The complex three-dimensional shape of the apparatus including its numerous muscle groups leads to directional changes that are not isometric. Craniofacial mechanics, the pharyngeal jaw apparatus, phenotypic plasticity, and hybridization may have facilitated diversification of cichlid fishes by generating functional, morphological, and/or genetic diversity. Liem (1973) hypothesized that morpholog-ical specialization of the cichlid PJA was a key evolution-ary innovation. Unusual tissues develop in these regions of cichlid … While pharyngeal jaw shape was unaffected by either head length or head depth, its disparity … Interestingly, mutagenesis screens within Danio rerio, the zebrafish, have suggested that the genetic architecture underlying the oral and pharyngeal jaws of teleosts (Piotrowski et al. Their highly integrated pharyngeal jaw appa-ratus permits cichlids to transport and process food, thus enabling the oral jaws to develop special-isations for acquiring a variety of food items. [10], The exceptional mobility of the moray eel's pharyngeal jaws was discovered in 2007 by UC Davis scientists. Finally, the pharyngeal jaw apparatus contains yet another set of teeth, which, as with the outer jaws, contain a wide variety of tooth types depending on the diet. Suggest some possible structural differences between teeth on the pharyngeal jaws of carnivorous and herbivorous minnows. We measure a suite of morphological and functional traits, characterizing both oral and pharyngeal jaws in 84 species of Neotropical cichlids and use phylogenetic comparative methods Register, Oxford University Press is a department of the University of Oxford. Whether dental divergence arises as a result of environmental induced plasticity or originates via trophic polymorphism as found in the species Herichthys minckleyi, there appear to be general rules that structure interspecific divergence in cichlid pharyngeal erupted and replacement dentitions. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. Cichlids are able to process a wide range of foods and having two sets of jaws enables them to process that food very efficiently. This allows them to crush, chew, and process whatever … Most users should sign in with their email address. Dental convergence is a hallmark of cichlid fish adaptive radiations. Introduction. Pharyngognathy is a widely recognized innovation of the pharyngeal jaws that enhances the ability to process hard and tough prey in several major radiations of fishes, including marine wrasses and freshwater cichlids. These are possibly a response to their inability to swallow as other fishes do by creating a negative pressure in the mouth, perhaps induced by their restricted environmental niche (burrows). These factors play a significant role... | … Evolutionary strategies and morphological innovations: Cichlid pharyngeal jaws. The lower pharyngeal bones in al cichlid species are fused into a single tooth-bearing structure controlled by a complex set of muscles which allows the structure to be used as a second set of jaws. Electric Blue Cichlid/Electric Blue Hap (Sciaenochromis fryeri) Difficulty: Easy. Syst. However, both sets of jaws operate in tandem such that the oral jaws are involved in food acquisition (i.e. In the aquarium, it can be observed sucking larger snails from their shells. To test several hypotheses regarding the evolution of cichlid crushing pharyngeal dentitions, we used X-ray computed tomography scans to comparatively examine dental evolution in the pharyngeal jaw of a diversity of New World Heroine cichlid lineages. Primary oral jaws contain teeth which are used to capture and hold food, while pharyngeal jaws have pharyngeal teeth which function as a chewing tool. They are believed to have originated as modified gill arches, in much the same way as oral jaws. Cichlids are therefore able to chew their food and exploit vegetable matter for food. citrinellus) between two different types of pharyngeal jaws, molariform and papilliform. The benthic–pelagic axis has been an important source of divergence during adaptive radiation. cichlid and labrid pharyngeal jaw development, and for functional studies of the molecular mechanisms underly-ing pharyngeal patterning and skeletogenesis in Nile tilapia and its relatives. They are believed to have originated as modified gill arches, in much the same way as oral jaws. MATERIAL AND METHODS Embryos and Staging Embryos were obtained as in Le Pabic et al. citrinellus) between two different types of pharyngeal jaws, molariform and papilliform. The invading cichlids successfully occupy contiguous and … We explored diversification of lower pharyngeal jaw shape, a key feature of pharyngognathy, and the extent to which it is influenced by head shape