Colour polymorphism in the intertidal snail Littorina rudis Maton

The tictors affecting the variation of shell colour morph frequencies of the intertidal snail Littorina rudis Maton have been examined using ordination and correlation techniques. The major gradient underlying variation in shell colour is related to habitat type and structure. Dark, patterned shells are characteristic of cliff and salt marsh habitats whilst light, unpatterned shells reach high frequencies on boulder shores. Colour morphs do not differ significantly in shell thickness and between-shore variation in colour morph frequencies cannot be directly attributed to habitat dilterences in the risk of shell injury, but to other factors associated with habitat type and structure, e.g. parasitism. It is suggested that the maintenance of shell colour polymorphism in L. rudis is probably mediated by selection on pleiotropic characters rather than shell colour alone.     

The taxonomy of the Littorina saxatilis speicies-complex, with particular reference to the systemaitc status of Littorina patula Jeffrys

Variation in shell shape and penis morphology of Littorina rudis Maton is examined using data from all parts of Britain. The shell shape variation within populations of L. rudis is shown to account for Liltorina patula Jeffrys at the only site where the latter species was recorded. In addition, the shell shape of I,. rudis varies with exposure, individuals on exposed shores having a relatively larger aperture than those on sheltered shores. Wave action and desiccation are considered the most likely factors maintaining this variation. The penis morphology of L. rudis varies within and between shores to an extent that renders the use of this character invalid for distinguishing L. patula from L. rudis. The radulae of adults of L. rudis, L. patula and Littorina nigrolineata (Gray) are similar in structure having blunt cusps, whilst adult Littorina neglecta Bean and juvenile L. rudis have pointed cusps. The possibility of a neotenous origin of L. neglecta from L. rudis is discussed. On the evidence presented here it is suggested that L. patula must be regarded as a synonym of L. rudis.     

The role of selection in the colour polymorphism of Littorina rudis Maton and Littorina arcana Hannaford-Ellis (Prosobranchia: Littorinidae)

Forty-one mixed samples of winkles containing the closely related species, Littorina rudis and L. arcana , were collected from different parts of the British Isles. Littorina rudis was the more ubiquitous species, with L. arcana being more-or-less confined to vertical cliffs and rocks. The frequencies of different shell colour patterns were determined for both species in each sample. Several colour morphs were diagnostic of one or other of the species over large areas, which confirms that these are separate species. The frequencies, in the two species, of the two commonest morphs, brown and fawn, were strongly correlated; in several other morphs their presence and absence in the two species was significantly associated; the levels of phenotypic diversity in the two species were also correlated. These patterns could not have arisen if the variation between shores was the result of genetic drift or founder effects, so some form of selection is implicated. In the brown morph there is some evidence for frequency-dependent selection, and it is suggested that the polymorphism may be maintained by visual predators through apostatic selection.     

Reproductive biology of two sympatric skates in the south‐west Atlantic: Psammobatis rudis and Psammobatis normani

The total lengths ( L _T) at which 50% were mature of Psammobatis rudis and Psammobatis normani , in the south‐west Atlantic were: P. rudis , 428 mm for males and 414 mm for females and P. normani , 443 mm for males and 403 mm for females. Clasper length in mature males was greater in P. normani than in P. rudis , whereas oviducal gland width was not different between species. Females of P. normani with egg cases were found in every month sampled, and in January, March, April and July in P. rudis , although insufficient samples were available to identify peak oviposition times. Geographic variation in size frequency and maturity were found. The effects of oceanographic conditions and fishing pressure are discussed. Size at 50% maturity in both species was >74% of the maximum L _T, indicating late sexual maturity and low potential stock recovery rate.     

Relative size of the foot of two species of Littorina on a rocky shore in Wales

Populations of the gastropods Littorina rudis and L. arcana on the rocky shore at Westdale, Pembroke, South Wales, have been investigated. Foot area has been related to shell length. It is shown that there are differences in the areas of the expanded foot of different groups of snails: those from more exposed sites have a relatively larger foot. Littorina arcana achieves the greatest foot area of all, and it extends on to the most exposed rocks investigated, where L. rudis is absent. It is suggested that this may be important in ecological separation of the two species. It is noted that there is still uncertainty over the taxonomic identity of what is referred to here as 'Littorina rudis' , and further work is in progress.     

