Environmental variation in physical and chemical cues used by the parasitic wasp,Aphytis melinus, for host recognition

O-caffeoyltyrosine is a host recognition kairomone forAphytis melinus DeBach (Aphelinidae) found in the covers of its host, California red scale,Aonidiella aurantii (Maskell) (Diaspidae). This study tests the hypothesis that the concentration ofO-caffeoyltyrosine and scale cover size are reliable indicators of scale body size, an important component of host quality forA. melinus, over a range of scale rearing conditions. Both scale cover area andO-caffeoyltyrosine concentrations were only qualitatively related to scale body size during the third instar, the scale life stage most suitable forA. melinus. Scale cover area andO-caffeoyltyrosine concentrations were reduced, relative to scale body size, when scale were reared on bark and leaves compared to fruits. Scale cover area andO-caffeoyltyrosine concentration were also relatively reduced when scales were reared in mid-summer compared to spring and fall, and when reared on orange cultivars compared to lemon cultivars in the field. Finally, scale cover area andO-caffeoyltyrosine concentration were reduced when scale were reared at 52% compared to 100% humidity in the laboratory. Scales appear to be chemically conspicuous toA. melinus for a short period of the time in which they are physiologically susceptible. Scales that minimize their cover size and maximize the incorporation rate ofO-caffeoyltyrosine into covers may minimize their conspicuousness toA. melinus. Minimizing scale cover size, but not necessarily incorporation rates, may make scales more vulnerable to predators, however.     

High molecular mass glycoproteins associated with the siliceous scales and bristles of Mallomonas splendens (Synurophyceae) may be involved in cell surface development and maintenance

The elaborate scale case of Mallomonas splendens (Synurophyceae) consists of an overlapping arrangement of siliceous scales. In addition, siliceous bristles are attached to specialized base plate scales located at both the anterior and posterior ends of the cells. We have generated monoclonal antibodies against molecules associated with the scale case of M. splendens. One of these antibodies, designated MsS.H9, labelled a proteinaceous epitope of high-molecular-mass cell surface glycoproteins. Immunofluorescence and immunoelectron microscopy demonstrated that only two regions of M. splendens scale cases were labelled by MsS.H9, namely, the upper surface of the scales that contact neighboring scales and the bases of the bristles. Immunoelectron microscopy using thin sections of M. splendens cells showed these labelling sites corresponded to the amorphous material at the sites of scale-to-scale overlap and to a fibrillar complex located at scale-to-bristle attachment sites. Scales and bristles of M. splendens are formed within the cell, in silica deposition vesicles. Immunolabelling of cell sections containing developing scales and bristles showed that MsS.H9 labelling sites were present very early in the formation of these cell surface components. MsS.H9 labelling was also found associated with developing flagellar hairs whereas no labelling was detected on these structures after their deployment onto the flagellum. The location of MsS.H9 labelling sites strongly suggests that the molecule(s) recognized by the antibody plays a role in the adhesion of the individual components making up the scale case of M. splendens.Abbreviations CER chloroplast endoplasmic reticulum - ER endoplasmic reticulum - SDV silica deposition vesicle This work was supported by a grant from the Australian Research Council to R.W. We thank Dr. P. L. Beech for Fig. 13, Dr. L. Perasso for technical assistance and the Plant Cell Biology Group for the use of their monoclonal facilities.     

Identification of a 41 kDa Protein Embedded in the Biosilica of Scales and Bristles Isolatedfrom Mallomonas splendens (Synurophyceae, Ochrophyta)

Cells of the photosynthetic protist Mallomonas splendens (Synurophyceae, Ochrophyta) are encased within a highly patterned wall or scale case that consists of silicified scales and bristles. In an effort to understand the mechanisms that unicellular protists utilize to produce elaborate, mineralized structures of great complexity and hierarchical structure, we identified and characterized a 41 kDa protein from purified scales/bristles isolated from M. splendens (SP41 for Scale Protein of 41 kDa). A cDNA encoding this protein was isolated and sequence analysis indicated that it is a novel protein. Polyclonal antibodies were generated against bacterially expressed SP41 and used to localize the protein throughout scale and bristle morphogenesis. Immunoelectron microscopy confirmed the biochemical data that SP41 is a component of mature scales and bristles, the protein localizing to silicified components of the purified extracellular matrix. During scale and bristle biogenesis within the cell, SP41 is deposited into a specialized Silica Deposition Vesicle (SDV) concomitant with silica deposition, a highly regulated event during scale and bristle formation. These results argue for SP41 playing a role in morphogenesis and/or silicification within the SDV during scale and bristle biogenesis.     

