Sexual conflict and polyspermy under sperm-limited conditions : in situ evidence from field simulations with the free-spawning marine echinoid Evechinus chloroticus

For free‐spawning organisms that release gametes into the sea, sperm limitation (too few sperm to fertilize all eggs) is a major factor limiting reproductive success. Given such circumstances, the presence of several mechanisms to prevent polyspermy (too many sperm) may seem paradoxical; however, a growing body of data suggests that natural fertilization levels, though variable, can routinely be high. Under such conditions, polyspermy is much more likely. The tension between sperm limitation and polyspermy represents sexual conflict because males, in competing to fertilize as many eggs as possible, can impose lethal costs on eggs if multiple sperm gain entry. Here we present data for a marine invertebrate indicating high levels of polyspermy under sperm‐limited conditions. When the sea urchin Evechinus chloroticus was induced to spawn in situ, mean rates of polyspermy were 17.3% ±3.4%, and polyspermy was recorded at rates as high as 62.7%. Polyspermy was nearly always present, even when fertilization rates were <50%, confirming predictions that it should be present under sperm‐limited conditions. Both sperm limitation and polyspermy imposed substantial reproductive costs, and we conclude that both sexual conflict related to polyspermy and sperm limitation have been simultaneous strong selective forces shaping the evolution of reproductive traits in the sea.

Sensitivity of Antarctic marine invertebrates and microalgae to metal exposure

Negative impacts from contaminants have occurred in Antarctic marine ecosystems resulting from human activities. To improve risk assessment procedures and develop site-specific environmental quality guidelines and remediation targets, this study successfully developed novel toxicity testing methods to determine the sensitivity of Antarctic marine invertebrate and microalgal species to metals.

A biogeographically mixed, Middle Permian brachiopod fauna from the Baoshan Block, Western Yunnan, China

A small brachiopod fauna is described from the carbonate rocks of the basal Shazipo Formation of the Baoshan Block, western Yunnan, south-west China, including significant new ventral and dorsal internal morphological features of Cryptospirifer omeishanensis Huang. This fauna is regarded as Wordian (Middle Guadalupian, Middle Permian) because of the presence of Cryptospirifer omeishanensis Huang and associated fusulinids (Neoschwagerina craticulifera Zone). Palaeobiogeographically, the brachiopod fauna is of considerable interest because of its admixed nature characterized by typical warm-water Cathaysian elements intermingled with temperate Peri-Gondwanan taxa. This in turn is interpreted to indicate that the Baoshan Block may have been situated in an intermediate palaeogeographical position between Gondwanaland to the south and Cathaysia to the north during the Mid Permian and, as such, it probably furnished an important 'stepping stone' for the dispersal of Mid Permian eastern Tethyan marine invertebrate taxa (e.g. Cryptospirifer) to the western Tethys.

Permian Chonetoidea and Spiriferoidea of Australasia : Gondwanan relationships, provincialism, palaeobiogeography

Members of the Spiriferoidea are dominant in the Permian marine invertebrate faunas of Australia. Genera can be endemic to a particular province, demonstrate a wider Gondwanan and peri-Gondwanan distribution and several indicate a bipolar distribution. Australasian spiriferoids are included within the families Spiriferidae, Neospiriferidae, Trigonotretidae and the Spiriferellidae. Several genera and species are the largest spirferoids ever recorded. Few genera are shared between the Westralian and Austrazean provinces but a higher proportion of genera are shared between the Westralian province and the Cimmerian Realm.
Representatives of the Chonetoidea are a less common but significant element of the faunas. They were used over 20 years ago to define the Westralian and Austrazean provinces of Australasia – concepts that are in widespread use today. The Paratinan and Cimmerian provinces were also defined at that time despite difficulties in their definition. Through more recent studies the Cimmerian Province has been upgraded to a Realm while the Paratinan Province is more clearly defined for the earliest Permian, based on the chonetoids of Patagonia and western and eastern central Argentina.
Distribution of the various genera is best explained by an interplay of factors including surface and deeper oceanic currents, marine water temperatures and tectonic events such as the clockwise rotation of Gondwanan and the dispersal of the peri-Gondwanan Cimmerian terranes. Austrazean faunas developed in isolation under the influence of cooler and cold waters during the early Permian. Late Permian faunas demonstrated more widespread linkages.

