Hybridization between pallid sturgeon Scaphirhynchus albus and shovelnose sturgeon Scaphirhynchus platorynchus

This study found that introgressive hybridization of the pallid sturgeon Scaphirhynchus albus with the common shovelnose sturgeon Scaphirhynchus platorynchus has probably occurred across the range of S. albus. Bayesian clustering found evidence of hybridization in all management units of S. albus. Some individuals were intermediate at both genetic and morphological characters, and some had discordant results. The results support introgressive hybridization throughout much of the range of S. albus, yet individuals consistent with being pure members of each species were detected in all management units. Simulations demonstrated that it would be very difficult to distinguish introgressed individuals from pure specimens after multiple generations of backcrossing with these microsatellite markers. Using hybrid or backcross fish as broodstock could artificially accelerate the loss of unique genetic variation in S. albus. Additional microsatellite loci or additional genetic markers, along with morphological data may be required to ensure that hybrid or backcross fish are not used. Introgressive hybridization requires at least two generations and generation lengths of S. albus are long, perhaps as long as 30 years. The proportion of individuals consistent with introgressive hybrid origins indicates that hybridization between S. albus and S. platorynchus probably has occurred for several generations and is not a recent phenomenon.     

Life history and status of the shovelnose sturgeon, Scaphirhynchus platorynchus

The shovelnose sturgeon, Scaphirhynchus platorynchus, is a freshwater sturgeon of the Mississippi and Missouri rivers and their tributaries. It is one of the smaller North American sturgeons, seldom weighing more than 2.5 kg over most of its range except in the upper Missouri River, where individuals of over 7 kg have been found. Spawning occurs in spring at temperatures between 17 and 21 °C over rock or gravel substrate downstream from dams, near rock structures, or in tributaries. most males reach sexual maturity at 5 years, most females at 7 years. Adults do not spawn every year. Shovelnose sturgeon are found in large, turbid rivers and frequently concentrate in areas downstream from dams or at the mouths of tributaries. Population densities range up to 2500 fish per km. They are commonly found in areas of current over sandy bottoms or near rocky points or bars, where they feed primarily on aquatic invertebrates. The shovelnose sturgeon is classified as a sport species in 12 of 24 states where it occurs. Commercial harvest is allowed in seven states, where fresh shovelnose sturgeon sell for 55 to 88 cents per kg, smoked shovelnose for about $5.75 per kg, and roe from 33 to 110 dollars per kg. About 25 tons of shovelnose sturgeon are harvested commercially each year. Shovelnose sturgeon are considered extirpated in three states, fully protected in four states, and rare, threatened, or of special concern in eight states. Populations are considered stable throughout most of the upper Mississippi, lower Missouri, Red, and Atchafalaya rivers. Three states, Wyoming, West Virginia, and New Mexico, have developed plans to reintroduce the species into rivers where it has been extirpated.     

Bycatch of the endangered pallid sturgeon (Scaphirhynchus albus) in a commercial fishery for shovelnose sturgeon (Scaphirhynchus platorynchus)

We quantified the bycatch of pallid sturgeon Scaphirhynchus albus in Tennessee's shovelnose sturgeon ( Scaphirhynchus platorynchus) fishery by accompanying commercial fishers and monitoring their catch on five dates in spring 2007. Fishers were free to keep or discard any sturgeon they collected in their gillnets and trotlines and we were afforded the opportunity to collect meristic and morphometric data and tissue samples from discarded and harvested specimens. Fishers removed 327 live sturgeon from their gear in our presence, of which 93 were harvested; we also obtained the carcasses of 20 sturgeon that a fisher harvested out of our sight while we were on the water with another fisher. Two of the 113 harvested sturgeon were confirmed pallid sturgeon based on microsatellite DNA analyses. Additionally, fishers gave us five, live pallid sturgeon that they had removed from their gear. If the incidental harvest rate of pallid sturgeon (1.8% of all sturgeon harvested) was similar in the previous two commercial seasons, at least 169 adult pallid sturgeon were harvested by commercial fishers in the Tennessee waters of the Mississippi River in 2005–2007. If fishers altered their behavior because of our presence (i.e. if they were more conservative in what they harvested), the pallid sturgeon take was probably higher when they fished unaccompanied by observers. While retrieving a gill net set the previous day, a fisher we were accompanying retrieved a gillnet lost 2 days earlier; this ghost net caught 53 sturgeon whereby one fish was harvested but most fish were dead, including one confirmed pallid sturgeon.     