Scale Arrangement Pattern in a Lepidopteran Wing. 1. Periodic Cellular Pattern in the Pupal Wing of Pieris rapae

In most species of lepidopteran insects, anteroposterior rows formed by scales are arranged at regular intervals in the adult wing; within each row two kinds of scales are alternately arranged. To investigate the cellular basis for the scale arrangement pattern, we examined cell arrangement in the epidermal monolayer of the pupal wing of a small white cabbage butterfly, Pieris rapae , by scanning electron microscopy and light microscopy.
The arrangement of scale precursor cells, closely resembling that of scales in the adult wing, was observed in the wing epidermis of the early pupa. Scale precursor cells are proximodistally elongated and form anteroposterior rows. Within a row two kinds of scale precursor cells are nearly alternately arranged, which is not so precise as the alternation of scales in the adult wing. Individual rows of scale precursor cells are separated by rows of single or double undifferentiated general epidermal cells. Occasionally, arrangement abnormalities occur both in the adult and the pupal wing. The cellular basis for the regular spacing of scale rows is discussed.     

A new sparid, Acanthopagrus akazakii, from New Caledonia with notes on nominal species of Acanthopagrus

A new sparid species, Acanthopagrus akazakii, is described on the basis of 12 types collected from Noumèa, New Caledonia. Acanthopagrus akazakii is most similar to Acanthopagrus berda in overall appearance but differs from the latter in having 4¹/₂ scale rows between the 5th dorsal fin spine base and lateral line, anteriormost head scales broadly rounded without small scales anteriorly, upward- and downward-oriented portions present on anteriormost part of upper lip, a downward-oriented portion on anteriormost part of lower lip in specimens over ca. 160 mm in standard length, and upper head profile gently convex from snout tip to above eye throughout growth (vs. 3¹/₂ scale rows between 5th dorsal fin spine base and lateral line, anteriormost head scales rounded with small scales anteriorly, no upward and downward portions in anteriormost upper and lower lips, and upper head profile becoming concave from snout tip to above eye with growth). Furthermore, A. berda develops a strong concavity of the ventral edge of the first two infraorbitals above the posterior part of upper jaw with growth, whereas A. akazakii has a generally straight series throughout growth. Nominal species in A. berda are reviewed, with notes on nominal species of Acanthopagrus.     

Golgi apparatus activity and membrane flow during scale biogenesis in the green flagellateScherffelia dubia (Prasinophyceae). I: Flagellar regeneration

Summary Cells ofScherffelia dubia regenerate flagella with a complete scale covering after experimental flagellar amputation. Flagellar regeneration was used to study Golgi apparatus (GA) activity during flagellar scale production. By comparing the number of scales present on mature flagella with the flagellar regeneration kinetics, it is calculated that each cell produces ca. 260 scales per minute during flagellar regeneration. Flagellar scales are assembled exclusively in the GA and abstricted from the rims of thetrans-most GA cisternae into vesicles. Exocytosis of scales occurs at the base of the anterior flagellar groove. The central portion of thetrans-most cisterna, containing no scales, detaches from the stack of cisternae and develops a coat to become a coated polygonal vesicle. Scale biogenesis involves continuous turnover of GA cisternae, and scale production rates indicate maturation of four cisternae per minute from each of the cells two dictyosomes. A possible model of membrane flow routes during flagellar regeneration, which involves a membrane recycling loop via the coated polygonal vesicles, is presented.     

