore, has been relatively little studied compared to tropical terrestrial communities. As such, new and quantitative methods are needed to study the immature stages (e.g., postlarvae) associated with amphidromy. In this paper, we introduce two modifications of the original Breder trap that have been designed specifically for standardized, quantitative monitoring of amphidromous postlarval migration. In addition, the two modified traps can be used in a variety of stream settings from natural bed substrates to modified channels with little heterogeneity. The first modification is made of acrylic, with a flat bottom useful in channelized streams or streams with relatively flat, unnatural benthic substrate (e.g., concrete). The second modification is an affordable trap made from polyvinylchloride (PVC) compression couplings, to be used in streams with natural benthic substrates, particularly those with large, embedded and immovable boulders and bedrock outcrops. Both traps were designed for continuous water flow through the traps, providing the necessary rheotactic cue for migrating amphidromous postlarvae, and ameliorating deficiencies of earlier traps not intended for tropical amphidromous fauna. We also make recommendations for standardized use of these traps to facilitate data comparisons among studies.     

Otolith microchemistry of tropical diadromous fishes: spatial and migratory dynamics

Otolith microchemistry was applied to quantify migratory variation and the proportion of native Caribbean stream fishes that undergo full or partial marine migration. Strontium and barium water chemistry in four Puerto Rico, U.S.A., rivers was clearly related to a salinity gradient; however, variation in water barium, and thus fish otoliths, was also dependent on river basin. Strontium was the most accurate index of longitudinal migration in tropical diadromous fish otoliths. Among the four species examined, bigmouth sleeper Gobiomorus dormitor, mountain mullet Agonostomus monticola, sirajo goby Sicydium spp. and river goby Awaous banana, most individuals were fully amphidromous, but 9–12% were semi‐amphidromous as recruits, having never experienced marine or estuarine conditions in early life stages and showing no evidence of marine elemental signatures in their otolith core. Populations of one species, G. dormitor, may have contained a small contingent of semi‐amphidromous adults, migratory individuals that periodically occupied marine or estuarine habitats (4%); however, adult migratory elemental signatures may have been confounded with those related to diet and physiology. These findings indicate the plasticity of migratory strategies of tropical diadromous fishes, which may be more variable than simple categorization might suggest.     

Delivery of hatching larvae to estuaries by an amphidromous river shrimp: tests of hypotheses based on larval moulting and distribution

1.  Amphidromous shrimps live and breed in freshwater rivers and streams, but their larvae require development in sea water. Larvae may hatch upstream and then drift to the sea, although in some species females have been reported to migrate to the coast before larvae are released. Here, we tested the relative importance of larval drift and female migration in Macrobrachium ohione (Decapoda: Palaemonidae) in a distributary of the Mississippi River in Louisiana, U.S.A.
2.  Newly hatched (stage-1) larvae are nonfeeding and will not moult to stage 2 (first feeding stage) without encountering salt water. A factorial experiment was conducted in the laboratory to test the effects on moulting to stage 2 of (i) time spent by stage-1 larvae in fresh water before (ii) exposure to and maintenance in water of different salinity. Larvae kept in fresh water for 1 or 3 days before a change to saline water at 6 or 10 ppt showed a greater frequency of moulting than those kept for longer (5 days) in fresh water or changed to less saline water (2 ppt). Non-moulting larvae died or were moribund within 11 days of hatching.
3.  The relative abundance of stage-1 larvae was measured with plankton tows at two locations in the river c. 150 km apart, one near the sea and one upstream. Larval abundances near the sea were significantly greater than those upstream.
4.  The results indicate that hatched larvae of M. ohione have a limited period in which to drift in fresh water before reaching water sufficiently saline to stimulate moulting to the first feeding stage. Female migrations may play an important role in delivering larvae of amphidromous species from large continental river systems in which distances to the sea are great, while larval drift alone may be sufficient in species living in short streams, like those found in many small mountainous tropical islands.     

