Mineral Preservatives in the Wood of Stradivari and Guarneri

Following the futile efforts of generations to reach the high standard of excellence achieved by the luthiers in Cremona, Italy, by variations of design and plate tuning, current interest is being focused on differences in material properties. The long-standing question whether the wood of Stradivari and Guarneri were treated with wood preservative materials could be answered only by the examination of wood specimens from the precious antique instruments. In a recent communication (Nature, 2006), we reported about the degradation of the wood polymers in instruments of Stradivari and Guarneri, which could be explained only by chemical manipulations, possibly by preservatives. The aim of the current work was to identify the minerals from the small samples of the maple wood which were available to us from the antique instruments. The ashes of wood from one violin and one cello by Stradivari, two violins by Guarneri, one viola by H. Jay, one violin by Gand-Bernardel were analyzed and compared with a variety of commercial tone woods. The methods of analysis were the following: back-scattered electron imaging, X-ray fluorescence maps for individual elements, wave-length dispersive spectroscopy, energy dispersive X-ray spectroscopy and quantitative microprobe analysis. All four Cremonese instruments showed the unmistakable signs of chemical treatments in the form of chemicals which are not present in natural woods, such as BaSO₄, CaF₂, borate, and ZrSiO₄. In addition to these, there were also changes in the common wood minerals. Statistical evaluation of 12 minerals by discriminant analysis revealed: a. a difference among all four Cremona instruments, b. the difference of the Cremonese instruments from the French and English antiques, and c. only the Cremonese instruments differed from all commercial woods. These findings may provide the answer why all attempts to recreate the Stradivarius from natural wood have failed. There are many obvious implications with regard to how the green tone wood should be treated, which chould lead to changes in the practice of violin-making. This research should inspire others to analyze more antique violins for their chemical contents.     

Investigating the Role of Auditory and Tactile Modalities in Violin Quality Evaluation

The role of auditory and tactile modalities involved in violin playing and evaluation was investigated in an experiment employing a blind violin evaluation task under different conditions: i) normal playing conditions, ii) playing with auditory masking, and iii) playing with vibrotactile masking. Under each condition, 20 violinists evaluated five violins according to criteria related to violin playing and sound characteristics and rated their overall quality and relative preference. Results show that both auditory and vibrotactile feedback are important in the violinists’ evaluations but that their relative importance depends on the violinist, the violin and the type of evaluation (different criteria ratings or preference). In this way, the overall quality ratings were found to be accurately predicted by the rating criteria, which also proved to be perceptually relevant to violinists, but were poorly correlated with the preference ratings; this suggests that the two types of ratings (overall quality vs preference) may stem from different decision-making strategies. Furthermore, the experimental design confirmed that violinists agree more on the importance of criteria in their overall evaluation than on their actual ratings for different violins. In particular, greater agreement was found on the importance of criteria related to the sound of the violin. Nevertheless, this study reveals that there are fundamental differences in the way players interpret and evaluate each criterion, which may explain why correlating physical properties with perceptual properties has been challenging so far in the field of musical acoustics.     

Wood Densitometry in 17th and 18th Century Dutch,German, Austrian and French Violins,Compared to Classical Cremonese and Modern Violins

Classical violins produced by makers such as Antonio Stradivari and Guarneri del Gesu have long been considered the epitome of the luthier''s art and the expressive tool of choice for the most celebrated violinists. It has been speculated these makers had access to wood that was unique in some way and that this was responsible for their acclaimed tonal characteristics. In an attempt to discern whether the above conjecture is true, we analyzed 17 modern and classical Dutch, German, Austrian and French violins by wood densitometry using computed tomography and correlated these results with our previous study of modern and Cremonese violins; in all studying 30 instruments of the violin family. In order to make this comparison possible we developed methods to cross calibrate results from different CT manufacturers using calibration wood pieces. We found no significant differences in median densities between modern and classical violins, or between classical violins from different origins. These results suggest that it is unlikely classical Cremonese makers had access to wood with significantly different wood density characteristics than that available to contemporaneous or modern makers.     