Structure,composition, and biogenesis of prasinophyte cell coverings

Summary The cell body and flagellar surfaces of certain green flagellates are covered by non-mineralized scales. Scale structure has been widely used in the systematics of this group of algae commonly known as the Prasinophyceae. The special importance of the flagellar hairs as a taxonomic marker is discussed. We summarize current knowledge about the structure and chemical composition of these scales with emphasis on thecate flagellates. Scales consist mainly of acidic polysaccharides involving unusual 2-keto sugar acids. Glycoproteins as minor components are mainly involved in mediating scale subunit and scale-membrane interactions and species specific glycosylation patterns exist. In thecate prasinophytes the elaboration of 3-deoxy-manno-2-octulosonic acid and galacturonic acid side chains presumably favours a complex of thecal scales with calcium ions and thus extracellular coalescence of the scales to a rigid cell wall. Scales are formed within the Golgi apparatus (GA) and especially in thecate prasinophytes scale formation (i.e., during flagellar regeneration) represents an excellent model system for GA function. Movement of developing scales through the GA requires cisternal progression. Biogenesis of scales involves mainly polysaccharide synthesis, whereas about 50% of the scale-associated glycoproteins are added from a pre-existing pool. Possible functions of prasinophyte scales are briefly discussed.Abbreviations Dha 3-deoxy-lyxo-2-heptulosaric acid - DSA Datura stramonium agglutinin - ER endoplasmic reticulum - GA Golgi apparatus - GNA Galanthus nivalis agglutinin - Kdo 3-deoxy-manno-2-octulosonic acid - MeKdo 3-deoxy-5-O-methyl-manno-2-octulosonic acid - SDS-PAGE sodium dodecyl sulphate polyacrylamide gel electrophoresis     

Squamatological differences between two closely related flatfish species of the genus <Emphasis Type="Italic">Pleuronichthys</Emphasis> (Pleuronectidae,Pleuronectiformes), with a proposal of unambiguous distinctive characters

Squamation and scale morphology were examined in two closely related species, Pleuronichthys cornutus and P. japonicus, in order to establish the unambiguous characters to clearly separate the two species. Scale counts on the lateral body surface were also examined, with new count definitions proposed. Both the longitudinal and vertical scale rows followed essentially regular patterns in P. cornutus, whereas the longitudinal scale rows were occasionally irregular in P. japonicus. Scales were basically fan-shaped and oval in P. cornutus and P. japonicus, respectively, although considerable variation in scale shape was apparent in both species. The blind side scales of P. cornutus had sharp posterior margins, whereas those of P. japonicus had round ones. Scale length (scales on the ocular side, area above the lateral line) in P. japonicus was relatively greater than in P. cornutus, a plot of total length versus scale length separating the species completely. Measurements of many ocular side scales from the area above the lateral line indicated high intraspecific variance in both species, being particularly prominent in P. cornutus. Nevertheless, considerable scale measurement differences were found between the two species in some body areas, supported by principal component analysis. Longitudinal and vertical row scale counts were higher in P. cornutus than in P. japonicus, with the ranges of central longitudinal scales (88–103 vs. 75–86) and scales below the lateral line counts (59–74 vs. 47–57), respectively, not overlapping between the species. The scale count difference could be caused by the difference of the scale size between the two species. The study demonstrated that the two Pleuronichthys species can be readily distinguished on the basis of scale morphology and size, in addition to scale counts.     

SILICEOUS SCALE PRODUCTION IN CHRYSOPHYTE AND SYNUROPHYTE ALGAE. I. EFFECTS OF SILICA-LIMITED GROWTH ON CELL SILICA CONTENT,SCALE MORPHOLOGY,AND THE CONSTRUCTION OF THE SCALE LAYER OF SYNURA PETERSENII1

The cells of synurophyte flagellates (algal class Synurophyceae, formerly included in the Chrysophyceae) are enclosed within a regularly imbricate layer of ornamented siliceous scales. Scale morphology is of critical taxonomic importance within this group of algae, and the scales are valuable indicator microfossils in paleolimnological studies. The data presented here demonstrate that scale morphology and the integrity of the scale layer can exhibit extreme variability in culture as a function of the cellular quota of silica under silica-limited growth. Silica-limited, steady-state populations of the colonial flagellate Synura petersenii Korsh. were maintained over a range of specific growth rates (μ= 0.11–0.69 days^?1) and silica cell quotas (Q_si= 0.13–2.40 pmoles Si · cell¹). Scale morphology and the organization of the scale layer became increasingly aberrant as silica stress increased. Under severe stress, scale deposition was completely suppressed so that cells appeared scale-free. This depression of scale deposition was reversible; populations of silica-starved, scale-free cells rapidly regenerated new scale layers when placed in batch culture and spiked with dissolved silica. During recovery from silica stress, cell division was repressed for 24 h while mean cell silica quota increased 25-fold. The first new scales appeared within 2 h after the silica addition, and development of the new scale layer proceeded in an approximately synchronous manner, residting in normal scale layers on virtually all cells after 48 h of recovery in Sirich medium. Silica content of silica-replete Synura cells is comparable to freshwater diatoms of siynilar size, but Synura has much greater potential quota variability than diatoms and no apparent threshold silica requirement. Silica-limited growth kinetics and competition between diatoms and Synura for silica are discussed. The results suggest that morphological variability of siliceous scales in natural populations of synurophyte flagellates may result from silica stress and that the experimental approach developed here has great potential value as a means for circumscribing ecotypic variation in scale morphology. Results also demonstrate that scale production can be uncoupled from cell division, suggesting that cell cycle regulation of silica biomineralization in the Synurophyceae may be fundamentally different from that of diatoms (algal class Bacillariophyceae). This experimental system has application in the future study of the intracellular membrane systems and the regulatory processes involved in silica biomineralization.     