Dispersal and colonisation of plants in lowland streams: success rates and bottlenecks

Plant dispersal and colonisation, including rates of dispersal, retention, colonisation and survival of dispersed propagules (shoots and seeds), were studied in a 300-m stream reach in a macrophyte-rich lowland stream during one growing season. Relationships between colonisation processes and simple flow parameters were tested. Each fortnight during a growing season, the number of dispersed plant propagules and the number of new and lost plant colonisations since the last sampling day were recorded. The retention of dispersing shoots was tested on two occasions during the growing season by releasing plant shoots and subsequently re-collecting the retained shoots in the study reach. The main bottleneck for plant colonisation in macrophyte-rich lowland streams is the primary colonisation (development of attached roots in the sediment from vegetative propagules or seedlings) of retained shoots and seeds, due in part to low retention success (1% of the dispersed shoots per 100-m reach) and to unsuccessful colonisation of retained shoots (3.4% of retained shoots colonised). The number of drifting shoots and seeds per day during the growing season were 650–6,950 and 2,970–62,780, respectively, and caused no constraint to colonisation. The survival rate of primary colonists was high with 80% surviving during the first growing season and about 50% surviving the first winter. There was no relationship between number of drifting shoots and flow, but the number of drifting seeds increased with flow. The number of colonisations between two consecutive sampling days correlated to the extent of low flow in the period. The loss rate of colonisations correlated to high flow events, but was low overall. My study strongly indicates that the number of propagules is not a constraint to colonisation in macrophyte-rich lowland streams. Rather, I conclude that primary colonisation is the main constraint to regaining vegetation in lowland streams in general and in vegetation-free rehabilitated streams in particular. Therefore, if plant colonisation is a target for stream rehabilitation, it is important to enhance retention and colonisation of propagules by creating areas of low flow and by providing physical obstacles to work as retention agents in the stream. Handling editor: L. M. Bini     

Dispersal and population expansion in a direct developing marine snail (Littorina saxatilis) following a severe population bottleneck

Most marine benthic invertebrate species have planktonic larvae, and in species in which juveniles and adults have low vagility a larva is obviously an efficient way of active dispersal. A minority of benthic invertebrate species develop without any pelagic phase at all. A largely unsolved question is how and at what rate do these species disperse. We have addressed this question using the marine littoral snail Littorina saxatilis (Olivi) as an example of a species that completely lacks larval dispersal. In the Koster archipelago (north part of the Swedish west coast), L. saxatilis occupies rocky island habitats of different sizes, from large islands to small intertidal skerries (islets). In 1988 an extremely dense bloom of a toxin-producing flagellate killed more than 99% of this snail species in this area. Populations of larger islands were reduced, often to less than 1%, but were restored over 2–4 yr. In contrast, populations of small intertidal skerries were completely wiped out and thus could not increase by local recruitment. Four years later, however, four of 33 skerries (12%) were successfully recolonized with relatively dense populations, and another five had received a few founder individuals. These results indicate recruitment through founder individuals, and are rough estimates of dispersal rate in a snail species that lacks a pelagic developmental stage.     

Skip of the routine habitat in an amphidromous migration of ayu

Ayu Plecoglossus altivelis is a typical amphidromous fish with ontogenic requirements for habitats among lower reaches of a river, the marine coastal area, and middle reaches of a river at the immature adult stage. Mature fish that had a small body size like ordinary young were found around the spawning ground. In comparison with standard adults, the migratory histories of the dwarf adults were examined using otolith microchemistry analysis. Sr:Ca ratio estimated that the dwarf fish were exposed to saline or brackish water after once moving into the river, whereas the standard-size fish migrated upstream.     

Dispersal,isolation and diversification with continued gene flow in an Andean tropical dry forest

The Andes are the world's longest mountain chain, and the tropical Andes are the world's richest biodiversity hot spot. The origin of the tropical Andean cordillera is relatively recent because the elevation of the mountains was relatively low (400–2500 m palaeoelevations) only 10 MYA with final uplift being rapid. These final phases of the Andean orogeny are thought to have had a fundamental role in shaping processes of biotic diversification and biogeography, with these effects reaching far from the mountains themselves by changing the course of rivers and deposition of mineral‐rich Andean sediments across the massive Amazon basin. In a recent issue of Molecular Ecology, Oswald, Overcast, Mauck, Andersen, and Smith (2017) investigate the biogeography and diversification of bird species in the Andes of Peru and Ecuador. Their study is novel in its focus on tropical dry forests (Figure 1) rather than more mesic biomes such as rain forests, cloud forests and paramos, which tend to be the focus of science and conservation in the Andean hot spot. It is also able to draw powerful conclusions via the first deployment of genomic approaches to a biogeographic question in the threatened dry forests of the New World.     

