Development of the paired fins in the paddlefish, Polyodon spathula

In Polyodon spathula, the pectoral fin radials, with the exception of the metapterygium, are derived from the decomposition of a single continuous cartilage fin plate that is continuous with the scapulocoracoid. This cartilage sheet develops two interior splits to form three precursor pieces, and these decompose in a predictable way to generate the propterygium and radials. The metapterygium is an extension of the scapulocoracoid that segments off of it during early development. To our knowledge, this has not been reported for acipenserids or other basal actinopterygians. In teleosts, the proximal radials also develop from the "break up" of an initially continuous paddle-like sheet of cartilage along the posterior edge of the scapulocoracoid, and in Polypterus and sharks a similar pattern holds. Thus, the pattern observed in Polyodon may represent the basal developmental condition for the gnathostome pectoral fin. The process underlying development of the superficially similar cartilages of the pelvic and pectoral fins is different. In the pectoral fin, the metapterygium is segmented off of the scapulocoracoid and other radials form from the decomposition of the cartilage plate. In contrast, individual rod-like basipterygial elements form in a close one-to-one correspondence with the middle radials of the pelvic fin, but later fuse to form an anterior element that is branched in appearance. To evaluate further claims of similarity among the pectoral and pelvic fin elements of various fishes, the course of the development of these structures must be observed. The pectoral fin and girdle in Polyodon ossifies in a different sequence than that proposed as ancestral (and highly conserved) for actinopterygians: the supracleithrum ossifies significantly before the cleithrum. The later ossification of the cleithrum in Polyodon may be related to the primary use of the caudal fin vs. the pectoral fins in their locomotion.     

Anatomy and early development of the pectoral girdle,fin, and fin spine of sturgeons (Actinopterygii: Acipenseridae)

Acipenseriformes hold an important place in the evolutionary history of bony fishes. Given their phylogenetic position as extant basal Actinopterygii, it is generally held that a thorough understanding of their morphology will greatly contribute to the knowledge of the evolutionary history and the origin of diversity for the major osteichthyan clades. To this end, we examined comparative developmental series from the pectoral girdle in Acipenser fulvescens, A. medirostris, A. transmontanus, and Scaphirhynchus albus to document, describe, and compare ontogenetic and allometric differences in the pectoral girdle. We find, not surprisingly, broad congruence between taxa in the basic pattern of development of the dermal and chondral elements of the pectoral girdle. However, we also find clear differences in the details of structure and development among the species examined in the dermal elements, including the clavicle, cleithrum, supracleithrum, posttemporal, and pectoral‐fin spine. We also find differences in the internal fin elements such as the distal radials as well as in the number of fin rays and their association with the propterygium. Further, there are clear ontogenetic differences during development of the dermal and chondral elements in these species and allometric variation in the pectoral‐fin spine. The characters highlighted provide a suite of elements for further examination in studies of the phylogeny of sturgeons. Determining the distribution of these characters in other sturgeons may aid in further resolution of phylogenetic relationships, and these data highlight the role that ontogenetic and comparative developmental studies provide in systematics. J. Morphol. 276:241–260, 2015. © 2014 Wiley Periodicals, Inc.     

Study of the pectoral girdle and fins of the Late Carboniferous sibyrhynchid iniopterygians (Vertebrata,Chondrichthyes, Iniopterygia) from Kansas and Oklahoma (USA) by means of microtomography,with comments on iniopterygian relationships

The latest works on iniopterygians question their monophyly when considering only the neurocranium of the two families (Sibyrhynchidae and Iniopterygidae), which have different conditions of preservation. Some of the synapomorphies of the Iniopterygia concern the pectoral girdle and fins. However, the anatomy of these different elements is still poorly known in this taxon. Here we describe in details three dimensionally preserved cartilages of the pectoral girdle and fins of the sibyrhynchid Iniopera sp. These structures have been extracted virtually from phosphatised nodules thanks to conventional and synchrotron microtomography, using absorption and phase contrast based techniques in the later case. The pectoral girdle of Iniopera sp. consists of three elements, which are, from dorsal to ventral, a paired suprascapular cartilage, a pair of robust scapulocoracoids and an unpaired intercoracoid cartilage. The scapular part of the scapulocoracoids is extremely reduced and the suprascapular cartilages link the scapulcoracoids to the rear of the neurocranium. These characters may be iniopterygian synapomorphies. Iniopterygians, stem and crown-holocephalans share a basipterygium that articulates with the pectoral girdle and bears an enlarged first pectoral fin radial. Posteriorly, the basipterygium articulates with either a well-defined metapterygium (in crown-holocephalans) or a metapterygial axis (in stem-holocephalans).     

