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A Review of Extant Taxa of the "Group of Balanus Conca Vus" (Cirripedia, Thoracica) and a Proposal for Genus-Group Ranks1)
Abstract
Les taxa caractéristiques du “Groupe Balanus concavus” sont connus de l’Oligocène d’Europe et de l’ouest de l’Amérique du Sud. Ses représentants étaient apparemment largement distribués au Miocène mais le groupe a ensuite décliné (Miocène du Pakistan et du Japon, Pleistocène d’Europe et Pliocène de l’Atlantique occidental). De la vingtaine de taxa décrits, quelques uns seulement existent encore, et, ce qui est intéressant, sont confinés au Pacifique oriental. Dans le présent travail sont revues toutes les espèces existantes qui ont été attribuées au groupe. Un statut générique est proposé pour celles qui peuvent actuellement y être rattachées et elles sont réparties en deux nouveaux sous-genres.
... The ORV Fm consists of clays and silty-sandy clays with local layers and lenses of coarse-grained material (usually siliciclastic conglomerates, but bioclastic intervals also occur) (Giammarino & Tedeschi 1983;Giammarino et al. 2010;Dalla Giovanna et al. 2016). The ORV Fm is generally characterised by a molluscan and foraminiferal assemblage suggesting deposition in circalittoral to bathyal settings, at a few hundreds of meters of water depth (Giammarino & Tedeschi 1976, 1982, 1983Robba 1981;Giammarino et al. 1984;Bernasconi 1989), but relevant differences exist between various outcrops (Boni et al. 1976;Marini 2000). The CMV Fm is comprised of grain-supported, poorly organised, and generally poorly sorted conglomerates with a sandy-silty matrix and siltymarly lenses (Boni et al. 1984;Gnaccolini 1998;Giammarino et al. 2010;Dalla Giovanna et al. 2016) (Fig. 1C). ...
... The sheath is largely appressed, displaying prominent transverse growth lines on the carinolatera only, and the alae are cleft. On the whole, these characters allow us to identify the specimens as belonging to the extinct species Concavus concavus of the balanid subfamily Concavinae (Newman 1982;Zullo 1992;Pitombo 2004), which is a common constituent of the Pliocene barnacle assemblages of the Mediterranean region (Davadie 1963;Menesini 1965). A single small-sized specimen from the lowermost calcirudites of section S3 does not conform to the above characterization and is here referred to cf. ...
... The lack of sorting and cross-lamination suggests a depositional environment below the SWWB, which is currently placed at around 30 m depth off the Ligurian coasts (Vacchi et al. 2012). Based on the molluscan and foraminiferal assemblages, the fine-grained Pliocene deposits of western Liguria are usually related to a deep water (~500 m) bathyal setting (Robba 1981;Giammarino & Tedeschi 1976, 1982, 1983. This interpretation can be rejected for the observed foramol facies exposed at Pairola. ...
... The ORV Fm consists of clays and silty-sandy clays with local layers and lenses of coarse-grained material (usually siliciclastic conglomerates, but bioclastic intervals also occur) (Giammarino & Tedeschi 1983;Giammarino et al. 2010;Dalla Giovanna et al. 2016). The ORV Fm is generally characterised by a molluscan and foraminiferal assemblage suggesting deposition in circalittoral to bathyal settings, at a few hundreds of meters of water depth (Giammarino & Tedeschi 1976, 1982, 1983Robba 1981;Giammarino et al. 1984;Bernasconi 1989), but relevant differences exist between various outcrops (Boni et al. 1976;Marini 2000). The CMV Fm is comprised of grain-supported, poorly organised, and generally poorly sorted conglomerates with a sandy-silty matrix and siltymarly lenses (Boni et al. 1984;Gnaccolini 1998;Giammarino et al. 2010;Dalla Giovanna et al. 2016) (Fig. 1C). ...
... The sheath is largely appressed, displaying prominent transverse growth lines on the carinolatera only, and the alae are cleft. On the whole, these characters allow us to identify the specimens as belonging to the extinct species Concavus concavus of the balanid subfamily Concavinae (Newman 1982;Zullo 1992;Pitombo 2004), which is a common constituent of the Pliocene barnacle assemblages of the Mediterranean region (Davadie 1963;Menesini 1965). A single small-sized specimen from the lowermost calcirudites of section S3 does not conform to the above characterization and is here referred to cf. ...
... The lack of sorting and cross-lamination suggests a depositional environment below the SWWB, which is currently placed at around 30 m depth off the Ligurian coasts (Vacchi et al. 2012). Based on the molluscan and foraminiferal assemblages, the fine-grained Pliocene deposits of western Liguria are usually related to a deep water (~500 m) bathyal setting (Robba 1981;Giammarino & Tedeschi 1976, 1982, 1983. This interpretation can be rejected for the observed foramol facies exposed at Pairola. ...
