Pyura Stolonifera Research Articles

A morphogenetic characterisation of a potentially dominant African marine species in Europe

Studying the early stages of the introduction of non-indigenous species (NIS) is crucial as it allows immediate management actions aimed at preventing NIS spread at a time when these actions are more likely to be effective. Recent species introductions present unique opportunities to study key aspects of the invasion process. However, comprehensive information on how and when NIS are first introduced remains rare. We assessed the characteristics of the introduction of a member of the widespread Pyura stolonifera species group (Chordata, Tunicata) in Europe. These ascidians are important bioengineer species that can attain amongst the highest benthic biomass per surface area ever reported. We collected introduced individuals on the coast of the northwestern Iberian Peninsula, an area with high shipping traffic and one of the world’s most important mussel farming regions. The specimens were analysed using taxonomic and genetic tools. Both field surveys and taxonomic analysis showed that one of the African members of the species group, Pyura herdmani, has recently been introduced to Europe and, although it does not yet form the large aggregates found in the native range, it is already well-established. Genetic data revealed that only the northwest African lineage of P. herdmani has been introduced to Europe. Considering the low dispersal capabilities of the early life-history stages of P. herdmani, the introduction of this species into Europe can only be explained via human mediated transport of NIS. In order to prevent any detrimental effects on both native biota and/or economically-important anthropogenic activities, regular monitoring of recent NIS introductions is urgently required.

Contemporary climate change hinders hybrid performance of ecologically dominant marine invertebrates.

Human activities alter patterns of biodiversity, particularly through species extinctions and range shifts. Two of these activities are human mediated transfer of species and contemporary climate change, and both allow previously isolated genotypes to come into contact and hybridize, potentially altering speciation rates. Hybrids have been shown to survive environmental conditions not tolerated by either parent, suggesting that, under some circumstances, hybrids may be able to expand their ranges and perform well under rapidly changing conditions. However, studies assessing how hybridization influences contemporary range shifts are scarce. We performed crosses on Pyura herdmani and Pyura stolonifera (Chordata, Tunicata), two closely related marine invertebrate species that are ecologically dominant and can hybridize. These sister species live in sympatry along the coasts of southern Africa, but one has a disjunct distribution that includes northern hemisphere sites. We experimentally assessed the performance of hybrid and parental crosses using different temperature regimes, including temperatures predicted under future climate change scenarios. We found that hybrids showed lower performance than parental crosses at the experimental temperatures, suggesting that hybrids are unlikely to expand their ranges to new environments. In turn, we found that the more widespread species performed better at a wide array of temperatures, indicating that this parental species may cope better with future conditions. This study illustrates how offspring fitness may provide key insights to predict range expansions and how contemporary climate change may mediate both the ability of hybrids to expand their ranges and the occurrence of speciation as a result of hybridization.

A short note on heavy cunjevoi growth observed on commercial scallops (Pecten fumatus)

ABSTRACTCommercial scallops (Pecten fumatus) were collected from Bass Strait, Australia from 41 dredge tows. Of these, four dredges undertaken in February 2016 at 46 m depth returned scallops that were covered by ascidians of the Pyura stolonifera species complex, commonly known as cunjevoi. There were no obvious signs of immediate ill health in the scallops, with meat being assessed as normal quality and all scallops requiring force to separate the shells. Ascidian-encrusted scallops were significantly smaller, and previous tows in the same location ten and eight months earlier returned far fewer scallops with clean shells and no signs of ascidians. This suggests that both both scallop and ascidian recruitment and growth occurred during the period between sampling events. Future research combining laboratory experiments and field observations is recommended to understand this relationship and its potential impacts on scallop populations.