Early scale development in Heterodontus (Heterodontiformes; Chondrichthyes): a novel chondrichthyan scale pattern

In two species of Heterodontus, H. portusjacksoni and H. galeatus, the first scales to develop form two opposing rows along the caudal fin axis on both the left and right sides of the fin. The opposing rows originate from an initial scale located on either side of the posterior tip of the caudal fin, with subsequent scales erupting in a posterior to anterior direction along the tail axis. These scale rows may strengthen tail movements, providing aeration in the egg case, but are lost later in ontogeny. Development of subsequent body scales shows a more irregular origin and arrangement, from anterior to posterior, to cover the dorsal and ventral lobes of the caudal fin. Although the early developmental pattern of the scale associated with the Heterodontus caudal fin has not been previously described, several chondrichthyan taxa, including chimeroids, likewise possess ordered rows of flank scales early in ontogeny that are subsequently lost. These ordered scales contrast with previous suggestions that chondrichthyan scale development is entirely random. Instead, regulated and sequential development of scales may be a plesiomorphic character for both chondrichthyans and osteichthyans, with the less organized arrangement in later ontogenetic stages being a derived condition within Chondrichthyes.     

<Emphasis Type="Italic">Sebastapistes taeniophrys</Emphasis> (Fowler 1943): a valid scorpionfish (Scorpaenidae) from the Philippines

The poorly known scorpionfish, Scorpaena taeniophrys, originally described from two specimens from the Philippines, is redescribed as a valid species of Sebastapistes. Sebastapistes taeniophrys differs from all other congeners in having a combination of 15 pectoral-fin rays, 31–33 scale rows in longitudinal series, 11–14 pored lateral-line scales, 3 predorsal scale rows, 12 gill rakers, 3 suborbital spines, absence of coronal spines, lower opercular spine with a median ridge and not covered with scales, ctenoid body scales, several dark transverse bands on ventral surface of mandible, a distinct elongate black blotch distally between the second or third and seventh dorsal-fin spines, and no black blotch on the nape.     

Ultrastructure of the flagellar apparatus inPyramimonas octopus (Prasinophyceae)

Summary While green algae usually lack one of the outer dynein arms in the axoneme, flagella of the octoflagellated prasinophytePyramimonas octopus possess dynein arms on all peripheral doublets. The outer dynein arm on doublet no. 1 is modified, and additional structures are associated with doublets no. 2 and 6. The flagellar scales are asymmetrically arranged. Thus the two rows of thick flagellar hairscales are displaced towards doublet no. 6,i.e., in the direction of the effective stroke of each flagellum. The underlayer of small scales includes two nearly opposite double rows scales, arranged in the longitudinal direction of the flagellum. The hairscales emerge from these rows. The double rows are separated on one side by 9, on the other by 11 rows of helically arranged scales. The central pair of microtubules twists, but the axoneme itself (represented by the 9 peripheral doublets), does not seem to rotate. The flagella are arranged in two groups, showing modified 180° rotational symmetry. The effective strokes of the two central flagella are exactly opposite, while the other flagella beat in six intermediate directions.     

Identification of a 41 kDa protein embedded in the biosilica of scales and bristles isolated from Mallomonas splendens (Synurophyceae, Ochrophyta)

Cells of the photosynthetic protist Mallomonas splendens (Synurophyceae, Ochrophyta) are encased within a highly patterned wall or scale case that consists of silicified scales and bristles. In an effort to understand the mechanisms that unicellular protists utilize to produce elaborate, mineralized structures of great complexity and hierarchical structure, we identified and characterized a 41 kDa protein from purified scales/bristles isolated from M. splendens (SP41 for Scale Protein of 41 kDa). A cDNA encoding this protein was isolated and sequence analysis indicated that it is a novel protein. Polyclonal antibodies were generated against bacterially expressed SP41 and used to localize the protein throughout scale and bristle morphogenesis. Immunoelectron microscopy confirmed the biochemical data that SP41 is a component of mature scales and bristles, the protein localizing to silicified components of the purified extracellular matrix. During scale and bristle biogenesis within the cell, SP41 is deposited into a specialized Silica Deposition Vesicle (SDV) concomitant with silica deposition, a highly regulated event during scale and bristle formation. These results argue for SP41 playing a role in morphogenesis and/or silicification within the SDV during scale and bristle biogenesis.     