Pennsylvanian brachiopods from the Geumcheon-Jangseong formation, Pyeongan supergroup, Taebaeksan Basin, Korea

We provide the first detailed systematic taxonomy and paleoecological investigation of late Paleozoic brachiopod faunas from Korea. Specifically, we focus on the brachiopods from the Geumcheon-Jangseong Formation, the lower part of the Pyeongan Supergroup in the Taebaeksan Basin. The formation yields a variety of marine invertebrate fossils, including brachiopods, molluscs, echinoderms, corals, fusulinids, and conodonts. Diverse brachiopods are described from six siliciclastic horizons of the formation at three localities, including 23 species belonging to 20 genera with two new species: Rhipidomella parva n. sp. and Stenoscisma wooi n. sp. Three brachiopod assemblages of the late Moscovian (Pennsylvanian) age are recognized based on their species compositions and stratigraphic distributions, namely the Choristites, Rhipidomella, and Hustedia assemblages. The brachiopod faunal composition varies within each assemblage as well as between the Assemblages, most likely reflecting local paleoenvironmental and hence paleoecological differences. The Choristites Assemblage includes relatively large brachiopods represented by Derbyia, Choristites, and Stenoscisma and may have inhabited open marine to partly restricted marine environments, whereas the Rhipidomella and Hustedia Assemblages consist of a small number of small-sized brachiopods living in lagoonal environments. The Choristites Assemblage shows a close affinity with Moscovian brachiopod assemblages in the eastern Paleo-Tethys regions, especially the Brachythyrina lata–Choristites yanghukouensis–Echinoconchus elegans Assemblage of North China, whereas the Rhipidomella and Hustedia assemblages both exhibit strong endemism.

The more pieces, the better the puzzle: sperm concentration increases gametic compatibility

The genetic benefits individuals receive from mate choice have been the focus of numerous studies, with several showing support for both intrinsic genetic benefits and compatibility effects on fertilization success and offspring viability. However, the robustness of these effects have rarely been tested across an ecologically relevant environmental gradient. In particular, sperm environment is a crucial factor determining fertilization success in many species, especially those with external fertilization. Here, we test the importance of sperm environment in mediating compatibility-based selection on fertilization using a factorial breeding design. We detected a significant intrinsic male effect on fertilization success at only one of four sperm concentrations. Compatibility effects were significant at the two highest sperm concentrations and, interestingly, the magnitude of the compatibility effect consistently increased with sperm concentration. This suggests that females are able to modify the probability of sperm-egg fusion as the amount of sperm available increases.