Dispersal influences genetic and acoustic spatial structure for both males and females in a tropical songbird

Animals exhibit diverse dispersal strategies, including sex‐biased dispersal, a phenomenon common in vertebrates. Dispersal influences the genetic structure of populations as well as geographic variation in phenotypic traits. Patterns of spatial genetic structure and geographic variation may vary between the sexes whenever males and females exhibit different dispersal behaviors. Here, we examine dispersal, spatial genetic structure, and spatial acoustic structure in Rufous‐and‐white Wrens, a year‐round resident tropical bird. Both sexes sing in this species, allowing us to compare acoustic variation between males and females and examine the relationship between dispersal and song sharing for both sexes. Using a long‐term dataset collected over an 11‐year period, we used banding data and molecular genetic analyses to quantify natal and breeding dispersal distance in Rufous‐and‐white Wrens. We quantified song sharing and examined whether sharing varied with dispersal distance, for both males and females. Observational data and molecular genetic analyses indicate that dispersal is female‐biased. Females dispersed farther from natal territories than males, and more often between breeding territories than males. Furthermore, females showed no significant spatial genetic structure, consistent with expectations, whereas males showed significant spatial genetic structure. Overall, natal dispersal appears to have more influence than breeding dispersal on spatial genetic structure and spatial acoustic structure, given that the majority of breeding dispersal events resulted in individuals moving only short distances. Song sharing between pairs of same‐sex animals decreases with the distance between their territories for both males and females, although males exhibited significantly greater song sharing than females. Lastly, we measured the relationship between natal dispersal distance and song sharing. We found that sons shared fewer songs with their fathers the farther they dispersed from their natal territories, but that song sharing between daughters and mothers was not significantly correlated with natal dispersal distance. Our results reveal cultural differences between the sexes, suggesting a relationship between culture and sex‐biased dispersal.     

Dispersal evolution in temporally variable environments: implications for plant range dynamics

Dispersal—the movement of an individual from the site of birth to a different site for reproduction—is an ecological and evolutionary driver of species ranges that shapes patterns of colonization, connectivity, gene flow, and adaptation. In plants, the traits that influence dispersal often vary within and among species, are heritable, and evolve in response to the fitness consequences of moving through heterogeneous landscapes. Spatial and temporal variation in the quality and quantity of habitat are important sources of selection on dispersal strategies across species ranges. While recent reviews have evaluated the interactions between spatial variation in habitat and dispersal dynamics, the extent to which geographic variation in temporal variability can also shape range-wide patterns in dispersal traits has not been synthesized. In this paper, we summarize key predictions from metapopulation models that evaluate how dispersal evolves in response to spatial and temporal habitat variability. Next, we compile empirical data that quantify temporal variability in plant demography and patterns of dispersal trait variation across species ranges to evaluate the hypothesis that higher temporal variability favors increased dispersal at plant range limits. We found some suggestive evidence supporting this hypothesis while more generally identifying a major gap in empirical work evaluating plant metapopulation dynamics across species ranges and geographic variation in dispersal traits. To address this gap, we propose several future research directions that would advance our understanding of the interplay between spatiotemporal variability and dispersal trait variation in shaping the dynamics of current and future species ranges.     

The fate of received sperm in the reproductive tract of a hermaphroditic snail and its implications for fertilisation

Multiple mating, sperm storage and internal fertilisation enhance sperm competition. The great pond snail can use stored sperm for over three months, and frequently mates with different partners. This hermaphrodite, Lymnaea stagnalis, can also self-fertilise and often produces egg masses containing both selfed and outcrossed eggs. Hence, a sperm recipient may exert considerable control over paternity. Using microsatellite markers, we show that when allosperm are present, all genotyped eggs are cross-fertilised. We also find that sperm have the opportunity to compete, because double matings lead on average to equal paternity for each sperm donor. This indicates that received sperm are randomly mixed in storage. To gain further insight into the mechanisms underlying the process of sperm storage, digestion and utilisation, we investigated the fate of donated sperm at different times after copulation. We find that within 3 h after transfer most sperm have been transported into the sperm-digesting organ. Fluorescent labelling of sperm in histological sections further reveals that allosperm are not stored in the fertilisation pouch, but upstream in either the hermaphroditic duct, seminal vesicles, or ovotestis. Besides contributing to the understanding of the mechanisms underlying sperm competition and/or cryptic sperm choice, this study shows that mixed mating cannot be treated as a separate issue in hermaphroditic animals.     