Current status and trends in shovelnose sturgeon (Scaphirhynchus platorynchus) management and conservation

The objective of this study was to update information regarding the status of shovelnose sturgeon fisheries. Although a substantial amount of shovelnose sturgeon research has been conducted in the past decade, the study purpose was not to provide a comprehensive review of the literature; the primary interest was in the status, trends, and management of Scaphirhynchus platorynchus fisheries in North America. Biologists were surveyed in all 24 states within the native distribution of the species; results indicate that commercial harvest is currently permitted in eight states, recreational harvest is allowed in 13 states, and that regulations vary within rivers and jurisdictional boundaries. Although recreational exploitation of shovelnose sturgeon is thought to be low and not a significant threat to populations, commercial harvest is a major concern in states with a commercial fishery. In the last decade harvest has increased in all states with commercial shovelnose sturgeon fisheries, but recent implementation of regulations has decreased harvest in some states. Approximately half of the states with extant shovelnose sturgeon populations conduct routine monitoring of the species, and the understanding of shovelnose sturgeon populations is increasing.     

Heteroplasmy in the mtDNA control region of sturgeon (Acipenser, Huso and Scaphirhynchus)

Data from 1238 fishes from 19 sturgeon species and 1 paddlefish were used to analyze heteroplasmy in sturgeon. Lengths of central repeat units ranged from 74 to 83 bp among sturgeon species. No repeat sequence was found in the paddlefish, Polyodon spathula. A general feature of the repeat units was the presence of termination associated sequence (TAS) motifs. About 50% of 138 interspecific mutations observed among the D-loop sequences are located 10 bp down- and upstream from these TAS motifs. Interestingly, most homoplasmic species showed deletions upstream to the TAS motifs, whereas deletions downstream to the TAS motifs observed in two species do not seem to preclude heteroplasmy. Calculations of secondary structures and thermal stabilities of repeat units showed DeltaG values for all heteroplasmic species to be <-8 and for most homoplasmic species DeltaG value to be >-8. Most heteroplasmic fishes had two and/or three repeat units. No homoplasmic sturgeon with >2 repeat units were observed. Molecular phylogeny based on the entire cytochrome b showed that heteroplasmy probably resulted from a single evolutionary event. Our data demonstrate that heteroplasmy is present in most sturgeon species and suggest that the thermal stability of the secondary structure of the repeat unit in combination with mutations downstream of the TAS sequences influences heteroplasmy.     

Reproductive traits of shovelnose sturgeon Scaphirhynchus platorynchus (Rafinesque, 1820) in the lower Platte River,Nebraska

We assessed reproductive status, fecundity, egg size, and spawning dynamics of shovelnose sturgeon Scaphirhynchus platorynchus in the lower Platte River. Shovelnose sturgeon were captured throughout each year during 2011 and 2012 using a multi‐gear approach designed to collect a variety of fish of varying sizes and ages. Fish were collected monthly for a laboratory assessment of reproductive condition. Female shovelnose sturgeon reached fork length at 50% maturity (FL₅₀) at 547 mm and at a minimum length of 449 mm. The average female spawning cycle was 3–5 years. Mean egg count for adult females was 16 098 ± 1103 (SE), and mean egg size was 2.401 ± 0.051 (SE) mm. Total fecundity was positively correlated with length (r² = 0.728; P < 0.001), mass (r² = 0.896; P < 0.001), and age (r² = 0.396; P = 0.029). However, fish size and age did not correlate to egg size (P > 0.05). Male shovelnose sturgeon reached FL₅₀ at 579 mm and at a minimum length of 453 mm. The average male spawning cycle was 1–2 years. Reproductively viable male and female sturgeon occurred during the spring (March–May) and autumn (September–October) in both years, indicating spring and potential autumn spawning events. Shovelnose sturgeon in the lower Platte River are maturing at a shorter length and younger age compared to populations elsewhere. Although it is unknown if the change is plastic or evolutionary, unfavorable environmental conditions or over‐harvest may lead to hastened declines compared to other systems.     