Invertebrate larval dynamics in seasonally closed estuaries

Estuarine benthic assemblages are often numerically dominated by polychaetes. The limits of these populations are determined by larval, and probably to a lesser extent adult movement. A previous study (Newton 1996), indicated that planktonic polychaete larvae were very abundant over the summer months in the Hopkins River; however, the identification and source of these larvae was not known. Defining the extent of a population, and therefore the likelihood of that population recovering following a perturbation, is crucial for effective estuarine management. This study investigated both the likely source of the larvae, (i.e. estuarine or marine) and the extent of larval dispersal within and between estuaries by addressing the following questions: Which taxa produced the planktonic larvae? Are these taxa resident estuarine species? Are the larvae of different taxa evenly distributed within the estuary or do physicochemical parameters or other factors influence their abundance? Are the same larvae found in other estuaries along the coast? and Is there exchange of these larval taxa with the marine environment and other estuaries? Larvae were identified and described by culturing commonly occurring planktonic larvae until adult characteristics appeared. The spionids, Carazziella victoriensis and Prionospio Tatura, numerically dominated the plankton in the Hopkins and the spionid, Orthoprionospio cirriformia was recorded from the Hopkins, Curdies and Gellibrand estuaries. Two spionids, Carazziella sp. and Polydora sp. were identified from tidal waters. Mouth status and physicochemical conditions (salinity, temperature and dissolved oxygen) were monitored in each estuary. Whereas the Merri and Gellibrand estuaries were predominantly stratified over the sampling period, the Curdies was more often well mixed and the Hopkins varied from well mixed to stratified. The duration of mouth opening and hence the opportunity for larval exchange also varied in each estuary. The Merri River was closed for 13.5% of days over the study period, the Gellibrand River for 18.4%, the Hopkins River for 49% and the Curdies River for 71.0%. The distributions of larvae at spatial scales of metres, 100s of metres and kilometres were investigated within a single estuary. While the same larvae, C. victoriensis, P. Tatura and bivalve larvae, were found along the length of the Hopkins estuary the abundances varied at different spatial scales suggesting different processes were influencing the distribution of P. Tatura larvae, and C. victoriensis and bivalve larvae. The distribution of larvae between several estuaries was investigated by monitoring meroplankton at two sites at the mouth of each of the four estuaries approximately monthly (except for winter months). Different meroplanktonic assemblages were found to distinguish each estuary. Further, C. victoriensis and P. Tatura larvae were only recorded in the Hopkins but larvae of the spionid, Orthoprionopio cirriformia were detected in the Hopkins, Curdies and Gellibrand estuaries. The extent of larval exchange with other estuaries and the marine environment was determined by monitoring tidal waters. Settlement trays were also deployed to determine if larvae were moving into estuaries and settling but not recruiting. P. tatura larvae were not detected in the tidal waters of any estuary and while C. victoriensis and O. cirriformia were found in both flood and ebb tides there was no evidence of movement of theses taxa to other estuaries. Larvae of the spionids, Carazziella sp. and Polydora sp., were found in tidal waters of each estuary but were rarely detected in the plankton within the estuaries. Neither species was found as an adult in background cores from any estuary, nor with the exception of a few individuals in the Merri, were they detected in settlement trays in any estuary. I conclude that the source of the larvae of C. victoriensis, P. Tatura and O. cirriformia is estuarine and while C. victoriensis, and O. cirriformia move in and outh of the source estuary in tidal waters there was no evidence for movement to other estuaries. The spionids, Carazziella sp. and Polydora sp were considered to be marine and while they moved in and out of estuaries in tidal waters they did not usually settle in the estuaries. The results of this study are a crucial first step in the development of ecological models to better understand dispersal in seasonally closed estuaries that are typical of southern Australia. This study emphasises the unique physicochemical characteristics and biological assemblages within these estuaries and the need for estuarine management to reflect these differences.

Invertebrate diversity of the unexplored marine western margin of Australia: taxonomy and implications for global biodiversity

However derived, predictions of global marine species diversity rely on existing real data. All methods, whether based on past rates of species descriptions, on expert opinion, on the fraction of undescribed species in samples collected, or on ratios between taxa in the taxonomic hierarchy, suffer the same limitation. Here we show that infaunal macrofauna (crustaceans and polychaetes) of the lower bathyal depth range are underrepresented among available data and documented results from Australia. The crustacean and polychaete fauna (only partially identified) of the bathyal continental margin of Western Australia comprised 805 species, representing a largely novel and endemic fauna. Overall, 94.6% of crustacean species were undescribed, while 72% of polychaete species were new to the Australian fauna, including all tanaidaceans, amphipods, and cumaceans, as well as most isopods. Most species were rare, and the species accumulation rate showed no sign of reaching an asymptote with increasing area sampled. Similar data are likely for the largely unexplored bathyal regions. This leads us to conclude that the numbers upon which extrapolations to larger areas are based are too low to provide confidence. The Southern Australian and Indo-West Pacific deep-sea regions contribute significantly to global species diversity. These regions and bathyal and abyssal habitats generally are extensive, but are so-far poorly sampled. They appear to be dominated by taxonomically poorly worked and species-rich taxa with limited distributions. The combination of high species richness among infaunal taxa-compared to better known taxa with larger individuals, higher endemism than presently acknowledged because of the presence of cryptic species, the low proportion of described species in these taxa, and the vast extent of unexplored bathyal and abyssal environments-will lead to further accumulation of new species as more and more deep sea regions are explored. It remains to be tested whether ratios of 10 or more undescribed to described species, found in this study for the dominant taxa and for the deep Southern Ocean and the Indo-West Pacific, are replicable in other areas. Our data and similar figures from other remote regions, and the lack of faunal overlap, suggest that Appeltans et al.'s (Current Biology 22:1-14, 2012) estimate that between one-third and two-thirds of the world's marine fauna is undescribed is low, and that Mora et al.'s (PLoS Biol 9(8):e1001127. doi:10.1371/journal.pbio.1001127, 2011) of 91% is more probable. We conclude that estimates of global species, however made, are based on limited data. © 2014 Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg.