淡水蜗牛视觉系统的输入与输出通路(英文)

通过神经生物素在视神经上的逆行性传输对淡水蜗牛(Planorbarius corneus)视网膜及中央神经节的输入、输出神经元进行标记。由于没有发现突触联系,所以至少一部分光感受细胞的轴突可被视为直接参与形成视神经。这些神经元的轴突进入大脑神经节形成密集的细传入神经纤维束-视神经堆。传出神经元则存在于除颊部以外的所有神经节。一些上行轴突在大脑神经节处分叉,通过脑-脑联合,到达对侧眼并在眼杯处形成分枝。部分传出神经元的轴突也投射于不同的外周神经,如:n.n.intestinalis,pallialis dexter,pallialis sinister internus et externus。五羟色胺能纤维和FMRF-酰胺能纤维均存在于视神经上,且这些纤维隶属于只投射在同侧眼的中央神经元。它们形成了位于眼杯处的丰富曲张结构及视网膜核心层,并且可能有助于调节视网膜对光的敏感性。     

Distribution of migratory fishes and shrimps along multivariate gradients in tropical island streams

Among the 16 species of fishes and shrimps studied at 51 sites along several small streams at Basse Terre, Guadeloupe, Lesser Antilles, more than 60% showed a spatial distribution significantly influenced by environmental conditions. These included altitude, basin size, terrestrial vegetation and land use. However, the range of the species habitat was generally high, except for three species limited to the downstream stretches (two fishes and one shrimp). The habitat characteristics of fish and shrimp species tended to differ, but the difference was not significant (P<0·05). On the contrary, the habitat characteristics of amphidromous and catadromous species clearly differed, with a lower occurrence at the most elevated sites of catadromous species than amphidromous species.     

Around the Mediterranean: an extreme example of loop migration in a long‐distance migratory passerine

An important issue in migration research is how small‐bodied passerines pass over vast geographical barriers; in European–African avian migration, these are represented by the Mediterranean Sea and the Sahara Desert. Eastern (passing eastern Mediterranean), central (passing Apennine Peninsula) and western (via western Mediterranean) major migration flyways are distinguished for European migratory birds. The autumn and spring migration routes may differ (loop migration) and there could be a certain level of individual flexibility in how individuals navigate themselves during a single migration cycle. We used light‐level loggers to map migration routes of barn swallows Hirundo rustica breeding in the centre of a wide putative contact zone between the northeastern and southernwestern European populations that differ in migration flyways utilised and wintering grounds. Our data documented high variation in migration patterns and wintering sites of tracked birds (n = 19 individuals) from a single breeding colony, with evidence for loop migration in all but one of the tracked swallows. In general, two migratory strategies were distinguished. In the first, birds wintering in a belt stretching from southcentral to southern Africa that used an eastern route for both the spring and autumn migration, then shifted their spring migration eastwards (anti‐clockwise loops, n = 12). In the second, birds used an eastern or central route to their wintering grounds in central Africa, shifting the spring migration route westward (clockwise loops, n = 7). In addition, we observed an extremely wide clockwise loop migration encompassing the entire Mediterranean, with one individual utilising both the eastern (autumn) and western (spring) migratory flyway during a single annual migration cycle. Further investigation is needed to ascertain whether clockwise migratory loops encircling the entire Mediterranean also occur other small long‐distance passerine species.     

The reliability of estimates of migration in the peppered moth Biston betularia and some implications for selection-migration models

Migration of adult males is one of the important variables involved in the mathematical models of industrial melanism in Biston betularia. Values for this variable are based on data from a capture-recapture performed by Bishop (1972) using both local and bred moths which were at least one night old at release. We carried out an experiment to compare the rate of recapture close to the point of release for moths allowed to fly away immediately after their emergence around dusk and those which were at least one night old at release. Unheld moths were less likely to be recaptured suggesting that males have an initial dispersal phase on their first night which results in a higher rate of emigration than on subsequent nights. Such a phase would have been largely missed in Bishop's experiment. The implications of this type of behaviour pattern for the models of spatial variation based on a selection-migration balance are discussed.     

