The occurrence and ecology of a marine hydrobiid mudsnail in the southern hemisphere: the Knysna Estuary, South Africa

Earlier published records of the gastropod Hydrobia from the warm temperate Knysna Lagoon on the Indian Ocean coast of South Africa are in error. Nevertheless, an animal hitherto identified as an Assiminea (Assimineidae) or as a Tomichia (Pomatiopsidae) is in fact a Hydrobia s.l., and it does occur intertidally within the Knysna system. The same species also occurs in salt pans in the cool temperate Great Berg Estuary on the Atlantic coast of South Africa. This extends the distribution of confirmed species of the dominant coastal hydrobiid mudsnails of the northern hemisphere into the southern hemisphere. H. knysnaensis (Krauss) occurs patchily at low density near the head of the Knysna Estuary in waters of low salinity. This appears to be a suboptimal habitat, and it is suggested that Hydrobia is restricted to this zone as a result of interference competition from potamidid mudwhelks and ocypodid crabs. The prevalence of potamidids and ocypodids in the tropics and in the temperate southern hemisphere may account for the rareness or absence of hydrobiids in these areas. Unlike its equally intertidal relative, the widespread North Atlantic/Arctic H. ulvae, H. knysnaensis develops directly like other Hydrobia.     

Are nest sites a limiting resource for speleophilic nesting fishes in sand dominated temporarily open/closed estuaries?

Southern African temporarily open/closed estuaries are often characterized by sand dominated mouth regions, a feature in contrast to many permanently open estuaries of the region which regularly have substantial rocky areas. Here we explore how habitat characteristics may reduce nesting opportunities and potentially explain population trends for a common estuarine fish species, the speckled sand goby Psammogobius knysnaensis, in many temporarily open/closed estuaries in South Africa. The present study is the first to document that P. knysnaensis is a speleophilic nester, utilizing the underside of rocks and other hard substrata for egg attachment. Furthermore, our results suggest that ideal nesting habitat for the speckled sand goby is potentially a limiting resource in sand-dominated temporarily open/closed estuaries. We postulate that this is a factor contributing to the relatively low numbers of adult P. knysnaensis found in sandy temporarily open/closed estuaries of the region, although the role of marine connectivity, food availability, predation pressure and other environmental factors in this regard still remain largely unassessed.     

WATER TEMPERATURE AND THE 1987 FISH KILL AT LAKE ST LUCIA ON THE SOUTH EASTERN COAST OF AFRICA

Summary

An unusually cold spell during the winter of 1987 caused a drop in water temperature at Lake St Lucia. This coupled with typical estuarine salinities in the lake resulted in a fish kill. The kill was one of the largest recorded in a South African estuarine environment and involved an estimated 250,000 fish comprising at least 21 species. Most fish that died belonged to small species. Densities of dead fish recorded along the lake's shoreline ranged from 0.1 to 16.6 fish per metre. It is concluded that the combination of low water temperature and near marine salinities was the major cause of the 1987 fish kill at Lake St Lucia.     

Monthly population density and structure patterns of an endangered seahorse Hippocampus capensis: a comparison between natural and artificial habitats

This study investigated aspects of the population ecology of the endangered Knysna seahorse Hippocampus capensis within different habitat types. High densities of H. capensis were found within artificial Reno mattress habitat, within the Knysna Estuary, South Africa. Monthly surveys at three sites were conducted from October 2015 to August 2016 to compare population densities of H. capensis in this artificial habitat with natural eel grass Zostera capensis habitat. Hippocampus capensis densities varied significantly across all sites and highest population densities were consistently observed within the Reno mattress habitat. Hippocampus capensis were also found to be significantly larger within the Reno mattress habitat and pooled data showed that males were significantly larger than females. The overall sex ratio for all three sites was female biased, although this varied across seasons at two sites. The findings suggest that artificial Reno mattresses provide novel habitat for this endangered species and consideration should be given to the usefulness of these structures in future conservation actions.     