The taxonomy of some British Littorina species, with notes on their reproduction (Mollusca: Prosobranchia)

The rough periwinkle Littorina saxatilis is considered, in current literature, to consist of various sympatric "subspecies" and "varieties". These, it has been suggested, are incipient species and are evidence for sympatric speciation.
This paper shows that in Wales the "saxatilis" aggregate consists of four separate, fully sympatric species: L. rudis, patula, nigrolineata and neglecta . Differences between these species include proportions, sculpture and size of the shell, range of colour patterns, the anatomy of the genitalia, and isozyme patterns. The possible evidence for incipient species is discussed and rejected.
The diversity of penis form amongst the British winkles is remarkable, and it is suggested that in these sympatric species the structure of the penis could be of importance in species recognition.
The four species occupy different zones on the shore and three certainly differ also in their methods of reproduction: L. rudis is ovoviviparous and contains embryos throughout the year, patula is ovoviviparous but contains embryos only in winter, nigrolineata is oviparous. The reproduction and dispersal methods of all British winkles are reviewed. The evidence available supports the view that non-planktonic dispersal is an adaptation which, in species whose shell colour varies considerably from one locality to another, enables each population to become adapted to its local environmental conditions. Ovoviviparity occurs in high-shore non-planktonic species, and may be an adaptation to avoid the dangers of egg desiccation. Planktonic dispersal is found in species whose shell colour does not vary as much as in non-planktonic ones. In these species, it is argued that a common planktonic pool enables a rapid re-colonization in the case of local decrease in population numbers, and also the rapid dispersal of successful mutations.     

The effects of exposure and predation on the shell of two British winkles

Changes in shell size and shell shape of the two British winkles Littorina nigrolineata and L. rudis were studied in relation to exposure and to crab-size. In both species, shells from exposed shores are smaller and more globose than those from sheltered shores. Also, in rudis of exposed shores the mouth is relatively wider. In shores of equally sheltered conditions, shells are bigger at those localities where crabs are large than at those localities where they are small. The largest shells are found in those localities where it is extremely sheltered, and the crabs are very large.
It is argued that on exposed shores, small shells are favoured because they have more possibilities than large ones to shelter in crevices and in barnacle interspaces, from the impact of winds and waves. A globose shell could accommodate more foot muscle and thus enable a stronger adherence to the rock; and an increased mouth diameter would increase the area of foot adherence to the rock. On sheltered shores, on the other hand, large, narrow-mouthed shells are favoured because they discourage crab predation, large crabs being abundant mainly on sheltered shores.
The possible significance of shell size and shape in relation to zonation is discussed, in view of the different predatory and physical conditions which prevail in different zones of the shore, and the different shell specializations which these conditions would require.     

盐地碱蓬盐沼与相邻泥质滩涂湿地迁徙期鸻鹬类的群落组成及行为差异

盐地碱蓬(Suaeda salsa)盐沼湿地是黄渤海地区河口区域的重要湿地类型, 是水鸟迁徙停歇期的重要栖息地。本研究以辽河口国家级自然保护区为研究地点, 通过对盐地碱蓬盐沼湿地和相邻泥质滩涂两个固定样区连续三年的水鸟组成调查和行为观察, 分析盐地碱蓬盐沼湿地在鸻鹬类多样性维持和栖息地利用中的作用。共记录到鸻鹬类水鸟28种6,348只次, 其中盐地碱蓬湿地记录到4科13种, 泥质滩涂记录到4科27种, 泥质滩涂的物种多样性显著高于盐地碱蓬盐沼湿地。此外, 盐地碱蓬盐沼湿地与相邻的泥质滩涂的鸻鹬类鸟类群落组成存在较大差异, 盐地碱蓬盐沼湿地的鸟类群落组成以体型较大的大杓鹬(Numenius madagascariensis)、白腰杓鹬(N. arquata)、灰鸻(Pluvialis squatarola)等为主, 而泥质滩涂以环颈鸻(Charadrius alexandrinus)、黑腹滨鹬(Calidris alpina)等小型鸻鹬类为主, 这说明两种生境在鸟类多样性维持中具有不同的功能。行为分析发现, 泥质滩涂中栖息鸟类的主要行为为取食(58.71%-93.26%), 而盐地碱蓬盐沼湿地鸟类的行为既包括较大比例的取食, 也包括休息, 特别是在春季迁徙期。这进一步说明, 两种生境在水鸟的栖息地利用中具有一定的生态功能差异。尽管盐地碱蓬盐沼湿地记录到的鸟类物种数和数量均低于泥质滩涂, 但是, 两种生境中存在较大比例的共同分布物种, 这说明其生态功能具有较强的生态互补性, 二者作为一种独特的湿地景观组合, 在鸻鹬类迁徙停歇期的栖息地利用和物种多样性维持中发挥着不可替代的作用。     

