Diapause in copepods (Crustacea) from ephemeral habitats with different hydroperiods in Everglades National Park (Florida,U.S.A.)

Water management practices in the Everglades have severely stressed the natural system, particularly by reducing the hydroperiods of much of the region. During the dry season of 1999, we investigated the influence of hydroperiod on the species composition and dormancy patterns of freshwater copepod communities in seasonal wetlands of Everglades National Park, Florida, U.S.A. The habitats were characterized by an annual dry season, from December through June. We sampled at two locations: the Long Pine Key area of the Rocky Glades region (short hydroperiod, ca. 4–5 months), and western Taylor Slough (intermediate hydroperiod, ca. 8–10 months). Both areas have experienced a reduction in natural hydroperiods and an increase in the frequency of dry-down. We collected weekly plankton samples from Rocky Glades solution holes to assess the potential species pool of copepods. To document the taxa capable of surviving dry-down by resting, we performed three immersion trials in which we rehydrated, in laboratory aquaria, sediment patches from solution holes and surface soils from all stations. Only a subset of the planktonic species collected emerged from the dried sediments. The cyclopoids Microcyclops rubellus and Paracyclops poppei were dominant. This is the first record of diapause for P. poppei. Species distributions from the different hydroperiod soil patches indicated that more diapausing species occurred at the sites that dried for shorter periods. Emerging individuals of M. rubellus and P. poppei were mainly ovigerous females, demonstrating a resting strategy seldom before recorded. The cyclopoid Diacyclops nearcticushad not been previously reported to diapause, but they emerged from the dried sediments in our trials. Our collections included six new records for Florida: Diacyclops nearcticus, Megacyclops latipes, Orthocyclops modestus, Elaphoidella marjoryae, Bryocamptus sp. and Bryocamptuscf. newyorkensis. Paracyclops poppei, Macrocyclops fuscus and Arctodiaptomus floridanus are new records for Everglades National Park. Clearly, diapause is an important strategy for the persistence of copepods in short-hydroperiod wetlands. The duration of the dry period appears to be inversely related to the number of species that emerge from diapause.     

Ecological succession,patch age and the evolution of social behaviour and terminal investment

Recent years have witnessed a growing interest in understanding the evolution of social behaviour in heterogeneous spatially structured populations. These studies, however, have neglected the impact of extinction–colonisation dynamics and ecological succession on the dynamical expression of social behaviour over time. Here, I present a kin‐selection model in which patches are structured into age‐classes. I show that ecological succession and patch age lead to highly plastic social phenotypes that vary dramatically as societies age since their initial establishment until their ultimate collapse. I find that the mode of colonisation following dispersal strongly influences the patch age‐dependent trajectories of social phenotypes. When patches are colonised by a random collection of immigrants, aggression is favoured during the build‐up of a society, but it slowly subsides until it eventually gives place to cooperation throughout the later stages of a society's lifespan. When newly established societies are formed by collectives of close relatives, cooperation is favoured during the build‐up of the society as well as when the society nears its eventual collapse. At intermediate societal ages, the genetic structure of the society is sufficiently resilient to the influx of immigrants such that cooperation remains relatively high. Moreover, I report a novel form of social terminal investment, whereby cooperative effort rises when patches approach their collapse. When dispersal is allowed to co‐evolve with cooperation, we observe a sudden rise in dispersal phenotypes before a patch's collapse, and the surprising result that clonal colonisation does not yield significantly higher levels of cooperation than the individual mode of colonisation. More generally, my results show that ecological succession strongly determines the dynamics of kin selection after colonisation, and therefore I expect that these findings will be valuable for understanding behavioural syndromes during range expansion or biological invasions.     