Potential effects of physiological plastic responses to salinity on population networks of the estuarine crab <Emphasis Type="Italic">Chasmagnathus granulata</Emphasis>

Chasmagnathus granulata is a South American crab occurring in estuarine salt marshes of the Brazilian, Uruguayan and Argentine coasts. Life history is characterized by an export strategy of its larval stages. I reviewed information on experimental manipulation of salinity during embryonic and larval development (pre- and posthatching salinities), and on habitat characteristics of C. granulata in order to determine potential effects of larval response to salinity in the field and to suggest consequences for the population structure. Local populations are spread over coastal areas with different physical characteristics. Benthic phases occupy estuaries characterized by different patterns of salinity variation, and release larvae to coastal waters characterized by strong salinity gradients. The zoea 1 of C. granulata showed a strong acclimatory response to low salinity. This response operated only during the first weeks of development (during zoeae 1 and 2) since subsequent larval survival at low posthatching salinities was consistently low. Larvae developing at low salinity frequently followed a developmental pathway with five instead of four zoeal stages. The ability to acclimate and the variability in larval development (i.e. the existence of alternative developmental pathways) could be interpreted as a strategy to buffer environmental variability at spatial scales of local or population networks. Early survivorship and production of larvae may be relatively high across a rather wide range of variability in salinity (5–32‰). Plastic responses to low salinity would therefore contribute to maintain a certain degree of population connectivity and persistence regardless of habitat heterogeneity. Electronic Publication     

Suppression of the aquatic weed Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) by a leaf-cutting moth Parapoynx diminutalis Snellen (Lepidoptera: Crambidae) in Jozini Dam,South Africa

A classical biological control programme against Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae) was initiated in South Africa following the discovery of extensive mats in Jozini Dam in northern KwaZulu-Natal (KZN) in 2006. However, in mid-2008, high densities of a phytophagous insect, Parapoynx diminutalis Snellen (Lepidoptera: Crambidae) were discovered on the H. verticillata infestation, causing high levels of defoliation and dieback of the weed. In order to quantify the impact of the moth and to assess the long-term threat of H. verticillata to South African water bodies, a monitoring programme was initiated in 2013. Although P. diminutalis did not demonstrate a preference for H. verticillata over a native aquatic plant, Potamogeton schweinfurthii A. Bennett (Potamogetonaceae), moth populations followed a cyclical pattern of abundance, with rapid population increases coinciding with increases in the abundance of H. verticillata. High damage levels to H. verticillata were associated with high densities of immature P. diminutalis, which consistently led to population crashes of H. verticillata. Parapoynx diminutalis appears to have great potential to contribute to the management of H. verticillata in South Africa. However, the moth's oligophagous habits will probably preclude its intentional introduction into water bodies with H. verticillata where it does not already occur.     

FISH CONSERVATION IN SOUTH AFRICAN ESTUARIES: PRESSURES,PROBLEMS AND PROSPECTS

Summary

A review of the conservation status of fish in the estuarine environment around the South African coastline reveals that some species face serious problems associated either with habitat destruction, and its associated biological, physical and chemical components or exploitation. The 65 species considered fall into three categories; truly estuarine species, which are dependent on estuaries for their entire life; marine species dependent on estuaries during the juvenile phase of their life cycle; and marine species whose juveniles occur mainly in estuaries but are also found at sea. Included in the first two categories are 14 species of fish which are on the South African Red Data list. These comprise one species which is endangered, five which are vulnerable and eight which are rare. All groups are considered in relation to factors in estuaries which are affecting their populations. A conservation strategy is suggested for certain estuarine types or for specific estuaries which could ensure the survival of the entire range of estuarine faunas.     

Conservation of Hipparchia autonoe (Esper) (Lepidoptera: Nymphalidae), Natural Monument in South Korea

Hipparchia autonoe, designated Natural Monument no. 458, is a species inhabiting Mt. Halla on Jeju Island, which is the only habitat of H. autonoe in South Korea. Recently, this species is in danger of extinction as the population has been remarkably reduced because its host plants and nectar plants are gradually disappearing due to plant succession in its habitat. On the climbing road of Jeju Island Eorimok, H. autonoe used to be observed at and above 1300 m a.s.l. However, since Sasa quelpaertensis has recently expanded its habitat up to 1400 m, H. autonoe is now observed at and above 1500 m. In Mongolia, the population of H. autonoe seems quite stable as there are dense populations of a host plant and nectar plants. Accordingly it is judged that we can maintain a stable community for support of H. autonoe if we successfully manage the host plant and nectar plant communities by controlling the density of S. quelpaertensis using biological and physical control methods. Comparative analysis of DNA barcode region of COI (658 bp) was done to check the homogeneity and the genetic diversity of H. autonoe collected in Korea and Mongolia. The DNA sequence difference among individuals collected in South Korea was 0.0–0.2%, and in Mongolia 0.0–1.4%. This confirmed a 0.06–1.2% barcoding gap. We believe that this result will provide basic information useful to guide conservation of H. autonoe in South Korea.     