The lower Pliocene deposits of Pairola (Liguria, Italy) display the otherwise rare occurrence of rock-forming amounts of barnacles (mostly belonging to the extinct Euromediterranean species Concavus concavus). Three main facies are recognised in the investigated succession: a barnacle-dominated facies, which formed along a shallow (<15 m deep) nearshore environment, a foraminifera-dominated facies from relatively deeper waters (40-100 m), and an intermediate facies forming at the boundary of the other two. These facies and their relationships suggest deposition in a flooded valley – a kind of setting that was common in the Mediterranean after the Messinian Salinity Crisis.Differing from other rias, the Pairola basin was exposed to strong waves, resulting in conditions favourable to barna-
cles. Sedimentological and stratigraphic observations indicate that the Pairola succession formed within a timespan covering both cold and warm phases. This is relevant because the sub-tropical foraminifer Amphistegina is ubiquitous throughout the succession. Amphistegina occurs in the Pliocene and lower Pleistocene (Gelasian) of Northern Italy, but not in the remainder of the Pleistocene, not even its warm portions. This genus is currently recolonizing the Mediterranean and is projected to reach the northern coast of the basin soon, foretelling that Anthropocene temperatures are going to overcome those of the late Pleistocene warm periods and reach those of the Pliocene.
... Sacco, 1897, and the balanid barnacle Con cavus sp. Newman, 1982. Other vertebrate fossils include sea turtles, whales, dolphins, and sea cows (Kellogg 1923;Domning 1997;Weems and Sanders 2014;Sanders and Geisler 2015;Boessenecker and Fordyce 2017;Geisler et al. 2017;Albright et al. 2019;Domning and Beatty 2019). ...
Paleogene dermochelyid species richness far exceeded that of today. Leatherback sea turtles were most species rich in the Paleogene, but their richness declined sharply during the Neogene with only one species existing today, Dermochelys coriacea. We describe the fossil remains of three leatherback genera (Natemys, Psephophorus, and Egyptemys) from the upper Oligocene Chandler Bridge Formation and two (Natemys and Psephophorus) from the lower Oligocene Ashley Formation of South Carolina, USA. The fossils consist of isolated and some associated carapacial ossicles. Several ossicles are referred to Natemys sp. because their scalloped edges are indicative of the carapacial sunflower pattern specific to this genus. Additionally, two Natemys morphotypes (Natemys sp. 1 and 2) are distinguished based on differences in ossicle thickness and internal structure. We refer two ossicles to cf. Psephophorus because of their internal diploic structure and because one has a dorsal radial pattern while the other has a prominent ridge that exhibits strong visceral concavity. Finally, we refer one ossicle to cf. Egyptemys because it has a shallow keel that shows little expression on the visceral surface, although we also acknowledge the ossicle’s similarity to some ridged ossicles of the genus Psephophorus. These ossicles represent the first multispecies assemblage of leatherback fossils reported worldwide. Furthermore, the specimens fill both temporal and geographic gaps for extinct leatherback genera and represent the first formally described dermochelyids from South Carolina and the Oligocene of the Atlantic Coastal Plain.
... By the Neogene, balanomorph cirripedes had become one of the defining taxa of the marine environment (Darwin 1854;Seguenza 1876;de Alessandri 1895;de Alessandri 1906;de Alessandri 1910;de Alessandri 1922;Davadie 1963;Menesini 1965;Bałuk & Radwański 1967;Newman et al. 1967;Newman et al. 1969;Newman & Ross 1971;Newman & Ross 1976;Newman 1979;Newman & Abbot 1980;Carriol 1982a;Carriol 1982b;Newman 1982;Buckeridge 1983;Bałuk & Radwański 1984;Menesini 1984;Newman 1991;Świerczewska-Gładysz 1994;Carriol 1996;Newman 1996;Wöhrer 1998). Since the turn of the millennium, an even greater diversification and distribution of the Balanomorpha has been uncovered through the work of cirripedologists such as Carriol (2000), Pitombo (2004), Carriol (2008), Carriol & Schneider (2008), Santos & Mayoral (2008), Carriol & Menkveld-Gfeller (2010), Santos et al. (2012), Carriol & Schneider (2016), Betancort et al. (2014), Buckeridge et al. (2014), Collins et al. (2014), Jagt et al. (2016), Chan et al. (2017), Buckeridge et al. (2018), Coletti et al. (2018), and Kočí et al. (2018), confirming that the Cenozoic was indeed the age of cirripedes as proclaimed by Darwin (1854). ...