Variation in elasmoid fish scale patterns is informative with regard to taxon and swimming mode

Variations in scales from nine regions on the flank of teleost fish were examined from the point of view of functional adaptation and with regard to which scales best differentiate species. Three teleost species were selected; two are from the genus Mugil, M. cephalus and M. curema, which are phylogenetically distant from the third, Dicentrarchus labrax. Scale form was described using seven landmarks, the coordinates of which were subjected to generalized Procrustes analysis followed by principal components analysis. Principal component scores were submitted to cross‐validated discriminant analysis to assess the utility of each scale in identifying species. The best discrimination (98%) was obtained with the scale from the central‐dorsal area. Scales from the anterior and central zones are relatively wide dorsoventrally and narrow anteroposteriorly. This appears to be related to the profile of the lateral body wall and with subcarangiform swimming. Scales from the posterior region are anteroposteriorly long and dorsoventrally narrow, this shape possibly being related to thrust. Despite the wide phylogenetic separation between mullets and D. labrax, the pattern of scale variation is similar. This may imply strong functional convergence, although studies of sister taxa with different swimming modes are required to confirm this. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 834–844.     

Biological Control of <Emphasis Type="Italic">Melanaspis Obscura</Emphasis> on Oaks in Northern California

In 1988, Encarsia aurantii (Howard) (Hymenoptera: Aphelinidae) was introduced into northern California to control an isolated infestation of obscure scale, Melanaspis obscura (Comstock) (Coccoidea: Diaspididae), on native and exotic oaks (Quercus spp.) in Sacramento’s Capitol Park. By 2002, there was no longer any need for chemical control of the scale (i.e., complete biological control). Both parasite and host are univoltine; peak emergence of adult parasites coincides with the peak of newly settled, first-instar scales. Increase of the parasite and concomitant decline of the scale from 1992 to 2004 are documented for one native and one exotic oak tree. During spring of 2004, mean density of female scales (based on 100 twigs per tree) on 12 previously infested oak trees was generally low, ranging from <1 (eight trees) to ∼15 (single tree) scales/twig. Mean percentage parasitization (per twig) ranged from ∼30 to ∼85%, and was density independent (spatial context) for each of five trees. Two refuges for the scale population were noted: some scale crawlers settled and developed under the parental scale cover (spatial refuge) and some female scales continued to produce crawlers into late summer and early fall, when adult parasites were no longer available (temporal refuge). This case illustrates how an introduction strategy (i.e., single-species release of E. aurantii), which was derived from an analysis of the parasite guild of the pest, was executed in the field and ultimately led to successful biological control.     

Gerres methueni Regan, 1920, a senior synonym ofG. rappi (Barnard, 1927) (Perciformes: Gerreidae)

Gerres methueni Regan, 1920, for many years identified asG. rappi (Barnard, 1927), is redescribed as a senior synonym of the latter species, following examination of two syntypes of the former and comparative material from South Africa and Madagascar.Gerres methueni is characterized by prominent dark stripes along the scale rows above the lateral line and the 4 or 5 rows immediately below it, 5–17 small scales on the preopercular flange, arranged in 1–3 scale row(s) at the corner, 42–44 pored lateral line scales+3–5 additional pored scales on the scaly sheath of the caudal fin base, 41/2–51/2 scales between the fifth dorsal fin spine base and lateral line, and second dorsal fin spine length equal to or slightly shorter than the third dorsal fin spine length.Gerres methueni is currently known from South Africa, southern Mozambique and Madagascar, being endemic in those areas.     