Contributions of airborne dispersal and dormant propagule recruitment to the assembly of rotifer and crustacean zooplankton communities in temporary ponds

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Differential dispersal rates in an intertidal meiofauna assemblage

Meiofaunal nematodes and copepods disperse passively with sediment bedload, and copepods also display active emergence and reentry behavior. Epigrowth-feeders may be the nematode feeding group most susceptible to passive transport because they live closest to the sediment surface. We used bottom traps at a nematode-dominated intertidal mudflat in Maine, USA, to test the hypotheses that (1) meiofauna taxa disperse in relative proportions different from those of the ambient community; (2) copepods have the highest relative dispersal rate (number of individuals trap⁻¹ day⁻¹ ambient individual⁻¹) and are not as tightly linked as other taxa to sediment flux; and (3) epigrowth-feeders have the highest nematode relative dispersal rate. Results supported all three hypotheses. Nematodes accounted for 95.8% of the individuals in cores, but only 38.9% of the individuals in traps. Copepods accounted for 1.5% of the individuals in cores, but 56.7% of the individuals in traps. Less common taxa also had different relative proportions in cores and traps, as did nematode feeding groups and individual species. The relative dispersal rate was far higher for copepods than for any other taxonomic group, and the absolute (number of individuals trap⁻¹ day⁻¹) and bulk (number of individuals g sediment⁻¹ trap⁻¹ day⁻¹) dispersal rates for copepods were equal to those of the 65-fold more abundant nematodes and higher than those for all other taxa. The non-selective deposit-feeders were the most abundant nematode feeding group in the ambient community, but the epigrowth-feeders as a group and as individual species had the highest absolute, relative, and bulk dispersal rates. Non-metric multidimensional scaling (MDS) using analysis of similarity (ANOSIM) and species similarity percentages (SIMPER) reflected these differences between ambient and dispersing nematode assemblages. Significant positive regression relationships between sediment weight and the number of individuals captured in traps for nematodes and some other taxa indicated that they moved passively in the bedload. Lack of a significant regression relationship for copepods suggested an active behavioral component to dispersal. Meiofauna populations in this soft-bottom community were highly dynamic, demonstrating that the role of dispersal must be included in any consideration of the ecology of soft-bottom systems at local and regional spatial scales.     

Patterns and drivers of Nothobranchius killifish diversity in lowland Tanzania

Temporary pools are seasonal wetland habitats with specifically adapted biota, including annual Nothobranchius killifishes that survive habitat desiccation as diapausing eggs encased in dry sediment. To understand the patterns in the structure of Nothobranchius assemblages and their potential in wetland conservation, we compared biodiversity components (alpha, beta, and gamma) between regions and estimated the role and sources of nestedness and turnover on their diversity. We sampled Nothobranchius assemblages from 127 pools across seven local regions in lowland Eastern Tanzania over 2 years, using dip net and seine nets. We estimated species composition and richness for each pool, and beta and gamma diversity for each region. We decomposed beta diversity into nestedness and turnover components. We tested nestedness in three main regions (Ruvu, Rufiji, and Mbezi) using the number of decreasing fills metric and compared the roles of pool area, isolation, and altitude on nestedness. A total of 15 species formed assemblages containing 1–6 species. Most Nothobranchius species were endemic to one or two adjacent regions. Regional diversity was highest in the Ruvu, Rufiji, and Mbezi regions. Nestedness was significant in Ruvu and Rufiji, with shared core (N. melanospilus, N. eggersi, and N. janpapi) and common (N. ocellatus and N. annectens) species, and distinctive rare species. Nestedness apparently resulted from selective colonization rather than selective extinction, and local species richness was negatively associated with altitude. The Nothobranchius assemblages in the Mbezi region were not nested, and had many endemic species and the highest beta diversity driven by species turnover. Overall, we found unexpected local variation in the sources of beta diversity (nestedness and turnover) within the study area. The Mbezi region contained the highest diversity and many endemic species, apparently due to repeated colonizations of the region rather than local diversification. We suggest that annual killifish can serve as a flagship taxon for small wetland conservation.     

Vicariance, dispersal and scale in the aquatic subterranean fauna of karst regions

1. The causes of distribution patterns of stygobionts (obligate subterranean-dwelling aquatic species) were examined with special emphasis on vicariance and dispersal.
2. Dispersal was investigated on the premise that if migration is important, then migration at small scales should predict patterns at larger scales. Data on the copepod fauna of epikarst in Slovenia were especially useful for the study of migration, because data on habitat occupancy could be collected at scales of individual drips located metres apart to the scale of individual caves to entire karst regions. Occupancy of drips in one cave was a remarkably good predictor of occupancy of caves in a region, although not of the overall range of a given species. These results were also supported by occupancy patterns of the general stygobiotic fauna of West Virginia caves, compared at different scales.
3. Vicariance was investigated by noting that proximity to marine embayments increases the likelihood of vicariant speciation. In the U.S.A., only the fauna of the Edwards Aquifer of Texas has a significant component of marine-derived species. Differences in shape of the relationship between species number and number of caves in a county indicated that the marine-derived component represented an addition to rather than a replacement of the other stygobiotic species.
4. Thus, we found evidence for the importance of both vicariance and dispersal. The techniques employed could be used to study these patterns more generally, as more data become available.     