A new species of acorn barnacle, Actinobalanus? sloveniensis, is recorded from Oligocene and Miocene deposits in Slovenia, representing the first record of extinct balanoids attached to plant remains from this country, and in fact from the entire central Paratethys. Assemblages available include more than 200 specimens attached to tree branches and/or roots from a cypress forest (Taxodioxylon) and from estuarine mangroves (Carapoxylon). A taphonomic interpretation is provided, suggesting that some of these remains represent fragments that were dislodged during a storm event and quickly embedded in what were likely to have been anoxic sediments. This paper is part of the special volume: Jagt, J.W.M., Fraaije, R.H.B., Van Bakel, B.W.M., Donovan, S.K., Mellish, C. & Schweigert, G. (eds.): A lifetime amongst fossil crustaceans: a tribute to Joseph S.H. Collins (1927–2019).
... Os tergos representados na jazida de Vale doFreixo com morfologias muito similares entre si (Fig. 1.14-15), caracterizam-se por apresentar o sulco do esporão mais fechado, pelo que não consideramos nenhum dos taxa representados conspecífico do material figurado porDarwin (1854). De facto,Zullo (1992) considerou que os espécimes figurados pelo famoso naturalista representam uma espécie de ArossiaNewman, 1982. ...
The preliminary study of the Cirripedia of the Pliocene of Vale de Freixo revealed four species of balanids previously unknown from the Portuguese Neogene, one of which with open nomenclature: Austromegabalanus (Notomegabalanus) decorus (Darwin), Balanus crenatus Bruguière, Concavus concavus (Bronn), Zulloconcavus sp. Two wall fragments collected show traces of bioerosion – Oichnus – being muricid gastropods the most likely producers.
... Concavine barnacles are known to occur in the Neogene deposits of the Alpine Molasse Basin. De Alessandri (1906) pointed out the presence of "Balanus" concavus Bronn, 1831 (=Concavus concavus) in lower Miocene strata of the Upper Marine Molasse of Switzerland, but under this specific name several taxa of specific and generic rank are lumped (Newman 1982;Zullo 1992;Carriol 2000). More recently, concavine barnacles (belonging to the genus Chesaconcavus Zullo, 1992) have been reported by Carriol & Schneider (2008) Nebelsick (1989: pl. 4, figs. ...
Acorn barnacles are sessile crustaceans common in shallow-water settings, both in modern oceans and in the Miocene geological record. Barnacle-rich facies occur from polar to equatorial latitudes, generally associated with shallow-water, high-energy, hard substrates. The aim of this work is to investigate this type of facies by analysing, from the palaeontological, sedimentological and petrographical points of view, early Miocene examples from Northern Italy, Southern France and Southwestern Peru. Our results are then compared with the existing information on both modern and fossil barnacle-rich deposits. The studied facies can be divided into two groups. The first one consists of very shallow, nearshore assemblages where barnacles are associated with an abundant hard-substrate biota (e.g., barnamol). The second one includes a barnacle-coralline algae association, here named "barnalgal" (= barnacle / red algal dominated), related to a deeper setting. The same pattern occurs in the distribution of both fossil and recent barnacle facies. The majority of them are related to very shallow, high-energy, hard-substrate, a setting that represents the environmental optimum for the development of barnacle facies, but exceptions do occur. These atypical facies can be identified through a complete analysis of both the skeletal assemblage and the barnacle association, showing that barnacle palaeontology can be a powerful tool for palaeoenvironmental reconstruction.
... The first maxilla has eight small spines between the major upper pair of spines and single lower spine, whereas Ren's specimens have 16 intermediate spines. Newman (1982) noted that in general the barnacles within the Balanus amphitrite complex have a relatively small shell with smooth or weakly ribbed walls, weakly developed scuta, and commonly inhabits shallow or intertidal brackish environments. Puspasari et al. (2001) and Puspasari (2001) summarized several diagnostic characteristics of the B. amphitrite complex including the structures between the sheath and inner lamina, parietal tubes, and armature of the thoracopods. ...
The intertidal barnacle Balanus zhujiangensis Ren, 1989, is reported from Okinawa, Japan, for the first time. A supplementary description of this species is provided, and the specimens are compared to those from the Zhujiang River estuary which empties into the South China Sea. Several morphological characteristics of the shell, mouthparts, and armature of cirrus III suggest that B. zhujiangensis is related to the B. amphitrite complex rather than to B. trigonus by consistently having but a single row of pits on the external surface of the scutum and lacking an adductor ridge. The presence of scutal pits is interpreted as a means of facilitating the transmission of light through the valve thereby allowing the photoreceptors to detect changes in light intensity while the operculum is closed.