in Neotropical cichlids. Photo by Juan Miguel Artigas Azas. Christoph J. Hellig, Michaela Kerschbaumer, Kristina M. Sefc, Stephan Koblmüller, Allometric shape change of the lower pharyngeal jaw correlates with a dietary shift to piscivory in a cichlid fish, Naturwissenschaften, 10.1007/s00114-010-0682-y, 97, 7, (663-672), (2010). Pharyngognathy is a widely recognized innovation of the pharyngeal jaws that enhances the ability to process hard and tough prey in several major radiations of fishes, including marine wrasses and freshwater cichlids. The level of significance determined from permutations is shown with a dotted line. CICHLID JAW MODULARITY 2097 FIG.1. Individuals with papilliform lower pharyngeal jaws are … 2006; Fraser et al. 27) Variations in the pharyngeal jaws between species of cichlids have contributed to an adaptive radiation by these fishes because ________. However, later studies based on Lake Victoria cichlids suggest that this trait may also become a handicap when competing with other predator species. Cichlids are one of the two known families of fish that possess movable pharyngeal jaws (the other is moray eels) that allow them to pre-process the food before it is swallowed. Apparently, such a trade-off exists in the Neotropical Midas Cichlid (Amphilophus cf. phenotypic plasticity in the pharyngeal jaw of Midas Cichlids. Search for other works by this author on: © The Author(s) 2020. Liem, Karel F. (Museum of Comparative Zoology, Harvard University, Cambridge, Mass. Relationships between diet and phylogenetic principal components (pPC) that reflect major shape variation among the lower pharyngeal jaws of Neotropical cichlid fishes. Individuals with papilliform lower pharyngeal jaws are … The pharyngeal jaws are furnished with sharp conical teeth that are straight rather than recurved (Fig. [4][5] A study conducted on cichlids showed that the pharyngeal jaws can undergo morphological changes in less than two years in response to their diet. In the present study, we explore the consequences of decou-pled prey capture and processing functions in an exemplar group of ray-finned fishes, the monophyletic cichlids of South and Middle America. Tooth number QTL in the oral and the pharyngeal jaws. Pharyngeal jaws are a second set of jaws contained within an animals throat, or pharynx, distinct from the primary or oral jaws. Syst. Introduction. tear- They share one thing in common, but even that feature is highly adaptable. Evolutionary strategies and morphological innovations: Cichlid pharyngeal jaws. grasp-ing prey) while the pharyngeal jaws process food (i.e. [2][3] Based on connections between musculoskeletal morphology and dentition, diet has been proposed as a main driver of the evolution of the pharyngeal jaw. 22:425441.-The percoid fish family Cichlidae possesses a phenomenal ability to colonize lakes and to diversify to an extent unmatched by any other vertebrate family in the presence of'predator pressure and strong competition. Dental convergence is a hallmark of cichlid fish adaptive radiations. This type of repeated evolution characterizes both the oral jaws of these fishes as well as their pharyngeal jaws that are modified gill arches used to functionally process prey like hard-shelled mollusks. The pharyngeal jaw apparatus of cichlids, a second set of jaws functionally decoupled from the oral ones, is known to mediate ecological specialization and often differs strongly between sister-species. A) pharyngeal jaws permit the cichlids to efficiently eat only other fish. Greater LPJ suturing, pharyngeal jaw splitting under compression and the forces used to crush molluscs in the wild suggest increased LPJ fusion in the trophically polymorphic Herichthys minckleyi operates to strengthen the pharyngeal jaw. The upper pharyngeal jaw may be moved up and down and dislocated and the lower pharyngeal jaw can be moved forwards and backwards. The substantial variation in erupted tooth sizes and numbers as well as replacement teeth found in these fishes showed several general patterns. Pharyngeal jaws in cichlids and in related pharyngognath families are a second pair of jaws located deep in the buccopharyngeal cavity and developmentally derived from the second to fifth branchial arches (Nelson, '69; Liem and Greenwood, '81; Vandewalle et al., '94). Syst. Evolutionary strategies and morphological innovations: Cichlid pharyngeal jaws. evolutionary innovation, are not universally beneficial. - "Evolutionary Strategies and Morphological Innovations: Cichlid Pharyngeal Jaws" FIG. pharyngeal jaws of cichlid fishes and several marine fish lineages, a classic example of. Dental convergence is a hallmark of cichlid fish adaptive radiations. This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (. The bottom pharyngeal bone fused into a single tooth-bearing structure manipulated by strong muscles to function as a second set of jaws. Functional decoupling of oral and pharyngeal jaws is widely considered to have expanded the ecological repertoire of cichlid fishes. [8] However, later studies based on Lake Victoria cichlids suggest that this trait may also become a handicap when competing with other predator species. Larger toothed species tended to have fewer teeth suggesting a potential role of spatial constraints in cichlid dental divergence. 