Sperm environment affects offspring quality in broadcast spawning marine invertebrates

The provisioning of offspring can have far-reaching consequences for later life in a wide range of organisms and generally this provisioning is thought to be under maternal influence or control. In experiments with a broadcast-spawning ascidian, we found that the size of offspring was determined by egg size and the abundance of sperm present during fertilization. Larger eggs were fertilized at low sperm concentrations, whilst smaller eggs were successfully fertilized at high sperm concentrations. These differences in fertilized egg size resulted in differences in the development rate, hatching success and mean size of the subsequent larvae. Our results suggest that, in contrast to females that reproduce by other mating systems, free-spawning mothers lack some control over the provisioning of offspring. Furthermore, because males can alter the sperm environment, they can exert paternal (non-genetic) control over key offspring characteristics.

Evaluation of the effectiveness of marine protected areas in temperate waters of Australasia

An evaluation of the effectiveness of Marine Protected Areas (MPAs) in temperate waters of Australasia has been conducted for this thesis. The aim was to identify key elements needed in a strategy for establishment and management of MPA’s in temperate waters of Australasia. This aim was achieved by assessing how effective a sample of MPA’s has been in meeting the conservation objectives for their establishment and by identifying factors that have contributed to success or failure of the MPA’s in meeting these objectives. Particular attention was paid to the objectives of ecological sustainability and biodiversity preservation. A MPA for the purposes of this research was defined as an area of coastal or marine environment, with a substantial subtidal component, set aside by law primarily for conservation purposes. The study region encompassed the coastal zones of Victoria, Tasmania and South Australia (Australia) and New Zealand. The questions posed in order to address the aim of this thesis were; a) Have existing MPA’s been effective in achieving conservation objectives? b) What have been the important influences on effectiveness of existing MPA’s? c) What are the key elements required for implementation of effective MPA’s? The thesis is divided into three parts. Part I is a review of the literature on implementation and effectiveness of MPA’s. Part II presents a detailed evaluation of one MPA: Point Lonsdale Marine Reserve (PLMR), Victoria, Australia. Part in is an evaluation of a number of Australasian temperate MPA’s based on information provided in a survey of people involved in management of MPA’s, and from a variety of published and unpublished documents. The MPA’s are described, evidence about ecological effectiveness is presented and factors that have enhanced and limited the ability of these MPA’s to achieve conservation objectives are derived. A substantial amount of scientific evidence was found for increases in abundance, mean size and size range of fish and invertebrate populations within the boundaries of ‘no-take’ MPA’s, Some evidence was found for ‘spillover’ of adults and juveniles into adjacent fishing grounds. Ecological effects detected within ‘no-take’ MPA’s in Australasia matched those described in the literature. The abundance and mean size of a number of previously exploited species have increased, migration into adjacent fishing grounds has been documented, and species richness has increased in at least one MPA. The PLMR was established primarily to protect the scientifically significant intertidal rock platforms. The results of the case study suggest that this objective has been achieved. Opinions about effectiveness were obtained for 28 MPA’s. Of these 19 were considered to be achieving some objectives and 10 were considered to be performing well in terms of overall ecological effectiveness. Positive effects on biodiversity were generally assumed as a result of reduction of damaging anthropogenic effects on habitat. Many questionnaire respondents noted an increase in community awareness about and support for marine conservation as a result of proclamation of MPA’s, Overall, the results support the value of MPA’s for sustainability of fish stocks and preservation of biodiversity, but there is substantial doubt over whether some of the MPA’s are too small to maintain benefits in the long-term. ‘No-take’ MPA’s, particularly those more remote from the impacts of human activities, have been the most effective in achieving objectives. A number of interacting factors important to eventual success of MPA’s were identified. The most important enhancing factors identified for the PLMR were physical attributes that limit the extent of human use and a strong conservation ethic amongst many of the visitors to this marine reserve. Limiting factors were far more numerous. Of most concern is the inadequacy of at-site management. Almost half of visitors to the area were not aware of its marine reserve status, despite the fact many were frequent visitors. The need for better educational and interpretative material on-site is highlighted by the results of the PLMR visitor survey. A total of 56 factors that have enhanced effectiveness and 46 factors that have limited effectiveness of Australasian temperate MPA’s were identified. A number of factors were important in more than one MPA and this was used to derive a set of critical, or key, factors. For example, a conclusion of all three approaches used in this study is that failure to implement day-to-day management through lack of resources was a major constraint on effectiveness. The importance of MPA’s to marine conservation depends, in part, on how well they are managed. The key factors that influence MPA effectiveness were used as the basis for derivation of the main requirements for implementation of MPA’s that will be capable of meeting the objectives for their establishment. The most important needs are: • that ‘no-take’ areas surrounded by buffer zones form the basis for a system of MPA’s; • that a high level of protection is bestowed by legislation and regulations; • that a minimum size be set for the ‘no-take’ core areas; • that the selection of sites for MPA’s takes into account land-based impacts; • that institutional arrangements are developed specifically for MPA’s; • that funding for MPA’s is increased to enable effective management; • that day-to-day management is implemented in all MPA’s, with enforcement and education programs as priority areas; • that a monitoring program for one or more MPA in each 'State' is established to provide evidence of ecological effects of reservation; • that public and stakeholder involvement in development and implementation of MPA’s is encouraged as this will influence the degree of public support and compliance; • that community-based programs to educate the general public, stakeholders, the media and decision-makers about the value of MPA’s are essential; and • that measures to reduce financial impact on affected stakeholders be implemented.