Sand-mediated divergence between shallow reef communities on horizontal and vertical substrata in the western Indian Ocean

Distinctions are rarely made between vertical and horizontal surfaces when assessing reef community composition, yet physical differences are expected because of hydrodynamic differences and sediment accumulation on flat surfaces. As sand often diminishes biotic cover, we hypothesised that vertical surfaces will support a greater biomass but have lower diversity due to domination by a few species. To test this, we quantified sessile communities on vertical and horizontal surfaces at three sites in the Delagoa Bioregion on the east coast of South Africa. Community composition consistently differed: vertical communities were dominated by various filter feeders, especially the ascidian Pyura stolonifera, whereas those on horizontal reef comprised a mixture of filter feeders and various algae. The total number of species and all diversity metrics were significantly greater for horizontal reef surfaces. Contrastingly, Simpson’s dominance and biomass were significantly greater for vertical reef surfaces. Percentage cover of sand explained much of the variation in community composition whereas depth did not. Small-scale topographic differences in substratum orientation associated with differences in sand inundation will therefore influence both α and β diversity. Coastal developments and activities that alter sand movements and delivery to the coastal zone are therefore likely to have a profound influence on the maintenance and diversity of shallow subtidal communities.

Riverine subsidies for inshore filter-feeder communities: potential influences on trophic patterns among bioregions

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 498:13-26 (2014) - DOI: https://doi.org/10.3354/meps10634 Riverine subsidies for inshore filter-feeder communities: potential influences on trophic patterns among bioregions Sean N. Porter1,4,*, Sven Kaehler2, George M. Branch1, Kerry J. Sink3 1Marine Research Institute, Zoology Department, University of Cape Town, Rondebosch 7701, South Africa 2Department of Botany, Rhodes University, Grahamstown 6140, South Africa 3South African National Biodiversity Institute, Claremont 7735, South Africa 4Present address: Oceanographic Research Institute, PO Box 10712, Marine Parade 4056, South Africa *Corresponding author: sporter@ori.org.za ABSTRACT: In the Natal Bioregion of eastern South Africa, biomass of marine subtidal filter feeders is particularly high and makes a central contribution to distinguishing this bioregion from adjacent ones. We analysed the trophic role of riverine suspended particulate organic matter (POM) and the extent to which subsidies from rivers may explain this high filter-feeder biomass. Using carbon, nitrogen and sulphur isotopes, and a 3-end-member Bayesian-mixing model, we determined (1) the proportions of various primary producers contributing to the inshore POM pool and (2) the relative amounts of marine seaweed, pelagic POM and riverine POM assimilated by filter-feeder communities at various distances from 4 river mouths during austral summer and winter. Of the inshore POM pool available to filter feeders, riverine POM contributed 17 to 62%, pelagic POM 18 to 77% and seaweed 6 to 53%. The contributions of riverine POM to inshore POM declined significantly with distance from river mouth, but were unaffected by season or river size. Most material assimilated by filter feeders was of marine origin, notably seaweed detritus (39-62%), but with a noteworthy uptake of riverine POM (9-33%). Only small seasonal differences (<10%) and no biologically meaningful spatial trends were detected in the proportional assimilation of the 3 food sources by filter feeders. Although important, the trophic contribution of riverine POM may be subordinate to other factors such as turbidity and productivity in explaining the high biomass of filter feeders. Collectively, however, these river-associated factors are likely to explain the contrasts in trophic organisation among marine bioregions. KEY WORDS: Rivers · Isotope · Sulphur · Seaweed · Perna perna · Pyura stolonifera Full text in pdf format PreviousNextCite this article as: Porter SN, Kaehler S, Branch GM, Sink KJ (2014) Riverine subsidies for inshore filter-feeder communities: potential influences on trophic patterns among bioregions. Mar Ecol Prog Ser 498:13-26. https://doi.org/10.3354/meps10634 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 498. Online publication date: February 17, 2014 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2014 Inter-Research.

Cryptic diversity in coastal Australasia: a morphological and mitonuclear genetic analysis of habitat-forming sibling species

Cryptic diversity represents a major challenge to the accurate assessment of biodiversity, but the combined use of genetic and morphological analyses has proven to be a powerful approach to detect it. This is especially important for groups for which genetic information is not yet available. Here, we studied the highly conspicuous habitat-forming Pyura stolonifera species complex (Tunicata), which, as has recently been revealed, shows surprising levels of cryptic diversity, but whose systematics and biogeographical patterns in Australasia nonetheless remain poorly understood. We first present detailed taxonomic information of all the species associated with the P.?stolonifera species complex. We then proceed to describe the results of an exhaustive survey that included south-east Australia, Tasmania, and New Zealand. Subsequently, we present morphological and mitonuclear genetic analysis of two unresolved lineages that comprise the species Pyura praeputialis and a species that is formally described here (Pyura doppelgangera sp. nov.). Although the ranges of these two species overlap on mainland Australia, we found no sites at which both species live in sympatry, and there was no morphological or genetic evidence of hybridization. Taken together, the present study illustrates the usefulness of a combined morphogenetic approach in unravelling overlooked marine diversity in a relatively well-studied region.