1996) may have been extensively decoupled before the novelties in the cichlid pharyngeal jaw originated. The innovation is thought to have played an important role in cichlids’ spectacular diversification throughout marine and freshwater ecosystems. [5] These rapid changes are an example of phenotypic plasticity, wherein environmental factors affect genetic expression responsible for pharyngeal jaw development. Instead, when the moray bites prey, it first bites normally with its oral jaws, capturing the prey. Instead, fish-eating cichlids must awkwardly chop large prey into pieces. [11], "Independent evolution of the specialized pharyngeal jaw apparatus in cichlid and labrid fishes", "The Evolution of Pharyngognathy: A Phylogenetic and Functional Appraisal of the Pharyngeal Jaw Key Innovation in Labroid Fishes and Beyond", "Morphological convergence of pharyngeal jaw structure in durophagous perciform fish", "Shaping development through mechanical strain: the transcriptional basis of diet-induced phenotypic plasticity in a cichlid fish", "Regulatory gene networks that shape the development of adaptive phenotypic plasticity in a cichlid fish", "Micro- and macroevolutionary decoupling of cichlid jaws: A test of Liem's key innovation hypothesis", "A pharyngeal jaw evolutionary innovation facilitated extinction in Lake Victoria cichlids", "If the First Bite Doesn't Do It, the Second One Will", "article explaining moray eel pharyngeal jaws", https://en.wikipedia.org/w/index.php?title=Pharyngeal_jaw&oldid=997529281, Creative Commons Attribution-ShareAlike License, This page was last edited on 31 December 2020, at 23:57. In cichlid pharyngeal jaws, shape changes from growth of the skull adjust the orientation of the acting muscles. This second set of jaws is situated in the pharynx of cichlid … A, B and C: sagittal sections through ossifying phb2‐UP2. To purchase short term access, please sign in to your Oxford Academic account above. Cichlid pharyngeal jaws have become very specialized in prey processing and may have helped cichlid fishes become one of the most diverse families of vertebrates. [9], Most fish species with pharyngeal teeth do not have extendable pharyngeal jaws. Positive and negative values of pPC1 are associated with widely and narrowly spaced lateral processes, respectively. The pharyngeal jaw also frees up the outer jaw from chewing, allowing more prey to be captured while the previous meal is being processed (See Food Habits for more details). Evolutionary strategies and morphological innovations: Cichlid pharyngeal jaws. Like cichlids, minnows use their pharyngeal jaws to process food. Temperament: Semi … C Darrin Hulsey, Axel Meyer, J Todd Streelman, Convergent Evolution of Cichlid Fish Pharyngeal Jaw Dentitions in Mollusk-Crushing Predators: Comparative X-Ray Computed Tomography of Tooth Sizes, Numbers, and Replacement, Integrative and Comparative Biology, Volume 60, Issue 3, September 2020, Pages 656–664, https://doi.org/10.1093/icb/icaa089. Species with larger numbers of erupted pharyngeal teeth also had larger numbers of replacement teeth. Oral and pharyngeal jaws are developmentally decoupled such that they are derived from the first (in part) and sev-enth pharyngeal arches, respectively (Fraser et al. The pharyngeal jaw apparatus of cichlids, a second set of jaws functionally decoupled from the oral ones, is known to mediate ecological specialization and often differs strongly between sister-species. Zool. Many fish have similar “pharyngeal jaws” but the cichlids’ ones are exceptionally strong. of Australia, some 70–80% of cichlids are found in Africa, with the greatest diversity found in the Great Lakes (lakes Victoria, Tanzania and Malawi). They are opportunistic feeders, but in the wild, when the competition for fish food gets tough, they can specialize on exotic food items because of their pharyngeal jaws. Pharyngeal jaw shape and head shape are known to evolve adaptively in cichlids, as both are related to feeding behav-iorandhabitatuse(López-Fernándezetal.2013;Seehausen 2015;Burressetal.2018a,2018b).Therelativedepth,length, Download with Google Download with Facebook. Like some other fish, they have two sets of jaws, one inside the other. Petenia splendida is the perfect example of a piscivorous cichlid. A particularly notable exception is the highly mobile pharyngeal jaw of the moray eels. The authors determine that head shape modulates diversification of pharyngeal jaws and subsequently the ecological diversity of cichlids. Ecology of Freshwater Fish, 2015. 