... The name Arossia was proposed Ьу Rasnitsyn et al. (1998) as а new genus for а new species of stephanoid apocritan (Insecta: Vespida=Hymenoptera) from the Barremian Weald Clay ofthe Auclaye Brickworks pit, Suпey, UК. lt has since been brought to our attention that the name is preoccupied: Arossia Newman, 1982 was proposed as а new subgenus for а living species of bamacle, Concavus panamensis (Rogers, 1948) Newman, 1982. Although the two taxa are never likely to Ье confused, this is nevertheless а case of homonomy at the genus-group level of zoological nomenclature. ...
... It also contains molluskan assemblages representing the Kadonosawa fauna (Chinzei, 1983) which is assigned to the early Middle Miocene (Masuda, 1969;Kitamura et al., 1986;Sato, 1991). The Hatatate Formation contains Middle (Newman, 1982). Miocene microfossil assemblages (Oda and Sakai, 1977;Shimamoto et al., 2001). ...
In situ fossil barnacle populations (Cirripedia, Balanomorpha) are described from the Miocene Natori Group of the Moniwa-Goishi area, northeast Japan. The Natori Group exhibits a transgressive sequence from subaerial lavas and volcaniclastics (Takadate Formation) to shoreface-shelf deposits (Moniwa and Hatatate formations) in ascending order. In the Takadate Formation, volcaniclastics locally intercalate bay deposits in which the fossil barnacle Balanus bisulcatus occurs on gravel surfaces in situ. The Takadate Formation is unconformably overlain by the basal conglomerate of the Moniwa Formation, which is interpreted as a ravinement deposit formed in an open coast. An extinct barnacle species, Concavus sendaicus, is found as gregarious clusters and solitary individuals on boulder surfaces in the basal conglomerate in situ. These features suggest that B. bisulcatus inhabited in bay environments at the early transgressive stage. On the other hand, C. sendaicus was distributed over open-coast environments during subsequent marine flooding. The in situ C. sendaicus provides strong evidence for the reconstruction of the extinct species habitat.
... The first maxilla has eight small spines between the major upper pair of spines and single lower spine, whereas Ren's specimens have 16 intermediate spines. Newman (1982) noted that in general the barnacles within the Balanus amphitrite complex have a relatively small shell with smooth or weakly ribbed walls, weakly developed scuta, and commonly inhabits shallow or intertidal brackish environments. Puspasari et al. (2001) and Puspasari (2001) summarized several diagnostic characteristics of the B. amphitrite complex including the structures between the sheath and inner lamina, parietal tubes, and armature of the thoracopods. ...
The intertidal barnacle Balanus zhujiangensis Ren, 1989, is reported from Okinawa, Japan, for the first time. A supplementary description of this species is provided, and the specimens are compared to those from the Zhujiang River estuary which empties into the South China Sea. Several morphological characteristics of the shell, mouthparts, and armature of cirrus III suggest that B. zhujiangensis is related to the B. amphitrite complex rather than to B. trigonus by consistently having but a single row of pits on the external surface of the scutum and lacking an adductor ridge. The presence of scutal pits is interpreted as a means of facilitating the transmission of light through the valve thereby allowing the photoreceptors to detect changes in light intensity while the operculum is closed.
... It also ¢ts with the group that forms the basis of Fistulobalanus as previously delimited by Henry & McLaughlin (1975) as Balanus pallidus group under Balanus amphitrite complex of the subfamily Balaninae Leach, 1817 on the nature of the shell structure has subsidiary tubes and multidenticulate labrum. And as noted by Newman (1982) it is consistent with the Balanus amphitrite complex in its small size, smooth or weakly ribbed shell wall and weakly or unornamented scutum, as well as being most commonly found in a shallow or intertidal brackish water environment. ...
The new mangrove barnacle, Fistulobalanus
sumbawaensis sp. nov. from Sumbawa Island, Indonesia shares the common morphological characteristics with the subfamily Amphibalaninae in having a shell wall of six, porous compartmental plates; radii solid; alae not cleft; basis calcareous, tubiferous in single layer. This new species belongs to the genus Fistulobalanus mainly in having parietal tubes arranged in two or more rows; primary tubes with transverse septa; subsidiary tubes formed between the primary septum and outer lamina. In particular, the new species is similar to F.
patelliformis in the shape of the shell, the transverse section of subsidiary tubes, the tergum and some characteristics of trophi and the cirri. However, this new species is easily distinguishable from all members of the genus Fistulobalanus in having no coloration or striations on the external surface of the shell, subsidiary tubes only appearing inside the ribbed part of the shell, and the scutum having a short adductor ridge.