Habitat-Associations of Turban Snails on Intertidal and Subtidal Rocky Reefs

Patchiness of habitat has important influences on distributions and abundances of organisms. Given the increasing threat of loss and alteration of habitats due to pressures associated with humans, there is a need for ecologists to understand species' requirements for habitat and to predict changes to taxa under various future environmental conditions. This study tested hypotheses about the generality of patterns described for one species of marine intertidal turban snail for a different, yet closely-related species in subtidal habitats along the coast of New South Wales, Australia. These two closely-related species live in similar habitats, yet under quite different conditions, which provided an opportunity to investigate how similar types of habitats influence patterns of distribution, abundance and size-structure in intertidal versus subtidal environments. For each species, there were similar associations between biogenically structured habitat and densities. The intertidal species, Turbo undulates, were more abundant, with greater proportions of small individuals in habitats formed by the canopy-forming alga, Hormosira banksii, the solitary ascidian, Pyura stolonifera or the turfing red alga, Corallina officinalis compared to simple habitat (bare rock). Similarly, more Turbo torquatus were found in biogenically structured subtidal habitat, i.e. canopy-forming algae, Ecklonia radiata, mixed algal communities (‘fringe’), or turfing red algae (Corallina officinalis and Amphiroa aniceps) than where habitat is simple (barrens). Small T. torquatus were more abundant in areas of turf and ‘fringe’, while large snails were more abundant in areas of kelp and barrens. These patterns were found at each location sampled (i.e. eight intertidal and two subtidal rocky reefs) and at all times of sampling, across each environment. This study highlighted the consistent influence of biogenically structured habitats on the distribution, abundance and size-structure of intertidal and subtidal turban snails and forms a basis for increasing the understanding of the potential underlying processes causing such patterns.

Environmental Conditions During Mass Mortalities of the AscidianPyura stolonifera(Heller) in the Tsitsikamma Marine Protected Area

Mass mortalities of the ascidian Pyura stolonifera occurred along the Tsitsikamma coast in May 1991 and again in February 2012, following infection of large proportions of the population with a white microbial growth. P. stolonifera appeared to be the only species affected, with the outer tunic the main site of infection. No signs of pollution or algal blooms were observed, but large variations in climatic conditions were associated with both events. The 1991 mortalities occurred during a warm spell in autumn, when there was a rapid increase in the daily air temperatures. Similarly, the February 2012 event occurred after abnormally high air temperatures of January 2012 declined sharply due to a cold-water upwelling event in early February. These findings are compared with those of other marine disease and mortality studies.

Environmental conditions during mass mortalities of the ascidian Pyura stolonifera (Heller) in the Tsitsikamma Marine Protected area

Mass mortalities of the ascidian Pyura stolonifera occurred along the Tsitsikamma coast in May 1991 and again in February 2012, following infection of large proportions of the population with a white microbial growth. P. stolonifera appeared to be the only species affected, with the outer tunic the main site of infection. No signs of pollution or algal blooms were observed, but large variations in climatic conditions were associated with both events. The 1991 mortalities occurred during a warm spell in autumn, when there was a rapid increase in the daily air temperatures. Similarly, the February 2012 event occurred after abnormally high air temperatures of January 2012 declined sharply due to a cold-water upwelling event in early February. These findings are compared with those of other marine disease and mortality studies.