A comparison of wood density between classical Cremonese and modern violins

Classical violins created by Cremonese masters, such as Antonio Stradivari and Giuseppe Guarneri Del Gesu, have become the benchmark to which the sound of all violins are compared in terms of their abilities of expressiveness and projection. By general consensus, no luthier since that time has been able to replicate the sound quality of these classical instruments. The vibration and sound radiation characteristics of a violin are determined by an instrument's geometry and the material properties of the wood. New test methods allow the non-destructive examination of one of the key material properties, the wood density, at the growth ring level of detail. The densities of five classical and eight modern violins were compared, using computed tomography and specially developed image-processing software. No significant differences were found between the median densities of the modern and the antique violins, however the density difference between wood grains of early and late growth was significantly smaller in the classical Cremonese violins compared with modern violins, in both the top (Spruce) and back (Maple) plates (p = 0.028 and 0.008, respectively). The mean density differential (SE) of the top plates of the modern and classical violins was 274 (26.6) and 183 (11.7) gram/liter. For the back plates, the values were 128 (2.6) and 115 (2.0) gram/liter. These differences in density differentials may reflect similar changes in stiffness distributions, which could directly impact vibrational efficacy or indirectly modify sound radiation via altered damping characteristics. Either of these mechanisms may help explain the acoustical differences between the classical and modern violins.     

Vertebral evolution and the diversification of squamate reptiles

Taxonomic, morphological, and functional diversity are often discordant and independent components of diversity. A fundamental and largely unanswered question in evolutionary biology is why some clades diversify primarily in some of these components and not others. Dramatic variation in trunk vertebral numbers (14 to >300) among squamate reptiles coincides with different body shapes, and snake-like body shapes have evolved numerous times. However, whether increased evolutionary rates or numbers of vertebrae underlie body shape and taxonomic diversification is unknown. Using a supertree of squamates including 1375 species, and corresponding vertebral and body shape data, we show that increased rates of evolution in vertebral numbers have coincided with increased rates and disparity in body shape evolution, but not changes in rates of taxonomic diversification. We also show that the evolution of many vertebrae has not spurred or inhibited body shape or taxonomic diversification, suggesting that increased vertebral number is not a key innovation. Our findings demonstrate that lineage attributes such as the relaxation of constraints on vertebral number can facilitate the evolution of novel body shapes, but that different factors are responsible for body shape and taxonomic diversification.     

The convergent evolution of snake‐like forms by divergent evolutionary pathways in squamate reptiles*

Convergent evolution of phenotypes is considered evidence that evolution is deterministic. Establishing if such convergent phenotypes arose through convergent evolutionary pathways is a stronger test of determinism. We studied the evolution of snake‐like body shapes in six clades of lizards, each containing species ranging from short‐bodied and pentadactyl to long‐bodied and limbless. We tested whether body shapes that evolved in each clade were convergent, and whether clades evolved snake‐like body shapes following convergent evolutionary pathways. Our analyses showed that indeed species with the same numbers of digits in each clade evolved convergent body shapes. We then compared evolutionary pathways among clades by considering patterns of evolutionary integration and shape of relationship among body parts, patterns of vertebral evolution, and models of digit evolution. We found that all clades elongated their bodies through the addition, not elongation, of vertebrae, and had similar patterns of integration. However, patterns of integration, the body parts that were related by a linear or a threshold model, and patterns of digit evolution differed among clades. These results showed that clades followed different evolutionary pathways. This suggests an important role of historical contingency as opposed to determinism in the convergent evolution of snake‐like body shapes.     

Musical string instruments: Potential and limitations of tree-ring dating and provenancing to verify their authenticity