Conservation value of Codium tenue habitat for the endangered Knysna seahorse Hippocampus capensis

We investigated the density and size structure of the endangered Knysna seahorse Hippocampus capensis in the macroalga Codium tenue at three sites in a residential marina estate in the Knysna Estuary, South Africa, over an 18 month period from March 2017 to August 2018. Seahorses were present in C. tenue throughout the year. Average (± SE) densities of 0.33 (± 0.03) and 0.23 (± 0.03) seahorses per kg of C. tenue were found for 2017 and 2018 respectively. Density did not vary across sites but varied significantly across months with a decreasing trend from summer to spring. The average (± SE) adult seahorse height was 75.16 mm (± 0.63 mm) and 69.09 mm (± 0.64 mm) and mass was 2.05 g (± 0.04 g) and 1.65 g (± 0.04 g) for males and females, respectively, and an even sex ratio was observed throughout the sampling period. Height varied across months, with significantly smaller seahorses found in June and August of 2017. Tail length was highly correlated with height and therefore can serve as a potential proxy for determining the height of H. capensis. This study showed that C. tenue is consistently utilised by Knysna seahorse and should therefore be considered an important habitat to protect and facilitate the ongoing conservation of this endangered seahorse species.     

WADERS (CHARADRII) AND OTHER WATERBIRDS AT LANGEBAAN LAGOON,SOUTH AFRICA, 1975–1986

Summary

Underhill, L. G. 1987. Waders (Charadrii) and other waterbirds at Langebaan Lagoon, South Africa, 1975–1986. Osrrich 58: 145–155.

Langebaan Lagoon was surveyed for waterbirds at midsummer and midwinter between 1975 and 1986. The median number of birds counted in summer was 37 500, of which 34 500 were waders (93% of the waders being Palaearctic migrants). Curlew Sandpiper (59,2%), Grey Plover (10,5%), Sanderling (8,3%), Knot (8,1%) and Turnstone (5,7%) were the major components of the summer wader population. The median number of birds in winter was 10 500, of which 4500 were flamingos and 4000 waders. For Palaearctic waders, the median winter population was 11,5% of the median summer population, but varied between 2,5% and 30,1%. For species of wader which breed in the Taimyr Peninsula, Siberia, a three-year cycle in the numbers of birds overwintering was detected, with large numbers in 1977, 1980, 1983 and 1986. Birds feeding on invertebrates consumed 126,9 kJ m^?2 yr^?1, or 24% of the total production of invertebrates. Greater Flamingos have a major impact on energy cycling at Langebaan Lagoon, accounting for 73,3% of the winter energy consumption by the avifauna. Langebaan Lagoon is the most important wetland for waders in South Africa, accounting for about 10% of the coastal wader population of South Africa. At midsummer, about 0,5% of the total wader population of the East Atlantic Flyway is at Langebaan Lagoon, which ranks about 20th in importance for waders on the flyway.     

Genetic differentiation in the molluscan species Littorina rudis and Littorina arcana (Prosobranchia: Littorinidae)

Six populations of Littorina rudis and ihree of L. arcana were screened for elcctrophorctically dctci table variation at 21 enzyme loci. These species reproduce by ovoviviparity and oviparily respectively, and both are members of the L. saxatilis species complex. The mean coefficient of genetic identity between the two species was high (0.957) and no diagnostic loci were observed. Allele frequency differences are maintained in sympatric populations of the two species. Intraspecific values of/ ranged from 0.944 to 0.995 and both species showed marked inter-population genetic heterogeneity, a feature consistent with their limited powers of dispersal. Heterozygosity is high in all populations of both species. Populations of I. rudis have higher mean heterozygosities (= 0.153) then L. arcane ( H = 0.132).     

