Barnacles

@article{maccabe1849barnacles,
    author = "Maccabe, W. B.",
    title = "Barnacles",
    year = "1849",
    journal = "Notes and Queries",
    url = "https://doi.org/10.1093/nq/s1-i.8.117b",
    doi = "10.1093/nq/s1-i.8.117b",
    number = "8",
    pages = "117-118",
    volume = "s1-I"
}

@article{kingsley1877barnacles,
    author = "Kingsley, J. S.",
    title = "Barnacles",
    year = "1877",
    journal = "The American Naturalist",
    url = "https://doi.org/10.1086/271830",
    doi = "10.1086/271830",
    number = "2",
    pages = "102-108",
    volume = "11"
}

@article{crossref1889barnacles,
    title = "Barnacles",
    year = "1889",
    journal = "Science",
    url = "https://doi.org/10.1126/science.ns-14.356.374",
    doi = "10.1126/science.ns-14.356.374",
    number = "356",
    pages = "374-375",
    volume = "ns-14"
}

1. Stimulation in the rock barnacle Balanus balanoides by hydrochloric, sulfuric, and nitric acids, and by the first seven members of the normal aliphatic acid series has been studied. The hydrogen ion concentrations of the solutions tested varied from 3.2 x 10–8 to 5.889 x 10–6. The criterion of response was percentage closure in groups of individuals, recorded at 1 minute intervals until maximum closure occurred. 2. The intensity of stimulation by these acids is proportional to the effects of two forces, one related to the change in the (H+), and the other to the field of force around the anion of the acid added to the environment. 3. A preliminary interpretation of the results led to the development of the following expression which fits approximately the data obtained at the end of 4 minutes: Per cent closure = 100 – 100e–0.1z+(0.003125)2–0.1z+(0.003125)2n(z–0.4) where z is the (H+) x 107 and n is the number of carbon atoms (if present) in the anion of the acid. This equation assumes that the anions of the mineral acids enter into the reaction stoichiometrically, and emphasizes the difference in the fields of force around the anion of the fatty acids, a difference which is correlated with the length of the carbon chain. 4. A further analysis of the data revealed the presence of three or more receptor groups which appeared to be differentially affected by forces originating from the anions of the acids. 5. The order of stimulating efficiency for the mineral acids was found to be: HCl>H2SO4>HNO3. 6. The order of stimulating efficiency for the fatty acids was found to be: heptylic>caproic>valeric>butyric = acetic>propionic = formic.

@article{cole1933stimulation,
    author = "Cole, William H. and Allison, J. B.",
    title = "STIMULATION BY MINERAL AND FATTY ACIDS IN THE BARNACLE BALANUS BALANOIDES",
    year = "1933",
    journal = "Journal of General Physiology",
    abstract = "1. Stimulation in the rock barnacle Balanus balanoides by hydrochloric, sulfuric, and nitric acids, and by the first seven members of the normal aliphatic acid series has been studied. The hydrogen ion concentrations of the solutions tested varied from 3.2 x 10–8 to 5.889 x 10–6. The criterion of response was percentage closure in groups of individuals, recorded at 1 minute intervals until maximum closure occurred. 2. The intensity of stimulation by these acids is proportional to the effects of two forces, one related to the change in the (H+), and the other to the field of force around the anion of the acid added to the environment. 3. A preliminary interpretation of the results led to the development of the following expression which fits approximately the data obtained at the end of 4 minutes: Per cent closure = 100 – 100e–0.1z+(0.003125)2–0.1z+(0.003125)2n(z–0.4) where z is the (H+) x 107 and n is the number of carbon atoms (if present) in the anion of the acid. This equation assumes that the anions of the mineral acids enter into the reaction stoichiometrically, and emphasizes the difference in the fields of force around the anion of the fatty acids, a difference which is correlated with the length of the carbon chain. 4. A further analysis of the data revealed the presence of three or more receptor groups which appeared to be differentially affected by forces originating from the anions of the acids. 5. The order of stimulating efficiency for the mineral acids was found to be: HCl\>H2SO4\>HNO3. 6. The order of stimulating efficiency for the fatty acids was found to be: heptylic\>caproic\>valeric\>butyric = acetic\>propionic = formic.",
    url = "https://doi.org/10.1085/jgp.16.6.895",
    doi = "10.1085/jgp.16.6.895",
    number = "6",
    pages = "895-903",
    volume = "16"
}