Fission-fusion and the evolution of hominin social systems

The course of hominin evolution has involved successive migrations towards higher absolute latitudes over the past three million years. Poorer habitat quality further from the equator has led to the necessity for groups occupying higher latitudes to live at lower population densities. Coupled with a trend towards increasing group size over this time period, this tendency towards expansion has led to exponential increases in the area requirements of hominin groups, and a concomitant need to adjust foraging patterns. The current analyses suggest that the development of increasingly complex, multi-level fission-fusion social systems could have freed hominins of the foraging constraints imposed by large group sizes and low population densities. Analyses of the fossil record suggest latitudinally-driven differences in area requirements of the australopithecines from East and South Africa, and African and Asian Homo erectus. In contrast, chronologically-driven differences appear between H. erectus as a whole and Homo heidelbergensis, and between H. heidelbergensis and the Neanderthals. These results are discussed in relation to studies of the foraging patterns of primates and hunter-gatherers.     

Biological effects of salinity gradient reversals in a southeast African estuarine lake

St Lucia Lake on the north coast of Natal, South Africa, has an area of 325 km² and is the largest estuarine complex in Africa. It consists of a 20 km tidal channel, averagingca. 400 m in width, linking the sea with the non-tidal lake which is H-shaped with a maximum length ofca. 40 km and width ofca. 20 km. Except during flood periods the depth of the lake does not exceed 2 m. The salinity gradient depends on evaporation, the configuration of the mouth and on the input of fresh water from four rivers which discharge into the northern and western areas of the lake. If fresh water input is high, the lake and much of the channel may be fresh. An intermediate stage features a normal salinity gradient while a third stage shows a reversed salinity gradient with salinities in excess of 100‰ in the upper reaches of the system. Changing salinities have marked effects on the biota. Aquatic macrophytes show cycles of appearance and disappearance depending on salinity tolerance and the presence of dormant stages. The resident benthic faunal species go through cycles of range expansion and contraction depending on prevailing salinities and recolonisation by dispersal phases. To date salinities in the southern part of the lake have approached, but not exceeded, lethal levels and this has therefore acted as a reservoir area. Catchment degradation and water abstraction are anticipated to exacerbate future salinity extremes. This has resulted in concern for the long term viability of this Ramsar site which has major southern African populations of hippopotamus and crocodile, provides breeding sites for South African Red Data water bird species and plays an important nursery role for marine fish and penaeid prawns.     

Effect of population density and presence of a potential competitor on the growth rate of the mud snail Hydrobia ulvae (Pennant)

The mud snail Hydrobia ulvae (Pennant) and the amphipod Corophium arenarium Crawford were used in a series of laboratory and field experiments to investigate the hypothesis that natural populations of such deposit-feeders live under conditions of resource limitation. Growth rates of Hydrobia confined at several different densities within cages in the field and in laboratory microcosms indicated that density-dependent reduction of growth rate and increased mortality occurred at densities found in natural populations. Similar experiments failed to provide any evidence that Corophium mortality or behaviour was modified by the presence of Hydrobia or members of its own species; nor did the presence of Corophium cause a reduction in growth rate of Hydrobia.     

The effects of salinity on swimming performance of two estuarine fishes,Fundulus heteroclitus and Fundulus majalis

Prolonged and high‐speed swimming performance measurements were used to explore the swimming abilities of two species of estuarine fishes, the mummichog Fundulus heteroclitus and the striped killifish Fundulus majalis, under different salinities. Critical swimming performance was significantly higher for F. majalis in high salinity than in low salinity, but no difference was observed in brief constant acceleration swimming trials in this species; however, the swimming performance of F. heteroclitus was not significantly affected by salinity changes, indicating that this species is well adapted to regular estuarine salinity oscillations. Fundulus majalis displayed higher swimming speeds than F. heteroclitus in both high and low salinities, and while this cannot be explained by their respective salinity preferences, the specific habitat preferences of F. majalis for sandy subtidal habitats and F. heteroclitus for vegetated marshes could explain the better swimming performance of F. majalis.     

Novel distribution pattern between coexisting sexual and obligate asexual variants of the true estuarine macroalga Ulva prolifera