Fossil fishes from china provide first evidence of dermal pelvic girdles in osteichthyans

Background

The pectoral and pelvic girdles support paired fins and limbs, and have transformed significantly in the diversification of gnathostomes or jawed vertebrates (including osteichthyans, chondrichthyans, acanthodians and placoderms). For instance, changes in the pectoral and pelvic girdles accompanied the transition of fins to limbs as some osteichthyans (a clade that contains the vast majority of vertebrates – bony fishes and tetrapods) ventured from aquatic to terrestrial environments. The fossil record shows that the pectoral girdles of early osteichthyans (e.g., Lophosteus, Andreolepis, Psarolepis and Guiyu) retained part of the primitive gnathostome pectoral girdle condition with spines and/or other dermal components. However, very little is known about the condition of the pelvic girdle in the earliest osteichthyans. Living osteichthyans, like chondrichthyans (cartilaginous fishes), have exclusively endoskeletal pelvic girdles, while dermal pelvic girdle components (plates and/or spines) have so far been found only in some extinct placoderms and acanthodians. Consequently, whether the pectoral and pelvic girdles are primitively similar in osteichthyans cannot be adequately evaluated, and phylogeny-based inferences regarding the primitive pelvic girdle condition in osteichthyans cannot be tested against available fossil evidence.

Methodology/Principal Findings

Here we report the first discovery of spine-bearing dermal pelvic girdles in early osteichthyans, based on a new articulated specimen of Guiyu oneiros from the Late Ludlow (Silurian) Kuanti Formation, Yunnan, as well as a re-examination of the previously described holotype. We also describe disarticulated pelvic girdles of Psarolepis romeri from the Lochkovian (Early Devonian) Xitun Formation, Yunnan, which resemble the previously reported pectoral girdles in having integrated dermal and endoskeletal components with polybasal fin articulation.

Conclusions/Significance

The new findings reveal hitherto unknown similarity in pectoral and pelvic girdles among early osteichthyans, and provide critical information for studying the evolution of pelvic girdles in osteichthyans and other gnathostomes.     

Development of the pectoral, pelvic, dorsal and anal fins in cultured sea bream

The pectoral fin girdle was the first element of the fins to develop in Sparus aurata. By 3·1mm L _N (notochord length) the cleithrum was ossified and the cartilaginous caracoid-scapula was present. The fin was fully developed at 11·6 mm L _S (standard length) and by 16·0 mm L _S most elements of the fin were ossified. The pelvic fins were the last pair to develop and rudiments of these were first detected at 7·9 mm L _S. The pelvic fin and girdle were completely formed and ossified at 16·0 mm L _S. The development of dorsal and anal fins began at c. 6·5–7·0 mm L _S with the formation of 10 cartilaginous dorsal proximal radials and eight cartilaginous ventral proximal radials. The three cartilaginous predorsals (supraneurals) appeared at 7·7 mm L _S and the ossification of dorsal and anal proximal and distal radials began, respectively, at 10·5 mm L _S and 11·3 mm L _S. Ossified structures in the fins were also classified according to their origin, as being either dermal or endochondral. Finally the chronology of appearance of fin structures in S. aurata was compared with that reported for other Sparidae, Engraulidae and Haemulidae.     

Fin duplications and deletions induced by disruption of retinoic acid signaling

 Retinoic acid (RA), a derivative of vitamin A, plays a critical role as a signaling molecule in axial patterning of vertebrates. Here we report that RA exposure of zebrafish (Danio rerio) and mummichog (Fundulus heteroclitus) embryos during gastrulation results in homeotic duplications of the pectoral fins in up to 94% of fish. We have observed three to four pairs of fins in an individual fish. Although some duplications are partial, many represent complete axial duplications of the pectoral girdle and fin and include coracoscapulae, proximal radials, and dermal fin elements. Fin duplications are observed only at a defined dose of RA. Inhibition of RA synthesis by exposure to citral during a narrow developmental window leads to fish which lack pectoral fins but can be rescued by addition of exogenous RA, suggesting that RA signaling is critical to fin specification during early development. The ability to consistently induce multiple fins in a large number of vertebrate embryos should contribute to the understanding of genetic regulation of the normal positioning of limbs during embryogenesis. Received: 30 August 1997 / Accepted: 6 December 1997     