Harpacticoid copepod communities of floating seaweed: controlling factors and implications for dispersal

The structure of harpacticoid copepod communities of floating detached intertidal algae was studied by surveys and experiments. Seaweed clumps were sampled from Faxaflói, western Iceland, at various distances from the shore (0.2–14 km). Altogether 20 species of harpacticoids were found in these clumps, most of them typical phytal species. Multidimensional scaling ordination (MDS) separated the samples into three distinct clusters in which the communities were significantly different from each other. The distance from shore did not seem to affect the community structure. However, diversity and abundance of harpacticoids increased with size and complexity of the clumps. Experimental Ascophyllum nodosum fronds placed 300–600 m offshore were rapidly colonised by harpacticoids. Density and species diversity were, after 20 days afloat, higher than in the free-floating clumps. Species of the genus Harpacticus were successful colonisers, in particular Harpacticus chelifer and Harpacticus uniremis. In a survey of macrofauna (0.5-mm sieve) on floating algae clumps from various locations around Iceland, H. chelifer was present in all clumps studied, up to 117 km offshore. Its sex ratio was skewed towards females, often with a high ratio of ovigerous females. There was, however, no evidence that this species is using the algal clumps as a platform for reproduction.     

Origin and dispersal of Potamophylax cingulatus (Trichoptera: Limnephilidae) in Iceland

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The influence of feeding strategies on trace element bioaccumulation in copepods (Calanoida)

Copepods are the most numerous taxonomic group in marine mesozooplankton communities. These planktonic organisms have an essential role in the function of marine trophic webs, as they are the link between phytoplankton and secondary consumers.The concentrations of 20 essential and non-essential trace elements were investigated by Inductively Coupled Plasma – Mass Spectrometry (ICP-MS) in calanoid copepods with different feeding behaviors. The sampling was performed in the Northwestern Mediterranean Sea, at the border between the Northern Tyrrhenian Sea and the Ligurian Sea (Italy). Aluminum, iron, zinc and copper were present in the highest concentrations in herbivorous calanoids (Temora stylifera, Nannocalanus minor, Neocalanus gracilis). Conversely, the nonessential element – arsenic – and the essential elements selenium and molybdenum – were present in the highest levels in carnivores (Pontella mediterranea, Candacia ethiopica). In the omnivorous copepod Centropages typicus, metal concentrations were found at an intermediate level between herbivores and carnivores, reflecting the importance of dietary pathways in metal intake and bioaccumulation.Finally, the bioaccumulation factors (BAFs), expressed as a ratio of the total metal levels in copepods compared to the seawater metal levels, were as follows: herbivores (83699) > omnivores (47855) > carnivores (41648).     

Effects of oxygen,carbon dioxide,and nitrogen on hatching behavior and hatching time in the katydid,Eobiana engelhardti subtropica Bey-Bienko (Orthoptera: Tettigoniidae)

The trigger for the hatching behavior and determination of hatching time of the katydids, Eobiana engelhardti subtropica (Orthoptera: Tettigoniidae) have been shown to be influenced by light–dark signals or temperature. In this study, I investigated the effects of oxygen, carbon dioxide, and nitrogen on the hatching behavior and hatching time of the katydid. Eggs rarely hatched under a constant temperature of 25°C and hatched sporadically at a constant temperature of 15°C under continuous light in the air. However, when eggs were exposed to 100% oxygen or a mixture of oxygen and nitrogen (2:1 or 1:1), hatching occurred within a few seconds. Hatching behavior was directly triggered by high concentrations of oxygen. It was inhibited by exposure to 100% carbon dioxide, 100% nitrogen, or a mixture of oxygen and nitrogen (1:2). The hatching time, determined by the temperature fall (transfer from 25°C to 15°C), was delayed by these gases, and was reset by the transfer back of eggs to the air. This suggests the existence of a time-measuring mechanism that is triggered by the transfer of eggs to the air. These results, indicating that hatching behavior was directly triggered by high concentrations of oxygen and that hatching time was set by the transfer from carbon dioxide or nitrogen to the air, are new findings to the best of my knowledge.     