Evidence of local short‐distance spawning migration of tropical freshwater eels,and implications for the evolution of freshwater eel migration

Freshwater eels have fascinated biologists for centuries due to the spectacular long‐distance migrations between the eels’ freshwater habitats and their spawning areas far out in the ocean and the mysteries of their ecology. The spawning areas of Atlantic eels and Japanese eel were located far offshore in the Atlantic Ocean and the Pacific Ocean, respectively, and their reproduction took place thousands of kilometers away from their growth habitats. Phylogenetic studies have revealed that freshwater eels originated in the Indonesian region. However, remarkably little is known about the life histories of tropical freshwater eels despite the fact that tropical eels are key to understanding the nature of primitive forms of catadromous migration. This study found spawning‐condition tropical freshwater eels in Lake Poso, central Sulawesi, Indonesia, with considerably high gonadosomatic index values and with histologically fully developed gonads. This study provides the first evidence that under certain conditions, freshwater eels have conditions that are immediately able to spawn even in river downstream. The results suggest that, in contrast to the migrations made by the Atlantic and Japanese eels, freshwater eels originally migrated only short distances of <100 kilometers to local spawning areas adjacent to their freshwater growth habitats. Ancestral eels most likely underwent a catadromous migration from local short‐distance movements in tropical coastal waters to the long‐distance migrations characteristic of present‐day temperate eels, which has been well established as occurring in subtropical gyres in both hemispheres.     

The determinants of land snail diversity along a tropical elevational gradient: insularity,geometry and niches

Aim We investigated the patterns of species richness in land snails and slugs along a tropical elevational gradient and whether these patterns correlate with area, elevation, geographic constraints, and productivity. We did so both at the scale at which land snail population processes take place and at the coarser scale of elevational zones. Location Mount Kinabalu (4096 m) and the adjacent Mount Tambuyukon (2588 m) in Kinabalu Park, Sabah, Malaysian Borneo. Methods We used an effort‐controlled sampling protocol to determine land snail and slug species richness in 142 plots of 0.04 ha at elevations ranging from 570 to 4096 m. Extents of elevational ranges were determined by interpolation, extended where appropriate at the lower end with data from lowlands outside the study area. We used regression analysis to study the relationships between species density and richness on the one hand and elevation and area on the other. This was done for point data as well as for data combined into 300‐m elevational intervals. Results Species density (based on the individual samples) showed a decline with elevation. Elevational range length profiles revealed that range lengths are reduced at greater elevations and that a Rapoport effect is absent. Diversity showed a mild mid‐domain effect on Kinabalu, but not on Tambuyukon. When the data were combined into 300‐m elevational intervals, richness correlated more strongly with elevation than with area. Ecomorphospace was seen to shrink with increasing elevation. Main conclusions The elevational species richness patterns show the combined effects of (1) reduced niche diversity at elevations with lower productivity and (2) historical events in which the upward migration of lowland species as well as the speciation of highland endemics took place.     

Dispersal and recruitment of fish in an intermittent stream network

Animal movement is an important process connecting habitats in heterogeneous landscapes, and can play a key role in population persistence. Laboratory swim trials were conducted to determine and compare the dispersal capabilities of two native Australian fish, mountain galaxias (Galaxias olidus, Family Galaxiidae) and southern pygmy perch (Nannoperca australis, Family Nannopercidae) that maintain populations in hydrologically variable and intermittently flowing streams in south‐eastern Australia. These experiments showed that G. olidus had significantly greater swimming endurance under a range of flow velocities. Concurrent field surveys were used to establish whether swimming abilities observed in laboratory studies were consistent with patterns of inferred movement from distribution and abundance patterns observed in the field. Data collected at multiple sites from headwater to lowland reaches along multiple streams revealed substantial temporal changes in the distribution of young‐of‐year (0+) G. olidus, with spawning occurring at upland sites in winter, followed by downstream larval migration and subsequent upstream movement in late spring. Observed spatial and temporal patterns in G. olidus abundances were consistent with a source‐sink population structure, which may be disrupted by prolonged cease‐to‐flow periods during drought years. In contrast, results for N. australis suggested limited dispersal, with restricted local populations that persist at sites with permanent surface water. These field and laboratory findings complement our understanding of the spatial population structure of these two species in intermittent streams, and highlight the importance of understanding the role of dispersal in species conservation and habitat restoration.