4.77B) and that are used to lacerate the flesh of the prey. Cichlids are one of the most striking examples of trophic diversity as they have evolved highly specialized pharyngeal jaws in addition to oral jaws . For permissions please email: journals.permissions@oup.com. 1. Zool. Zool. Please check your email address / username and password and try again. Immediately thereafter, the pharyngeal jaws are brought forward and bite down on the prey to grip it; they then retract, pulling the prey down the moray eel's gullet, allowing it to be swallowed. Small snails are cracked between the pharyngeal jaws (within its mouth) while large ones are tackled with the mouth. Edward D Burress, Peter C Wainwright, A peacock bass (Cichla) functional novelty relaxes a constraint imposed by the classic cichlid pharyngeal jaw innovation, Biological Journal of the Linnean Society, 10.1093/biolinnean/blaa050, (2020). However, at that time pharyngeal jaws in other fishes were already known. 1). The pharyngeal and oral jaws of Herichthys minckleyi.The levator externi IV and levator posterior (LEIV/LP) are almost inseparably associated in many fish and function as the primary pharyngeal crushing muscles that form the novel labroid muscular sling. (2007) Advances in imaging and modeling facilitate the calculation of biomechanical forces in biological specimens. Conclusions: The pharyngeal jaw apparatus of Midas Cichlids can be expressed plastically if stimulated mechanically during feeding. Originally hypothesized to have evolved only once,[1] current morphological and genetic analyses suggest at least two separate points of origin. It has long been thought that functional decoupling of the oral and pharyngeal jaws in cichlids has facilitated their explosive evolution. determiner Juan Miguel Artigas Azas While cichlids differ, they all share one common feature: pharyngeal jaws. PDF | Advances in imaging and modeling facilitate the calculation of biomechanical forces in biological specimens. Syst. The paired upper pharyngeal jaws and The pharyngeal jaws of carnivorous minnows are sharp teeth specialized for … Based on … Rates of piscivory predict pharyngeal jaw morphology in a piscivorous lineage of cichlid fishes. 22:425-441.-The percoid fish family Cichlidae possesses a phenomenal ability to colonize lakes and to diversify to an extent unmatched by any other vertebrate family in the presence of … B) pharyngeal jaws permit enhanced respiration in the Rift Lakes of East Africa. Originally hypothesized to have evolved only once, current morphological and genetic analyses suggest at least two separate points of origin. Introduction. Here we report the in-depth examination of the spectacular adaptive radiation of cichlid fishes in Lake Tanganyika. MicroCT imaging of radiopaque-stained vertebrate embryos were used to accurately capture the spatial relationships of the pharyngeal jaw apparatus in two cichlid species (Haplochromis elegans and Amatitlania nigrofasciata) for the purpose of creating a time series of developmental stages using finite element models, which can be used to assess the effects of biomechanical forces present … It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide, This PDF is available to Subscribers Only. If you originally registered with a username please use that to sign in. Dental convergence is a hallmark of cichlid fish adaptive radiations. The pharyngeal and oral jaws are shown in a diagrammatic lateral view (A).The dorsal view of a non-molluskivorous papilliform H. minckleyi jaw is shown (B).CT scans layers produced of this jaw (C) allow the visualization of the number of erupted teeth as well as the number of replacement teeth as one toggles through … 22:425-441.