H. Barnes, H. T. Powell, The Development, General Morphology and Subsequent Elimination of Barnacle Populations, Balanus crenatus and B. Balanoides, After a Heavy Initial Settlement, Journal of Animal Ecology, Vol. 19, No. 2 (Nov., 1950), pp. 175-179

@article{doi1023071526,
    author = "Barnes, H. and Powell, H. T.",
    title = "The Development, General Morphology and Subsequent Elimination of Barnacle Populations, Balanus crenatus and B. Balanoides, After a Heavy Initial Settlement",
    year = "1950",
    journal = "Journal of Animal Ecology",
    abstract = "H. Barnes, H. T. Powell, The Development, General Morphology and Subsequent Elimination of Barnacle Populations, Balanus crenatus and B. Balanoides, After a Heavy Initial Settlement, Journal of Animal Ecology, Vol. 19, No. 2 (Nov., 1950), pp. 175-179",
    url = "https://doi.org/10.2307/1526",
    doi = "10.2307/1526",
    openalex = "W2331426927"
}

There are in the literature a number of conflicting statements regarding the effect of various stimuli on the cypris stage of barnacle larvae, before and at the time of metamorphosis, and little experimental work has been done to elucidate the type of reactions involved. Visscher (1928), working with Balanus amphitrite, B. improvisus and Chthamalus fragilis, stated that during the earlier part of their free-swimming stage the cypris larvae were positively phototropic, but later the reaction to light became erratic, and at the time of attachment was decidedly negatively phototropic. He further stated that in a settlement experiment they orientated with the anterior end (i.e. with the paired eyes) away from the light source. However, MacDougall (I943) working with two of the above species (Balanus improvisus and Chthamalus fragilis) and also Balanus eburneus, and Pyefinch (I948) working with B. balanoides and B. crenatus, both state that the cypris larvae retain their positive reaction to light up to the time of settlement. There also appears to be some disagreement regarding the orientation at, and subsequent to, metamorphosis of several species of barnacles. Moore (1935), in considering B. balanoides, has suggested that if a barnacle is settling in a current, then at the time of settlement it is likely to be orientated with the current. A general orientation at right angles to the current was supposed to be due to rotation after settlement. This rotation was considered to be a response to the orientation of the beating cirri in relation to the current; it was suggested that feeding would be more effective with the animal orientated across the current.

@article{doi1023071542,
    author = "Barnes, H. and Crisp, D. J. and Powell, H. T.",
    title = "Observations on the Orientation of Some Species of Barnacles",
    year = "1951",
    journal = "Journal of Animal Ecology",
    abstract = "There are in the literature a number of conflicting statements regarding the effect of various stimuli on the cypris stage of barnacle larvae, before and at the time of metamorphosis, and little experimental work has been done to elucidate the type of reactions involved. Visscher (1928), working with Balanus amphitrite, B. improvisus and Chthamalus fragilis, stated that during the earlier part of their free-swimming stage the cypris larvae were positively phototropic, but later the reaction to light became erratic, and at the time of attachment was decidedly negatively phototropic. He further stated that in a settlement experiment they orientated with the anterior end (i.e. with the paired eyes) away from the light source. However, MacDougall (I943) working with two of the above species (Balanus improvisus and Chthamalus fragilis) and also Balanus eburneus, and Pyefinch (I948) working with B. balanoides and B. crenatus, both state that the cypris larvae retain their positive reaction to light up to the time of settlement. There also appears to be some disagreement regarding the orientation at, and subsequent to, metamorphosis of several species of barnacles. Moore (1935), in considering B. balanoides, has suggested that if a barnacle is settling in a current, then at the time of settlement it is likely to be orientated with the current. A general orientation at right angles to the current was supposed to be due to rotation after settlement. This rotation was considered to be a response to the orientation of the beating cirri in relation to the current; it was suggested that feeding would be more effective with the animal orientated across the current.",
    url = "https://doi.org/10.2307/1542",
    doi = "10.2307/1542",
    openalex = "W2316112678"
}