A guide to the embryonic development of the shovelnose sturgeon (Scaphirhynchus platorynchus), reared at a constant temperature

Described is the sequence and timing of embryologic development of shovelnose sturgeon, (Scaphirhynchus platorynchus) reared at a constant temperature 20 ± 0.5°C. Artificially spawned, fertilized eggs were held in a recirculating system. Embryos were sampled hourly during the first 48 h of development and every 3 h thereafter. Embryos were viewed and imaged at 35× magnification. The first cleavage furrow appeared 2 h post‐fertilization; early (synchronous) cleavage was completed after 7 h. Blastulation concluded at 16 h when the dorsal blastopore lip formed. The slitlike blastopore appeared at 29 h, signifying the completion of gastrulation. At 33 h, the rudiments of the excretory system emerged, followed by closing of the neural tube at 36 h and formation of the s‐shaped heart at 60 h. The body continued to elongate with mass hatch occurring at 102 h. After hatch, larvae swam into the water column and drifted in the flow for approximately 2 days after which the larvae became positively rheotaxic. After expulsion of the pigment plug, the larvae began exogenous feeding and other structures continued to develop. Metamorphosis was completed after 26 days of development. Because the shovelnose sturgeon possesses developmental patterns similar to those of other sturgeon, we can use this species as a model for the closely related pallid and Alabama sturgeon.     

The fishery,biology, and management of Atlantic sturgeon,Acipenser oxyrhynchus,in North America

Synopsis The Atlantic sturgeon supported major fisheries along the entire Atlantic coast of North America. These fisheries peaked about 1890 and then suffered almost total collapse by 1905. The Atlantic sturgeon is anadromous and highly susceptible to capture during spawning migrations. Further, this species biological characteristics makes it very vulnerable to man-induced changes in natural habitat and slow to recover. Atlantic sturgeon mature at an advanced age (7–27 year for females, depending on latitude), exhibit a long interspawning period (2–5 year), and require suitable riverine, estuarine, and coastal environments for successful completion of their life cycle. Today, only remnant stocks exist in areas of former abundance. Management regulations vary considerably from state to state and range from full protection to no protection. Biological data are needed to: identify and characterize specific spawning and nursery areas; delineate migratory patterns and recruitment to various stocks; establish stock abundance; and, assess effects of various management strategies. In order to protect remaining stocks, the imposition of a total harvesting moratorium is recommended.     

Effects of norfloxacin on the drug metabolism enzymes of two sturgeon species (Acipenser schrencki and Acipenser ruthenus)

The enzyme activities of 7‐ethoxycoumarin O‐deethylase (ECOD) and glutathione S‐transferase (GST), and on glutathione (GSH) content, involved in metabolism of the antibiotic Norfloxacin (NFLX), were investigated in Acipenser schrencki and Acipenser ruthenus. Sturgeons weighing 45–55 g were kept in an aquarium (0.5 m × 0.5 m × 0.9 m) for two weeks under controlled conditions (fish density 88 individuals per m³, 18°C) before the experiment. The two species of sturgeon were divided into five groups each (n = 15 in each group), with each group subdivided into three replicates of five fish per tank. A control group in which distilled water was administered orally was also tested. NFLX was forced into the stomachs of the fish at a concentration of 20, 40, 60, 80 and 100 mg kg^?1 body weight, respectively. ECOD activities in liver microsomes, and GST activity and GSH content in liver microsomes and blood plasma, were measured and compared. Results indicate that ECOD activity is progressively inhibited with increasing NFLX concentrations. ECOD activity varied from 0.12 nmol mg^?1 min^?1 to 0.07 nmol mg^?1 min^?1, demonstrating an inhibition rate of 60.83% in A. schrencki and 65.14% in A. ruthenus. In both species tested, GST and GSH levels exhibited a trend of first increasing, and then decreasing with increasing NFLX levels, reaching a peak value at 40 mg per kg^?1 body weight. Thus, the presented results indicate that NFLX can induce a change in the activity of some drug metabolism related enzymes such as ECOD and GST in vivo.     