Ocean currents influence the genetic structure of an intertidal mollusc in southeastern Australia – implications for predicting the movement of passive dispersers across a marine biogeographic barrier

Major disjunctions among marine communities in southeastern Australia have been well documented, although explanations for biogeographic structuring remain uncertain. Converging ocean currents, environmental gradients, and habitat discontinuities have been hypothesized as likely drivers of structuring in many species, although the extent to which species are affected appears largely dependent on specific life histories and ecologies. Understanding these relationships is critical to the management of native and invasive species, and the preservation of evolutionary processes that shape biodiversity in this region. In this study we test the direct influence of ocean currents on the genetic structure of a passive disperser across a major biogeographic barrier. Donax deltoides (Veneroida: Donacidae) is an intertidal, soft-sediment mollusc and an ideal surrogate for testing this relationship, given its lack of habitat constraints in this region, and its immense dispersal potential driven by year-long spawning and long-lived planktonic larvae. We assessed allele frequencies at 10 polymorphic microsatellite loci across 11 sample locations spanning the barrier region and identified genetic structure consistent with the major ocean currents of southeastern Australia. Analysis of mitochondrial DNA sequence data indicated no evidence of genetic structuring, but signatures of a species range expansion corresponding with historical inundations of the Bassian Isthmus. Our results indicate that ocean currents are likely to be the most influential factor affecting the genetic structure of D. deltoides and a likely physical barrier for passive dispersing marine fauna generally in southeastern Australia.

Use of anthropogenic sea floor structures by Australian fur seals: potential positive ecological impacts of marine industrial development?