@article{barnes1953on,
    author = "Barnes, H.",
    title = "On the Southern Limits of the Intertidal Barnacle Balanus Balanoides",
    year = "1953",
    journal = "Ecology",
    url = "https://doi.org/10.2307/1930912",
    doi = "10.2307/1930912",
    number = "2",
    pages = "429-430",
    volume = "34"
}

Laboratory experiments showed that cyprids of Balanus balanoides settled readily only when offered surfaces which bore settled barnacles of their own species, or the cemented bases left on surfaces from which these barnacles had been removed. Without such surfaces they proved to be capable of postponing setting for at least 2 weeks, and they chose such surfaces in preference to similar surfaces which bore related species. Cyprids of B. crenatus and Elminius modestus were also capable of making this choice. Setting usually occurred in the immediate vicinity of barnacles or bases, but some cyprids crawled short distances away during their final reconnaissances. When they encountered barnacles attached to glass, they were stimulated to proceed with setting, but they swam away from this unfavourably smooth surface and settled on bare stones nearby. Fragments from various parts of the body, placed upon surfaces which had never borne barnacles, made these somewhat more favourable for setting. Since attached bases alone produced a strong reaction, the cyprids probably respond to contact between their groping antennules and some substance in the cuticle. They did not settle readily when prevented from making contact with adults by a barrier of bolting silk, or with bases by a thin film of nitrocellulose; chemosensory perception of a water-soluble substance, emanating from settled barnacles, is therefore unlikely to be involved. The power of the bases to induce setting was retained after heating to over 200° C., but not after heating to 275° C., when charring occurred. It withstood prolonged washing in water, heating with fat solvents and dilute acids, and treatment in the cold with caustic alkalis, concentrated acids, pepsin, formaldehyde, benzoquinone, sodium sulphide, phenol, urea, and diazonium solution. It was destroyed in the cold by sodium hypochlorite, which was observed to dissolve the bases, and also by hot concentrated acids and alkalis. The chemical properties of the active substance are thus entirely consistent with those of quinone-tanned proteins, which are known to form the epicuticle and attachment cement. A field experiment showed that similar behaviour occurs under natural conditions. It leads cyprids to suitable habitats and it facilitates breeding.

@article{knightjones1953laboratory,
    author = "Knight-Jones, E. W.",
    title = "Laboratory Experiments on Gregariousness During Setting in Balanus Balanoides and Other Barnacles",
    year = "1953",
    journal = "Journal of Experimental Biology",
    abstract = "Laboratory experiments showed that cyprids of Balanus balanoides settled readily only when offered surfaces which bore settled barnacles of their own species, or the cemented bases left on surfaces from which these barnacles had been removed. Without such surfaces they proved to be capable of postponing setting for at least 2 weeks, and they chose such surfaces in preference to similar surfaces which bore related species. Cyprids of B. crenatus and Elminius modestus were also capable of making this choice. Setting usually occurred in the immediate vicinity of barnacles or bases, but some cyprids crawled short distances away during their final reconnaissances. When they encountered barnacles attached to glass, they were stimulated to proceed with setting, but they swam away from this unfavourably smooth surface and settled on bare stones nearby. Fragments from various parts of the body, placed upon surfaces which had never borne barnacles, made these somewhat more favourable for setting. Since attached bases alone produced a strong reaction, the cyprids probably respond to contact between their groping antennules and some substance in the cuticle. They did not settle readily when prevented from making contact with adults by a barrier of bolting silk, or with bases by a thin film of nitrocellulose; chemosensory perception of a water-soluble substance, emanating from settled barnacles, is therefore unlikely to be involved. The power of the bases to induce setting was retained after heating to over 200° C., but not after heating to 275° C., when charring occurred. It withstood prolonged washing in water, heating with fat solvents and dilute acids, and treatment in the cold with caustic alkalis, concentrated acids, pepsin, formaldehyde, benzoquinone, sodium sulphide, phenol, urea, and diazonium solution. It was destroyed in the cold by sodium hypochlorite, which was observed to dissolve the bases, and also by hot concentrated acids and alkalis. The chemical properties of the active substance are thus entirely consistent with those of quinone-tanned proteins, which are known to form the epicuticle and attachment cement. A field experiment showed that similar behaviour occurs under natural conditions. It leads cyprids to suitable habitats and it facilitates breeding.",
    url = "https://doi.org/10.1242/jeb.30.4.584",
    doi = "10.1242/jeb.30.4.584",
    number = "4",
    openalex = "W2342878346",
    pages = "584-598",
    volume = "30",
    references = "doi101017s0025315400012716, doi101017s0025315400014594, doi101017s0025315400055375, doi1010381711109a0, doi101093icesjms14186, doi101098rspb19400018, doi101111j109636421949tb00409x, doi1023071542, doi1023073001536, openalexw1528170061"
}