Zooplankton on the move: first results on the quantification of dispersal of zooplankton in a set of interconnected ponds

In systems of interconnected ponds or lakes, the dispersal of zooplankton may be mediated by the active population component, with rivulets and overflows functioning as dispersal pathways and the dispersal being unidirectional. Such systems offer the possibility to study the impact of dispersal rates on local population dynamics and community structure, and provide opportunities to quantify dispersal in the field in a straightforward manner. In this study, dispersal of active zooplankton populations among interconnected ponds was quantified directly in the field by sampling the small waterways connecting the ponds. The number of dispersing zooplankton sampled in connecting elements was on average high (almost 7000 ind h^–1). However, the contribution of dispersing individuals to total population size in the target ponds was very limited (< 1% 24 h^–1.). Only a weak diel pattern in dispersal rates was observed.     

Expression of a dispersal trait in a guild of mites colonizing transient habitats

Summary Dispersal as a means of escape from deteriorating habitats is of particular ecological relevance for organisms such as certain astigmatic mites that colonize habitats which vary unpredictably in space and time. The mites meet these ecological challenges by a facultative dispersal morph, the heteromorphic deutonymph, also called hypopus. The appearance or absence of hypopodes in natural populations is attributable to two fundamentally different, albeit interacting, causes. Genetic polymorphism for the propensity to induce a hypopus provides for heritable variation within the population and allows selection to favor or eliminate certain genotypes. The genotypic composition of a population reflects selection forces previously acting on the population. But it holds no predictive power. Rather, it adapts the population to cope with unpredictably varying living conditions because it ensures instantaneous fit of certain genotypes of the population (those displaying hypopus-free development) to favorable (moist) environmental conditions, and others (those expressing a hypopus) to detrimental (dry) conditions. In contrast, environmentally cued inducibility allows mites to anticipate food quality inasmuch as it allows each genotype of the population to adjust its development rapidly to impending adversity or benefit. Inducibility occurs by means of a developmental switching mechanism and leads either to a developmental pathway with a hypopus or else one without. The expression of a hypopus depends on interacting genetic and environmental (trophic) factors. High levels of additive genetic variation combine with considerable genetic-trophical interaction (comprising a threshold for phenotypic expression of the trait) to control hypopus induction. The results are consistent with a variable threshold whose level depends on diet quality. Different trophic conditions set the threshold at different points along the genetic scale resulting in different proportions of hypopus-forming and directly developing individuals within the population. The threshold, therefore, converts the concealed continuous genetic variation underlying the trait into a discontinuous response of the mite.     

不同扰动生境中动物对酸苔菜种子的捕食和散布

有效的种子散布是木本植物形成入侵种需要经历的过程之一,但在预测入侵种时却常常被忽略.紫金牛科东方紫金牛(Ardisia elliptica)原产热带亚洲而在北美成为入侵植物,分布在云南南部的其同属种酸苔菜(A.solariacea)与之具有相似的生物学特征.本文以酸苔菜为研究对象,于2004年12月至次年2月分别在人为干扰轻的野象谷和人为干扰重的植物园进行酸苔菜的种子散布及捕食研究,试图了解生境变化对其种子散布和种子捕食的影响.结果表明,酸苔菜在两地的种子散布者均为白喉冠鹎(Alophoixus pallidus)、黑冠黄鹎(Pycnonotus melanicterus)和灰眼短脚鹎(Iole propinqua),但3种食果实鸟类的组成比例、拜访行为、频率及种子捕食者的影响在两地均不相同.人为干扰轻的野象谷生境中白喉冠鹎、黑冠黄鹎与灰眼短脚鹎的拜访频率分别为25%、32%和26%.取食后的第一次停栖地点有4%在10 m以外;人为干扰重的植物园生境中3种鸟的拜访频率分别为67%、8%、5%,取食后的第一次停栖地点有26%在10 m以外.人工摆放种子试验表明,地面上种子捕食者主要是啮齿类;在两生境中种子捕食率均较低(2-6%),但野象谷生境中种子捕食率仍显著高于植物园生境.野象谷生境中种子还受到象鼻虫幼虫的危害,危害率为17.9±3.5%(n=512);而植物园生境中未发现种子被象鼻虫危害(n=489).干扰对生境中的动物组成及行为造成了明显影响,并可能通过种子散布与捕食的改变而间接影响与其有密切关系植物的种群动态.     