-The percoid fish family Cichlidae possesses a phenomenal ability to colonize lakes and to diversify to an extent unmatched by any other vertebrate family in the presence of predator pressure and strong competition. Our study group is the monophyletic ra-diation of about 500 cichlid species found in the Americas. But the second set (called pharyngeal jaws) can morph into vastly different shapes. These factors play a significant role during ontogenetic development of cichlid pharyngeal jaws, a key innovation responsible for one of … This fish eats mollusks. Similarly, all cichlids possess modified pharyngeal jaws that provide a strong biting mechanism, which provided access to previously inaccessible resources. Marcelo Loureiro. Download Free PDF. Linkage group numbers are given as the homologized regions of the Tilapia cichlid genome. All rights reserved. B: Intramembranous ossification has formed between the tooth … Development of pharyngeal jaws and their associated skeletal apparatus in the visceral arches of the cichlid Astatotilapia (Haplochromis) elegans is known in some detail, as is the microstructure of the bone of the pharyngeal jaws in two morphotypes (morphologically distinguishable stable variants) of the cichlid Astatoreochromis alluaudi (Huysseune et al., *1994). Don't already have an Oxford Academic account? Since this trait is commonly differentiated - among other traits - between Midas Cichlid species, its plasticity might be an important factor in Midas Cichlid speciation. The jaws of cichlids have been key to their diversification. 02138) 1974. Fish that ate softer prey, on the other hand, exhibited a more slender jaw with thin, curved teeth used for tearing apart fleshy prey. Both the oral (A) and pharyngeal jaws (B) have a large-effect QTL on chromosome 11. Cichlid pharyngeal jaws have become very specialized in prey processing and may have helped cichlid fishes become one of the most diverse families of vertebrates. Functional decoupling of oral and pharyngeal jaws is widely considered to have expanded the ecological repertoire of cichlid fishes. Zool. Don't already have an Oxford Academic account? CICHLID JAW MODULARITY 2097 FIG.1. A large-scale analysis of dietary. This second set of jaws is situated in the pharynx of cichlid fishes and is used to masticate food, allowing the fish to specialize in very specific nutritional niches. [6][7] Studies of the genetic pathways suggest that receptors in the jaw bone respond to the mechanical strain of biting hard-shelled prey, which prompts the formation of a more robust set of pharyngeal jaws. Under this hypothesis, the presence of such an efficient second set of jaws in the cichlid feeding apparatus would have released oral jaws from the task Almost three decades before (1979), the fictional xenomorph creature from Alien series was first depicted showing a second set of jaws for attacking its prey. Mollusk feeders, eg, the five-barred cichlid ( Neolamprologus tretocephalus ), crushes its prey between large, flat crowned pharyngeal teeth ( Fig. Free PDF. Create a … Sections through the ossifying lower pharyngeal jaws and phb2‐UP2. You could not be signed in. How these events and traits altered cichlids trajectory across the adaptive landscape motivates much of my current research. plosive adaptive radiation of cichlid fishes in East African lakes. actions by the pharyngeal jaws, or high force-transmitting oral jaws paired with highly kinetic pharyngeal jaws (Fig. A: 126 hpf, mineralization of tooth bud alone is observed. Some species capture snails and have strong muscles and specialized mollariform teeth on their pharyngeal jaws to help them crush them, like Herichthys labridens in North America. or. We present evidence that the modified pharyngeal jaws of cichlid fishes and several marine fish lineages, a classic example of evolutionary innovation, are … For full access to this pdf, sign in to an existing account, or purchase an annual subscription. ... Bolivian rams are members of the dwarf cichlid group. Pharyngeal jaw Last updated November 23, 2019 The pharyngeal jaws of the moray eel Oral and pharyngeal jaws of a cichlid.The photographs show a Malawi eyebiter (Dimidiochromis compressiceps).. Pharyngeal jaws are a "second set" of jaws contained within an animal's throat, or pharynx, distinct from the primary or oral jaws.They are believed to have originated as … It crushes snails mainly with its mouth. Lines denote significant relationships. The invading cichlids Replacement tooth size is almost exactly predicted (r = 0.99) from the size of erupted teeth across all of the species. But, the degree to which the evolution of these jaw systems is decoupled and whether decoupling has impacted trophic diversification remains unknown. 