Authenticity is the prime factor affecting the market value of a work of art. String instruments are among the most valued works of art, particularly those made by the old violin-making masters of northern Italy in the late 17th and early 18th centuries. However, it is difficult to verify the authenticity of string instruments on the basis of style and design alone, as these are often copied or forged. Uncertainties related to craftsmanship can lead to financial and legal controversy, sometimes with even millions of dollars at stake. The authenticity of the Stradivari “Messiah” has long been disputed. Controversies at the end of the 1990s concerning its craftsmanship have enhanced interest in dating this violin. After different dendrochronological analyses provided conflicting tree-ring dates for the front of the violin, a scientifically-sound dendrochronological study eventually established 1682 as terminus post quem, i.e., the year when the last ring of the violin front was formed, before which the violin could not have been made. This date is consistent with the attributed date of manufacture, 1716, supporting Antonio Stradivari as the maker of the “Messiah”. However, this controversial dating of the “Messiah” sent shockwaves through the violin community. Here, we present the main facts which played a role in this controversy and we show how dangerous the use of dendrochronology can be if investigators do not adhere to well-established techniques and are not versed scholars in the literature. Such controversies threaten the reputation of dendrochronology. Today, many false theories and conceptual mistakes continue to circulate in the violin community. A thorough and scientifically-sound dendrochronological analysis of the wood used to make the instrument is the only analysis that can objectively indicate, if not the exact year an instrument was made, at least the date before which it certainly was not made. Here, we describe the importance, in terms of acoustics, of the anatomical characteristics of the wood with which instruments are made and its possible geographical provenance. We review the dendrochronological studies undertaken to assess the authenticity of the instruments made by the old Italian masters. Such studies help to establish the earliest date the tree from which the wood was taken could have been felled, and to determine the source region of the wood. We present the main achievements and challenges that have arisen in the past 50 years of studying the authenticity of string instruments, and discuss the limitations and advantages of using dendrochronological methods to establish the provenance and time period in which a work of art was created. Finally, we describe needs of research in history, wood anatomy and dendrochronology, proposing several new methods that may open up new avenues of research and aid in the assessment of the authenticity of old string instruments.     

A newly recognized fossil coelacanth highlights the early morphological diversification of the clade

Previously considered an actinopterygian or an osteichthyan incertae sedis, the Devonian (Givetian-Frasnian) Holopterygius nudus is reinterpreted as a coelacanth. This genus is among the oldest coelacanths known from articulated remains, but its eel-like morphology marks a considerable departure from the conventional coelacanth body plan. A cladistic analysis places Holopterygius as the sister taxon of the Carboniferous (Serpukhovian) genus Allenypterus. Despite the specialized morphology of these genera, they occupy a surprisingly basal position in coelacanth phylogeny; only Diplocercides and Miguashaia are further removed from the crown. A morphometric analysis reveals that coelacanths were anatomically disparate early in their history. Conflicts between this result and those of previous studies challenge the adequacy of systematic character sets for describing historical patterns of morphological variety. Coelacanths have long had an iconic place in the study of vertebrate evolution for their apparent anatomical conservatism over geological time, but Holopterygius provides clear evidence for rapid morphological evolution early in the history of this clade.     

Darwin, historical biogeography, and the importance of overcoming binary opposites

It is well known that Darwin and Wallace came to discover the phenomenon of evolution through a historical approach to the geographical distribution of organisms. Before Darwin, evolution was a mere speculation that could be invoked to explain some facts. Darwin's biogeographical argument for evolution is based largely on three main explanatory hypotheses. The first is that the geographical distribution of organisms is historically informative. The second hypothesis is that long-distance dispersal over barriers is one main force (extinction is the other) that modifies the distribution of organisms. The third of Darwin's biogeographical hypotheses is that the factors that shape the distribution of organisms are mainly historical (large, often global and long temporal scales) rather than ecological (small spatial and short temporal scales). From the time of Darwin until now, a wide spectrum of biogeographical schools have provided new insights that challenge the central role of space, dispersal and history as the main explanatory hypotheses for the distribution of organisms, generating three binary opposites: (1) the spatial dimension of evolution: geographical distribution of organisms as historically informative vs. historically uninformative; (2) the processes that modify the geographical distribution of organisms: dispersal vs. vicariance; and (3) the explanation of geographical distribution: history vs. ecology. We analyse these three binary opposites to show that the components of each are complementary rather than antagonistic approaches to the study of biogeography.     

Tree balance,time slices,and evolutionary turnover in cretaceous planktonic foraminifera

Studies of phylogenetic tree shape often concentrate on the balance of phylogenies of extant taxa. Paleontological phylogenies (which include extinct taxa) can contain additional useful information and can directly document changes in tree shape through evolutionary time. Unfortunately, the inclusion of extinct taxa lowers the power of direct examinations of tree balance because it increases the range of tree shapes expected under null models of evolution (with equal rates of speciation and extinction across lineages). A promising approach for the analysis of tree shape in paleontological phylogenies is to break the phylogeny down into time slices, examining the shape of the phylogeny of taxa alive at each time slice and changes in that shape between successive time slices. This method was illustrated with 57 time slices through a stratophenetic phylogeny of the Cretaceous planktonic foraminiferal superfamily Globotruncanacea. At 3 of 56 intervals between time slices, 93-92.5 million years ago (MYA), 89-88.5 MYA, and 85.5-84 MYA, the group showed steep increases in imbalance. Although none of these increases were significant after Bonferroni correction, these points in the history of the Globotruncanacea were nevertheless identified as deserving of further macroevolutionary investigation. The 84 MYA time slice coincides with a peak in species turnover for the superfamily. Time slices through phylogenies may prove useful for identifying periods of time when evolution was proceeding in a nonstochastic manner.     