A revision of the Pyura stolonifera species complex (Tunicata, Ascidiacea), with a description of a new species from Australia

A revision of the Pyura stolonifera species complex (Tunicata, Ascidiacea), with a description of a new species from Australia

"Nested" cryptic diversity in a widespread marine ecosystem engineer: a challenge for detecting biological invasions

BackgroundEcosystem engineers facilitate habitat formation and enhance biodiversity, but when they become invasive, they present a critical threat to native communities because they can drastically alter the receiving habitat. Management of such species thus needs to be a priority, but the poorly resolved taxonomy of many ecosystem engineers represents a major obstacle to correctly identifying them as being either native or introduced. We address this dilemma by studying the sea squirt Pyura stolonifera, an important ecosystem engineer that dominates coastal communities particularly in the southern hemisphere. Using DNA sequence data from four independently evolving loci, we aimed to determine levels of cryptic diversity, the invasive or native status of each regional population, and the most appropriate sampling design for identifying the geographic ranges of each evolutionary unit.ResultsExtensive sampling in Africa, Australasia and South America revealed the existence of "nested" levels of cryptic diversity, in which at least five distinct species can be further subdivided into smaller-scale genetic lineages. The ranges of several evolutionary units are limited by well-documented biogeographic disjunctions. Evidence for both cryptic native diversity and the existence of invasive populations allows us to considerably refine our view of the native versus introduced status of the evolutionary units within Pyura stolonifera in the different coastal communities they dominate.ConclusionsThis study illustrates the degree of taxonomic complexity that can exist within widespread species for which there is little taxonomic expertise, and it highlights the challenges involved in distinguishing between indigenous and introduced populations. The fact that multiple genetic lineages can be native to a single geographic region indicates that it is imperative to obtain samples from as many different habitat types and biotic zones as possible when attempting to identify the source region of a putative invader. "Nested" cryptic diversity, and the difficulties in correctly identifying invasive species that arise from it, represent a major challenge for managing biodiversity.

A revision of the Pyura stolonifera species complex (Tunicata, Ascidiacea), with a description of a new species from Australia

Pyura stolonifera is a large solitary ascidian found in Africa, Australasia and South America. The taxonomic status of different populations of this species is disputed, especially since there is evidence for several distinct morphological and genetic units that point towards the existence of multiple cryptic species. While some researchers still recognize P. stolonifera as a single species, others treat the different populations as distinct species. Here, we present a revision of the P. stolonifera species complex based on the examination of samples from all regions where there are reliable reports of this taxon. We recognize four species that are both morphologically and genetically distinct, one of which is new to science and is formally described here. This species is morphologically distinct from the other three members of the species complex in terms of the colour and texture of the tunic, the arrangement of the gonads within the gut and the shape of the dorsal tubercle, among other characters. We name the new species Pyura dalbyi after Dr. J.E. Dalby Jr., whose research on its ecology and distribution provided the incentive for examining this species more closely.

Variability of hair cells in the coronal organ of ascidians (Chordata, Tunicata)

The tunicate ascidians are nonvertebrate chordates that possess mechanoreceptor cells in the coronal organ in the oral siphon, which monitor the incoming water flow. Like vertebrate hair cells, the mechanoreceptor–coronal cells are secondary sensory (axonless) cells accompanied by supporting cells and they exhibit morphological diversities of apical specialisations: they are multiciliate in ascidians of the order Enterogona, whereas they are more complex and possess one or two cilia accompanied by stereovilli, also graded in length, in ascidians of the order Pleurogona. In morphology, embryonic origin, and arrangement, coronal sensory cells closely resemble vertebrate hair cells. We describe here the coronal organs of five ascidians ( Pyura haustor (Stimpson, 1864), Pyura stolonifera (Heller, 1878), Styela gibbsii (Stimpson, 1864), Styela montereyensis (Dall, 1872), and Polyandrocarpa zorritensis (Van Name, 1931)), belonging to Pleurogona, also comprising species of one family (Pyuridae), not yet considered, and thus completing our overview of the order. Each species possesses at least two kinds of secondary sensory cells, some of them characterized by stereovilli graded in length. In some species, the coronal sensory cells exhibit secretory activity; in P. haustor, a mitotic sensory cell has also been found. We compare the coronal organ in both ascidians and with other chordate sensory organs formed of secondary sensory cells, and discuss their possible homologies.

Context-dependent genetic benefits of polyandry in a marine hermaphrodite.