The two commonest species of intertidal barnacles indigenous to the British Isles* have very different geographical distributions, and belong to different faunal groups. Chthamalus stellatus Poli is a Tropical-Lusitanian form which reaches its northern limit in Scotland, while Balanus balanoides L. is a Boreal-Arctic form which reaches its southernmost European limits in south-west England and northern France. Over a large part of the British Isles the two species occur together, but Chthamalus is dominant in the south and west while Balanus alone is present in the north and east (see Darwin 1854, pp. 272 and 456). Since both Chthamalus stellatus and Balanus balanoides are at the limits of their respective distributions in the British Isles, these limits may be particularly sensitive to any general change in the environment. Such a change might also affect many other organisms, and will well repay careful study.

@article{doi1023071665,
    author = "Southward, A. J. and Crisp, D. J.",
    title = "Recent Changes in the Distribution of the Intertidal Barnacles Chthamalus stellatus Poli and Balanus balanoides L. in the British Isles",
    year = "1954",
    journal = "Journal of Animal Ecology",
    abstract = "The two commonest species of intertidal barnacles indigenous to the British Isles* have very different geographical distributions, and belong to different faunal groups. Chthamalus stellatus Poli is a Tropical-Lusitanian form which reaches its northern limit in Scotland, while Balanus balanoides L. is a Boreal-Arctic form which reaches its southernmost European limits in south-west England and northern France. Over a large part of the British Isles the two species occur together, but Chthamalus is dominant in the south and west while Balanus alone is present in the north and east (see Darwin 1854, pp. 272 and 456). Since both Chthamalus stellatus and Balanus balanoides are at the limits of their respective distributions in the British Isles, these limits may be particularly sensitive to any general change in the environment. Such a change might also affect many other organisms, and will well repay careful study.",
    url = "https://doi.org/10.2307/1665",
    doi = "10.2307/1665",
    openalex = "W2313550652"
}

It is well known that towards the limits of an animal's geographical range both its abundance and exact boundaries may fluctuate from time to time. These fluctuations can sometimes be correlated with environmental changes, and thus may help to assess the relative importance of the factors that control distribution. For example the distribution of the common intertidal barnacles Chthamalus stellatus (Poli) and Balanus balanoides (Linnaeus) underwent changes which were attributed to a general rise in temperature over several years (Southward & Crisp, 1954a). The abundance of these barnacles has fluctuated further since 1951–52, and we are now able to analyse more closely the relation between the population changes and environmental variations.

@article{doi101017s0025315400009073,
    author = "Southward, A. J. and Crisp, D. J.",
    title = "Fluctuations in the distribution and abundance of intertidal barnacles",
    year = "1956",
    journal = "Journal of the Marine Biological Association of the United Kingdom",
    abstract = "It is well known that towards the limits of an animal's geographical range both its abundance and exact boundaries may fluctuate from time to time. These fluctuations can sometimes be correlated with environmental changes, and thus may help to assess the relative importance of the factors that control distribution. For example the distribution of the common intertidal barnacles Chthamalus stellatus (Poli) and Balanus balanoides (Linnaeus) underwent changes which were attributed to a general rise in temperature over several years (Southward \& Crisp, 1954a). The abundance of these barnacles has fluctuated further since 1951–52, and we are now able to analyse more closely the relation between the population changes and environmental variations.",
    url = "https://doi.org/10.1017/s0025315400009073",
    doi = "10.1017/s0025315400009073",
    openalex = "W2096259324"
}