Dormancy, dispersal and the survival of cyclopoid copepods (Cyclopoida, Copepoda) in a lowland floodplain

1. The survival of cyclopoid copepods was investigated in a floodplain for 2 years where flooding occurred during the cold season. The cyclopoid community was studied in three waterbodies with distinct hydroperiods: a permanent pond connected to the flooded area during inundation, a temporary pool that is part of the flooded area and an isolated temporary pool.
2. Field studies, including data obtained from samples of water, sediment and soil, showed the overall predominance of species with a summer diapause over those with a winter diapause or without diapause. Emergence of cyclopoid copepods at the onset of flooding, examined using emergence traps and data from recently filled or still isolated temporary pools, showed that some species can survive several months of drying.
3. The ability of the diapausing fourth copepodid stages of Cyclops strenuus and C. insignis , the two cyclopoids most abundant during winter and spring flooding, to survive terrestrial conditions was tested in laboratory experiments. Both species survived for several months, but rates differed among the species. A higher percentage of C. strenuus survived for a longer period, possibly explaining why this species was relatively more abundant in more temporary habitats.
4. Both dormancy and dispersal facilitated survival of cyclopoid copepods in transient habitats connected to each other during flood periods. Dormancy was the most important survival strategy, whereas dispersal could be more important following prolonged periods without flooding.     

Habitat use and dispersal of the invasive crayfish Procambarus clarkii in ephemeral water bodies of Portugal

We used radio-telemetry to analyze habitat use and dispersal of the invasive crayfish Procambarus clarkii in the extreme environmental conditions of a temporary river in southern Portugal. The rationale of this study was that an understanding of the properties that make this species a successful invader can be of help for the conservation of freshwater biodiversity. The results showed that: (1) crayfish displaced at a similar rate (ranging 2.5-38 m d^-1) as other European populations; (2) movement was faster at night; (3) crayfish movement had no clear environmental correlates, except for water temperature; (4) peaks of more intense locomotion were intercalated with longer periods of slow or null speed; and (5) P. clarkii did not excavate burrows in the study habitat, but took refuge under boulders and most often occupied complex microhabitats (e.g., vegetated sections of the river). The conclusion was that, notwithstanding their extreme conditions, ephemeral water bodies in southern Europe are highly susceptible to invasion by P. clarkii.     

Variation in dispersal mortality and dispersal propensity among individuals: the effects of age, sex and resource availability

1.  Dispersal of individuals between habitat patches depends on both the propensity to emigrate from a patch and the ability to survive inter-patch movement. Environmental factors and individual characteristics have been shown to influence dispersal rates but separating the effects of emigration and dispersal mortality on dispersal can often be difficult. In this study, we use a soil mite laboratory system to investigate factors affecting emigration and dispersal mortality.
2.  We tested the movement of different age groups in two-patch systems with different inter-patch distances. Differences in immigration among age groups were primarily driven by differences in emigration but dispersal mortality was greater for some groups. Immigration declined with increasing inter-patch distance, which was due to increasing dispersal mortality and decreasing emigration.
3.  In a second experiment, we compared the dispersal of recently matured males and females and tested the impact of food availability during the developmental period on their dispersal. Dispersal was found to be male biased but there was no significant sex bias in dispersal mortality. There was some evidence that food availability could affect emigration and dispersal mortality.
4.  These results demonstrate that both emigration and dispersal mortality can be affected by factors such as individual age and resource availability. Understanding these effects is likely to be important for predicting the fitness costs and population consequences of dispersal.     