Numerous studies emphasize the potential indirect (genetic) benefits of polyandry in animals with resource-free mating systems. In this paper, we examine the potential for these benefits to fuel sexual selection and polyandry in the hermaphroditic ascidian Pyura stolonifera. Individuals were designated either sire (sperm producers) or dam (egg producers) at random and crossed in a North Carolina II breeding design to produce both paternal and maternal half siblings for our quantitative genetic analysis. We then partitioned the phenotypic variance in fertilization and hatching rates into additive and non-additive variance components. We found significant additive variance attributable to sire and dam effects at fertilization and hatching, suggesting the potential for selection to favour individuals carrying intrinsically 'good genes' for these traits. In separate analyses involving monandrous and polyandrous clutches, we found that both traits were elevated under polyandry, but the difference in hatching rates was due entirely to the difference in fertilization rates between treatments. When the hatching rates were standardized to account for variance at fertilization, there was no overall net benefit of polyandry for this trait. Despite this, we found that hatching success declined with increasing embryo densities, and that the slope of this decline was significantly greater in monandrous than polyandrous clutches. Hence, selection on embryo viability may still favour polyandry under restricted environmental conditions. Nevertheless, our results caution against interpreting elevated hatching success as an indirect genetic benefit of polyandry when variance in fertilization is not controlled.

Observations on non‐didemnid ascidians from Australian waters (1)

Thirty‐five species (including eight that are new) of non‐didemnid aplousobranch species and 25 phlebobranch (including one new) and stolidobranch species are discussed and a further 42 species in non‐didemnid taxa in all suborders are newly recorded. Material was taken by scuba diving in the waters around Kangaroo I. and other southern Australian and Tasmanian locations; and further material was taken by dredge from canyons off the southeastern Continental Slope and from inter‐reefal areas in the northern Great Barrier Reef region. The known geographic ranges of many southern Australian species are extended into Tasmanian waters and a new species of Rhopalaea is described from the Tasmanian Canyons. The genera Sigillina and Pseudodistoma are found to be diverse in Australian temperate waters, and they have many characters in common, suggesting a probable phylogenetic relationship that formerly was not recognised. The greatest diversity of species reported is from the vicinity of the northern Great Barrier Reef. Almost without exception these are non‐indigenous tropical species often extending into Australian temperate locations where they are sympatric with indigenous species found mainly around the southern half of the continent. Pyura stolonifera has a similar range in temperate Australian waters but appears also to occur on the South African and Chilean coasts in similar habitats and may represent a Gondwana relict.

Passive Flow Through an Unstalked Intertidal Ascidian: Orientation and Morphology Enhance Suspension Feeding in Pyura stolonifera

Passive flow is believed to increase the gains and reduce the costs of active suspension feeding. We used a mixture of field and laboratory experiments to evaluate whether the unstalked intertidal ascidian Pyura stolonifera exploits passive flow. We predicted that its orientation to prevailing currents and the arrangement of its siphons would induce passive flow due to dynamic pressure at the inhalant siphon, as well as by the Bernoulli effect or viscous entrainment associated with different fluid velocities at each siphon, or by both mechanisms. The orientation of P. stolonifera at several locations along the Sydney-Illawarra coast (Australia) covering a wide range of wave exposures was nonrandom and revealed that the ascidians were consistently oriented with their inhalant siphons directed into the waves or backwash. Flume experiments using wax models demonstrated that the arrangement of the siphons could induce passive flow and that passive flow was greatest when the inhalant siphon was oriented into the flow. Field experiments using transplanted animals confirmed that such an orientation resulted in ascidians gaining food at greater rates, as measured by fecal production, than when oriented perpendicular to the wave direction. We conclude that P. stolonifera enhances suspension feeding by inducing passive flow and is, therefore, a facultatively active suspension feeder. Furthermore, we argue that it is likely that many other active suspension feeders utilize passive flow and, therefore, measurements of their clearance rates should be made under appropriate conditions of flow to gain ecologically relevant results.

A complement component C3a-like peptide stimulates chemotaxis by hemocytes from an invertebrate chordate—the tunicate, Pyura stolonifera

Recent evidence suggests that the complement system evolved as a critical host defence mechanism among invertebrates, long before the origin among vertebrates of adaptive immune responses mediated by somatically re-arranging antibodies. The current study supports that contention by identifying a complement component C3a-like peptide in the tunicate, Pyura stolonifera. Activation of P. stolonifera serum with common inflammatory elicitors (lipopolysaccharide and zymosan) resulted in the proteolytic generation of an 8.5 kDa peptide, and concomitantly conferred chemoattractant activity on the serum. The 8.5 kDa peptide shares substantial amino acid sequence homology with a previously characterised tunicate complement component C3-like protein (72% amino acid identity in an 18 amino acid overlap). It is also recognised by an anti-C3 antiserum that is known to cross react with tunicate C3 homologues. Hemocyte migration assays performed with the 8.5 kDa peptide that had been partially purified by gel filtration confirmed that the molecule acts as a powerful chemotactic agent. This suggests that the proteolytic activation of tunicate C3-like molecules can initiate inflammatory responses involving cellular recruitment by liberating a pro-inflammatory peptide akin to the vertebrate anaphylatoxin, C3a.