Disturbance and contrasting patterns of population structure in the brachiopod Terebratulina septentrionalis (Couthouy) from two subtidal habitats

The population structures of Terebratulina septentrionalis (Couthouy) from exposed upper rock surface and semi-cryptic rock wall habitats at 33 m depth in the Gulf of Maine differ. Over a 3-yr period, population densities were consistently higher in rock wall habitats. Although both populations were dominated by juveniles (1–4 mm shell length), size-frequency distributions constructed from upper rock surface and rock wall populations were significantly different, as a result of a greater frequency of large brachiopods (> 20 mm shell length) in rock wall populations. Prominent modes occurred at 14–15 mm shell length in upper surface populations and at 19–20 mm length in rock wall populations. Recruitment was higher in rock wall habitats where ambient light intensities were significantly lower than on upper rock surfaces. Differences in recruitment are either the result of larval selection for shaded rock walls or differential juvenile mortality between habitats. The larvae of Terebratulina settle on a diverse array of substrata. These include bedrock, sandy polychaete tubes and algae in upper surface habitats and bedrock, calcareous polychaete tubes, and ascidians in rock wall habitats. Individuals attached to polychaete tubes and algae in upper surface habitats do not attain large body size (> 13 mm shell length). It is suggested that these differences in population structure reflect the greater intensity of disturbance in upper surface habitats. For example, the cod, Gadus morhua (Linnaeus), ingests brachiopods attached to algae and polychaete tubes in this habitat. Gastropod predation affects brachiopods in upper surface habitats but not in rock wall habitats. Predation by gastropods and asteroids is not size-specific. These results are consistent with the hypothesis that predation contributed to the decline in the abundance and diversity of articulate brachiopods since the Mesozoic, and suggest that the restriction of recent populations to semi-cryptic rock wall and crevice habitats is, in part, controlled by disturbance.     

Effects of exotic <Emphasis Type="Italic">Spartina alterniflora</Emphasis> on the habitat patch associations of breeding saltmarsh birds at Chongming Dongtan in the Yangtze River estuary,China

Smooth cordgrass (Spartina alterniflora) is one of the most invasive exotic plants of saltmarshes worldwide. To understand the effects of smooth cordgrass invasion on the habitat use and selection by breeding saltmarsh birds, we compared species number and abundance of breeding birds in native reed (Phragmites australis) and smooth cordgrass-invaded habitats (reed-cordgrass mixed habitats and cordgrass monocultures) at Chongming Dongtan in the Yangtze River estuary, China. We further examined the similarity of bird communities in different habitats and habitat selection by dominant bird species. For saltmarsh generalists, species number and abundance did not differ among the habitats. For saltmarsh specialists, species number and abundance did not differ in reed monocultures and reed-cordgrass mixed habitats but were significantly lower in cordgrass monocultures than in reed monocultures and reed-cordgrass mixed habitats. ANOSIM indicated that the difference in bird communities was larger between cordgrass monocultures and the habitats with reed than between the habitats with reed. The saltmarsh specialists preferred reed monocultures, while saltmarsh generalists avoided reed monocultures. Most species indicated no selection (neither preferred nor avoided) on reed-cordgrass mixed habitats, and no species preferred the cordgrass monocultures. The use of cordgrass monocultures by the common saltmarsh birds was negatively related to their body size. This study suggests that the spread of exotic smooth cordgrass has greatly affected the species composition and structure of local bird communities and has been especially disadvantageous to the saltmarsh specialists.     