The morphology of the cephalic lobes and anterior pectoral fins in six species of batoids

Many benthic batoids utilize their pectoral fins for both undulatory locomotion and feeding. Certain derived, pelagic species of batoids possess cephalic lobes, which evolved from the anterior pectoral fins. These species utilize the pectoral fins for oscillatory locomotion while the cephalic lobes are used for feeding. The goal of this article was to compare the morphology of the cephalic lobes and anterior pectoral fins in species that possess and lack cephalic lobes. The skeletal elements (radials) of the cephalic lobes more closely resembled the radials in the pectoral fin of undulatory species. Second moment of area (I), calculated from cephalic lobe radial cross sections, and the number of joints revealed greater flexibility and resistance to bending in multiple directions as compared to pectoral fin radials of oscillatory species. The cephalic lobe musculature was more complex than the anterior pectoral fin musculature, with an additional muscle on the dorsal side, with fiber angles running obliquely to the radials. In Rhinoptera bonasus, a muscle presumably used to help elevate the cephalic lobes is described. Electrosensory pores were found on the cephalic lobes (except Mobula japonica) and anterior pectoral fins of undulatory swimmers, but absent from the anterior pectoral fins of oscillatory swimmers. Pore distributions were fairly uniform except in R. bonasus, which had higher pore numbers at the edges of the cephalic lobes. Overall, the cephalic lobes are unique in their anatomy but are more similar to the anterior pectoral fins of undulatory swimmers, having more flexibility and maneuverability compared to pectoral fins of oscillatory swimmers. The maneuverable cephalic lobes taking on the role of feeding may have allowed the switch to oscillatory locomotion and hence, a more pelagic lifestyle. J. Morphol. 274:1070–1083, 2013. © 2013 Wiley Periodicals, Inc.     

Fish fingers: digit homologues in sarcopterygian fish fins

A defining feature of tetrapod evolutionary origins is the transition from fish fins to tetrapod limbs. A major change during this transition is the appearance of the autopod (hands, feet), which comprises two distinct regions, the wrist/ankle and the digits. When the autopod first appeared in Late Devonian fossil tetrapods, it was incomplete: digits evolved before the full complement of wrist/ankle bones. Early tetrapod wrists/ankles, including those with a full complement of bones, also show a sharp pattern discontinuity between proximal elements and distal elements. This suggests the presence of a discontinuity in the proximal-distal sequence of development. Such a discontinuity occurs in living urodeles, where digits form before completion of the wrist/ankle, implying developmental independence of the digits from wrist/ankle elements. We have observed comparable independent development of pectoral fin radials in the lungfish Neoceratodus (Osteichthyes: Sarcopterygii), relative to homologues of the tetrapod limb and proximal wrist elements in the main fin axis. Moreover, in the Neoceratodus fin, expression of Hoxd13 closely matches late expression patterns observed in the tetrapod autopod. This evidence suggests that Neoceratodus fin radials and tetrapod digits may be patterned by shared mechanisms distinct from those patterning the proximal fin/limb elements, and in that sense are homologous. The presence of independently developing radials in the distal part of the pectoral (and pelvic) fin may be a general feature of the Sarcopterygii.     

Batoid wing skeletal structure: novel morphologies, mechanical implications, and phylogenetic patterns

The skeleton of the "wings" of skates and rays consists of a series of radially oriented cartilaginous fin rays emanating from a modified pectoral girdle. Each fin ray consists of small, laterally oriented skeletal elements, radials, traditionally represented as simple cylindrical building blocks. High-resolution radiography reveals the pattern of calcification in batoid wing elements, and their organization within the fin ray, to be considerably more complex and phylogenetically variable than previously thought. Calcification patterns of radials varied between families, as well as within individual pectoral fins. Oscillatory swimmers show structural interconnections between fin rays in central areas of the wing. Morphological variation was strongly predictive of locomotor strategy, which we attribute to oscillatory swimmers needing different areas of the wing stiffened than do undulatory swimmers. Contributions of various forms of calcification to radial stiffness were calculated theoretically. Results indicate that radials completely covered by mineralized tissue ("crustal calcification") were stiffer than those that were calcified in chain-like patterns ("catenated calcification"). Mapping this functionally important variation onto a phylogeny reveals a more complicated pattern than the literature suggests for the evolution of locomotor mode. Therefore, further investigation into the phylogenetic distribution of swimming mode is warranted.     