Isolation and phenotypic characterization ofChlamydomonas reinhardtii mutants defective in chloroplast DNA segregation

Summary Each wild-typeChlamydomonas reinhardtii cell has one large chloroplast containing several nuclei (nucleoids). We used DNA insertional mutagenesis to isolate Chlamydomonas mutants which contain a single, large chloroplast (cp) nucleus and which we namedmoc (monokaryotic chloroplast). DAPI-fluorescence microscopy and microphotometry observations revealed thatmoc mutant cells only contain one cp-nucleus throughout the cell division cycle, and that unequal segregation of cpDNA occurred during cell division in themoc mutant. One cell with a large amount of cpDNA and another with a small amount of cpDNA were produced after the first cell division. Unequal segregation also occurred in the second cell division, producing one cell with a large amount (about 70 copies) of cpDNA and three other cells with a small amount (only 2–8 copies) of cpDNA. However, most individualmoc cells contained several dozen cpDNA copies 12 h after the completion of cell division, suggesting that cpDNA synthesis was activated immediately after chloroplast division. In contrast to the cpDNA, the mitochondrial (mt) DNA of themoc mutants was observed as tiny granules scattered throughout the entire cell. These segregated to each daughter cell equally during cell division. Electron-microscopic observation of the ultrastructure ofmoc mutants showed that a low-electron-density area, which was identified as the cp-nucleus by immunoelectron microscopy with anti-DNA antibody, existed near the pyrenoid. However, there were no other structural differences between the chloroplasts of wild-type cells andmoc mutants. The thylakoid membranes and pyrenoid were identical. Therefore, we propose that the novelmoc mutants are only defective in the dispersion and segregation of cpDNA. This strain should be useful to elucidate the mechanism for the segregation of cpDNA.Abbreviations DAPI 4,6-diamidino-2-phenylindole - VIMPCS video-intensified microscope photon-counting system     

Effect of divalent cations on flagellar scales in the green flagellateTetraselmis cordiformis

Summary The structure and topography of flagellar scales (underlayer scales, rodshaped scales, hair-scales) in the green flagellateTetraselmis cordiformis has been studied in detail and the effect of divalent cations and fixation conditions on scale structure and topography was followed quantitatively. Hair-scales occur in two rows on opposite sides of a flagellum and are linked to the flagellar membrane and to two axonemal doublets by B-tubule-flagellar membrane connectives. Underlayer scales form about 24 longitudinal rows along the flagellum and occur in two distinctive shapes (pentagonal and square). The square shaped underlayer scales are related in position to the attachment sites of the hair-scales. Rod-shaped scales occur in about 20 longitudinal rows along the flagellum and are characteristically positioned as double scales. Calcium in the culture medium is necessary to retain rod-shaped scales on the flagellum, absence of calcium or chelation with EGTA or pyrophosphate leads to disappearance of rod-shaped scales from the flagellum. Other divalent cations can only partially substitute for calcium. It is suggested that calcium provides the linkage between underlayer scales and rod-shaped scales inTetraselmis. Flagellar scales inTetraselmis apparently fall into two categories: a) hair-scales (not affected by fixation or absence of divalent cations, firmly bound to axonemal microtubules via the flagellar membrane), b) underlayer scales and rod-shaped scales (affected by fixation and absence of divalent cations, kept on the flagellum mainly by electrostatic forces). The function of flagellar scales inTetraselmis is discussed.     

Formation and structure of scales in the Australian lungfish,neoceratodus forsteri (Osteichthyes: Dipnoi)

The large elasmoid scales of the Australian lungfish, Neoceratodusforsteri, are formed within the dermis by unpigmented scleroblasts, growing within a collagenous dermal pocket below a thick glandular epidermis. The first row of scales, on the trunk of the juvenile lungfish, appears below the lateral line of the trunk, single in this species, at around stage 53. The scales, initially circular in outline, develop anteriorly and posteriorly from the point of initiation in the mid‐trunk region, and rows are added alternately below the line, and above the line, until they reach the dorsal or ventral midline, or the margins of the fins. Scales develop later on the ventral surface of the head, from a separate centre of initiation. Scales consist of three layers, all produced by scleroblasts of dermal origin. The outermost layer of interlocking plates, or squamulae, consists of a mineralised matrix of fine collagen fibrils, covered by unmineralised collagen and a single layer of cells. Squamulae of the anterior and lateral surfaces are ornamented with short spines, and the mineralised tissue of the posterior surface is linked to the pouch by collagen fibrils. The innermost layer, known as elasmodin, consists of bundles of thick collagen fibrils and cells arranged in layers. An intermediate layer, made up of collagen fibrils, links the outer and inner layers. The elasmoid scales of N. forsteri can be compared with scale types among other osteichthyan groups, although the cellsand canaliculi in the mineralised squamulae bear littleresemblance to typical bone. J.Morphol., 2012. © 2011 Wiley Periodicals, Inc.