H. Barnes, Balanus balanoides (L.) in the Firth of Clyde: The Development and Annual Variation of the Larval Population, and the Causative Factors, Journal of Animal Ecology, Vol. 25, No. 1 (May, 1956), pp. 72-84

@article{doi1023071851,
    author = "Barnes, H.",
    title = "Balanus balanoides (L.) in the Firth of Clyde: The Development and Annual Variation of the Larval Population, and the Causative Factors",
    year = "1956",
    journal = "Journal of Animal Ecology",
    abstract = "H. Barnes, Balanus balanoides (L.) in the Firth of Clyde: The Development and Annual Variation of the Larval Population, and the Causative Factors, Journal of Animal Ecology, Vol. 25, No. 1 (May, 1956), pp. 72-84",
    url = "https://doi.org/10.2307/1851",
    doi = "10.2307/1851",
    openalex = "W2318891473"
}

Synopsis The distribution and zonation of Chthamalus stellatus and Balanus balanoides on British coasts is reviewed, and it is shown that the very sheltered waters of Loch Sween and West Loch Tarbert are unusual in that Chthamalus occurs there in greater quantity than in any other sheltered (British) locality, and the upper limit of Chthamalus is consistently higher, relative to other species, than elsewhere in shelter. The lower limit of Chthamalus is generally at a rather high level on these shores even when bare rock occurs below. Balanus balanoides, on the other hand, is often scarce or absent at the same sheltered sites. The physical conditions in the lochs (water temperatures, tides, winds, sunshine, rainfall) and their seasonal variations are examined and it is shown that there probably is a unique succession of environmental factors that would favour Chthamalus and operate against Balanus at critical times of the year. It is suggested that Chthamalus occurs higher than Pelvetia in the lochs because of a greater tolerance of irregular and often prolonged periods of desiccation. The failure of Chthamalus to occupy the middle and lower shore when Balanus is absent indicates that competition between the two barnacles does not alone determine their vertical distribution on the shore. A tentative hypothesis is developed showing how the very local distribution of Chthamalus and the level occupied on the shore might be related to responses of the cyprids to light and to a supposed requirement of emersion during the early stages on the shore.

@article{doi101017s0080456800100067,
    author = "Lewis, J. R. and Powell, H. T.",
    title = "IV.—Aspects of the Intertidal Ecology of Rocky Shores in Argyll, Scotland. II. The Distribution of Chthamalus stellatus and Balanus balanoides in Kintyre",
    year = "1960",
    journal = "Earth and Environmental Science Transactions of the Royal Society of Edinburgh",
    abstract = "Synopsis The distribution and zonation of Chthamalus stellatus and Balanus balanoides on British coasts is reviewed, and it is shown that the very sheltered waters of Loch Sween and West Loch Tarbert are unusual in that Chthamalus occurs there in greater quantity than in any other sheltered (British) locality, and the upper limit of Chthamalus is consistently higher, relative to other species, than elsewhere in shelter. The lower limit of Chthamalus is generally at a rather high level on these shores even when bare rock occurs below. Balanus balanoides, on the other hand, is often scarce or absent at the same sheltered sites. The physical conditions in the lochs (water temperatures, tides, winds, sunshine, rainfall) and their seasonal variations are examined and it is shown that there probably is a unique succession of environmental factors that would favour Chthamalus and operate against Balanus at critical times of the year. It is suggested that Chthamalus occurs higher than Pelvetia in the lochs because of a greater tolerance of irregular and often prolonged periods of desiccation. The failure of Chthamalus to occupy the middle and lower shore when Balanus is absent indicates that competition between the two barnacles does not alone determine their vertical distribution on the shore. A tentative hypothesis is developed showing how the very local distribution of Chthamalus and the level occupied on the shore might be related to responses of the cyprids to light and to a supposed requirement of emersion during the early stages on the shore.",
    url = "https://doi.org/10.1017/s0080456800100067",
    doi = "10.1017/s0080456800100067",
    openalex = "W2948891525",
    references = "doi101017s0025315400009073, doi101017s0025315400014168, doi101017s0025315400014909, doi101111j109636421957tb00295x, doi1023071526, doi1023071665, doi1023071851, doi1023072255831, doi1023072256147, doi1023072256672"
}