The origins of adipose fins: an analysis of homoplasy and the serial homology of vertebrate appendages

Adipose fins are appendages found on the dorsal midline between the dorsal and caudal fins in more than 6000 living species of teleost fishes. It has been consistently argued that adipose fins evolved once and have been lost repeatedly across teleosts owing to limited function. Here, we demonstrate that adipose fins originated repeatedly by using phylogenetic and anatomical evidence. This suggests that adipose fins are adaptive, although their function remains undetermined. To test for generalities in the evolution of form in de novo vertebrate fins, we studied the skeletal anatomy of adipose fins across 620 species belonging to 186 genera and 55 families. Adipose fins have repeatedly evolved endoskeletal plates, anterior dermal spines and fin rays. The repeated evolution of fin rays in adipose fins suggests that these fins can evolve new tissue types and increased structural complexity by expressing fin-associated developmental modules in these new territories. Patterns of skeletal elaboration differ between the various occurrences of adipose fins and challenge prevailing hypotheses for vertebrate fin origin. Adipose fins represent a powerful and, thus far, barely studied model for exploring the evolution of vertebrate limbs and the roles of adaptation and generative biases in morphological evolution.     

Necessity of an adequate nerve supply for regeneration of the amputated pectoral fin in the teleost Fundulus

The present work deals with determination of the threshold of nerve fibers per unit of amputation surface necessary for regeneration of the pectoral fins of a teleost, Fundulus. Partial denervation of the amputated pectoral fins, i.e., resection of one or two of the three nerves of the brachial (=pectoral) plexus revealed that the presence of a single one allows the amputated fin to regenerate. From these data and others obtained previously, it is concluded that the nervous requirements for a teleost fin to regenerate are similar or slightly lower than those for tetrapods, for example in the newt, which are capable of appendage regeneration.     

Stem sarcopterygians have primitive polybasal fin articulation

Among osteichthyans, basal actinopterygian fishes (e.g. paddlefish and bowfins) have paired fins with three endoskeletal components (pro-, meso- and metapterygia) articulating with polybasal shoulder girdles, while sarcopterygian fishes (lungfish, coelacanths and relatives) have paired fins with one endoskeletal component (metapterygium) articulating with monobasal shoulder girdles. In the fin–limb transition, the origin of the sarcopterygian paired fins triggered new possibilities of fin articulation and movement, and established the proximal segments (stylopod and zeugopod) of the presumptive tetrapod limb. Several authors have stated that the monobasal paired fins in sarcopterygians evolved from a primitive polybasal condition. However, the fossil record has been silent on whether and when the inferred transition took place. Here we describe three-dimensionally preserved shoulder girdles of two stem sarcopterygians (Psarolepis and Achoania) from the Lower Devonian of Yunnan, which demonstrate that stem sarcopterygians have polybasal pectoral fin articulation as in basal actinopterygians. This finding provides a phylogenetic and temporal constraint for studying the origin of the stylopod, which must have originated within the stem sarcopterygian lineage through the loss of the propterygium and mesopterygium.     

On the osteology and myology of catfish pectoral girdle, with a reflection on catfish (Teleostei: Siluriformes) plesiomorphies

The configuration of the pectoral girdle bones and muscles of numerous catfishes was studied in detail and compared with that of other siluriforms, as well as of other teleosts, described in the literature. The pectoral girdle of catfishes is composed of only three bones, which probably correspond to the posttemporo-supracleithrum (posttemporal + supracleithrum), scapulo-coracoid (scapula + coracoid), and cleithrum of other teleosts. These latter two bones constitute the place of origin of the pectoral girdle muscles. Two of these muscles are related to the movements of the pectoral fin. These two muscles correspond, very likely, to the abductor superficialis and to the adductor superficialis of other teleostean fishes. In relation to the pectoral spine (thickened first pectoral fin ray), it is usually moved by three well-developed muscles, which are probably homologous with the arrector ventralis, arrector dorsalis, and abductor profundus of nonsiluriform teleosts. The morphological diversity and the plesiomorphic configuration of these muscles, as well as of the other catfish pectoral girdle structures, are discussed.     