Historical biology and the problem of design

Organisms are historical entities in that their past history plays an important role in shaping the properties they exhibit today. While this fact is widely acknowledged, little attempt has been made to develop a testable approach to the analysis of the historical factor in evolutionary morphology. The analysis of extrinsic environmental factors may reveal the limits imposed by the environment on biological design, but the intrinsic phylogenetic component of design may severely constrain the directions of structural modification that can occur. The importance of history can be assessed with (1) a phylogenetic hypothesis of genealogical relationship, (2) the use of emergent structural or functional attributes with general properties, and (3) the testing of historical hypotheses by the comparison of general properties between monophyletic lineages. The synthesis of a structural/phylogenetic approach to historical morphology with the analysis of extrinsic limits to form may provide the level of resolution needed to generate testable mechanistic hypotheses regarding the distribution of extant organismal forms in the hyperspace of possible morphologies.     

Correlated evolution of body and fin morphology in the cichlid fishes

Body and fin shapes are chief determinants of swimming performance in fishes. Different configurations of body and fin shapes can suit different locomotor specializations. The success of any configuration is dependent upon the hydrodynamic interactions between body and fins. Despite the importance of body–fin interactions for swimming, there are few data indicating whether body and fin configurations evolve in concert, or whether these structures vary independently. The cichlid fishes are a diverse family whose well‐studied phylogenetic relationships make them ideal for the study of macroevolution of ecomorphology. This study measured body, and caudal and median fin morphology from radiographs of 131 cichlid genera, using morphometrics and phylogenetic comparative methods to determine whether these traits exhibit correlated evolution. Partial least squares canonical analysis revealed that body, caudal fin, dorsal fin, and anal fin shapes all exhibited strong correlated evolution consistent with locomotor ecomorphology. Major patterns included the evolution of deep body profiles with long fins, suggestive of maneuvering specialization; and the evolution of narrow, elongate caudal peduncles with concave tails, a combination that characterizes economical cruisers. These results demonstrate that body shape evolution does not occur independently of other traits, but among a suite of other morphological changes that augment locomotor specialization.     

Head shape evolution in monitor lizards (Varanus): interactions between extreme size disparity,phylogeny and ecology

Characterizing patterns of observed current variation, and testing hypotheses concerning the potential drivers of this variation, is fundamental to understanding how morphology evolves. Phylogenetic history, size and ecology are all central components driving the evolution of morphological variation, but only recently have methods become available to tease these aspects apart for particular body structures. Extant monitor lizards (Varanus) have radiated into an incredible range of habitats and display the largest body size range of any terrestrial vertebrate genus. Although their body morphology remains remarkably conservative, they have obvious head shape variation. We use two‐dimensional geometric morphometric techniques to characterize the patterns of dorsal head shape variation in 36 species (375 specimens) of varanid, and test how this variation relates to size, phylogenetic history and ecology as represented by habitat. Interspecific head shape disparity is strongly allometric. Once size effects are removed, principal component analysis shows that most shape variation relates to changes in the snout and head width. Size‐corrected head shape variation has strong phylogenetic signal at a broad level, but habitat use is predictive of shape disparity within phylogenetic lineages. Size often explains shape disparity among organisms; however, the ability to separate size and shape variation using geometric morphometrics has enabled the identification of phylogenetic history and habitat as additional key factors contributing to the evolution of head shape disparity among varanid lizards.     