... Both taxa are abundant, well-known inhabitants of intertidal environments of the present-day Mediterranean (e.g. Norris and Crisp 1953; Davidae 1963; Ross 1964; Relini and Giordano 1969; Newman 1982; Zullo 1992; Herbert et al. 2003). In general, the physical and biological processes controlling population dynamics presented in this paper appear to conform to those of present-day shore barnacles, allowing reasonable palaeobiological deductions about the formation of the dense barnacle concentrations in the Lower Pliocene deposits of basins of southeastern Spain. ...
Sessile barnacle assemblages, dominated by Concavus concavus (Bronn) and Balanus perforatus Bruguière, are very abundant in the Lower Pliocene deposits of the Almería-Níjar and Carboneras basins (southeastern Spain).
They occur in shallow-coastal siliciclastic and mixed siliciclastic-carbonate sediments, forming dense concentrations in two
contexts: (1) sheltered shallow-marine depressions and (2) the mouth of distributary channels feeding a delta lobe. Extensive
colonization took place during periods of quiescence with a high nutrient and food supply inducing the formation of hummocks.
Crowding of high morphotypes was presumably triggered by a high larval supply and recruitment rate. The barnacles are exceptionally
well preserved, often as in situ clusters, many with opercula, and include a range of ontogenetic stages with some specimens
preserving original colouration. Good preservation is probably due to sudden burial without further reworking.
Palaeoenvironmental data are fundamental in determining the manifold impacts of climate change. This paper argues that sessile barnacles are an excellent palaeoenvironmental proxy: they are present in nearly all nearshore environments, and their shell consists of diagenetically stable low-magnesium calcite and grows fast enough to record short-term variations. To demonstrate their utility, specimens from several Western Mediterranean Pliocene-Pleistocene barnacle-rich deposits are analysed herein, using for the first time a combination of sedimentology, taphonomy, stable isotope geochemistry and detailed comparisons with modern counterparts. Within shelf carbonate systems, barnacle diversity is highest in the shallow, nearshore waters and decreases moving offshore, thus providing a good proxy for the reconstruction of water depth and distance from the coastline. Barnacle taphonomy is also informative. Well-preserved complete specimens are characteristic of protected areas, whereas less well-preserved specimens occur in high-energy areas. The presence/absence of opercular plates is also particularly relevant for evaluating hydrodynamic conditions. As regards the C and O stable isotope ratios, due to the porous and coarse-grained nature of the deposits in which barnacle remains are usually found, the shells are often exposed to meteoric water percolation during diagenesis. Among the analysed specimens, only those collected from fine-grained deposits displayed no evidence of alteration and had isotopic ratios in line with those of their modern counterparts. These fossils display intra shell variations that in modern barnacles have been related to short-term environmental changes (e.g., seasonal cycles). These results demonstrate that barnacles can always be useful for detailed paleoenvironmental reconstructions based on skeletal assemblages. Furthermore, with further research aimed at gathering more data and assessing potentially interfering signals, they could become useful proxies for paleoseasonality.
Plio-Pleistocene cirripeds from eastern Sicily and a review of Mediterranean Cainozoic species
The new balanid subfamily Concavinae is based on the genus Concavus Newman, 1982, and includes five genera and 29 species-group taxa. The subgenera Concavus, Tamiosoma Conrad, 1856 (senior synonym of Menesiniella Newman, 1982), and Arossia Newman, 1982, are raised to generic rank. Two new genera, Chesaconcavus (type species Balanus concavus chesapeakensis Pilsbry, 1916) and Paraconcavus (type species Balanus concavus pacificus Pilsbry, 1916), are proposed, based on fossil and extant species from North America. New species include Tamiosoma advena from the Pliocene of southern Florida, Chesaconcavus rossi and C. santamaria from the middle Miocene of Maryland, C. myosulcatus from the upper Miocene of Virginia, Arossia newmani from the middle Miocene of Maryland, A. rubra from the upper Miocene of central California, A. aurae from the lower Pliocene of North Carolina, and Paraconcavus margaritanus from the upper Miocene of southern California. Concavines, a predominantly Northern Hemisphere group, appear in the late Oligocene in North America and Europe, diversify and spread rapidly in the Neogene, and, with the exception of a few eastern Pacific relicts, become extinct before the late Pleistocene.