Effects of temperature and hydrology on growth of shovelnose sturgeon Scaphirhynchus platorynchus (Rafinesque, 1820) in the lower Mississippi River

We evaluated the effects of thermal and hydrologic conditions on the growth of shovelnose sturgeon (Scaphirhynchus platorynchus) in the lower Mississippi River, USA. Duration of water temperatures 12–24°C had a positive influence and temperatures below 11°C had a negative influence on annual growth increment, but these two variables accounted for less than 15% of the variation in growth. Duration of water temperatures above 28°C, duration of floodplain inundation, duration of low water, and minimum and maximum river stage did not influence annual growth increment. Growth of shovelnose sturgeon in the lower Mississippi River appears to be positively influenced by duration of moderate water temperatures but minimally influenced by hydrologic conditions. The low variation accounted for by thermal and hydrologic variables suggests annual growth increment may be largely influenced by additional abiotic or biotic factors.     

A laboratory examination of substrate, water depth, and light use at two water velocity levels by individual juvenile pallid (Scaphirhynchus albus) and shovelnose (Scaphirhynchus platorynchus) sturgeon

We investigated the influence of substrate type, water depth, light, and relative water velocity on microhabitat selection in juvenile pallid (Scaphirhynchus albus) and shovelnose (Scaphirhynchus platorynchus) sturgeon. Individual sturgeon were placed in an 18 927 L elliptical flume, and their location was recorded after a 2‐h period. Data were analyzed using exact chi‐square goodness of fit tests and exact tests of independence. Both sturgeon species used substrate, depth, and light in similar proportions. (all comparisons; P > 0.05). Specifically, pallid and shovelnose sturgeon did not use substrate in proportion to its availability (pallid: P = 0.0026; shovelnose: P = 0.0199). Each species used sand substrate more and gravel substrate less than expected based on availability. Additionally, neither species used woody structure. Both species used deep areas in greater proportion than availability while shallow areas were used less than expected based on availability (pallid; P < 0.0001; shovelnose; P = 0.0335). Pallid and shovelnose sturgeon used very dark areas in greater proportion than expected based on availability; however, very light areas were used in lower proportion than expected (P < 0.0001). Overall, neither species changed their use of habitat in relation to a change in water velocity (pallid, all comparisons P > 0.05; shovelnose, all comparisons P > 0.05). This study is the first investigation of juvenile pallid and shovelnose sturgeon habitat selection in a large‐scale artificial stream system. Field studies of microhabitat selection by juvenile pallid and shovelnose sturgeon should be carried out to substantiate the results of this study, and to identify critical habitat for recovery and management of sturgeon species. Due to the extensive range, longevity, and migratory behavior of these fishes, proper management likely requires river improvements that provide sturgeon with access to a broad range of habitat conditions over time, including system‐wide habitat diversity; natural variation in flow, velocity, temperature, and turbidity; high water quality; a broad prey base; free‐flowing river sections which provide suitable spawning and rearing sites, as well as protection from recreational and commercial harvesting.     

Genome duplication events and functional reduction of ploidy levels in sturgeon (Acipenser, Huso and Scaphirhynchus).