2009). PDF. But pharyngeal jaws don’t open widely enough to efficiently swallow large prey items such as fish. A notable example are fish from the family Cichlidae. With their many differences in form and function, you might wonder what unifying trait makes a cichlid a cichlid. Most cichlids can eat anything that is available, which is welcome in the aquarium. But, the degree to which the evolution of these jaw systems is decoupled and whether decoupling has impacted trophic diversification remains unknown. The zoologists were particularly interested in the three-dimensional structure of the pharyngeal jaw. Pharyngognathy is a widely recognized innovation of the pharyngeal jaws that enhances the ability to process hard and tough prey in several major radiations of fishes, including marine wrasses and freshwater cichlids. Pharyngeal jaws are a "second set" of jaws contained within an animal's throat, or pharynx, distinct from the primary or oral jaws. The pharyngeal jaws of cichlid fish have been hypothesized to be the mechanism responsible for sound production (Lanzing, 1974; Lobel, 2001), with the pharyngeal muscles used to maneuver the pharyngeal jaws together, facilitating stridulation of the pharyngeal teeth.Courtship and feeding sounds in the cichlid Tramitichromis intermedius have the same … To test several hypotheses regarding the evolution of cichlid crushing pharyngeal dentitions, we used X-ray computed tomography scans to comparatively examine dental evolution in the pharyngeal jaw … You do not currently have access to this article. Variation in the Sensory Space of Three-spined Stickleback Populations, Introduction to the Symposium: New Frontiers in Antarctic Marine Biology, Epigenetics of Animal Personality: DNA Methylation Cannot Explain the Heritability of Exploratory Behavior in a Songbird, Comparative study of snake lateral undulation kinematics in model heterogeneous terrain, Molecular Plasticity in Animal Pigmentation: Emerging Processes Underlying Color Changes, About Integrative and Comparative Biology, About the Society for Integrative and Comparative Biology, https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model, Receive exclusive offers and updates from Oxford Academic, Copyright © 2021 The Society for Integrative and Comparative Biology. 2009). The pharyngeal apparatus serves very much like our teeth, whereas the teeth of the lips serve more like our hands. The African cichlids of the East-African rift-lakes provide one of the most dramatic examples of adaptive radiation known. Apparently, such a trade-off exists in the Neotropical Midas Cichlid (Amphilophus cf. This type of repeated evolution characterizes both the oral jaws of these fishes as well as their pharyngeal jaws that are modified gill arches used to functionally process prey like hard-shelled mollusks. [6] Fish that ate hard shelled prey had a robust jaw with molar-like teeth fit for crushing their durable prey. Mollusk crushing was, therefore, highly associated with not only larger pharyngeal teeth, but also larger replacement teeth. (27-Mar-2014). This study provides an example of how a simple aspect of morphology can have broad evolutionary and ecological implications by interacting with a major innovation. 12.-Dorsal view of dissected and isolated branchial n~uscles and apparatus of the generalized perciform Badis badis. oral and pharyngeal jaw traits in cichlids, and studies in this area have focused on a limited number of traits (Hulsey et al. In cichlids generally, the oral and pharyngeal teeth differ with different species in ways that allow them to process different kinds of prey. 22:425–441.—The percoid fish family Cichlidae possesses a phenomenal ability to colonize lakes and to diversify to an extent unmatched by any other vertebrate family in the presence of predator pressure and strong competition. This type of repeated evolution characterizes both the oral jaws of these fishes as well as their pharyngeal jaws that are modified gill arches used to functionally process prey like hard-shelled mollusks. [7], A notable example are fish from the family Cichlidae. pharyngeal jaw. This article is also available for rental through DeepDyve. This type of repeated evolution characterizes both the oral jaws of these fishes as well as their pharyngeal jaws that are modified gill arches used to functionally process prey like hard-shelled mollusks. The pharyngeal jaws of cichlid fish have been hypothesized to be the mechanism responsible for sound production (Lanzing, 1974; Lobel, 2001), with the pharyngeal muscles used to maneuver the pharyngeal jaws together, facilitating stridulation of the pharyngeal teeth.Courtship and feeding sounds in the cichlid Tramitichromis intermedius have the same dominant frequency of 517 … The pharyngeal and oral jaws of Herichthys minckleyi.The levator externi IV and levator posterior (LEIV/LP) are almost inseparably associated in many fish and function as the primary pharyngeal crushing muscles that form the novel labroid muscular sling. The innovations in the cichlid pharyngeal jaw may have facilitated the utilization of novel prey and ultimately led to the unparalleled trophic diversification of these fishes ().Although most bony fishes have pharyngeal gill arches modified to process prey (Liem 1986; Wainwright 1989), the cichlid pharyngeal jaw uniquely exhibits novel upper pharyngeal jaw …
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