The purpose of this paper is to document recent additions to the South African barnacle (Cirripedia) fauna. New species records were obtained by examining accumulated collections of unidentified material in the Iziko South African Museum, as well as via material collected directly by the authors. Fourteen species, none of which are new to science, are added to the known fauna, raising the total number of barnacle species recorded from South Africa by over 20%, from 69 to 83 species. Ten of the additional species are offshore benthic forms and represent deep-water taxa whose ranges are extended into South African waters. The remaining four are coastal inshore or surface-dwelling species, of which three are considered recent anthropogenic introductions. The large number of additions to the fauna can be attributed to a long absence of taxonomic expertise on this group in the region and consequent accumulation of unidentified samples. A high proportion of the deep-water samples examined contained previously unreported species, suggesting that additional sampling of hard abyssal substrata in the region is certain to yield yet more new cirripede records.
A checklist of the barnacles (Cirripedia: Thoracica) of Singapore and its neighbouring waters is presented. Currently 48 species are recognised from Singaporean waters. Singapore is the type locality for three genera (Hoekia recorded in Singaporean waters are marked and the locality Singapore is noted in bold type in the distribution data. Species not specifically recorded from Singaporean waters but whose presently recorded distributions encompass them are also included in the checklist. The checklist accommodates contemporary information, such as cladistic, molecular, ontogenetic, larval, ecological and phylogeographic data, as well as recent revisions and new species records of intertidal, shallow water and deep water species. Additionally, it also provides information on type species and details of specific hosts where applicable. The checklist and its comprehensive bibliography will inform future workers, aid identification of barnacle species and allow the thoracican cirripede fauna of Singapore to be considered in a broader regional context.
The large- to giant-sized balanids, mass-aggregated in a tempestite of the Lower Pliocene ( Zanclean) section at Rafina near Pikermi, Greece, represent a single species, Concavus ( Concavus) concavus ( DARWIN, 1854) [ non BRONN, 1831], the taxonomy of which is revised. The peculiarly shaped forms 'raphanoides' and 'scutorum' are ecophenotypes of this species; the same applies to the 'tulipiformis' specimens reported previously from this section. Discussed are dynamic events controlling the life and death conditions in the nearshore ( offshore) environment of Rafina, where the giant specimens of Concavus ( Concavus) concavus ( DARWIN, 1854) flourished through several successive generations. As opportunistic species they adopted the r-selection reproduction strategy in order to dominate over other biota. The intermittent action of high-energy agents was responsible for the production of balanid-shell hash that involved taphonomic feedback. This enabled further colonization of the biotope and the growth of multiphased ( bouquet-like and pyramidal) clusters. The demise of the monospecific balanid communities is ascribed to a heavy storm which stirred-up the whole balanid-bearing sequence, to produce a proximal tempestite. This final burial and subsequent depositional lull in the Rafina environment was favoured by a temporary deepening of the whole Lower Pliocene ( Zanclean) basin beyond the bathymetric range in which the balanid population could survive.
The association of the Galápagos Islands with the birth of Darwinian evolutionary theory has inspired many detailed studies of their fauna and flora. Yet the cirripeds, the only invertebrates to receive Darwin’s personal attention as a systematist, are virtually unknown for the region. His monographs on the Cirripedia (Darwin, 1851,1854) contain references to three species collected during the voyage of the Beagle: Balanus tintinnabulum (Linnaeus) and Tetraclita porosa (Gmelin) from rocky intertidal areas, and Platylepas decorata Darwin from marine turtles.
Two Concavinae occur in the Miocene of the Aquitaine Basin (France): Zulloconcavus CARRIOL, 2000 (type species Z. miosensis CARRIOL, 2000) from the Middle Miocene and Alessandriella n. gen. (type species A. saucatsensis n. sp.) from the Lower Miocene. This new genus possesses externally striate scuta, simple alar sutural edge, parietal tubes with transverse septa and sheath with tubes.
A new mangrove barnacle, Balanus thailandicus sp. nov., is described from the Puyu Canal, Tamalung Bay, Satun, southwest Thailand. The new species has the common characteristics of the Balanus amphitrite complex and is more or less sympatric with B. reticulatus, B. amphitrite, B. patelliformis, and B. variegatus. Balanus thailandicus is most similar to B. reticulatus, but it is clearly distinguishable from the last three mentioned species of the complex in having the parietes folded or ribbed in adults but smooth in juveniles, parietal tubes solid in the upper half and interrupted by transverse septa in the lower part, the large number of articles in the anterior ramus of cirrus I, and erect hooks below the posterodistal angles of the articles of cirri III and IV.
Since Leach’s (1817) original description, the concept of the family Balanidae has undergone many modifications, nowadays comprising about 90 recent species distributed in three subfamilies and twelve genera. Most of the recent suggestions were directed toward an attempt to render the classification of the family more natural. However, there is still a lack of comparative anatomical studies dealing with the whole family, although some subfamilies and genera have been studied in more detail (e.g., Balanus amphitrite complex; Megabalaninae, Concavinae and Fistulobalanus). The monophyly of the Balanidae and their subgroups also need to be ascertained.