Recent introduction of the dominant tunicate, Pyura praeputialis (Urochordata, Pyuridae) to Antofagasta, Chile.

The large sessile tunicate Pyura stolonifera (Pleurogona: Stolibranchiata: Pyuridae), has been regarded as a complex taxon with disjointed distributions, including Australia (Pyura stolonifera praeputialis), South Africa (Pyura stolonifera stolonifera) and South America (Chile, Antofagasta: Pyura sp., the 'piure de Antofagasta'), and has been cited under at least five taxonomic combinations. The 'piure de Antofagasta' is a competitively dominant species in rocky intertidal habitats and shows a limited geographical range (60-70 km) exclusively inside the Bay of Antofagasta. Using cytochrome oxidase I (COI) mitochondrial sequence data from Pyura specimens of the three taxa we tested whether the Chilean taxon represents: (i) a Gondwana relict; (ii) a more recently divergent species; or (iii) a recently introduced species. The results suggest that the Chilean taxon is a recent introduction to Chile from Australian populations and that Pyura stolonifera praeputialis, from Australia, and the 'piure de Antofagasta' are geographical populations of a single species: Pyura praeputialis; whereas the South African taxon represents a second species: Pyura stolonifera.

Patches of the ascidian Pyura stolonifera (Heller, 1878): structure of habitat and associated intertidal assemblages

Beds of the ascidian Pyura stolonifera on rocky shores around Sydney, Australia, provide an important habitat for many organisms, such as algae, chitons, limpets and snails. Fishermen collect Pyura for bait, which can change the structure of those beds. This may, in turn, cause changes in their associated biota. To predict the effects of such disturbances, it is necessary to understand the relationship between the structure of the habitat provided by beds of Pyura and their associated assemblages. Beds of Pyura can provide two types of habitat: Clumped, with >50% cover of closely packed individuals or Sparse, with <50% cover, where individuals are mostly isolated from contact with others. The spaces amongst the Pyura can be bare rock or rock covered by numerous species of algae and sessile animals. Nineteen algal species and 45 species of animals were identified in the field in Clumped and Sparse habitats. Assemblages differed significantly between patches of Clumped and Sparse habitat and much of the difference was attributable to relatively few species. In addition, a number of species of algae and animals live on the surface of the Pyura themselves. Eighty-four taxa were found at the scale of individual Pyura, many of which were relatively small and cryptic. Organisms on Pyura differed between individual ascidians that were isolated from other Pyura (i.e. a more common situation in Sparse habitat) and those that were surrounded by and in contact with other Pyura (i.e. more common in Clumped habitat). These differences were mostly attributed to a few families of small molluscs. It appears that changes in the structure of beds of Pyura have the potential to make significant alterations to intertidal assemblages at the scale of individual Pyura and at the scale of the habitats formed by beds of Pyura.

In situ measures of spawning synchrony and fertilization success in an intertidal, free-spawning invertebrate

The rocky intertidal zone has the potential to be one of the harshest environments for free-spawning organisms, but empirical data on fertilization success are scarce. Here, I report on an intertidal, solitary ascidian, Pyura stolonifera, which was observed to spawn at low tide. At a scale likely to be most important to gametes (metres, duration of tide), approximately 30% of individuals in the population were spawning synchronously. Spawned gametes remained in a viscous matrix and this appeared to minimise their dilution. Fertilization success varied greatly among individuals (0 to 92%) and was related to the distance to the nearest neighbouring spawner. Occasional wave wash facilitated the movement of sperm between spawners. Fertilization success in some individuals was limited by the scarcity of sperm whilst the experimental addition of sperm did not increase success in others.