The effect of phylogeny on interspecific body shape variation in darters (Pisces: Percidae)

We conducted a geometric morphometric analysis of interspecific body shape variation among representatives of 31 species of darters (Pisces: Percidae) to determine whether there is evidence of a phylogenetic effect in body shape variation. Cartesian transformation grids representing relative shape differences of individual species and subspecies revealed qualitative similarities within most traditionally recognized taxonomic groups (genera and subgenera). Canonical variates analysis and a UPGMA cluster analysis were conducted to explore further the relationships among body shapes of species; both analyses revealed patterns of variation consistent with the interpretation that shape is associated with taxonomic affinities. Normalized Mantel statistics revealed a significant positive association between body shape differences and phylogenetic interrelatedness for each of four recent phylogenetic hypotheses, providing evidence of a phylogenetic effect. This result is somewhat surprising, however, given the largely incompatible nature of these four phylogenies. We provide evidence that this result may be due to (1) the inclusion of multiple sets of closely related species to represent the traditionally recognized genera and subgenera within each phylogeny and/or (2) the inclusion of several species with relatively divergent shapes and their particular positions within the phylogenies relative to one another or to the other species of darters.     

THE SHAPE OF CONTENTION: ADAPTATION,HISTORY, AND CONTINGENCY IN UNGULATE MANDIBLES

Mandibles and teeth of ungulates have been extensively studied to discern the functional significance of their design. Grazing ungulates have deeper mandibles, longer coronoid processes, flatter incisor arcades, and more hypsodont molars in comparison to browsers. If the functional significance of both mandible and teeth shapes is well‐established, it remains uncertain to what extent mandible shapes are really adapted to grazing, meaning that they evolved either to serve their current biological function or just as a structural requirement to accommodate higher crowned molars. Here, we address this question by studying the contribution of phylogeny, hypsodonty, and body size to mandibular shape variation. The mandible shape appeared to be significantly influenced by hypsodonty but not by body size. Interestingly, hypsodonty‐related changes influenced the tooth row in artiodactyls and perissodactyls significantly but in the opposite directions, which is ultimately related to their different digestive strategies. Yet, we obtained a strong phylogenetic effect in perissodactyls, suggesting that their mandible shape should be strongly inherited. The strength of this effect was not significant within artiodactyls (where hypsodonty explained much more variance in mandible shape). Digestive strategy is deemed to interplay with hypsodonty to produce different paths of adaptation to particular diets in ungulates.     

Contemporary evolutionary divergence for a protected species following assisted colonization

Background

Contemporary evolution following assisted colonization may increase the probability of persistence for refuge populations established as a bet-hedge for protected species. Such refuge populations are considered “genetic replicates” that might be used for future re-colonization in the event of a catastrophe in the native site. Although maladaptive evolutionary divergence of captive populations is well recognized, evolutionary divergence of wild refuge populations may also occur on contemporary time scales. Thus, refuge populations may lose their “value” as true genetic replicates of the native population. Here, we show contemporary evolutionary divergence in body shape in an approximately 30-year old refuge population of the protected White Sands pupfish (Cyprinodon tularosa) resulting in a body-shape mismatch with its native environment.

Methodology/Principal Findings

Geometric morphometic data were collected from C. tularosa cultures raised in experimental mesocosms. Cultures were initiated with fish from the two native populations, plus hybrids, in high or low salinity treatments representing the salinities of the two native habitats. We found that body shape was heritable and that shape variation due to phenotypic plasticity was small compared to shape variation due to population source. C. tularosa from the high salinity population retained slender body shapes and fish from the low salinity population retained deep body shapes, irrespective of mesocosm salinity. These data suggest that the observed divergence of a recently established pupfish population was not explained by plasticity. An analysis of microsatellite variation indicated that no significant genetic drift occurred in the refuge population, further supporting the adaptive nature of changes in body shape. These lines of evidence suggest that body shape divergence of the refuge population reflects a case of contemporary evolution (over a 30-year period).

Conclusions/Significance

These results suggest assisted colonization can introduce novel, and/or relaxed selection, and lead to unintended evolutionary divergence.     

ALTERNATE PATHWAYS OF BODY SHAPE EVOLUTION TRANSLATE INTO COMMON PATTERNS OF LOCOMOTOR EVOLUTION IN TWO CLADES OF LIZARDS

Body shape has a fundamental impact on organismal function, but it is unknown how functional morphology and locomotor performance and kinematics relate across a diverse array of body shapes. We showed that although patterns of body shape evolution differed considerably between lizards of the Phrynosomatinae and Lerista, patterns of locomotor evolution coincided between clades. Specifically, we found that the phrynosomatines evolved a stocky phenotype through body widening and limb shortening, whereas Lerista evolved elongation through body lengthening and limb shortening. In both clades, relative limb length played a key role in locomotor evolution and kinematic strategies, with long‐limbed species moving faster and taking longer strides. In Lerista, the body axis also influenced locomotor evolution. Similar patterns of locomotor evolution were likely due to constraints on how the body can move. However, these common patterns of locomotor evolution between the two clades resulted in different kinematic strategies and levels of performance among species because of their morphological differences. Furthermore, we found no evidence that distinct body shapes are adaptations to different substrates, as locomotor kinematics did not change on loose or solid substrates. Our findings illustrate the importance of studying kinematics to understand the mechanisms of locomotor evolution and phenotype‐function relationships.     