The present study furnishes a comparative analysis of various morphological structures from the opercular plates, parietes, mouth and appendages in order to establish primary homologies (sensu Pinna, 1992) between states found among the balanids and three functional outgroups (Chirona, Semibalanus, and Membranobalanus). Twenty-three species representing each of the currently accepted taxa of Balanidae were compared. This analysis demonstrated that there are many underestimated characters, such as appendages, and others that need a more accurate evaluation because to their high interspecific variation (e. g. scutal and tergal sculpture) among the Balanidae. Some morphological patterns are described and Homologies are proposed.
A new genus and a new species of Concavinae, Zulloconcavus miosensis,
is described from the Serravallian of Mios (Gironde, France). This genus possesses
externally nodose scuta with tergal segment not inflected and terga externally
without radial striae but with a long spur.
Résumé: Description d'un nouveau genre et d'une nouvelle espèce de Concavinae,
Zulloconcavus miosensis, provenant du Serravallien de Mios (Gironde, France).
Ce genre présente un scutum dont la face externe est parcourue par de profondes
stries rayonnantes entre lesquelles, sur les bourrelets d'accroissement qu'elles
incisent, se développent des nodosités. Le très étroit segment tergal du scutum
n'est pas plié. Le tergum a un long éperon et une face externe dépourvue de stries
longitudinales.
Zusammenfassung: Beschreibung einer neuen Art und Gattung von Concavinae,
Zulloconcavus miosensis, aus dem Serravallien aus Mios (Gironde, Frankreich)
stammend. Die Art besitzt ein Scutum mit Knoten an der äußeren Seite und einen
nicht gefaIteten Abschnitt, dazu noch ein Tergum mit langem Sporn und eine äußeren
Seite ohne Längsrillen.
A new species of Concavinae, Chesaconcavus gurlarnensis, is described from the Upper Burdigalian (Ottnangian) Upper Marine Molasse deposits of Lower Bavaria (Germany). This species is characterized by narrow scuta with truncate basiscutal angle and ridge for lateral depressor muscle extending from apex to base of lateral depressor muscle pit, and terga with short, truncate spur, and open spur furrow. It is the first Chesaconcavus found in Europe.
German
Eine neue Spezies der Concavinae, Chesaconcavus gurlarnensis, aus dem Oberen Burdigal (Ottnangium) der Oberen Meeres-Molasse in Niederbayern (Deutschland) wird beschrieben. Die Art besitzt schmale Scuta mit abgestutztem basiscutalem Winkel. Die Rippe für den lateralen Depressor-Muskel erstreckt sich vom Apex bis zur Basis des lateralen Depressor-Muskeleindrucks. Die Terga zeigen eine kurze, abgestutzte Rippe mit offener Rippen-Furche. Es handelt sich hierbei um den Erstnachweis der Gattung Chesaconcavus in Europa.
French
Description d'une nouvelle espèce de Concavinae, Chesaconcavus gurlarnensis, provenant de la molasse marine supérieur du Burdigalien supérieur (Ottnangien) de Basse-Bavière (Allemagne). Cette espèce présente d'étroits scuta à l'angle basi-scutal tronqué et à l'arête du muscle dépresseur latéral rayonnant de l'apex à la base de la cavité du muscle dépresseur latéral, ainsi que des terga à court éperon tronqué dont le sillon est ouvert. Il s'agit du premier Chesaconcavus trouvé en Europe.
Geometric design in the barnacle genus Balanus has been studied in relation to variation in adult shell form, that includes differences among species, and size‐related changes in shape. The genus comprises 40 Recent species, and as a group these display a more or less constant morphology over an extraordinary size range (10 to 200 mm in basal length).
Linear and volumetric measurements were collected from 232 adult individuals of 14 species representing the variation in size, shell form and shell design thought to occur in the genus. Specimens were chosen to represent the size ranges of the species. Only isolated individuals growing on planar surfaces were used; shells were complete, undamaged and undistorted.
Shell form differs among taxa, and no two species scale alike; intraspecific variation for five ratio variables shows strong allometry over the adult size range of each species. As size increases, there is a trend for the basis and orifice to maintain their shapes or to become slightly more elliptical, and for shells to become more conical and proportionately taller.