Human-induced changes to habitats can have deleterious effects on many species that occupy them. However, some species can adapt and even benefit from such modifications. Artificial reefs have long been used to provide habitat for invertebrate communities and promote local fish populations. With the increasing demand for energy resources within ocean systems, there has been an expansion of infrastructure in near-shore benthic environments which function as de facto artificial reefs. Little is known of their use by marine mammals. In this study, the influence of anthropogenic sea floor structures (pipelines, cable routes, wells and shipwrecks) on the foraging locations of 36 adult female Australian fur seals (Arctocephalus pusillus doriferus) was investigated. For 9 (25%) of the individuals, distance to anthropogenic sea floor structures was the most important factor in determining the location of intensive foraging activity. Whereas the influence of anthropogenic sea floor structures on foraging locations was not related to age and mass, it was positively related to flipper length/standard length (a factor which can affect manoeuvrability). A total of 26 (72%) individuals tracked with GPS were recorded spending time in the vicinity of structures (from <1% to >75% of the foraging trip duration) with pipelines and cable routes being the most frequented. No relationships were found between the amount of time spent frequenting anthropogenic structures and individual characteristics. More than a third (35%) of animals foraging near anthropogenic sea floor structures visited more than one type of structure. These results further highlight potentially beneficial ecological outcomes of marine industrial development.

Drought survival strategies, dispersal potential and persistence of invertebrate species in an intermittent stream landscape

Intermittent stream systems create a mosaic of aquatic habitat that changes through time, potentially challenging freshwater invertebrate dispersal. Invertebrates inhabiting these mosaics may show stronger dispersal capacity than those in perennial stream systems. To relate different combinations of dispersal and drought survival strategies to species persistence, we compared the distribution and dispersal potential of six invertebrate species across all streams in a montane landscape where drying is becoming increasingly frequent and prolonged. Invertebrates were collected from seventeen streams in the Victoria Range, Grampians National Park, Victoria, Australia. The species analysed were as follows: the caddisflies Lectrides varians Moseley (Leptoceridae) and Agapetus sp. (Glossosomatidae); the mayflies Nousia AV1 and Koorrnonga AV3 (Leptophlebiidae); the water penny beetle Sclerocyphon sp. (Psephenidae); and a freshwater crayfish Geocharax sp. nov. 1 (Parastacidae). These species were widespread in the streams and varied in their dispersal and drought survival strategies. The distribution of each species across the Victoria Range, their drought responses and within-stream habitat associations were determined. Hypotheses of the dispersal capacity and population structure for each species were developed and compared to four models of gene flow: Death Valley Model (DVM), Stream Hierarchy Model (SHM), Headwater Model (HM) or panmixia (PAN). Molecular genetic methods were then used to infer population structure and dispersal capacity for each species. The large caddisfly Lectrides resisted drought through aestivation and was panmictic (PAN) indicating strong dispersal capacity. Conversely, the small caddisfly Agapetus relied on perennially flowing reaches and gene flow was limited to short distances among stream headwaters, resembling the HM. Both mayflies depended on perennial surface water during drying and showed evidence of gene flow among streams: Koorrnonga mainly dispersed along stream channels within catchments, resembling the SHM, whereas Nousia appeared to disperse across land by adult flight. Sclerocyphon relied on perennial water to survive drying and showed an unusual pattern of genetic structure that indicated limited dispersal but did not resemble any of the models. Geocharax survived drought through aestivation or residence in perennial pools, and high levels of genetic structure indicated limited dispersal among streams, resembling the DVM. Despite good knowledge of species' drought survival strategies, the population structure of four species differed from predictions. Dispersal capacity varied strongly among species; most species were poor dispersers and only one species showed panmixia. Therefore, intermittent stream species may not necessarily be better dispersers than those in perennial streams. Species showing strong drought resistance strategies differed in dispersal capacity. Knowledge of life-history characteristics, distribution and refuge use does not necessarily enable successful prediction of invertebrate dispersal pathways or population structure. Dispersal among intermittent streams may be restricted to relatively short distances (km) for most invertebrate species. Thus, frequent drought refuges (perennial water) that provide strong connectivity to subpopulations through stream flow (hydrological dispersal), or continuous terrestrial vegetation (flight dispersal), will be critical to maintain genetic diversity, adaptability and population persistence.