Sturgeon (order Acipenserformes) provide an ideal taxonomic context for examination of genome duplication events. Multiple levels of ploidy exist among these fish. In a novel microsatellite approach, data from 962 fish from 20 sturgeon species were used for analysis of ploidy in sturgeon. Allele numbers in a sample of individuals were assessed at six microsatellite loci. Species with approximately 120 chromosomes are classified as functional diploid species, species with approximately 250 chromosomes as functional tetraploid species, and with approximately 500 chromosomes as functional octaploids. A molecular phylogeny of the sturgeon was determined on the basis of sequences of the entire mitochondrial cytochrome b gene. By mapping the estimated levels of ploidy on this proposed phylogeny we demonstrate that (I) polyploidization events independently occurred in the acipenseriform radiation; (II) the process of functional genome reduction is nearly finished in species with approximately 120 chromosomes and more active in species with approximately 250 chromosomes and approximately 500 chromosomes; and (III) species with approximately 250 and approximately 500 chromosomes arose more recently than those with approximately 120 chromosomes. These results suggest that gene silencing, chromosomal rearrangements, and transposition events played an important role in the acipenseriform genome formation. Furthermore, this phylogeny is broadly consistent with previous hypotheses but reveals a highly supported oceanic (Atlantic-Pacific) subdivision within the Acipenser/Huso complex.     

Realization of the project for Acipenser oxyrhynchus oxyrhynchus restoration in the basin of the Neman River

The results of the works on the international project for the Baltic sturgeon restoration in the basin of the Neman River are considered. The mutual goal of the project is the restoration of the Baltic sturgeon population in its whole former area in the Baltic Sea. Successful works on the restoration of the Baltic sturgeon population are conducted in the Oder and the Vistula basins. The stockings of the Neman River with the sturgeon fry were started from 2012 year. It is contemplated that the sturgeon stockings of the main rivers of the Baltic Sea will enable to restore local sturgeon populations in these river basins. Till the beginning of the XXI century it was supposed that the Atlantic sturgeon Acipenser sturio had inhabited the Baltic Sea, but numerous genetic analysis of the samples of the sturgeon bones from the archeological finds in the museums showed that the Baltic Sea had been inhabited with Acipenser oxyrhynchus and its population still inhabits several rivers in Canada. This discovery permitted to start the creation of the Baltic sturgeon broodstock. Sturgeon fry from the eggs imported from Canada were used for stocking at the first stage of the project. Later it is contemplated to use the fry from the broodstocks in Germany and Poland. Sturgeon fry hatched from Canadian eggs in Poland and reared in Lithuania was used for the Neman River stocking. The fry were tagged and released in the Neris and the Sventoji Rivers in the Neman River basin. The sturgeon migrated into the Curonian lagoon where the monitoring of the catches was fulfilled. The results of sturgeon catches monitoring are discussed.     

Comparative neuromorphology of the telencephalon of sturgeon of the genera Acipenser, Huso and Scaphirhynchus (Actinopterygii; Acipenseridae)

Brain anatomy and structure, composition of the neurons, and cytoarchitectural maps were compared between five species of Acipenser, one species of Huso and two species of Scaphirhynchus. Morphological differences in the forebrain structure between species and genera were apparent. The evolutionary history of the Acipenseridae has been marked by an extensive mosaic radiation in the evolutionary development of the inner structure of their brains. Members of Scaphirhynchus have a very simple telencephalon structure differing significantly from that of Huso and Acipenser. It is clear that neuromorphological variation of sturgeon species will serve as an additional data useful in understanding the evolutionary processes and history associated with the speciation in Acipenseridae.     

Morphologic comparisons of hatchery-reared specimens of Scaphirhynchus albus, Scaphirhynchus platorynchus, and S. albus × S. platorynchus hybrids (Acipenseriformes: Acipenseridae)