@article{connell1961effects,
    author = "Connell, Joseph H.",
    title = "Effects of Competition, Predation by Thais lapillus, and Other Factors on Natural Populations of the Barnacle Balanus balanoides",
    year = "1961",
    journal = "Ecological Monographs",
    url = "https://doi.org/10.2307/1950746",
    doi = "10.2307/1950746",
    number = "1",
    pages = "61-104",
    volume = "31"
}

@misc{connell1961the1,
    author = "Connell, J. H",
    title = "The effects of competition, predation by Thais lapillus and other factors on natural populations of the barnacle Balanus balanoides",
    year = "1961",
    howpublished = "Ecological Monographs, v. 31, p. 61-104",
    note = "talkorigins\_source = {true}; raw\_reference = {Connell, J. H., 1961, The effects of competition, predation by Thais lapillus and other factors on natural populations of the barnacle Balanus balanoides: Ecological Monographs, v. 31, p. 61-104.}"
}

@misc{connell1961the2,
    author = "Connell, J. H",
    title = "The influence of interspecific competition and other factors on the distribution of the barnacle, Chthamalus stellatus",
    year = "1961",
    howpublished = "Ecology, v. 42, p. 710-723",
    note = "talkorigins\_source = {true}; raw\_reference = {Connell, J. H., 1961, The influence of interspecific competition and other factors on the distribution of the barnacle, Chthamalus stellatus: Ecology, v. 42, p. 710-723.}"
}

@article{doi1023071933500,
    author = "Connell, Joseph H.",
    title = "The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus",
    year = "1961",
    journal = "Ecology",
    url = "https://doi.org/10.2307/1933500",
    doi = "10.2307/1933500",
    openalex = "W1973688100",
    references = "beauchamp1932competitive, connell1961effects, doi101017s0025315400009073, doi101017s002531540001047x, doi1023071665, doi1023071851, doi1023071931034, doi1023071931600, doi1023071950746, doi1023072256872, doi1023072273"
}

@article{doi1023071950746,
    author = "Connell, Joseph H.",
    title = "Effects of Competition, Predation by Thais lapillus, and Other Factors on Natural Populations of the Barnacle Balanus balanoides",
    year = "1961",
    journal = "Ecological Monographs",
    url = "https://doi.org/10.2307/1950746",
    doi = "10.2307/1950746",
    openalex = "W1988997784",
    references = "doi101038180560a0, doi101086394476, doi101086395888, doi101093icesjms115, doi1023071664, doi1023073498751, doi1023073564801, doi105479si03629236931, knightjones1953laboratory, openalexw1528170061"
}

@misc{connell1970a3,
    author = "Connell, J. H",
    title = "A predator-prey system in the marine intertidal region. I. Balanus glandula and several predatory species of Thias",
    year = "1970",
    howpublished = "Ecological Monographs, v. 40, p. 49-78",
    note = "talkorigins\_source = {true}; raw\_reference = {Connell, J. H., 1970, A predator-prey system in the marine intertidal region. I. Balanus glandula and several predatory species of Thias: Ecological Monographs, v. 40, p. 49-78.}"
}

An understanding of community structure should be based on evidence that the growth and regulation of the component populations in the community are affected in a predictable manner by natural physical disturbances and by interactions with other species in the community. This study presents an experimental evaluation of the effects of such disturbances and competitive interactions on populations of sessile organisms in the rocky intertidal community, for which space can be demonstrated to be the most important limiting resource. This research was carried out at eight stations on the Washington coastline which have been ranked according to an exposure/desiccation gradient and subjected to comparable manipulation and observation. Physical variables such as wave exposure, battering by drift logs, and desiccation have important effects on the distribution and abundance of many of the sessile species in the community. In particular, wave exposure and desiccation have a major influence on the distribution patterns of all the algae and of the anemone Anthopleura elegantissima. The probability of damage from drift logs is very high in areas where logs have accumulated along the intertidal. Log damage and wave exposure have complementary effects in the provision of free space in a mussel bed, as wave shock enlarges a patch created by log damage by wrenching the mussels from the substratum at the periphery of the bare patch. Competition for primary space results in clear dominance hierarchies, in which barnacles are dominant over algae. Among the barnacles, Balanus cariosus is dominant over both B. glandula and Chthamalus dalli; B. glandula is dominant over C. dalli. The mussel Mytilus californianus requires secondary space (certain algae, barnacles, or byssal threads) for larval settlement, but is capable of growing over all other sessile species and potentially is the competitive dominant of space in the community.