Where sexual and asexual forms coexist within a species, the asexuals are often found to be prevalent in marginal habitats. This asexual distribution pattern has received evolutionary attention linked to the paradox of sex. In many marine species, there is a distributional trend of asexual modes being more common in lower salinity waters regarded as the ecogeographic marginal, being explained by negative effects of low salinities on sexual reproductive success. However, the distribution pattern of estuarine species recently adapted to low salinity waters has remained unknown. The brackish macroalga Ulva prolifera being a major benthic component of estuarine ecosystems includes a sexual variant and obligate asexual variants by means of motile spores. We examined the sexual–asexual distribution pattern of this alga along a salinity gradient in river estuaries. Surprisingly, opposite to the distributional trend of marine organisms, the results clearly showed the persistent predominance of sexuals in the lower salinity reaches than the asexuals. In addition, a frequent alternating of dioecious gametophytes and sporophytes in the sexual population was observed, suggesting the sexual reproductive process would be robustly performed in the low salinity waters. Considering U. prolifera had evolved from the ancestral marine species to become a true estuarine species of which the core habitat is the low salinity reaches, in a broad sense its sexual–asexual distribution pattern would be involved in asexual marginal occupations of the species range previously reported in other organisms. Based on the frozen niche variation model, we can give a concise explanation for the evolutionary process of this pattern; multiple asexuals with frozen genotypic variation had arisen from sexual ancestors undergoing low salinity adaptation and share the estuarine habitat with the sexuals at present.     

Modelling the effectiveness of contraception for controlling introduced populations of elephant in South Africa

Re‐introduced African elephant (Loxodonta africana Blumenbach) populations are growing at very high rates in many of southern Africa’s reserves, have attained densities higher than previously thought possible and may be exhibiting irruptive growth. Active management of such populations is necessary to prevent the potentially negative effects on habitat and biodiversity that are associated with elephant overpopulation. One potentially feasible method of elephant management is immunocontraception, but very little is known about the long‐term effectiveness of this method. Using demographic data from three South African elephant populations, we made model projections of the effects of contraception on population growth rates to determine whether contraception may be a feasible management tool for elephant. In comparison with noncontracepted populations, realistic reductions in population growth rate after 20 years of contraception were projected to be up to c. 64%, with 50% being a very feasible target. Through its ability to reduce population growth rates, immunocontraception should be an effective tool for preventing or minimizing irruption in elephants and, perhaps, other introduced ungulate species.     

Genetic diversity of <Emphasis Type="Italic">Bradysia difformis</Emphasis> (Sciaridae: Diptera) populations reflects movement of an invasive insect between forestry nurseries

The fungus gnat, Bradysia difformis (Sciaridae: Diptera) has recently been recorded for the first time from South Africa where it has been found in forestry nurseries. The presence of this insect in all the major forestry nurseries as the dominant and only sciarid species raises intriguing questions regarding its origin and population genetic structure. A 395 bp portion of the mitochondrial COI gene was analysed from B. difformis individuals collected from four nursery populations in South Africa and three nursery populations in Europe. Shared haplotypes between South African and European populations indicated a historical connection. South African populations showed high genetic diversity and low genetic differentiation. These patterns most likely reflect multiple and/or relatively large introductions of B. difformis into South Africa from its origin combined with subsequent and continued movement of plants between nurseries.     

Invasion history, habitat preferences and phenology of the invasive ladybird Harmonia axyridis in Belgium

Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is native to Asia, and was widely introduced as a biocontrol agent of aphids and coccids in Europe and North America. In Europe, H. axyridis is considered to be an invasive alien species because of its potential to disrupt native ladybird communities. Since 1999, the Belgian Ladybird Working Group mapped all Belgian Coccinellidae and recorded data on substratum plants and habitat. The first feral H. axyridis population in Belgium was recorded in 2001, but the expansion rate is decreasing because it now colonised the whole country. Recorded occupancy in Belgium showed an average rate of increase of 189% between 2002 and 2006. In Belgium, H. axyridis occurred in a wide range of habitats, including those of high conservation value. However, habitat and land cover analysis showed that H. axyridis is more frequently found in urbanised landscapes than in semi-natural landscapes. A niche overlap analysis based on plant use data showed that the potential to affect native species is higher for generalist, deciduous and coniferous tree ladybird species than for heathland and wetland specialist species. Phenology data showed that H. axyridis is able to reproduce later in the year than native species. Based on recorded distribution, ecology and phenology, we discuss the potential of H. axyridis to disrupt native ladybird assemblages in Belgium.     

Evidence of a landlocked reproducing population of the marine pejerrey Odontesthes argentinensis (Actinopterygii; Atherinopsidae)

In South America, the order Atheriniformes includes the monophyletic genus Odontesthes with 20 species that inhabit freshwater, estuarine and coastal environments. Pejerrey Odontesthes argentinensis is widely distributed in coastal and estuarine areas of the Atlantic Ocean and is known to foray into estuaries of river systems, particularly in conditions of elevated salinity. However, to our knowledge, a landlocked self-sustaining population has never been recorded. In this study, we examined the pejerrey population of Salada de Pedro Luro Lake (south-east of Buenos Aires Province, Argentina) to clarify its taxonomic identity. An integrative taxonomic analysis based on traditional meristic, landmark-based morphometrics and genetic techniques suggests that the Salada de Pedro Luro pejerrey population represents a novel case of physiological and morphological adaptation of a marine pejerrey species to a landlocked environment and emphasises the environmental plasticity of this group of fishes.     