The relative importance of dispersal and local processes in structuring phytoplankton communities in a set of highly interconnected ponds

1. The recognition that both local and regional processes act together in shaping local communities makes determining their relative roles in natural communities central to understanding patterns in community structure. 2. We investigated the relative influence of these processes on the phytoplankton communities of a highly interconnected pond system. We sampled the phytoplankton communities of 28 ponds concurrently with 20 local environmental variables. 3. We found that phytoplankton community variation, in terms of both phytoplankton community composition (PCC) and diversity, was only significantly explained by local environmental variables. These were mainly associated with the contrasting clear‐water and turbid ecological states of the shallow ponds studied. Clear‐water conditions favoured only a few taxa, resulting in a significantly lower taxon diversity and richness under these conditions. 4. The failure to explain variation in PCC by a dispersal model based on the water flow between ponds points at very effective species sorting. This is attributed to the high population turn‐over rates and sensitivity to environmental conditions of phytoplankton communities. Some evidence was found, however, that dispersal influences local communities through mass effects between neighbouring ponds. 5. Overall, our results emphasize both the strong selection pressure that components of the food web exert on phytoplankton communities and the high potential of these communities to respond to such environmental change, thereby effectively opposing the homogenizing effects of continuous dispersal.     

Avian seed dispersal and seedling distribution of the endangered tree species,Taxus chinensis,in patchy habitats

Background: There is limited understanding about bird dispersal behaviour and seedling distribution of endangered tree species in patchy environments, although these processes are important for plant species persistence.

Aims: We tested how patch features affected bird behaviour and seed dispersal, and thus seedling distribution of the endangered Chinese yew tree (Taxus chinensis).

Methods: In the present study, we combined field data of bird dispersal behaviour and GIS-based information to elucidate the influence of spatial features of habitat patches on bird dispersal behaviour, and the resulting effects on the seedling distribution of the endangered Chinese yew in two patchy habitats.

Results: Our results showed that the only seed source patch could attract eight bird species for dispersal at the two sites. Post-foraging movements of bird dispersers was strongly related to both topography and the relative locations of habitat patches. Yew seedlings aggregated only at the seed source and bamboo recruitment patches, which was affect by both the spatial distribution of recruitment patches and patch use by dispersers.

Conclusions: Our results emphasise that bamboo patches in both patchy environments provide the necessary conditions for germination of yew seeds, and the post-foraging behaviour of dispersers determines seed deposited in these patches. Our study highlights the importance of the dispersal behaviour of frugivorous birds in the successful regeneration and colonisation of yew populations in patchy habitats.     

Locomotion in copepods: pattern of movements and energetics of Cyclops

The cost of swimming in copepods has generally been estimated through the application of fluid dynamics theory to data on velocity and acceleration obtained by means of movies. It has also been estimated through the changes in fat content of copepods after sustained swimming (i.e. vertical migration). However, the range of estimated costs of locomotion is exceedingly large (from 0.1% to 95% of total metabolism). This communication studies the pattern of swimming movements and the work done by Cyclops, using high speed cinematographic techniques. The contribution of swimming to the energy expenditure of the individual is estimated, and consideration of the possible role of rubber-like proteins in the cuticle of copepods is made.     

Patterns of ecological diversification in thelodonts

Here we explore the spatial, temporal and phylogenetic patterns of ecological diversification for the entire clade of thelodonts, one of the earliest groups of vertebrates and longest lasting of the Palaeozoic agnathans in the fossil record. Parsimony and maximum‐likelihood methods are used to reconstruct ancestral states of their geographical distributions, habitats and lifestyles. Our results support the concept that thelodonts originated during the Middle?–Late Ordovician probably in marine open waters of Laurasia, with a demersal lifestyle on hard substrates being the ancestral condition for the whole clade. Later, thelodonts underwent a complex ecological diversification and palaeobiogeographical history, comparable in many aspects to those of some major groups of living fishes. Different modes of life evolved repeatedly and a wide range of habitats were colonized by distinct groups, including deep waters and brackish marine and/or freshwater environments. Diadromous strategies presumably appeared on nine different occasions. The palaeobiogeographical history of thelodonts reveals significant differences in the dispersal potential of some major groups. Dispersal of thelodontiforms entailed displacements over long distances and the crossing of deep‐water biogeographical barriers, whereas those of furcacaudiforms were always limited to areas interconnected by shallow platforms. We propose that the evolution of pelagic larval stages in thelodontiforms might explain this biogeographical pattern and could satisfactorily account for the greater evolutionary success of this group.