Duplicated Abd-B class genes in medaka hoxAa and hoxAb clusters exhibit differential expression patterns in pectoral fin buds

Hox genes form clusters. Invertebrates and Amphioxus have only one hox cluster, but in vertebrates, they are multiple, i.e., four in the basal teleost fish Polyodon and tetrapods (HoxA, B, C, D), but seven or eight in common teleosts. We earlier completely sequenced the entire hox gene loci in medaka fish, showing a total of 46 hox genes to be encoded in seven clusters (hoxAa, Ab, Ba, Bb, Ca, Da, Db). Among them, hoxAa, hoxAb and hoxDa clusters are presumed to be important for fin-to-limb evolution because of their key role in forelimb and pectoral fin development. In the present study, we compared genome organization and nucleotide sequences of the hoxAa and hoxAb clusters to these of tetrapod HoxA clusters, and found greater similarity in hoxAa case. We then analyzed expression of Abd-B family genes in the clusters. In the trunk, those from the hoxAa cluster, i.e., hoxA9a, hoxA10a, hoxA11a and hoxA13a, were expressed in a manner keeping the colinearity rule of the hox expression as those of tetrapods, while those from the hoxAb cluster, i.e., hoxA9b, hoxA10b, hoxA11b and hoxA13b, were not. In the pectoral fins, the hoxAa cluster was expressed in split domains and did not obey the rule. By contrast, those from the hoxAb and hoxDa clusters were expressed in a manner keeping the rule, i.e., an ancestral pattern similar to those of tetrapods. It is plausible that this differential expression of the two clusters is caused by changes occurred in global control regions after cluster duplications.     

The pectoral anatomy of selected Ostariophysi. I. The Characiniformes.

The muscles and bones of the pectoral fin of Serrasalmus nattereri, the piranha, resemble those of generalized, lower teleosts with specializations related to a body shape adapted for high-speed carnivory; the pectoral fins being highly mobile with strong ligaments to the rays. The presence of two occipital nerves appears primitive, while the emergence of the subclavian artery within the branchial cavity, as in Gasteropelecus sternicla, appears specialized. The muscles and bones of the latter fish, a fresh-water flying fish, are specialized for self-propelled, aerial flight in the fusion of the right and left girdles greatly expanded for insertions of complex appendicular (flight) muscles, and in the consolidation of the rays and radials into one functional unit moving vertically in flight through contraction of vertical, massive ventral flight muscles. The bony pectoral anatomy of Electrophorus electricus, the electric eel, is specialized in having a mobile joint between the primary girdle and the cleithrum, the former being suspended vertically from the cleithrum by ligaments. The proximal radials and rays are very numerous and vertically aligned. The cleithrum is shaped to accommodate the extensive sternohyoid and pharyngocleithral muscles. The sheet-like appendicular muscles extend beyond the special joint and control its movement. The deeper muscles do not cross this joint. The arterial system is specialized in lacking a deep brachial artery.     

Development of zebrafish (Danio rerio) pectoral fin musculature

During posthatching development the fins of fishes undergo striking changes in both structure and function. In this article we examine the development of the pectoral fins from larval through adult life history stages in the zebrafish (Danio rerio), describing in detail their pectoral muscle morphology. We explore the development of muscle structure as a way to interpret the fins' role in locomotion. Genetic approaches in the zebrafish model are providing new tools for examining fin development and we take advantage of transgenic lines in which fluorescent protein is expressed in specific tissues to perform detailed three-dimensional, in vivo fin imaging. The fin musculature of larval zebrafish is organized into two thin sheets of fibers, an abductor and adductor, one on each side of an endoskeletal disk. Through the juvenile stage the number of muscle fibers increases and muscle sheets cleave into distinct muscle subdivisions as fibers orient to the developing fin skeleton. By the end of the juvenile period the pectoral girdle and fin muscles have reoriented to take on the adult organization. We find that this change in morphology is associated with a switch of fin function from activity during axial locomotion in larvae to use in swim initiation and maneuvering in adults. The examination of pectoral fins of the zebrafish highlights the yet to be explored diversity of fin structure and function in subadult developmental stages. J. Morphol. (c) 2005 Wiley-Liss, Inc.     