Responses to simulated herbivory and water stress in two tropical C4 grasses

Summary The African grass Hyparrhenia rufa has established itself successfully in South American savannas (Llanos) and displaced dominant native grasses such as Trachypogon plumosus from the wetter and more fertile habitats. Several ecophysiological traits have been related to the higher competitive capacity of H. rufa. To further analyze the behavior of both species, their growth, biomass allocation, physiological and architectural responses to defoliation and water stress were compared under controlled conditions. Although total, aerial and underground biomass decreased under defoliation in both grasses, increases in clipped-leaf biomass and area compensated for defoliation in H. rufa but not in T. plumosus. This difference was due mainly to a higher proportion of assimilates being directed to leaf and tiller production and a higher leaf growth rate in the African grass as compared to T. plumosus, which showed incrased senescence under frequent defoliation. In both species, water stress ameliorated the effects of defoliation. The ability to compensate for defoliated biomass in H. rufa is possibly related to its long coevolution with large herbivores in its original African habitat and is apparently one of the causes of its success in Neotropical savannas.     

Ontogenetic and sex‐based differences in habitat preferences and site fidelity of White's seahorse Hippocampus whitei

The aim of this study was to determine and compare habitat preferences for male and female adult and juvenile White's seahorse Hippocampus whitei and assess their movements and site fidelity over 4 years. Data were collected from three sites along 1·5 km of estuarine shoreline in Port Stephens, New South Wales, Australia, from 2006 to 2009 using H. whitei that had been tagged with visible implant fluorescent elastomer. Relative availability of 12 habitats and habitat preferences of H. whitei was determined, based on the habitat that H. whitei used as a holdfast. Hippocampus whitei occurred in nine different habitats; adults preferred sponge and soft coral Dendronephthya australis habitats with no difference between male and female habitat preferences whilst juveniles preferred gorgonian Euplexaura sp. habitat. There was a significant preference by adults for D. australis colonies with height >40 cm and avoidance of colonies <20 cm. Neither adults nor juveniles used sand or the seagrasses Zostera muelleri subsp. capricorni and Halophila ovalis. Hippocampus whitei showed cryptic behaviour with c. 50% of adult sightings cryptic and c. 75% for juveniles with crypsis occurring predominantly in Sargassum sp. for adults and Euplexaura sp. habitat for juveniles. Within sites, females moved significantly longer distances (maximum of 70 m) than males (maximum of 38 m) over 20 months. Strong site fidelity was displayed by H. whitei with males persisting at the same site for up to 56 months and females for 49 months and no H. whitei moved between sites. The longest period that an H. whitei was recorded on the same holdfast was 17 months for a male and 10 months for a female. As this species displays strong site fidelity, specific habitat preferences and has a limited distribution, future management needs to minimize the risk of habitat disturbance as loss of key habitats could have a negative effect on species abundance and distribution.     

Macrofaunal Loss and Microhabitat Destruction: The Impact of Trampling in a Temperate Mangrove Forest, NSW Australia

Trampling paths are a feature of estuarine habitats in southeastern Australia. An experimental investigation quantified the impact of trampling over a 3 year period on the microhabitat features and macrofaunal assemblages in a temperate mangrove forest in New South Wales, Australia. The magnitude of the impact varied between 0 and 25 passes (representing a visit by 25 people), which reduced the biomass of the Bostrychia–Caloglossa algal association by 50%. The frequency of trampling varied between one and four, simulating a single and seasonal visits by a group of people to the mangrove forest. The main impact of trampling was an alteration to the microhabitat structures of the forest floor including a decrease in the number and vertical height of pneumatophores, an increase in the proportion of bent and broken pneumatophores in trampled lanes and a decrease in the biomass of the epiphytic Bostrychia–Caloglossa algal association. The macrofauna most impacted by trampling were the gastropods most commonly associated with the pneumatophores and algal assemblages and included Ophicardelus spp. and Assiminea buccinoides. Those gastropods least impacted by trampling were commonly associated with the surface of the sediment or were known to move over the surface and included Bembicium auratum and Cassidula zonata. There was little impact on the main burrowing crab, Heloecius cordiformis. There were some indirect impacts of trampling with the recruitment of Ophicardelus spp. Assiminea buccinoides and Salinator solida reduced in the subsequent year after trampling had ceased. People can create walking paths in mangrove forests that have long term effects on the habitat. An appropriate management emphasis may lie in maintaining the structural features of the habitat.