@article{doi1023071948498,
    author = "Dayton, Paul K.",
    title = "Competition, Disturbance, and Community Organization: The Provision and Subsequent Utilization of Space in a Rocky Intertidal Community",
    year = "1971",
    journal = "Ecological Monographs",
    abstract = "An understanding of community structure should be based on evidence that the growth and regulation of the component populations in the community are affected in a predictable manner by natural physical disturbances and by interactions with other species in the community. This study presents an experimental evaluation of the effects of such disturbances and competitive interactions on populations of sessile organisms in the rocky intertidal community, for which space can be demonstrated to be the most important limiting resource. This research was carried out at eight stations on the Washington coastline which have been ranked according to an exposure/desiccation gradient and subjected to comparable manipulation and observation. Physical variables such as wave exposure, battering by drift logs, and desiccation have important effects on the distribution and abundance of many of the sessile species in the community. In particular, wave exposure and desiccation have a major influence on the distribution patterns of all the algae and of the anemone Anthopleura elegantissima. The probability of damage from drift logs is very high in areas where logs have accumulated along the intertidal. Log damage and wave exposure have complementary effects in the provision of free space in a mussel bed, as wave shock enlarges a patch created by log damage by wrenching the mussels from the substratum at the periphery of the bare patch. Competition for primary space results in clear dominance hierarchies, in which barnacles are dominant over algae. Among the barnacles, Balanus cariosus is dominant over both B. glandula and Chthamalus dalli; B. glandula is dominant over C. dalli. The mussel Mytilus californianus requires secondary space (certain algae, barnacles, or byssal threads) for larval settlement, but is capable of growing over all other sessile species and potentially is the competitive dominant of space in the community.",
    url = "https://doi.org/10.2307/1948498",
    doi = "10.2307/1948498",
    openalex = "W1974072473",
    references = "connell1961effects, doi101086282455, doi1023071933500, doi1023071942327, doi1023071950746"
}

Sex allocation theory for simultaneous hermaphrodites predicts increases in relative allocation to male-specific function as competition for fertilizations increases. Theoretical models developed specifically for competing acorn barnacles predict that the proportional allocation to male function increases toward an asymptote of 50% as the number of competitors for fertilizations increases. Experimental manipulations were used to investigate how mate competition affected both relative and absolute allocation to the sex functions for two species of acorn barnacle: Semibalanus balanoides and Balanus glandula. The ratio of male to female allocation did not increase with the number of competitors for either species. However, both species showed increased allocation to male function (estimated as total mass of sex-specific tissues) with increased crowding. Allocation to female function seemed to be limited by other factors and did not vary with mating group size as predicted. Allocation to male and female function were both positively related to body size, but a trade-off between male and female function, a key assumption of prior models, was not observed.

@article{doi101111j15585646201101548x,
    author = "Hoch, J. Matthew and Levinton, Jeffrey S.",
    title = "EXPERIMENTAL TESTS OF SEX ALLOCATION THEORY WITH TWO SPECIES OF SIMULTANEOUSLY HERMAPHRODITIC ACORN BARNACLES",
    year = "2011",
    journal = "Evolution",
    abstract = "Sex allocation theory for simultaneous hermaphrodites predicts increases in relative allocation to male-specific function as competition for fertilizations increases. Theoretical models developed specifically for competing acorn barnacles predict that the proportional allocation to male function increases toward an asymptote of 50\% as the number of competitors for fertilizations increases. Experimental manipulations were used to investigate how mate competition affected both relative and absolute allocation to the sex functions for two species of acorn barnacle: Semibalanus balanoides and Balanus glandula. The ratio of male to female allocation did not increase with the number of competitors for either species. However, both species showed increased allocation to male function (estimated as total mass of sex-specific tissues) with increased crowding. Allocation to female function seemed to be limited by other factors and did not vary with mating group size as predicted. Allocation to male and female function were both positively related to body size, but a trade-off between male and female function, a key assumption of prior models, was not observed.",
    url = "https://doi.org/10.1111/j.1558-5646.2011.01548.x",
    doi = "10.1111/j.1558-5646.2011.01548.x",
    openalex = "W1817706633",
    references = "connell1961effects, doi101016s0169534797010781, doi101017cbo9780511542053, doi101073pnas7652480, doi101086406066, doi101111j15585646200900669x, doi101126science1563774477, doi1015159780295743240, doi1015159781400832019, doi105860choice321533"
}