Revealing the mechanisms of the rostral shift of pelvic fins among teleost fishes

In teleost fishes, the position of the pelvic fins shift during evolution; this positional shift seems to have diversified their locomotion and feeding behavior, thereby expanding the habitats of these fishes. Thus, such a positional shift of the pelvic fins is one of the significant features of teleost fishes from evolutionary, embryological, and taxonomic viewpoints, but no studies to date have investigated the mechanism for the rostral shift of the pelvic fins from the anal region in teleosts. Examining the fate of the prospective pelvic fin cells of the zebrafish Danio rerio and the Nile tilapia Oreochromis niloticus embryos demonstrates that the prospective pelvic fin cells are originally located near the anus, as seen in tetrapods, but their position shifts with respect to the body trunk after its protrusion from the yolk surface. In this article, we highlight such recent findings and discuss the mechanisms of pelvic fin evolution among teleost fishes.     

The distribution of fibronectin in developing zebrafish (Danio rerio) cartilage

The extracellular matrix (ECM) plays a complex and vital role throughout the process of cartilage formation. Fibronectin is a large ECM glycoprotein with an important role in various developmental processes, including skeletogenesis. Taking advantage of the known sequence of cartilage development in zebrafish and using an immunohistochemical stain for collagen type II to identify differentiation phase cartilage, we evaluate the distribution of fibronectin in various cartilaginous elements of the zebrafish (elements of the splanchnocranium, and of the dorsal, caudal, pelvic and pectoral fins). Contrary to what is observed in tetrapods, our data on zebrafish indicate the apparent lack of fibronectin during the condensation phase of cartilage development. This lack is possibly linked to the high developmental rate of the zebrafish and the small size of the condensations, which brings different needs for the extracellular environment to ensure cell survival. Furthermore, the fin disk cartilage of the pectoral fin develops an ECM with a strong fibronectin signal, whereas other cartilage elements show only a weak fibronectin signal in early differentiation, which gradually disappears. Thus, the pectoral fin disk cartilage is unique not only because of its specific way of development (subdivision of a continuous plate into four elements, the proximal radials), but also because of its strong fibronectin‐positive ECM.     

Expression of Hoxa-11 and Hoxa-13 in the pectoral fin of a basal ray-finned fish, Polyodon spathula: implications for the origin of tetrapod limbs

Summary Paleontological and anatomical evidence suggests that the autopodium (hand or foot) is a novel feature that distinguishes limbs from fins, while the upper and lower limb (stylopod and zeugopod) are homologous to parts of the sarcopterygian paired fins. In tetrapod limb development Hoxa-11 plays a key role in differentiating the lower limb and Hoxa-13 plays a key role in differentiating the autopodium. It is thus important to determine the ancestral functions of these genes in order to understand the developmental genetic changes that led to the origin of the tetrapod autopodium. In particular it is important to understand which features of gene expression are derived in tetrapods and which are ancestral in bony fishes. To address these questions we cloned and sequenced the Hoxa-11 and Hoxa-13 genes from the North American paddlefish, Polyodon spathula, a basal ray-finned fish that has a pectoral fin morphology resembling that of primitive bony fishes ancestral to the tetrapod lineage. Sequence analysis of these genes shows that they are not orthologous to the duplicated zebrafish and fugu genes. This implies that the paddlefish has not duplicated its HoxA cluster, unlike zebrafish and fugu. The expression of Hoxa-11 and Hoxa-13 in the pectoral fins shows two main phases: an early phase in which Hoxa-11 is expressed proximally and Hoxa-13 is expressed distally, and a later phase in which Hoxa-11 and Hoxa-13 broadly overlap in the distal mesenchyme of the fin bud but are absent in the proximal fin bud. Hence the distal polarity of Hoxa-13 expression seen in tetrapods is likely to be an ancestral feature of paired appendage development. The main difference in HoxA gene expression between fin and limb development is that in tetrapods (with the exception of newts) Hoxa-11 expression is suppressed by Hoxa-13 in the distal limb bud mesenchyme. There is, however, a short period of limb bud development where Hoxa-11 and Hoxa-13 overlap similarly to the late expression seen in zebrafish and paddlefish. We conclude that the early expression pattern in tetrapods is similar to that seen in late fin development and that the local exclusion by Hoxa-13 of Hoxa-11 from the distal limb bud is a derived feature of limb developmental regulation.