Morphological correlates with diet of fish assemblages in brush park fisheries of tropical estuaries

Brush park fishery in Negombo estuary, Sri Lanka is a traditional fishing practice which relies on fishes attracted to artificial woody fish aggregation devices. This study investigates whether constituent species in these brush parks exhibit morphological variations in relation to their dietary habits. Fishes caught in brush parks were sampled from April 2014 to April 2016 covering rainy, intermediate and dry seasons. There were 817 specimens of 46 species belonging to 24 families. From each specimen, 17 morphological attributes were determined and diet composition of each species was analyzed in terms of relative biovolume. Trophic index of each species estimated from the proportions of dietary items and their possible trophic level in the community was significantly related to two body proportions (Maximum body height/Maximum body width and Total length/ Maximum body height) which described shape of fish. Principal component analysis of morphometric attributes and dietary habits indicated that the species in the higher trophic levels are characterized by slender, long-body shapes and those occupy lower trophic levels are predominantly laterally compressed with deep body shapes. As such, structure of coexisting species in brush parks of Negombo estuary is predominantly along the trophic dimension and is related to morphological traits of constituent species. The predictive power of ecomorphological correlates with diets of fish species other than mugilids which are attracted to brush parks, can therefore be considered as a useful tool for conducting rapid ecological assessment.     

Surface electromyography of forearm and shoulder muscles during violin playing

The aims of this study were to explore muscle activity levels during different violin repertoires, quantify the general levels bilaterally in upper extremity muscles, and evaluate associations between muscle activity and anthropometrics characteristics. In 18 skilled violin players surface EMG was recorded bilaterally from trapezius (UT), flexor digitorum superficialis (FDS), extensor carpi ulnaris (ECU), extensor digitorum cummunis (EDC), and extensor carpi radialis (ECR) during A and E major scales played in three octaves and Mozart’s Violin Concerto no. 5. To compare side differences the static, median and peak levels of muscle activity were calculated from an amplitude probability distribution function (APDF). This study demonstrated that scales played as standardized tasks can be used to estimate the average muscle activity during violin playing. Comparing results from scales and the music piece revealed a similar muscle activity across all muscles in the music piece and E major scales. The static, median and peak EMG levels were higher in left than in right forearm muscles with left ECU presenting the highest peak load of 30 %MVE. Females demonstrated a higher muscle activity than males, but this was in accordance with differences in anthropometric measures.     

Body shape, diet and ontogenetic diet shifts in young fish of the Sinnamary River, French Guiana, South America

A total of 1468 young fish representing 66 taxa from the Sinnamary River, French Guiana was classified by complete cluster analysis of mean relative body width and mean relative body height into four groups. These had anguilliform, disciform, flat or intermediate body shapes and belonged chiefly to Gymnotiformes, Perciformes, Siluriformes and Characiformes, respectively. Several of the taxa shifted from one to another body shape during ontogenesis. Seven diet groups were defined by complete cluster analysis. Among these, six groups were represented by carnivorous fish. The three most frequent groups had diets of (1) mainly insect larvae and small crustaceans, (2) insect larvae, and (3) predominantly terrestrial insects. The majority of the fish taxa showed ontogenetic diet shifts. Carnivorous fish usually switched from small-size prey, such as small crustaceans, to intermediate-size prey, such as insect larvae and/or to large-size prey, such as insects and/or fish. However, taxa differed in their capacities to switch from small prey to intermediate and/or to large prey. Taxa of different body shapes had significantly different diets. Disciform fish fed mainly on aquatic insect larvae and terrestrial insects but also, in small amounts, on small curstaceans. Most anguilliform taxa ate insect larvae. Individuals belonging to the depressiform or intermediate morphotype had varied diets ranging from plant debris and substratum to fish.