Extensive habitat modifications within the Mississippi and Missouri rivers have presumably interfered with the reproductive isolating mechanisms between the endangered pallid sturgeon, Scaphirhynchus albus, and the sympatric shovelnose sturgeon, Scaphirhynchus platorynchus, causing hybridization between these two species. Several character indices were developed to assist fisheries biologists in identifying specimens of S. albus, S. platorynchus, and their putative hybrids. The indices have numerous assumptions, including that pure strains of both parental species are within the sample analyzed and that hybrids are morphologically intermediate relative to their parents. If these indices have produced inaccurate identifications, then previous work on Scaphirhynchus studies in the Mississippi and Missouri rivers are questionable, including status surveys, captive propagation efforts, or the harvesting of tissues for genetic studies. In this study we tested indices by examining progeny of ‘known’ pallid, shovelnose, and hybrid sturgeon propagated, raised, and preserved at hatcheries. These 60 specimens [78–600 mm standard length (SL)] were propagated with breeding stock from the upper Missouri River drainage, where hybridization between these two species presumably does not occur. Existing indices did not correctly identify small (<250 mm SL) or a combination of small and large (>250 mm SL) sizes of S. albus, S. platorynchus, and their hybrids. Indices worked fairly well in identifying large S. platorynchus, but not in differentiating large S. albus from hybrids. We used principal components analysis (PCA) as an alternative approach to character indices. No a priori knowledge of the identity of the specimen is required with this multivariate technique, which avoids potential circular reasoning present in indices. We were able to completely or almost completely separate both sturgeon species and their hybrids by extracting principal components from a correlation matrix of 13 meristic characters in a standard PCA and extracting size‐corrected principal components from a covariance matrix of 51 morphometric variables using a sheared PCA. Additionally, we demonstrated that first generation hybrids were intermediate with respect to their parental species. Multivariate analyses with a reduced character set of six meristic and 12 morphometric variables also led to accurate and reliable specimen identification. Head spines and numerous qualitative characters are also extremely useful in differentiating between Scaphirhynchus species and their hybrids. In addition to all morphometric characters, some meristic characters and the degree of head spine fusion vary significantly with SL of sturgeons. Recording appropriate data from released specimens, including photovouchers, and making this information available is essential for researchers to have any scientific or legal basis for genetic or any other studies involving these sturgeons.     

Similarities and differences in 13C and 15N stable isotope ratios in two non‐lethal tissue types from shovelnose sturgeon Scaphirhynchus platorynchus (Rafinesque, 1820)

We tested the hypothesis that δ¹³C and δ¹⁵N signatures of pectoral spines would provide measures of δ¹³C and δ¹⁵N similar to those obtained from fin clips for adult shovelnose sturgeon Scaphirhynchus platorynchus. Thirty‐two shovelnose sturgeon (fork length [FL] = 500–724 mm) were sampled from the lower Mississippi River, USA on 23 February 2013. Isotopic relationships between the two tissue types were analyzed using mixed model analysis of covariance. Tissue types differed significantly for both δ¹³C (P < 0.01; spine: mean = ?23.83, SD = 0.62; fin clip: mean = ?25.74, SD = 0.97) and δ¹⁵N (P = 0.01; spine: mean = 17.01, SD = 0.51; fin clip: mean = 17.19, SD = 0.62). Neither FL nor the FL × tissue type interaction had significant (P > 0.05) effects on δ¹³C. Fin clip δ¹³C values were highly variable and weakly correlated (r = 0.16, P = 0.40) with those from pectoral spines. We found a significant FL‐tissue type interaction for δ¹⁵N, reflecting increasing δ¹⁵N with FL for spines and decreasing δ¹⁵N with FL for fin clips. These results indicate that spines are not a substitute for fin clip tissue for measuring δ¹³C and δ¹⁵N for shovelnose sturgeon in the lower Mississippi River, but the two tissues have different turnover rates they may provide complementary information for assessing trophic position at different time scales.     

Sound Production in Sturgeon Scaphirhynchus albus and S. platorynchus (Acipenseridae)

Sound production has been recently discovered in several species of Acipenser. Our work has focused on testing for sound production in species of sturgeon in the genus Scaphirhynchus. We discovered that pallid sturgeon Scaphirhynchus albus and shovelnose sturgeon, S. albus produce sounds during the breeding season. These signals may be used as part of efforts to localize populations of sturgeon in the field, including the Alabama sturgeon, S. suttkusi.     

Afferent projections of the trigeminal nerve in the goldfish,Carassius auratus

The horseradish peroxidase (HRP) histochemical technique was used to examine the peripheral distribution and afferent projections of the trigeminal nerve in the goldfish, