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The Spheniopsidae are today represented by five living species of Spheniopsis and nine of Grippina, distributed in the Pacific and Atlantic Oceans. Little is known of their anatomy and the phylogenetic position of the family within the Bivalvia is debated. In order to investigate these questions, the functional morphology of Grippina coronata obtained from the continental shelf off Rio de Janeiro and Espírito Santo States, Brazil, has been examined. Less than 2 mm in shell length, the siphonal apparatus of G. coronata is complex, with seven sensory papillae, and the ctenidia are reduced to transverse septa pierced by four pairs of ciliated pores. There are no labial palps and the stomach is of Type II with epibenthic harpacticoid and ostracod prey identified inside it. Although there is an intestine producing faeces, the stomach also possesses a unique waste storage pouch for exoskeletal remains of digested prey. Collectively, these features suggest that the Spheniopsidae comprise carnivorous taxa belonging to the Cuspidarioidea within the Anomalodesmata. Grippina coronata is a self-fertilizing simultaneous hermaphrodite that, uniquely, broods fertilized oocytes within the ovarian follicles and, thereby, provides the first example of intrafollicular fertilization and brooding in the Bivalvia. Release of the encapsulated oocytes must be by parental death, which coincidentally releases the exoskeletal remains from the storage pouch. Such post mortem semelparity creates a life-history trait hitherto unrecognized in the Bivalvia.
The Anomalodesmata, lying within the basal Heterodonta (Bieler et al., 2014), comprises a richly diverse assemblage of exclusively marine bivalves, which vary widely both in overall form and in the highly specialized niches they occupy. As a consequence, they exhibit a high diversity of unique anatomies. The Anomalodesmata comprises some eight superfamilies (Runnegar, 1974; Morton, 1981a, 2012), one of which is extinct (the Ceratomyoidea), although most putative families of the Palaeozoic Pholadomyoidea also occur only as fossils (Runnegar, 1974). The adaptive radiation expressed by the living representatives of the Anomalodesmata is spectacular but still poorly appreciated, because most extant taxa occur typically as solitary individuals in highly specialized niches. Two anomalodesmatan clades stand out, for mention, however. The first includes the adventitious tube-dwelling representatives of the Clavagelloidea (Morton, 2007). The second comprises the remarkable deeper-water predatory bivalves often collectively referred to as the septibranchs. The classification schemes erected for these latter bivalves have undergone many changes over the last 50 years and these were reviewed by Morton (1982: Table II). The septibranchs are today thought to be represented by the fossil Orthonotoidea (Bieler, Carter & Coan, 2010) and three extant superfamilies: the Verticordioidea (Verticordiidae, Euciroidae and Lyonsiellidae), Poromyoidea (Poromyidae) and the Cuspidarioidea (Keen, 1969; Bernard, 1979; Bieler et al., 2010). Hitherto, this last superfamily has been considered to comprise but one extant family—the Cuspidariidae—although, as will be described and discussed, the most recent classification by Carter et al. (2011) considered the Cuspidarioidea to also include the (unstudied) Halonymphidae, Protocuspidariidae and the Spheniopsidae (similarly unstudied). A fourth family of deep-water predatory bivalves, the Parilimyidae, is believed to be affiliated with the Pholadomyoidea (Morton, 1982).
The predatory anomalodesmatan bivalves have received some attention, focusing largely on anatomy. Prior to a greater appreciation of their familial diversity, such bivalves were judged to belong, collectively, to the Septibranchia, characterized by the presence of a horizontal muscular septum in place of the typical bivalve ctenidium. This generalization, however, is not true—Halicardia nipponensis Okutani, 1957 (Verticordiidae), for example, possesses a ctenidum (but no labial palps), a funnel-shaped mouth and a stomach for the processing and digestion of its copepod prey (Nakazima, 1967). Yonge (1928) was perhaps the first to bring the septibranch bivalves, Cuspidaria and Poromya, to wider scientific attention although Pelseneer (1888, 1911) and Ridewood (1903) had earlier made comprehensive studies of numerous septibranchs, the latter author describing the structure of the septum in detail.
Later, Reid & Reid (1974) examined aspects of the morphology and behaviour of Cuspidaria rostrata (Spengler, 1793) and C. obesa (Lovén, 1846) and Reid & Crosby (1980) subsequently undertook a study of how Cardiomya planetica (Dall, 1908) captured its prey. Reid (1978) made the only physiological study to date of any carnivorous bivalve, again C. planetica, demonstrating the presence of proteolytic enzymes in the stomach to digest its prey. Purchon (1956) examined the stomach architecture of Cuspidaria cuspidata (Olivi, 1792), classifying it, mistakenly, alongside that of the Protobranchia in his Type II, the Gastrodeuteia (Purchon, 1958) and 'sub-class' Oligosyringia (Purchon, 1962, 1987). The anatomies of the deep-water representatives of the Verticordiidae (Allen & Turner, 1974) and Cuspidariidae and Poromyidae (Allen & Morgan, 1981) have been described, the latter paper also discussing the evolution of the septibranch condition. In a series of studies, Morton (1981b, 1982, 1984, 2003) undertook investigations of the anatomies of Poromya granulata (Nyst & Westendorp, 1839) (Poromyidae), Parilimya fragilis (Grieg, 1920) (Parilimyidae), Lyonsiella formosa (Jeffreys, 1881) and Bentholyonsia teramachii (Habe, 1952) (Lyonsiellidae), also reviewing statocyst and siphon structure in representatives of the various families (Morton, 1985, 1987). Oliveira & Sartori (2013) examined the arenophilic radial glands in the siphons of Cuspidaria obesa and a species of Cardiomya.
In his comprehensive reviews of the abyssal and hadal bivalves collected from many parts of the earth's oceans, Knudsen (1967, 1970) described the shells of numerous species of the Septibranchia, while Simone & Cunha (2008), Oliveira & Absalão (2010) and Absalão & Oliveira (2011) have reviewed the species of Verticordiidae, Lyonsiellidae and Cuspidariidae, respectively, occurring in the deeper waters off Brazil.
The Spheniopsidae Gardner, 1928, known since the Oligocene, are represented today by a number of small bivalves with shells usually less than 5 mm in length (Boss, 1982). Two genera occur in the Recent fauna, Spheniopsis Sandberger, 1861 and GrippinaDall, 1912, the first with five and the second with nine species, both recorded from the warm, tropical waters of the Americas, Africa and New Zealand (Verril & Bush 1898; Dall, 1912; Coan, 1990; Cosel, 1995; Coan, Valentich-Scott & Bernard, 2000; Marshall, 2002; Coan & Valentich-Scott, 2012; Machado & Passos, 2015). From the Atlantic Ocean, Spheniopsis senegalensisCosel, 1995 is recorded from off the coast of West Africa (Cosel, 1995), Spheniopsis triquetra (Verril & Bush, 1898) from North Carolina, Florida and the Bahamas (Coan & Valentich-Scott, 2012; Machado & Passos, 2015), and Spheniopsis brasiliensisMachado & Passos, 2015 and Grippina coronataMachado & Passos, 2015 were recently described from the southeastern Brazilian coast. In addition to Recent species, the Spheniopsidae have five fossil species, belonging exclusively to the genus Spheniopsis (Coan, 1990).
The anatomy of representatives of the Spheniopsidae is virtually unknown, with little snippets of information available on muscle scars on the shells and the dried tissues of Grippina californicaDall, 1912 (Coan et al., 2000; Mikkelsen & Bieler, 2008). Associated with this, the phylogenetic position of the Spheniopsidae has been the subject of recent debate (Marshall, 2002; Harper, Dreyer & Steiner, 2006; Bieler et al., 2010). The family was considered to be allied to the Heterodonta (Gardner, 1928; Keen, 1969; Coan, 1990; Cosel, 1995), notably between the Corbulidae and the Hiatellidae, i.e. within the Myoidea (Boss, 1982; Amler, 1999; Coan et al., 2000), Bieler & Mikkelsen (2006) considering the Spheniinae to be a subfamily of the Myidae. Based on the observations that species of Grippina have sunken resilifers with a lithodesma and some have an exterior commarginal microsculpture most similar to that seen in some anomalodesmatans, Marshall (2002) concluded that the Spheniopsidae should be placed in the Cuspidarioidea. Bieler et al. (2010) accepted this view, albeit questioningly, and suggested that "the family would profit from molecular analyses" (p. 133). Despite such information not being available, Carter et al. (2011) agreed with Marshall's view and placed the Spheniopsidae firmly within the Cuspidarioidea.
In the absence of such a molecular analysis and comprehensive information regarding the anatomy of any spheniopsid, however, uncertainties concerning their relationship with the Myoidea (Amler, 1999; Coan et al., 2000) or the Cuspidarioidea still remain. This work is a study of the anatomy of the minute (<2 mm) spheniopsid G. coronata—recently collected off the coast of Brazil, described as new and the shell described in detail by Machado & Passos (2015). The aim of the study, therefore, was to reveal those characters that would assist in identifying the systematic affiliations of the Spheniopsidae. In addition, however, it also examines aspects of the species' anatomy that show how such a minute bivalve is adapted to life on the continental shelf and deeper sea bed, and to a carnivorous mode of feeding.
Individuals of Grippina coronata were obtained from bottom samples collected by a box corer during research conducted under the 'Habitats Project—Campos Basin Environmental Heterogeneity', in February and July 2009. Sampling was undertaken on the shelf and continental slope of the Campos Basin, an area of oil and natural gas exploitation off the southeastern coast of the Brazilian states of Rio de Janeiro and Espírito Santo. Numerous bottom samples were obtained from depths of 12 to 3200 m and sieved through a 0.5-mm mesh. From 28 of these samples collected at depths of 21 to 53 m, 67 empty shells and 20 living individuals of G. coronata were obtained. These were initially fixed in 4% formalin and then preserved in 70% alcohol. Preserved individuals were dehydrated in an ascending series of ethanol, critical-point dried and mounted on aluminum stubs for examination by a scanning electron microscope (SEM). For histological purposes, two individuals of G. coronata were decalcified in a solution of 100 ml distilled water containing 0.88 g of NaCl and 1.02 g of ascorbic acid. These were dehydrated in an ascending series of ethanol and embedded in methyl methacrylate (Historesin®) to obtain serial transverse and sagittal sections 5 μm thick. Histological slides and SEM stubs have been deposited in the Museum of Zoology of the State University of Campinas (ZUEC) under accession numbers ZUEC-BIV 6181, 6183, 6203 and 6204.
Abbreviations used in figures
- aam
Anterior adductor muscle (or scar)
- an
- aprm
Anterior pedal retractor muscle (or scar)
- asm
Anterior septal muscle (or scar)
- at
Anterior hinge tooth
- atl
Anterior hinge lock
- au
Auricle (of heart)
- bo
Brooded oocytes
- cc
Collagen coat
- ci
- co
Collar around crystalline style sac
- cs
Crystalline style
- css
Crystalline style sac
- d
- dc
Digestive cell
- dd
Digestive diverticulae
- ddc
Dehiscing digestive cell
- do
Developing oocyte
- dt
Digestive tubule
- eo
Encapsulated oocyte
- es
Exhalant siphon
- f
- fimmf
Fused inner and inner surfaces of the middle mantle folds
- g
Gland in the foot
- gs
Gastric shield
- ha
- hg
- iec
Inner epithelium cells
- ilm
Inner lip of the mouth
- imf
Inner mantle fold
- ipi
Ingested prey item
- i
- is
Inhalant siphon
- isc
Infraseptal chamber
- l
- li
- m
- ma
- mc
Mantle cavity
- mg
- mi
- mm
Mantle margin
- mmf
Middle mantle fold
- mo
Mature oocyte
- o
- obe
Ooctye being encapsulated
- obf
Oocyte being fertilised
- oc
Oocyte capsule
- odd
Openings to the digestive diverticulae
- of
Ovarian follicle
- omf
Outer mantle fold
- ommf
Outer surface of the middle mantle fold
- ov
- p
- pam
Posterior adductor muscle (or scar)
- peg
Pedal groove
- pega
Pedal ganglia
- pg
Periostracal groove
- pga
Pedal gape
- pl
Pallial line
- pprm
Posterior pedal retractor muscle (or scar)
- ps
Pallial sinus
- psm
Posterior septal muscle (or scar)
- pr
- pr (I)
Prodissoconch I
- pr (II)
Prodissoconch II
- prm
Pallial retractor muscle
- pt
Posterior hinge tooth
- ptl
Posterior hinge lock
- r
- sa
Sorting area in the stomach
- sc
Secretory cell
- se
- selm
Septal elevator muscle
- sem
Septal membrane
- sep
Septal pore
- sep (1)–(4)
Septal pores (1–4)
- so
Siphonal open
- sp
Siphonal papilla
- spb
Siphonal papilla base
- spc
Sphincter cells
- spe
- spg
Septal pedal gape
- sph
- spm
Sperm morula
- spt
- srp
Skeletal remains of prey
- ssc
Supraseptal chamber
- st
- sta
- stl
- t
- tf
Transverse fibres
- v
Ventricle (of heart)
- vm
Visceral mass
- wsp
Waste-storage pouch
In side view, the shell of Grippina coronata (Fig. 1A) is subtrigonal, as in G. californica (Coan et al, 2000: 481, pl. 103), with the posterior face foreshortened and distinctively pointed posterodorsally, but more flattened posteroventrally. The anterior face is more smoothly rounded and oval. The shell is transparent, as illustrated in Figure 2A, which also shows oocytes being brooded (bo) internally, and its surface is smooth with only fine commarginal growth lines. The most characteristic feature is the dorsal prodissoconchs, described below. From the dorsal aspect (Fig. 1B), the shell is slightly sinusoidal with the anteroposterior greatest width (a–b) located through the dorsal umbones to the midventral border. In ventral view (Fig. 1C), the slightly inequivalve shape seen dorsally is not obvious and the left and right valve borders fit together tightly. The posterior shell margin from this aspect is variably flattened, as noted above. Seen from the posterior aspect (Fig. 1D), the slight sinusoidal form of the valve margins is again obvious and the greatest shell width, left to right (x–y), is somewhat dorsal to the midpoint of the dorsoventral axis of the shell. This is also obvious when seen from the anterior aspect (Fig. 1E). There are also indications of a lunule anteriorly and an escutcheon posteriorly (Machado & Passos, 2015), as identified for G. californica (Coan et al., 2000), although such structures might have been designated when the Spheniopsidae were considered to have heterodont affinities, since they are only expressed subtly.
The prodissoconchs of G. coronata (Fig. 3) are large in relation to the overall size of the shell (<2 mm). Apically, there is a prodissoconch I (pr (I)), about 260 μm in length (254 ± 14 μm, Machado & Passos, 2015), comprising a biconical apex surrounded by a raised rim. Surrounding this is a plain prodissoconch II (pr (II)) about 350 μm in diameter. There does not appear to be a nepioconch and thus the dissoconch (d) forms smoothly around prodissoconch II in a uniformly blended manner. Machado & Passos (2015: fig. 3) provided photographic images of the larval shell stages of G. coronata.
The internal hinge plates of the right and left shell valves of G. coronata (Fig. 4A, B) were also described by Machado & Passos (2015). That of the right bears anterior and posterior hinge teeth (at, pt), which fit beneath equivalent locking points (atl, ptl) that are merely indentations on the thickened hinge of the left valve. The two hinge teeth flank a central amphidetic ligament (l) located upon a resilifer and ventrally formed into a lithodesma (li), the structure of which has been described for a number of species of the Cuspidariidae by Bernard (1974) and for New Zealand representatives of the Spheniopsidae by Marshall (2002: fig. 21). Yonge & Morton (1980: fig. 11) described the ligament and lithodesma of Cuspidaria cuspidata and showed how it was formed. The lithodesma of G. coronata resembles those of the other spheniopsids described above, although it is too tiny to be dissected out and illustrated here.
An interior view of the left shell valve of G. coronata (Fig. 5) shows the muscle scars. These are simple and comprise approximately equal-sized anterior and posterior adductor muscles (aam, pam), each internally and dorsally flanked by small septal retractor muscle scars (asm, psm). These are indistinguishable as scars from the anterior and posterior pedal retractor muscle scars (aprm, pprm). The pallial line (pl) is thin and posteriorly forms a shallow pallial sinus (ps).
The withdrawn siphons of G. coronata (Figs 2D, 6A), as seen from the posterior aspect, are separate and formed by fusion of the inner folds and the inner surfaces of the middle mantle folds and are thus of Type B (Yonge, 1957). They comprise a conical exhalant siphon (es), only 100 μm in diameter, and a larger, 200 μm, inhalant one (is). The former siphon has a single sensory papilla (identified as tentacles by Reid & Crosby, 1980, for Cardiomya planetica) dorsally and two others laterally (sp). The latter siphon has four such papillae all located ventral to the siphon and sitting upon a raised base (spb). A single papilla is illustrated in Figure 6B and has an embayed tip from which arise up to approximately 12 long cilia, each with a paddle-shaped tip (Fig. 6C).
The ventral mantle margin at the pedal gape of G. coronata (Fig. 7A) is simple. Each lobe comprises an elongate inner fold (imf), a tiny middle fold (mmf) and a similarly small outer fold (omf). From the periostracal groove (pg) between the latter two folds arises an exceedingly thin periostracum (p). A pallial retractor muscle is not obvious here but, instead, there is a capacious haemocoel (ha), which is characterized by transverse muscle fibres (tf) that cross-connect the outer and inner mantle surfaces, presumably helping to sustain tonus. The inner surface of the mantle is characterized by an epidermis that has strangely vacuolated cells and one of these is illustrated in greater detail in Fig. 7B. No function can as yet be ascribed to these.
Posterior to the pedal opening, the left and right mantle margins fuse (Fig. 8); Figure 2B shows this posterior fusion forming the siphonal opening (so). In transverse section, fusion is seen to be of the inner folds and the inner surfaces of the middle mantle folds (fimmf). This, therefore, as with the siphons, is of Type B (Yonge, 1948, 1982). In this location, the pallial retractor muscles are more obvious (prm), extending through the haemocoelic spaces (ha) of the outer mantle fold (omf). The middle mantle folds are here still small and comprise the outer surfaces only (ommf). From between these outer surfaces and the inner surfaces of the outer mantle folds, within the periostracal grooves (pg), arises the periostracum (p). Oliveira & Sartori (2013) demonstrated the presence of arenophilic radial mantle glands in some species of Cuspidaria (e.g. C. obesa) and Cardiomya (e.g. C. cleryana (d'Orbigny, 1842)), but not others. Such glands have not been identified for G. coronata.
Organs of the mantle cavity
The organs of the mantle cavity of G. coronata, as seen from the right side after removal of the right shell valve and mantle lobe, are illustrated in Figure 9. The septum (se), with its four pairs of pores (sep), is suspended in the mantle cavity by anterior (asm) and posterior septal retractor muscles (psm), the latter larger than the former. As shown earlier, these muscles attach to the shell internal to the anterior (aam) and posterior (pam) adductor muscles and share an attachment with the minute and equivalent anterior and posterior pedal retractor muscles. The visceral mass (vm) is situated above the septum and has a posterior foot (f) that extends through a gape in the septum anteriorly and can, presumably, also be extended out from the similarly anterior pedal gape (pga) in the mantle margin (mm) to effect burrowing. Posteriorly, there are the siphons, the inhalant (is) opening into the infraseptal chamber (isc), the exhalant (es) into the supraseptal (ssc). Within the visceral mass, the digestive diverticulae (dd) are located anterodorsally with brooded oocytes (bo) present virtually everywhere else, left and right. The heart with a central ventricle (v), penetrated by the rectum (r), with left and right auricles (au), is situated posterodorsally between the visceral mass and the posterior adductor muscle. The rectum terminates in an anus (an) on the posterior face of this muscle, so that faeces exit from the individual via the exhalant siphon.
Also shown in Figure 9 (arrows) are the postulated cleansing currents of the infraseptal chamber. The positions and orientations of these cleansing currents are based on the locations of ciliary activity in this chamber, shown in subsequent figures. Having such short siphons, G. coronata must be feeding on prey items close to the sediment-water interface (discussed below) and it is inevitable, therefore, that some unwanted inorganic particles will be taken into the infraseptal chamber of the mantle cavity and need to be removed from it. This must be via the inhalant siphon, as in all bivalves.
Superficially, the presence of pedal retractor muscles within the visceral mass of G. coronata is undetectable. However, a transverse section (Fig. 10) through the left half of the visceral mass just posterior to the septum shows the posterior adductor muscle (pam) and the left posterior pedal retractor muscle (pprm). In this region of the body too, the inner fold (imf) of the mantle (ma) margin cavity extends across the mantle cavity (mc). The stomach (st), lined by a gastric shield (gs) secreted from the stomach epithelium (sc), is here approaching its posterior extremity and associated with it is a waste-storage pouch (wsp) containing the skeletal remains of prey (srp), which will be described in greater detail below. The base of the foot possesses glands (g) and this region of the visceral mass also sees the posterior termination of the gonads with, most obviously, mature oocytes (mo) and oocytes being encapsulated (obe), also to be described subsequently.
The septum of G. coronata is illustrated from the ventral aspect in Figures 2B and 11A. The muscular septum (se) is narrow posteriorly and wide anteriorly and has an overall length of 500 to 800 μm and a maximum width of 150 to 200 μm. Left and right halves are united by a septal membrane (sem), which separates anteriorly to create an opening (spg) through which the foot (f) can protrude. Along the length of the septum are four pairs of pores (sep(1) to sep(4)), which each have diameters of about 20 μm. Anteriorly, the septum ends at the mouth (m). There are no labial palps. Figure 11B shows a single septal pore in greater detail. Each one (sep) opens into the supraseptal chamber and is surrounded by a fringe of cilia (ci). The margin of the septal membrane (sem) is also fringed by cilia. It is probable that these cilia around the septal pores and along the margins of the septal membrane serve a cleansing function to keep the prey-capturing septum and its valves, the septal pores, free of accidentally inhaled sediment, as described and discussed above.
The foot (Figs 2B, E, F, 11C; f) of G. coronata extends through the septal pedal gape (spg) in the septal membrane (sem). Ventrally, the densely ciliated foot, with an overall length (when contracted) of 240 to 300 μm, has a pedal groove (peg) that may be the remnant of a juvenile byssal groove if, as in many bivalves, such a structure is produced to assist in the establishment of the juvenile in its chosen habitat. Morton (1981b) suggested for Poromya granulata that the foot, in addition to being responsible for burrowing, probably also served to push captured prey items into the mouth and seal the opening; it is likely that the same functions are served by the foot of G. coronata.
Figure 12 is a transverse section through the septum (se) of G. coronata. The central septal membrane (sem) divides the septum into left and right halves, which are connected to the shell by a rich array of septal elevator muscles (selm), responsible for the lifting of the septum to create the negative pressure inside the infraseptal chamber and thereby facilitate prey capture, as originally described for cuspidariids by Yonge (1928) and elaborated upon by Reid & Reid (1974).
The mouth (m) of G. coronata is shown in more detail in Figures 2C and 13. It has dimensions of 60 by 160 μm and is pleated dorsally and densely ciliated (ci) internally at its entrance, both ventrally and laterally. It has a structure similar to that described for Halicardia nipponensis by Nakazima (1967). There are no labial palps, which in the suspension-feeding bivalves process particles of collected seston or detritus prior to ingestion (Stasek, 1963). The anteriormost septal pores (sep(4)) are situated close to the mouth as is the medial foot (f). Figure 14 is a sagittal section through the mouth and oesophagus of G. coronata. Externally, the mouth has an inner lip (ilm), but not an outer. As noted above, the mouth (m) is lined internally by ciliated cells (ci) and opens into an oesophagus (o). This is surrounded by elongate (c. 50 μm) sphincter cells (spc) that seem to be lined distally by microvilli (mi). The gastric shield (gs) lining the stomach and secreted by underlying cells (sc) abuts the mouth closely, but is separated from it by these swollen sphincter cells. The function of the cilia within the mouth thus seems to keep it clean, whereas the oesophagus can be closed by the sphincter cells from contact with the mouth, possibly when prey items have been pushed into the stomach, to stop them getting out before being digested. The sphincter may also create a closed sac—the stomach—in which digestion can proceed more efficiently.
Alimentary system
The organs of the mantle cavity and visceral mass of G. coronata are illustrated in sagittal section in Figure 2G, H and from the right side in Figure 15 (vm). The dotted line (Fig. 15) shows the approximate position of the septum (se) with the foot (f) penetrating it anteriorly at the septal gape. The visceral mass itself is situated between and defined by the anterior and posterior adductor muscles (aam, pam). The mouth (m) opens into the stomach (st), which is located anterodorsally in the visceral mass and is surround by its digestive diverticulae (dd). From the stomach's ventral border arises the crystalline style sac (css), which for a short distance is conjoined with the midgut (mg). The narrow midgut (c. 40 μm) makes a complete loop in the visceral mass and dorsally becomes the somewhat wider (c. 50 μm) hindgut (hg). This penetrates the ventricle (v) of the heart, passes between left and right posterior septal muscles (psm), over the posterior adductor muscle and terminates in an anus (an) on this structure's posterior face. This figure also shows brooded oocytes (bo), occupying most of the visceral mass and parts of the ovaries (ov) and testes (t) although, as will be shown later, these germinal tissues are, in fact, scattered widely in the visceral mass.
Figure 16A is a transverse section through the stomach of G. coronata. It shows how the stomach is almost entirely covered internally by the gastric shield (gs) and partly surrounded here by digestive tubules (dt). Inside the stomach of this individual was an ingested prey item (ipi), possibly a copepod. The cells of the digestive tubules are illustrated at higher resolution in Figure 16B. They show more darkly staining, pyramidal, secretory cells (sc) interspersed with digestive cells (dc) full of absorbed globules of different sizes and structure. Some of these cells appear to be dehiscing (ddc), i.e. releasing back into the tubule lumen vacuoles of, presumably, unwanted material that will eventually be returned to the stomach and midgut for eventual defaecation. The gastric shield (gs) (Fig. 16C) is produced by specialized secretory cells (sc) that are vacuolated basally and have darkly staining granules apically. A few studies, but not of cuspidarioids, have shown that the gastric shield is itself a source of digestive enzymes that are incorporated into its structure by such secretory cells (Halton & Owen, 1968; McQuiston, 1970).
A transverse section (Fig. 17A) through the stomach (also Fig. 2G, H, in sagittal section), here less than 1 mm in diameter, in the region of the conjoined crystalline sac and midgut shows once again the gastric shield (gs) overlying its secretory cells (sc). The opening to the crystalline style sac (css) is surrounded by a collar (co) of raised cells and flanked on the right side by a sorting area (sa) of convoluted ridges comprising ciliated cells (described for C. cuspidata by Purchon, 1956, 1958). The style sac (css) itself contains a stubby crystalline style (cs) and is composed of vacuolated cells that seem to be lined by microvilli (mi). The midgut (mg) separates quickly and basally from the crystalline style sac to pursue an independent course within the visceral mass (Fig. 15). Figure 17B illustrates the separated midgut in transverse section and shows it to comprise tall columnar cells, which appear vacuolated and have apical nuclei such that the lumen of this region of the intestine is exceedingly narrow. The hindgut (Fig. 17C) is wider and surrounded by a coat of smooth muscle and collagen (cc). Its epithelium is convoluted into about eight ridges and is ciliated. The rectum is narrower, but also surrounded by a thick coat of muscle and collagen (cc) and, like the hindgut, has a ciliated and convoluted epithelium (Fig. 17D).
Figure 18A is a transverse section through the stomach in the region of the waste-storage pouch (wsp) arising from its left posterior end. This is situated directly under the mantle (ma) and contains the exoskeletal remains of ingested prey (ipi). In this location too, posterior to the sorting area (sa) and the collar (co) around the crystalline style sac, the stomach continues to be lined by the gastric shield (gs), secreted from specialized cells beneath it (sc) and contains an identifiable ingested prey item (ipi)—in this case a harpacticoid copepod. Also seen ventrally in the visceral mass (vm) is an ovarian follicle (of) containing mature oocytes (mo). Figure 18B is a detail of the junction between the stomach (st) and the waste-storage pouch (wsp). The two organs are connected by a sphincter (sph), which must regulate passage of the skeletal remains of ingested and digested prey from the stomach into the pouch. Such a structure has never before been identified for any other carnivorous (or protobranch) bivalve, although a left pouch is characteristic of all other bivalves, (Purchon, 1962). Whether or not these structures are analogous or homologous is unknown.
Histological sections through prey items in the stomachs of G. coronata (Fig. 2G, H; ipi) show benthic harpacticoid copepods (Fig. 19A) and benthic ostracods (Fig. 19B). These taxa (hc, os) can also be identified from their skeletal remains in the waste-storage pouch (Figs 2H, 19C; wsp), along with other unidentifiable skeletal remains (srp).
Figure 20 is a reconstruction of the internal architecture of the stomach as seen from the right side. The gastric shield (gs) covers most of the dorsal and posterior surfaces of the stomach. The crystalline style sac (css) with its stubby style (cs) and the conjoined midgut (mg) depart the stomach from its ventral border. Anterior to the collared entrance (co) to the crystalline style sac (css) is an anteroventral series of ciliated ridges that constitutes a sorting area (sa) and within this are two, left and right, openings into the digestive diverticulae (odd). The mouth (m) is located anteroventrally and opens into the stomach as the oesophagus (o), which has internal sphincters (sph). Recently, Tëmkin & Strong (2013) described the internal architecture of the stomach of Bathyneaera demistriata (Allen & Morgan, 1981) (Cuspidariidae) and showed an essentially similar structure to that of G. coronata, although without identifying a sorting area as such. Regardless, their illustration (Fig. 3) of B. demistriata shows extensive internal folds in the same region as seen in G. coronata.
It is not possible to detail the structure of the nervous system of G. coronata. Transverse sections through the tiny (50 μm) paired pedal ganglia (pega) in the visceral mass are illustrated in Figure 21. Separate from these is a pair of statocysts (50 μm) (sta) that are capsules formed by 4–5 cells (in transverse section), a few possessing cilia (ci) and each having a single darkly staining statolith (stl), about 20 μm in diameter, in the centre. Morton (1985) examined statocyst structure in numerous representatives of the Anomalodesmata and those of G. coronata are closely similar to those of other representatives of the Cuspidarioidea, i.e. of Type C in which they are distinctively separate from the pedal ganglia.
Reproductive system
A transverse section through the visceral mass (Fig. 22) shows the intestine (i), the tubules of the digestive diverticulae (dt), the foot (f) with a gland (g) at its base and elements of the reproductive system. Grippina coronata is a simultaneous hermaphrodite. The paired female ovaries are located virtually everywhere in the visceral mass, including the epithelium within which it is enclosed and, within its central region, separating two exceptionally large, what are here termed ovarian follicles (of). As will be seen, however, the tissues of the testes (t) are also present as mesodermal elements scattered similarly throughout the visceral mass and occur in association with the developing oocytes in the walls of the ovarian (and thus also testicular) follicles and elsewhere (Fig. 2G, H).
The paired ovarian follicles contain oocytes at various stages of maturity, including fertilized ones (Fig. 22). These developmental stages include developing oocytes (do), some of which are being encapsulated (eo) and measure up to c. 50 μm in diameter. The number of encapsulated oocytes (oc) within each ovarian follicle varies but, generally, there are about ten each, or a maximum of 20 (not including the many developing oocytes in the follicle walls). No oviducts or seminal ducts connecting the paired gonads to the supraseptal chamber have been identified. Similarly, no fertilized oocytes have been identified within the supraseptal chamber.
In a histological section through a single ovarian follicle (Fig. 23; of) the production of oocytes and spermatozoa and fertilization process can seen. In this section is a mature, fertilized and encapsulated, oocyte (mo). The fertilized oocytes of any anomalodesmatan have only been identified and described hitherto for Entodesma cuneata (Gray, 1828) by Campos & Ramorino (1981) and Laternula elliptica (King & Broderip, 1832) by Ansell & Harvey (1997) and for both of which a capsule (as in G. coronata) is surrounded by a fine sheet of mucus that, upon release from the parent, swells to become a thick, still mucoid, envelope. Figure 23 also shows how the capsule is formed around the oocyte (obe) as it develops from the follicular wall. In addition, Figure 23 shows that the follicle wall possesses germinal areas for spermatozoa. These groupings of testis tissues (t) comprise spermatocytes that are seen to develop into strings of spermatids (spt), which develop, in turn, into spermatozoa (spe) that are occasionally seen as clusters or newly divided sperm morulae (spm). The spermatozoa have elongate acrosomal heads. The spermatozoa of Cuspidaria latesulcata (Tenison-Woods, 1878), also with long acrosomal heads, have been described by Healey, Bieler & Mikkelsen (2008). Finally, Figure 23 shows an oocyte that is being fertilized (obf) as it buds off from the germinal follicle wall—which it must be, before the capsule can be laid down around the enlarging oocyte.
Until recently, representatives of the Spheniopsidae had only been recorded from Pacific and North Atlantic waters. The occurrence of Grippina coronata in Brazilian waters (Machado & Passos, 2015), however, expanded the distribution of this family to the shallow and tropical waters of the South Atlantic. Major interest in the hitherto poorly studied Spheniopsidae, however, centres around four other aspects of the family: (1) taxonomic affinities, (2) lifestyle, (3) reproductive strategy and (4) life history.
Taxonomic affinities
Based on dried specimens of G. californica, Coan et al. (2000) described the Spheniopsidae as being characterized by a fused mantle margin with a small pedal gape, two short conjoined siphons and with incubation of fertilized eggs in the mantle cavity. Only the description of the mantle margin is correct, however, as this study shows. The same authors considered that the Spheniopsidae were allied with the Heterodonta and placed them between the Corbulidae and Hiatellidae, i.e. within the Myoidea. The reason for this, it is suggested, is because spheniopsids possess hinge teeth, which are characteristically absent in all, save a few, anomalodesmatans (Morton, 1981a), including representatives of the Cuspidariidae. Similarly, the myoid ligament is typically amphidetic and located on either resilifers or chondrophores as in many anomalodesmatans such as species of Laternula (e.g. L. truncata, Morton, 1973), although no myoid possesses a ligamental lithodesma. Though this classification was later amended to place the Spheniopsidae in the Anomalodesmata—principally and firstly in the Cuspidarioidea by Marshall (2002) and subsequently by Bieler et al. (2010), Carter et al. (2011) and Coan & Valentich-Scott (2012)—there has not been any anatomical (or molecular) evidence provided in support of this until the present research.
Table 1 provides a comparison between the anatomy of G. coronata and of representatives of the Cuspidariidae. There are a few significant differences, notably with regard to a nonrostrate shell and the presence of hinge teeth in G. coronata, an inhalant siphon that is relatively short and the absence of any vestiges of labial palps. Nevertheless, the overriding picture confirms a closer affinity with the Cuspidariidae than with any of the other septibranch bivalves (Morton, 1981a). Importantly, Allen & Morgan (1981) noted that all species of Cuspidaria have seven siphonal tentacles, four along the ventral edge of inhalant siphon and three around the exhalant, an arrangement identical to that seen in G. coronata (Fig. 6).
|Inequilateral, sub-trigonal
|Typically rostrate
|Coan & Valentich-Scott (2012), Absalão & Oliveira (2011)
|Hinge teeth
|Present, right valve only
|Present, right valve only
|Allen & Morgan (1981)
|Internal, amphidetic
|External, opisthodetic or amphidetic
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Marshall (2002)
|Short stubby, pedal/byssal groove present
|Short, stubby, pedal/byssal groove present
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Byssal groove
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Possibly present in the post-larva
|Post-larva only
|Siphons (category)
|Separate (Type B)
|Separate (Type B)
|Reid & Reid (1974), Yonge (1982)
|Siphonal papillae/tentacles
|7: 3 around the exhalant and 4 around the inhalant
|7: 3 around the exhalant and 4 around the inhalant
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Ciliated sensory papillae
|Reid & Reid (1974)
|Arenophilic glands
|Present (in some)
|Oliveira & Sartori (2013)
|Mantle fusion
|Type B
|Type B
|Yonge (1982)
|Allen & Morgan (1981)
|Septal retractor muscles
|Yonge (1928), Reid & Reid (1974)
|Septal elevator muscles
|Yonge (1928), Reid & Reid (1974)
|Septal pores
|Four pairs
|Typically, four pairs
|Yonge (1928), Morton (1987)
|Pedal musculature
|Labial palps
|Yonge (1928), Coan & Valentich-Scott (2012)
|Stomach type
|Type II
|Type II
|Purchon (1958)
|Style sac and mid gut
|Morton (1987)
|Penetrates ventricle of heart
|Penetrates ventricle of heart
|Yonge (1928)
|Waste-storage pouch
|Bernard (1974)
|Present, Type C
|Present, Type C
|Morton (1985)
|Simultaneous hermaphrodite
|Bernard (1974), Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Bernard (1974)
|Internal, in visceral mass
|Not known
|Inequilateral, sub-trigonal
|Typically rostrate
|Coan & Valentich-Scott (2012), Absalão & Oliveira (2011)
|Hinge teeth
|Present, right valve only
|Present, right valve only
|Allen & Morgan (1981)
|Internal, amphidetic
|External, opisthodetic or amphidetic
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Marshall (2002)
|Short stubby, pedal/byssal groove present
|Short, stubby, pedal/byssal groove present
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Byssal groove
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Possibly present in the post-larva
|Post-larva only
|Siphons (category)
|Separate (Type B)
|Separate (Type B)
|Reid & Reid (1974), Yonge (1982)
|Siphonal papillae/tentacles
|7: 3 around the exhalant and 4 around the inhalant
|7: 3 around the exhalant and 4 around the inhalant
|Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Ciliated sensory papillae
|Reid & Reid (1974)
|Arenophilic glands
|Present (in some)
|Oliveira & Sartori (2013)
|Mantle fusion
|Type B
|Type B
|Yonge (1982)
|Allen & Morgan (1981)
|Septal retractor muscles
|Yonge (1928), Reid & Reid (1974)
|Septal elevator muscles
|Yonge (1928), Reid & Reid (1974)
|Septal pores
|Four pairs
|Typically, four pairs
|Yonge (1928), Morton (1987)
|Pedal musculature
|Labial palps
|Yonge (1928), Coan & Valentich-Scott (2012)
|Stomach type
|Type II
|Type II
|Purchon (1958)
|Style sac and mid gut
|Morton (1987)
|Penetrates ventricle of heart
|Penetrates ventricle of heart
|Yonge (1928)
|Waste-storage pouch
|Bernard (1974)
|Present, Type C
|Present, Type C
|Morton (1985)
|Simultaneous hermaphrodite
|Bernard (1974), Allen & Morgan (1981), Coan & Valentich-Scott (2012)
|Bernard (1974)
|Internal, in visceral mass
|Not known
In the deep sea a carnivorous lifestyle has been adopted by representatives of three bivalve clades, the Propeamussiidae (e.g. Propeamussium lucidum, Morton & Thurston, 1989), the Bathymodiolinae (e.g. Idas argenteus, Ockelmann & Dinesen, 2011) and four families of the Anomalodesmata—the Parilimyidae (Morton, 1982) and the three extant superfamilies of the clade Septibranchia: Verticordioidea, Poromyoidea and Cuspidarioidea (Bieler et al., 2010). This study adds another family, the Spheniopsidae, to this list of predatory bivalves, albeit at the most miniature (<2 mm) of scales.
Having determined the shell structure and form of the siphonal apparatus of G. coronata, it is possible to reconstruct not just its life position, shallowly buried in the sediment (Fig. 24A), but also to understand how it captures its prey (Fig. 24B, C). As described, there is incontrovertible evidence that G. coronata is a predator, because within the mantle cavity there is a septum typical of, particularly, cuspidariids (Reid & Reid, 1974) and a stomach that has, again, a structure typical of cuspidariids (Type II, Purchon, 1956) and that contains prey items, identifiable among which are a harpactocoid copepod and an ostracod. These two taxa are typically benthic (Bell, Walters & Hall, 1987; Gage & Tyler, 1991; Huys & Boxshall, 1991; Mackiewicz, 2006), the harpacticoids either excavating into (endopsammic) or living on the surface of the sediment (epipsammic), in both cases being accessible to G. coronata as prey.
Accordingly, given the short siphons of G. coronata and their associated ciliated sensory apparatus, we can hypothesize that such prey items are detected by these (Fig. 24B) and captured by the rapid eversion of the inhalant siphon (Fig. 24C), as described for Cuspidaria rostrata by Reid & Reid (1974: fig. 1), although that species has a much longer siphonal structure, encased within the shell's posterior rostrum. Moreover, Reid & Crosby (1980: fig. 5a) showed that the siphonal papillae (or tentacles) of Cardiomya planetica possess bunches of sensory cilia at their apex, as in G. coronata, and which were presumed to be mechanoreceptors.
An important distinction between cuspidariids and G. coronata is that the shells of the former are invariably rostrate to accommodate the longer siphons and create a tube, like a gun barrel, to focus rapid siphonal eversion and the capture of swimming prey, e.g. copepods, as illustrated for C. rostrata by Reid & Reid (1974: fig. 1). Lacking this shell rostrum and possessing only short siphons, it seems probable that G. coronata, and possibly other spheniopsids with a similarly foreshortened posterior shell face, feeds on surface-dwelling taxa such as the benthic harpacticoid and ostracod identified from the stomach contents. They therefore exploit a different food resource from that of the Cuspidariidae. Spheniopsids and cuspidariids are both, however, immobile predators that 'lie-in-wait' to ambush prey.
A unique feature of the intestine and stomach of G. coronata is the presence in the latter of a waste-storage pouch. As described, the intestinal tract itself is narrow (midgut lumen <40 μm) with no evidence of the skeletal remains of these relatively large prey items within it. Instead, such remains appear to be stored in a pouch accessed via a sphincter from the left wall of the stomach. As described, harpacticoid copepods and ostracods are found in the stomach and their exoskeletal remains in this storage pouch, thereby confirming their digestion. Fine, digested remains must therefore be transported from the stomach to the intestine and expelled from the anus as in all, more typical, suspension-feeding bivalves. The skeletal remains of prey in such a tiny spheniopsid as G. coronata cannot, however, be removed in this manner and, accordingly, are stored in the waste-storage pouch from which they will never be evacuated in life. As will be described below, G. coronata can only release large oocytes after death and the same strategy seems to have been adopted for defaecation also, creating an altogether remarkable life-history trait.
Reproductive strategy
Representatives of the Cuspidariidae are generally considered to be dioecious (Bernard, 1974). This is clearly not the case with G. coronata, which is here demonstrated to be a simultaneous hermaphrodite. Oocytes are budded off from mesodermal epithelia into large left and right follicular lobes within the visceral mass. The same germinal epithelia that produce the oocytes also produce spermatocytes, so that fertilization is not just internal, it is intrafollicular. As far as is known, this is unique in the Bivalvia. Fertilized oocytes are enclosed in a thick capsule. It is unknown if G. coronata possesses gonopores, the sectioned individuals being simply too small to determine this. Regardless, however, such large (>50 μm) fully-mature oocytes would be far too big to exit the parent in the usual manner. Indeed, they would even have trouble being discharged from the exhalant siphon. The oocytes are, in fact, so well developed when they are released from the parent that the prodissoconch I, at least, is fully formed and probably prodissoconch II also (Machado & Passos, 2015: fig. 3).
Grippina coronata thus exhibits direct development to produce a lecithotrophic larva (as defined by Ockelmann, 1965) that almost certainly has an exceedingly short benthopelagic life, if at all. Since it would seem impossible for G. coronata to discharge fertilized oocytes in any conventional manner, it is suggested that fully mature individuals of this this species die, thereby releasing their fertilized and encapsulated oocytes within the protection of their mucoid outer capsule, post-mortem. Fertilized but unencapsulated oocytes would die at this time. The only other anomalodesmatans that are known to produce such encapsulated oocytes are Entodesma cuneata (see Campos & Ramorino, 1981) and the stenothermal Laternula elliptica from Antarctic waters (Ansell & Harvey, 1997). The oocytes of both these species, further surrounded by a thick, gelatinous envelope, are released (by a means unknown) and remain in the plankton for only a few days. In the water column, the gelatinous layer surrounding the eggs forms into a strong, sticky and elastic capsule within which development proceeds. It seems possible that, since its intrafollicular oocytes are essentially the same as those described above for the other anomalodesmatans, a similar developmental strategy is adopted by G. coronata, albeit following parental death.
Life history
The pioneering research of Ockelmann (1965) showed that at greater depths in the Arctic North Sea an increasing number of bivalve species produce lecithotrophic fertilized eggs with either a short pelagic larval life or with maternal, typically ctenidial, brooding. The greater incidence of species showing direct development with brood protection was, however, more common in shallower-shelf species. This generalization seems applicable to G. coronata, recorded by Machado & Passos (2015) from depths of 21 to 53 m off the coast of Brazil. An example of miniaturization having a dramatic effect upon reproduction was provided by Sanders & Allen (1973) in a deep-water species of Microgloma (Protobranchia: Pristoglomidae) which, at any one time, only has two mature oocytes in left and right ovarian follicles. These authors attributed this reduction in egg numbers to the small size of these protobranchs, which consequently are unable to produce a sufficient number of planktotrophic larvae and therefore produce a small number of eggs that undergo nonplanktotrophic development. The life history of this minute septibranch, G. coronata, however, releasing fertilized and encapsulated oocytes following parental death, is even more extreme. This post-mortem semelparity and similarly post-mortem release of undigested food remains, represent life-history traits hitherto unseen in the Bivalvia.
This work was carried out with logistical and financial support provided by CENPES/Petrobras, under the Habitats Project. We thank Dr A.P. Falcão for kindly inviting us to participate in this project. Financial support was provided by a scholarship from CAPES to F.M. Machado. We thank Dr L.F. Tallarico for encouraging F.M. Machado to establish international partnerships and special thanks to Dr P. Valentich-Scott who introduced the authors to each other. Thanks are also due to A.C.S. Sprogis and S.M.F. Ferraz (Laboratory of Microscopy, IB/UNICAMP), who provided assistance with microscopy.


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694 posts
295 followers
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Greg Pollick
This oneMy happy Beauty#collegegirl
Great job Ashley! So #proudofmywife !#smartwoman
Great #run today#Slushy but good
Nailed it #JCrew !#spokane #grey
#SuperBowl ribs smile!
I stumbled upon this photo this morning, and it made me chuckle. I'm definitely guilty of putting words in Jesus' mouth by elevating some things as absolutely important, and then expecting them of others (or myself). But perhaps the even bigger danger is the opposite: not caring about the things he actually did say (Matthew 28:20). Sometimes Jesus' words are uncomfortable, yet I've never known them to not be liberating...or simple. Here's to the simple life. 🎉#Repost @chrislazo with @repostapp
Celebrating this one today. Love how God made him, and having fun being his dad#thisguy
Working in the middle of nowhere today.
Papier-mâché before church? It's a thing#marriedtoateacher
When you're sick at home with mom, you get to make quail crafts
Happy New Year!!#bestnine2016 It was a year the Lord enriched our life in every way. Jesus is good to me.

subset: commoncrawl


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About Diet Center
Diet Center is a unique brand in the sense that it caters to an inexhaustible number of customers at a time. The company has powerful weight loss results which make it appealing to customers who can find veritable proof of success cases. The company also has an online presence with its websites and online sales. The company has its own exclusive software, signature programs and signature products which make it stand out from others.


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When changing the air filter, it is easy to knock off a hose on the air filter housing, which is part of the emission control system. If the vehicle is driven with this hose disconnected, the Check Engine Light will illuminate.
Problem Description and Possible Solution
Problem Data
RepairPal Verified
Average mileage: 116,380 (17,000–250,000)
Engines affected: 2.4L 4 Cylinder, 3.0L V6, 3.3L V6
47 people reported this problem
Viewing 1 - 20 of 21 comments
I changed the air filter and this happened to me. A neighbor is a Toyota mechanic and told me about it when I called him about the check engine light.
I really hate this issue!! It happens most oil changes, except at the dealer, and it's a big time waster to get it resolved. I need a picture of this so I can fix it myself. Advance Auto parts store is happy to run the diagnostic and reset the CE light for me.
reattached hose.
every time a tech looks at the air filter, the hose in back disconnects
oil change place knocked hose off air filter housing...check engine light went off a few weeks after I fixed it.
Check engine light & VSC light come on together every time my air filter is changed. Hose comes off. Put hose on and disconnect battery to get the check engine lights to go off.
Every time someone checks the air filter this hose comes off.
I had serviced the car (which I have been for years) check the air cleaner, then the engine light came on few days later. I brought to my Toyota dealership to see what triggered the light, they found this same hose came disconnected from the air cleaner housing. I asked how much for the repair- they smiled and said "all set" no charge.Good people.
Got car serviced at local Jiffy Lube - where I was duped into a $75 engine flush. Next day the check engine light came on. Fortunately a mechanic friend was able to inspect the car before I spent a million dollars at the dealership and found that there were several loose and/or incorrectly attached hoses and connectors..
Main freg light to turn off
technicians repeatedly know hose loose during servicing. check hose each time it is serviced. check engine light will go off on its own in about 50 start/ stops
This problem has occurred numerous times. Even when getting oil changed a a Toyota dealership! It happens whenever there is an oil change and I don't explain the problem to the mechanic. This has probably happened over ten times over the last 7 years. It is a real bothersome issue in states where emissions require no check engine light be on when tested.
This happens nearly every time I get my oil changed. I have finally learned to mention it to the technician when I drop the car off, and they generally can reattach it before I pick up.
same thing happened to me.
This problem has occurred twice. Once was after a regular oil change, and the second was after I replaced the air filter myself. The air filter mechanism is hard to seat.
Discovered the hose was not attached after my check engine light appeared. I had gotten the air filter replaced several days before from a different shop. I was able to continue driving for a couple days until I could bring it into the shop but noticed the car lacked power.
Same thing others describe. Filter change with 30 point checkup. Someone replaced the air filter with wrong version. Did not latch the cleaner housing down correctly, and hose was off the back. Weird, deep throaty/gutteral sound when accelerating. Now have to reset the LED and hope that's all that went wrong.
After oil change, the check engine light came on. Hose was not connected.
Happened during an oil change. Gas mileage was awful until I brought it back to the oil change shop.


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in plus sizes
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Coral Collage Venus Large Vase
Ceramic Vase(ID: A90432)
Created by Judi TavillOne of a Kind
This bellied one of a kind vertical vase is both decorative and functional. It's coral-like textures and undulating lines are inspired by the enticing patterns and movement discovered during the artist's coastal adventures. Its deep repeating sections of texture are created by hand carving and detailing using both manmade and natural tools in the artist's ceramic studio. The artist throws sandy stoneware clay on the pottery wheel, alters the form and later carves and designs the pattern. Porcelain slip is added to create a white skin and then she hand textures the surface with various tools. A signature stamp is impressed into the recessed foot upon completion during the greenware stage. The piece is substantial in weight due to the necessity for thickness to achieve such texture. The first firing secures the carving and texture and then Tavill coats the interior (and the recessed foot portion) with clear glaze and delicately sprinkle portions of the exterior texture with the same clear glaze, creating a subtle glisten upon observation. The final firing takes the kiln to over 2185 degrees as the clay is fully vitrified (no longer porous).


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Is this restaurant appropriate for Kids?
Is this a diner?
Are the prices at this restaurant mid-range / moderate?
Is this restaurant good for dinner?
Is this an American restaurant?
Is this restaurant good for breakfast?
Is this restaurant good for business meetings?
Is this restaurant family-friendly?
Is this restaurant good for lunch?
Does this restaurant have waiters and waitresses?
Thanks for helping!
This little place, a bit aged, has really good food, and the prices are cheap. Plus, lots of antique stores in that area to check out. Kind of place where you will find the locals eating.
It's hard to describe this restaurant. It's a little like an old-school diner, but the menu is pretty small and focused more on soups and sandwiches. The prices are a little high for sandwiches, but the food is pretty good. I'd recommend the white bean soup.
This Cafe has nice choices to choose from. On our visit passing through the area our thoughts were something light to eat.
Boy were we surprised by our choices for a cup or bowl of soup and a piece of home made cornbread this was the best. Along with a warm and cozy atmosphere from the owners and service.
While antique shopping in Old Central City we worked up an appetite so thought we'd try the cafe. We were so pleasantly surprised! We go back whenever we're in the neighborhood. Everything we've ever had was delicious home cooking with little extras that make it more special such as a little slice of watermelon on the plate. The homemade soups...
Awesome! Wonderful People! Vegetarian Friendly too! You must visit Old Central Antiques District! A great destination.
Stopped by on a lark. Ended up bringing parents back the next day it was so good. Simple food with great taste. Recommend meatloaf, cheeseburger, and hot ham and cheese sandwiches. And the pie is to die for!!! Cherry and apple was what we had. Slices are huge and served with 2 scoops of ice cream.
I am in town this week on business and saw that the Central City Cafe had been on DD&Ds a few years ago and figured I'd give it a try. I was not disappointed! I got the soup and sandwich combo and it was great. I am hoping that I get to go back one more time this week and...
We stopped in after visiting some of the antique shops, this is a great place for a "home cooked" meal!
It is family owned and operated, they are great! Friendly and welcoming. Big portions too.
I highly recommend.
Plenty to choose from and what I ordered was very good. This was my second trip to the Cafe and I will return next time I have the chance. Waiter was very friendly and knowledgeable.
We had lunch today. My husband got the meatloaf and I got a Rueben. He said meatloaf maybe not his favorite but my Rueben was terrific. They serve the thousand island on the side ...glad they did cause I like mustard. Lots of meat and most delicious sauerkraut. A little expensive lunch...but dinner sized😋
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Greenville has 56,002 residents. It is located in Greenville county.
For South Carolina residents, the average tuition for all Greenville gerontology schools for the 2009 - 2010 school year was $14,841.00. For non-residents, the average tuition for Greenville gerontology schools was $15,445.00.
Books and supplies for gerontology schools in Greenville cost an average of $658.00. Students who live on campus while attending a gerontology school in Greenville, can expect to pay an average of $7,135.00 for room and board. Students who live at home can expect average costs to be $4,131.00. There is one gerontology college in Greenville. 3 students were reported to have graduated from gerontology programs in Greenville in the 2008 - 2009 school year.
|Average Salary
|Number Employed in City
|Mental Health Counselors
|Dietitians and Nutritionists


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Vitamin Shoppe New York CityStore's name : Union Sq., NY
Address & Contacts
(212) 254-8816
Vitamin Shoppe
24-30 Union Square E
10003 New York City
Opening Times
|08:00am - 8:00pm
|08:00am - 8:00pm
|08:00am - 8:00pm
|08:00am - 8:00pm
|08:00am - 8:00pm
|10:00am - 7:00pm
|11:00am - 6:00pm
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Vitamin Shoppe New York City is usually open Sunday.
These times do not include public holidays. To see if your store is open these days, contact them.
- Health & beauty shops
- Vitamin Shoppe in New York City 156 5th Avenue
- Vitamin Shoppe in New York City 100 E 23rd Street
- Vitamin Shoppe in New York City 655 Avenue Of The Americas
- GNC in New York City (0.18 km)
- Walgreens in New York City (0.19 km)
- Walgreens in New York City (0.25 km)
- Latitude : 40.7364039
- Longitude : -73.98870649999
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Vitamin Shoppe
568 stores Vitamin Shoppe in the U.S. registered on Stores-Locator.com
This is a store that specifically deals in the distribution of vitamin supplements. It has about 600 outlets and still growing. The first store in the chain was opened in 1977 and many more have sprung up since that time bringing the number of stores to the current number of about 600. The store believes that everyone is important and… more about Vitamin Shoppe
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A floral Spring to come with Liberty London and UniqloAfter numerous previous collaborations, Japanese Uniqlo has teamed with British Liberty London to release a floral capsule collection this Spring.
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The Biba Experience
By (author) Alwyn W. Turner
Format: Paperback
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Biba by Alwyn W. Turner
Book DescriptionBiba remains one of the most evocative names in British design history; it pioneered a new style, mixing the contemporary with Art Nouveau, Art Deco and the golden age of Hollywood, dressing itself in the richly luxuriant colours of a bygone time. This book showcases Biba clothing and much of the Biba memorabilia, merchandising and interior design, presenting an entire history of the company. "Biba: The Biba Experience" centres on commissioned photographs of one of the largest collections of Biba in the world, put together by Pari, who has accumulated around 500 pieces of Biba fashion items and ephemera covering the entire history of the company. Everything from dresses and trouser suits through original sketches and patterns to playing cards and soap powder is represented. Biba brought the cutting edge of couture to the masses. Brigitte Bardot, Yoko Ono and Princess Anne shopped there, whilst Sonny and Cher, Mick and Marianne, and David and Angie were regular visitors, but the store was never the exclusive preserve of the rich and famous: prices were kept deliberately low, and anyone who could tolerate the disdainful inefficiency of the staff was encouraged to soak up the glamour of a unique shopping experience. This lavishly illustrated and beautifully produced book is intended for those interested in the fashion and graphic design of the '60s and '70s, from a creative and historical standpoint, plus the huge number of people who remember Biba from the period with such affection.
Buy Biba book by Alwyn W. Turner from Australia's Online Bookstore, Boomerang Books.
Book DetailsISBN: 9781851495412
(287mm x 247mm x 22mm)
Pages: 240
Imprint: ACC Art Books
Publisher: ACC Art Books
Publish Date: 8-Jun-2007
Country of Publication: United Kingdom
Other Editions...
Books By Author Alwyn W. Turner
Last Post, Hardback (October 2014)
A moving and sometimes provocative exploration of the music, rituals and monuments used to remember the fallen of the Great War from one of Britain's most admired new historians.
Classless Society, Paperback (April 2014)
The first popular history of a decade that is yet to be defined or anatomised as the 1960s or 1970s have been, A Classless Society goes in search of a Britain still reeling from the conflicts of the Thatcher years.
Glam Rock, Hardback (October 2013)
Drawing on the collections of the V&A, Glam Rock narrates the story of glam & explores its impact on fashion, theatre & film. In the early 1970s, glam rock changed the face of popular culture in Britain and, against a backdrop of a nation racked by economical and social crises, its flamboyancy provided an escapist dream for musicians and fans alike
Rejoice! Rejoice!, Paperback (September 2013)» View all books by Alwyn W. Turner
An acclaimed history of the biggest, brashest and most tumultuous decade in recent British history.
» Have you read this book? We'd like to know what you think about it - write a review about Biba book by Alwyn W. Turner and you'll earn 50c in Boomerang Bucks loyalty dollars (you must be a member - it's free to sign up!)
Author Biography - Alwyn W. Turner
Alwyn W Turner is a widely published writer on the popular culture of post-war Britain. He is the author of 'Cult Rock Posters' and 'Welcome to Big Biba,' and the editor of 'Portmeirion' (Antique Collectors' Club, 2006). His work has also appeared in 'The Rough Guide to Rock, Mojo' and on BBC Online.
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Meaning and origin of the Latin baby name Eurus. Meaning of Eurus. What does Eurus mean? Eurus origin. Information about Eurus.
|Meaning of Eurus:
|East wind.
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Get best timeing (Muhurta) for all your auspacious deeds like marriage, house warming etc..
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text:
Big Fish Audio & Adam Smeaton (aka Dubkiller) präsentieren mit Dirty Dubstep eine Library mit 13 Construction Kits. Inbegriffen sind: Drums, Risers, Synth Leads, FX, und die für Dubstep typsichen Wobble Basses.
Weitere Informationen in Englischer Sprache:
Over 1.5 GB of nothing but the highest quality 24bit Apple Loops, Acidized WAV and REX files makes Mayhem: Dirty Dubstep an instant win for any Dubstep production imaginable. In addition to the demo mixes and stereo files, Big Fish Audio included Multi Drum tracks and one shots for every kit, giving you complete control over the beat and direction of the track. Mayhem Dirty Dubstep is the new standard for Dubstep loop libraries and will keep you coming back begging for more!
KLI Series
This product is part of Big Fish Audio's KLI series, which includes a custom Kontakt interface. The Kontakt format features patches of each loop. Customize each loop with the "sliced loops" patches. The Sliced Loop patches lay out each individual slice of a loop across the keyboard, making it easy to create new and original custom parts. With the "Kit Combos" you'll find each loop as part of a complete construction kit laid out across the keyboard for easy and quick arrangements.
Big Fish Audio included a Wobble Basses section that has patches from each kit that include all wobble bass slices combined in one file. These patches were intentionally set to "one shot mode and not intended to be used as a loop but rather as sample menus of all the bass wobbles from each kit, allowing you to create your own custom parts. Rearrange, tweak, shorten or go crazy with these wobble bass patches, the possibilities are endless.
Instruments Included:
Drums, Wobble Bass, Sub Bass, Sub Booms, Synth, Synth Pad, Risers, Sweeps, Guitar, Percussion FX & FX.
MULTI FORMAT Details:
KLI FORMAT Details:
The Full version of Kontakt 4.2 or higher is required to use the Kontakt 4 Instrument Patches, and the Full Version of Kontakt 5.3.1 is required to use the Kontakt 5 instrument patches in the KLI version of this product. This product does not work with the free kontakt player.
Diese Multi Format Library (enthaltene Formate sind oben gelistet) enthält keine Software zum Abspielen der enthaltenen Sounds. Sie benötigen auf Ihrem Computer eine Software, die eines der angezeigten Formate laden und abspielen kann.


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Viking Saga: The Cursed Ring
Viking Saga: The Cursed Ring - a great game on Android with fun adventures with Viking Ingolf. Once you have a spectacular opportunity to feel the full force of magic and enchantment, brave deeds and silly betrayals. Embark on a journey and have an unforgettable experience.
Gripping story of the game begins in the fairy kingdom of ancient Viking Arnar. It was learned that at the time when Arnar was young, he was cursed by a terrible evil dwarf, because the precious ring. How - what Arnar found the ring and did not want to sell it. After some time, or rather, a few years later, the curse has overtaken the king. From this moment begins your journey. Arnar requested such an important mission to find and return the ring dwarf, his youngest son.
Embark on a journey with the main character through the colorful places, solve issues of local population, become a wise friend
attention! all files hosted with permission of the authors or applications found in the public domain in the internet, if some of the files violates your rights, please let us know


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Apartheid's Festival
Contesting South Africa's National Pasts
By (author) Leslie Witz
Format: Paperback
Normal Price: $57.04
Your Price: $51.34 AUD, inc. GST
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Availability: Available, ships in 8-11 days
Apartheid's Festival by Leslie Witz
Book DescriptionApartheid's Festival highlights the conflicts and debates that surrounded the 1952 celebration of the 300th anniversary of the landing of Jan Van Riebeeck and the founding of Cape Town, South Africa. Taking place at the height of the apartheid era, the festival was viewed by many as an opportunity for the government to promote its nationalist, separatist agenda in grand fashion. Leslie Witz's fine-grained examination of newspapers, brochures, pamphlets, and advertising materials reveals the expectations of the festival planners as well as how the festival was engineered, historical figures were reconstructed, and the ANC and other anti-apartheid organizations mounted opposition to it. While laying open the darker motives of the apartheid regime, Witz shows that the production of local history is part of a global process forged by the struggle between colonialism and resistance. Readers interested in South Africa, representations of nationalism, and the making of public history will find Apartheid's Festival to be an important study of a society in transition.
Buy Apartheid's Festival book by Leslie Witz from Australia's Online Bookstore, Boomerang Books.
Book DetailsISBN: 9780253216137
(238mm x 150mm x 23mm)
Imprint: Indiana University Press
Publisher: Indiana University Press
Publish Date: 6-Oct-2003
Country of Publication: United States
Other Editions...
» Have you read this book? We'd like to know what you think about it - write a review about Apartheid's Festival book by Leslie Witz and you'll earn 50c in Boomerang Bucks loyalty dollars (you must be a member - it's free to sign up!)
Author Biography - Leslie Witz
Leslie Witz is Associate Professor of History at the University of the Western Cape.
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About Us | Contact Us | Help Desk | Security | Privacy | Terms of Use | Copyright
Phone: 1300 36 33 32 (9am-5pm Mon-Fri AEST) - International: +61 2 9960 7998 - Online Form
Address: Boomerang Books, 878 Military Road, Mosman Junction, NSW, 2088
© 2003-2017. All Rights Reserved. Eclipse Commerce Pty Ltd - ACN: 122 110 687 - ABN: 49 122 110 687
For every $20 you spend on books, you will receive $1 in Boomerang Bucks loyalty dollars. You can use your Boomerang Bucks as a credit towards a future purchase from Boomerang Books. Note that you must be a Member (free to sign up) and that conditions do apply.


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Sony GTKX1BT
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Product Features
|Audio technology
|Number of speakers
|Wireless or wired speakers
Manufacturer's Description
Your music, your party, your way
From hanging out at home to impromptu parties, this Home Audio system has got it covered. Play music straight from selected smartphones and turn it up loud with 500W1 of power. With a stylish, versatile design that fits in with your living room, you'll enjoy rich, full-spectrum sound wherever you place the system.
Two-way speaker system
From booming beats to mellow acoustic mixes, you'll enjoy full-spectrum sound across your entire music collection with the GTK-X1BT. Two 20cm woofers deliver warm, rich mid-range and low-end sound, while two 6cm tweeters deliver soaring highs and clear vocals – so you'll never miss a moment of your music.
Mica Reinforced Cellular woofers
Hear powerful audio with two 20cm MRC woofers. The cones' rigid design helps them keep their shape, even when vibrations are strong, for clean, undistorted bass.
Digital Sound Enhancement Engine (DSEE)
DSEE upscales compressed digital music files (MP3, ACC, ATRAC, and WMA). It restores highs and lows lost in compression, producing rich sound closer to CD quality.
Superior sound, whatever the setup
Stand it up or lay it down – the GTK-X1BT's versatile setup gives you great sound no matter where you place it.
Portable party starter
The GTK-X1BT's lightweight, compact, box-style design features built-in handles for maximum portability and convenience. Simply pick it up and party anywhere.
Pump up the party
Link systems together for booming sound
Really get the dance-floor jumping with party features. Get creative with four DJ effects – from panning sound across speakers and isolating vocals to adding flanger and WAH. With Party Chain, you can hook up and sync multiple sound systems with audio cables for an even bigger sound.
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no one is selling it, we're sorry to say,
not 421 retailers nor anyone on eBay,
but please don't worry even if it hurts,
you can look for alternatives or


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text: Canadian Mortgage Awards names first group of finalists It's that time of year again – time for the finalists for the Canadian Mortgage Awards to be announced. Beginning today, we will be releasing the names of the finalists in several categories, including Best Customer Service, Best Community Service Effort and Employer of Choice, with the naming of the Mortgage Broker of the Year category finalists later this week. "We would like to congratulate all the finalists," said CMP Editor John Tenpenny. "This year's group is the best of the best in the mortgage industry and was selected from thousands of nominations." The sixth annual Canadian Mortgage Awards, presented by Home Trust, will take place on Friday, June 1 in Toronto at The Carlu. The move to a new venue will allow for more than 600 attendees as the event continues to expand. You don't want to miss what is sure to be one of the biggest events of the year in the Canadian mortgage industry, so reserve your table now as the event will sell out (www.canadianmortgageawards.com). The event closes out The Mortgage Summit (www.themortgagesummit.ca), a two-day Toronto conference powered by CMP and MortgageBrokerNews.ca, also being held at The Carlu. The fate of the finalists in now in the hands of the judging panel, with the entire process being overseen by PwC, the official auditor of the Canadian Mortgage Awards. "As the nomination, finalist, and judging process grows in size and complexity we felt it was important to bring in a third party to verify the process and validate the results of this process," said Chris Davis, events manager for KMI Media. "To that end, we are happy to announce that PwC is now the official auditor of the Canadian Mortgage Awards." It is sure to be a night to remember, with a Roaring 20s theme, including live entertainment, prizes, giveaways, a delicious four course gourmet meal, drinks, and dancing. Join more than 600 of your colleagues, friends, and industry professionals from across Canada, for the biggest night on the industry calendar, and celebrate and honour the achievements and successes of 2011. HomEquity Bank Employer of Choice Centum Financial Group Dominion Lending Centres First National Financial Fisgard Capital Corporation Pacific Mortgage Group Paradigm Quest Street Capital Financial VERICO Canada Merix Financial Best Customer Service - Individual office Averbach Mortgages TMG The Mortgage Group, Vancouver, B.C. Dominion Lending Centres West Coast Mortgages, Coquitlam, B.C. VERICO Jessi Johnson Mortgage Team, South Surrey, B.C. Mortgage Architects (formerly MortgageBrokersAlberta.com), Calgary, Alta. VERICO Coastal Mortgages, Surrey, B.C. Best Community Service Effort Grant Mackenzie, Canadiana Financial, Toronto, Ont. Scott Ede, Mortgage Alliance S&R Mortgage Group, Calgary, Alta. Brian Nason, Mortgage Architects, Hamilton, Ont. Geoff Parkin, VERICO Select Mortgage Corp., Surrey, B.C. Deb White, Dominion Lending Centres White House Mortgages, Vernon, B.C. Diana Zitko, Dominion Lending Centres West Coast Mortgages, Coquitlam, B.C. Invis Mortgage Intelligence Best Industry Service Provider Superstar Mortgage Broker/Power of Choice Coaching Inc. First National Financial JOLT Marketing MCAP Service Corporation Paradigm Quest Strategic Information Technology Limited Street Capital Financial Best Internet Presence Canadian Mortgage Trends Dominion Lending Centres VERICO Jessi Johnson Mortgage Team Merix Financial Mortgage Architects VERICO Paragon Mortgage Group Best Advertising Centum Financial Group Dominion Lending Centres The Equitable Trust Company Mortgage Architects VERICO Canada FNF Canada Best Branding Centum Financial Group Dominion Lending Centres INVIS-Mortgage Intelligence VERICO Jessi Johnson Mortgage Team Mortgage Architects Syndicate Mortgages TMG The Mortgage Group VERICO Boomerang Financial VERICO Canada Leave a comment SHARE


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This Black Hills Gold Necklace features a guardian angel elegantly depicted on a blue agate cameo. Swirling vines of 10K yellow gold and 12K rose and green leaves gracefully frame the image. The Angel Cameo Pendant hangs on an 18" gold-filled chain and measures 1.141" H x 0.711" W. Made in USA.
Manufacturer model #: 25060.