Sex allocation theory for simultaneous hermaphrodites predicts increases in relative allocation to male-specific function as competition for fertilizations increases. Theoretical models developed specifically for competing acorn barnacles predict that the proportional allocation to male function increases towards an asymptote of 50% as the number of competitors for fertilizations increases. Experimental manipulations were used to investigate how mate competition affected both relative and absolute allocation to the sex functions for two species of acorn barnacle: Semibalanus balanoides and Balanus glandula. The ratio of male to female allocation did not increase with the number of competitors for either species. However, both species showed increased allocation to male function (estimated as total mass of sex-specific tissues) with increased crowding. Allocation to female function seemed to be limited by other factors and did not vary with mating group size as predicted. Allocation to male and female function were both positively related to body size, but a trade-off between male and female function, a key assumption of prior models, was not observed.

@misc{hoch2011data,
    author = "Hoch, J. Matthew",
    title = "Data from: Experimental tests of sex allocation theory with two species of simultaneously hermaphroditic acorn barnacles",
    year = "2011",
    publisher = "Dryad",
    abstract = "Sex allocation theory for simultaneous hermaphrodites predicts increases in relative allocation to male-specific function as competition for fertilizations increases. Theoretical models developed specifically for competing acorn barnacles predict that the proportional allocation to male function increases towards an asymptote of 50\% as the number of competitors for fertilizations increases. Experimental manipulations were used to investigate how mate competition affected both relative and absolute allocation to the sex functions for two species of acorn barnacle: Semibalanus balanoides and Balanus glandula. The ratio of male to female allocation did not increase with the number of competitors for either species. However, both species showed increased allocation to male function (estimated as total mass of sex-specific tissues) with increased crowding. Allocation to female function seemed to be limited by other factors and did not vary with mating group size as predicted. Allocation to male and female function were both positively related to body size, but a trade-off between male and female function, a key assumption of prior models, was not observed.",
    url = "https://datadryad.org/dataset/doi:10.5061/dryad.39np8rd7",
    doi = "10.5061/dryad.39np8rd7"
}

@incollection{crossref2015barnacles,
    title = "BARNACLES",
    year = "2015",
    booktitle = "Evenings at the Microscope",
    url = "https://doi.org/10.1017/cbo9781316146545.013",
    doi = "10.1017/cbo9781316146545.013",
    pages = "232-244"
}

@incollection{mehlhorn2015barnacles,
    author = "Mehlhorn, Heinz",
    title = "Barnacles",
    year = "2015",
    booktitle = "Encyclopedia of Parasitology",
    url = "https://doi.org/10.1007/978-3-642-27769-6\_4627-1",
    doi = "10.1007/978-3-642-27769-6\_4627-1",
    pages = "1-1"
}

@incollection{crossref2017barnacles,
    title = "Barnacles",
    year = "2017",
    booktitle = "The Quotable Darwin",
    url = "https://doi.org/10.2307/j.ctvc77m14.19",
    doi = "10.2307/j.ctvc77m14.19",
    pages = "84-89"
}

@incollection{mary2020barnacles,
    author = "Mary, Avelin and Sarojini, R.",
    title = "Barnacles",
    year = "2020",
    booktitle = "Fouling Organisms of the Indian Ocean",
    url = "https://doi.org/10.1201/9781003077992-14",
    doi = "10.1201/9781003077992-14",
    pages = "391-439"
}

@incollection{crossref2025barnacles,
    title = "Barnacles",
    year = "2025",
    booktitle = "Fossils on the Seashore",
    url = "https://doi.org/10.2307/jj.20945013.15",
    doi = "10.2307/jj.20945013.15",
    pages = "69-75"
}