Importance of matrix permeability and quantity of core habitat for persistence of a threatened saltmarsh bird

Species habitat preferences can be obscured when individuals have been recorded in non‐core habitats because of dispersal, spillover effects or spatial errors in observation locations. Disentangling the direct effects of the habitats species are observed in from the effects of proximity to other nearby habitats is especially challenging in fragmented landscapes, as many fragmentation metrics are correlated and it is difficult to prove independent effects. In this paper we addressed this issue by comparing a number of models based on predefined ecological theories. We compared models based on quantity of core habitat surrounding observations, proximity to core habitat, or a combination of the two to explain the observed distribution of the saltmarsh inhabiting white‐fronted chat (Epthianura albifrons) in coastal New South Wales, Australia. Proximity to core habitat was considered as either Euclidean distance or cost distance, and models were assessed using Akaike's information criterion and the area under the receiver operator characteristic curve on 10 random subsets of the data. We found that all models performed similarly, with the combination of cost distance and the quantity of saltmarsh performing better, but not significantly so. We compared the advantages and disadvantages of different models and also present a model averaged result. Our models suggested that the majority of saltmarshes in New South Wales were too small to have a large effect on probability of occurrence. As climate change is expected to further reduce the amount of available saltmarsh through continued mangrove incursion, coastal populations of the white‐fronted chat are expected to come under increasing threat. The conversion of grasslands to urban areas may also increase the effective distance between different populations of the species and reduce gene flow and rescue effects.     

The Knysna elephants: a population study conducted using faecal DNA

The elephants of the Knysna region continue to survive, despite fears that there was only a single surviving female. Their range is larger than previously believed, and includes the Afromontane forest and mountain fynbos. The five individuals detected in this study were all females, and share a single mitochondrial DNA control region haplotype with individuals from Addo Elephant National Park. At least two of these elephants appear to be first‐order relatives, and the others may be part of a single matrilineal group. The genetic diversity detected is lower than that found in most African savanna populations, but is higher than that found at Addo, where individuals represent the descendents of a severe population size bottleneck. Levels of genetic diversity are more similar to those detected at Kruger National Park, suggesting that the Knysna elephants represent a remnant of the once widespread populations of South Africa.     

Sediment characteristics of a restored saltmarsh and mudflat in a managed realignment scheme in Southeast England

Sediment characteristics and vegetation composition were measured in a restored and natural saltmarsh and mudflat at Wallasea Island managed realignment scheme (Essex, UK) from January to December 2007. The similar sediment characteristics in the restored and natural mudflat indicated that the sediment in the restored mudflat was approaching natural conditions. However, the sediment characteristics in the restored saltmarsh were not becoming similar to those in the natural saltmarsh. The sediment moisture content, organic matter content and porosity were lower while the sediment bulk density, salinity and pH were higher in the restored compared to the natural saltmarsh. The dissimilarities were mainly due to differences in the vegetation abundance and organic matter content. Although, 18 months after restoration the restored saltmarsh was only sparsely vegetated and there was no net change in the sediment characteristics, the occurrence of Salicornia europaea L. demonstrated that pioneer saltmarsh vegetation establishment preceded the development of sediment characteristics.     

Linkages between seagrass,mangrove and saltmarsh as fish habitat in the Botany Bay estuary,New South Wales

Shallow-water vegetated estuarine habitats, notably seagrass, mangrove and saltmarsh, are known to be important habitats for many species of small or juvenile fish in temperate Australia. However, the movement of fish between these habitats is poorly understood, and yet critical to the management of the estuarine fisheries resource. We installed a series of buoyant pop nets in adjacent stands of seagrass, mangrove and saltmarsh in order to determine how relative abundance of fishes varied through lunar cycles. Nets were released in all habitats at the peak of the monthly spring tide for 12 months, and in the seagrass habitat at the peak of the neap tide also. The assemblage of fish in each habitat differed during the spring tides. The seagrass assemblage differed between spring and neap tide, with the neap tide assemblage showing greater abundances of fish, particularly those species which visited the adjacent habitats when inundated during spring tides. The result supports the hypothesis that fish move from the seagrass to the adjacent mangrove and saltmarsh during spring tides, taking advantage of high abundances of zooplankton, and use seagrass as a refuge during lower tides. The restoration and preservation of mangrove and saltmarsh utility as fish habitat may in some situations be linked to the proximity of available seagrass.     