Throughout their size ranges, species can be described by these geometries: paraboloid (6 species), frustum of an ellipsoidal cone (5 species), frustum of a cone (2 species) and a cone (1 species). Shell geometry is not a function of species size, but there does appear to be a correlation between shell geometry and shell volume. Species with relatively small shell volumes are described by a frustum of an ellipsoidal cone, or by a cone, while those with a relatively large shell volume are described by a paraboloid, or by the frustum of a cone.
Barnacles are some of the most conspicuous and well-known ship fouling organisms in the world and thus many species no doubt
owe parts of their modern distribution to human-mediated translocations over the past several centuries. Reviewed here, as
a window into global patterns, are the introduced, cryptogenic, and range expanding barnacles of the Atlantic and Pacific
coasts of North and South America. Five species of thoracic barnacles have invaded the Pacific coasts of the Americas: Amphibalanus improvisus, A. eburneus, and A. subalbidus, all from the Atlantic, and A. amphitrite and A. reticulatus from the Indo-West Pacific. Seven species have invaded the Atlantic coasts of the Americas; six of these are from the Pacific:
A. amphitrite and A. reticulatus (shared as invaders with the Pacific coast) , and Balanus trigonus, B. glandula, Striatobalanus amaryllis, and Megabalanus coccopoma. The Western North Atlantic A. subalbidus has invaded the Western South Atlantic. Striking are the few barnacle invasions that have occurred on the Pacific coast of
South America and these species (A. improvisus, A. amphitrite and A. reticulatus) are reported only from northernmost locations (Ecuador, Colombia, and Peru) . For the first 100 years (1853–1955) two species,
A. amphitrite and A. improvisus, constituted the majority of invasion events in the Americas, the sole exception being the arrival of the Pacific Balanus trigonus in the 1860s and 1870s in the Atlantic. After 1955, the first records of invasions of A. reticulatus, A. eburneus, B. glandula, M. coccopoma, and S. amaryllis appear, an increased diversity of introductions in close concert with general observations of increasing invasions globally
of marine organisms after World War II. Known since the 1970s in Brazil, M. coccopoma appears to be responding to warming northern latitudes and has expanded to North Carolina as of 2005. The native Western
Atlantic barnacle Chthamalus fragilis arrived in New England in the 1890s, a range expansion perhaps facilitated by an earlier coastal warming period concomitant
with the decline in abundance of its colder-water competitor Semibalanus balanoides, although the latter also appears to have expanded south on the North American Atlantic coast in the twentieth century due
to increased habitat availability. Chthamalus is predicted to move north, and Semibalanus is predicted to return to its historical range, both due to continued warming. In turn, the native Eastern North Pacific
barnacle Tetraclita rubescens is expanding north due to coastal warming as well. Future invasion scenarios include increased introductions facilitated
through a newly expanded Panama Canal, the potential arrival of Austrominius modestus on the North American Atlantic coast (despite its failure to do so throughout the last half of the twentieth century) , and
the arrival on the warmer North and Central American Pacific coasts of Chthamalus proteus.
KeywordsCirripedia-barnacles-invasive-introduced-range expansion-climate change-
Balanus
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Amphibalanus
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Megabalanus
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Loxothylacus
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Austrominius
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Chthamalus
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Tetraclita
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Semibalanus
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Fistulobalanus
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Paraconcavus
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Striatobalanus
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: 1-18, pl. 1. Pilsbry, H. A., 1909. Report on barnacles of Peru, collected by Dr. R. E. Coker and others. Proc. U.S. Nati. Mus., 37: 63-74, pis. 16-19. -, 1916. The sessile barnacles
Symposium, Caracas, 12-16 July Cirripedia of the Iranian Gulf A new species belonging to the Balanus amphitrite Darwin group (Cirri pedia, Balanomorpha) from the late Pleistocene of Japan; an example of peripheral specia tion
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, Cooperative Investigations of the Caribbean and Adjacent Regions-II: F.A.O. Fish. Rep., 200: 407-425 (Symposium, Caracas, 12-16 July, 1976). Utinomi, H., 1969. Cirripedia of the Iranian Gulf. Vidensk. Meddr. dansk. naturh. Foren., 132: 79-94. Yamaguchi, T., 1980. A new species belonging to the Balanus amphitrite Darwin group (Cirri pedia, Balanomorpha) from the late Pleistocene of Japan; an example of peripheral specia tion. Journ. Paleontol., 54 (5): 1084-1101. -,
A comparison with the tethyan Balanus concavus group Zoogeographie affinities of the Balanomorpha (Cirripedia: Thoracica) of the Eastern Pacific
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Thoracic Cirripedia of the San Diego Formation, San Diego County, California. Contrib. Sei. Los Angeles County Museum Thoracican Cirripedia of the Lower Pliocene Pancho Rico Formation, Salinas Valley
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