Microgeographic variation in allozyme and shell characters in Littorina saxatilis Olivi (Prosobranchia: Littorinidae)

Twenty-three enzymes and five shell parameters were screened in 11 subpopulations of Littorina saxatilis Olivi (= L. rudis Maton) occupying different habitats over a 1 km stretch of coastline. Shell morphology varied considerably and consistently with respect to degree of exposure, and since there is evidence that such morphology is at least partly under genetic control, it is likely that natural selection selects particular genotypes at particular locations. There was significant allozyme heterogeneity between neighbouring subpopulations, sometimes only metres apart, but little of the allozyme variability could be related directly to environmental pressures. Thus, with the exception of the Odh locus, the considerable morphological differentiation between snails from exposed and sheltered sites was not reflected in differentiation of those genes coding for electrophoretically assayed enzymes. At the Odh locus, virtually all the genetic differentiation between subpopulations was attributable to differentiation between habitat types. Two loci, Sod-1 and Aat-1, showed highly significant genetic disequilibrium, and possible reasons for this are explored. The population structure as assessed electrophoretically accords well with the stepping-stone model which permits greater differentiation of neighbouring populations than the island model, and which seems realistic in the ovoviviparously reproducing L. saxatilis, where the greater part of gene flow is likely to occur through the occasional migration of adults between contiguous populations.     

Character displacement, frequency-dependent selection, and divergence of shell colour in land snails Mandarina (Pulmonata)

Endemic land snails of the genus Mandarina of the oceanic Bonin Islands offer an exceptional example of habitat and character divergence among closely related species. In this study, microhabitat differences between sympatric ground-dwelling species were studied by distinguishing habitats on the basis of vegetation and types of litter. In all sites where two ground species coexisted, segregation occurred with each species showing preference for the microhabitat in which they were found. When they were in sympatry, one species was predominant in relatively wet and sheltered sites and the other in relatively dry and exposed sites. Although most species can live in both types of habitat, occupation by one species is inhibited by occupation by another. This suggests that competitive interaction between sympatric species caused segregation. Except for populations that have undergone interspecific hybridization, no examples were found of sympatric populations of two ground species sharing a similar shell colour. Species that were predominant in relatively wet and sheltered sites possessed shells with dark coloration and their colour patterns were mostly of one type. Species that were predominant in relatively dry and exposed sites possessed shells with bright coloration and their color patterns were polymorphic. Most populations from areas in which single species were distributed had shells with medium coloration. Microhabitat differentiation between sympatric species possibly caused diversification of shell colour, because bright shells are advantageous in sites where snails are largely exposed, and dark shells are advantageous in sites in where they are mostly sheltered from sunlight. In addition, frequency-dependent selection by predators hunting by sight may have operated to maintain colour polymorphism in the populations which are restricted to exposed habitats by competition with other sympatric species. This reveals the importance of interaction among closely related species as a cause of diversification in ecological and morphological traits.     

Measuring, mapping and modelling: an integrated approach to the management of mangrove and saltmarsh in the Minnamurra River estuary, southeast Australia

Mangrove and saltmarsh ecosystems appear particularly vulnerable to the impacts of climate change, and their effective management will require forecasts of how these wetland habitats are likely to respond to sea-level rise through the twenty-first century. We describe a preliminary study of a small stand of mangrove and saltmarsh that involves measuring of elevation change and accretion, mapping of wetland communities, and modelling of their potential response to sea-level rise. The wetland occurs on the banks of the Minnamurra River estuary in southern New South Wales and has been the focus of several studies over recent decades. The research includes empirical measurements of sedimentation at sites in both mangrove and saltmarsh vegetation using the surface elevation table-marker horizon technique. This is a site at which mapping has been undertaken to delineate the extent of each vegetation community from a time-series of aerial photographs using geographical information systems; the gradual incursion of the mangrove, Avicennia marina, into more landward saltmarsh communities, observed over past decades for many systems in southeastern Australia, has continued into the twenty-first century. The observed patterns of change are compared with simulations of how this wetland system might respond to future sea-level rise, adopting several different approaches and the upper and lower bounds of Intergovernmental Panel on Climate Change sea-level rise projections. The model results show considerable variability in response depending on the parameters adopted. We advocate the need for the integration of these three approaches, measuring, mapping and modelling, as a basis for future management and adaptation. Our study demonstrates the considerable opportunities to refine the data input and model outputs as part of adaptive management, as more sophisticated technologies and data become available.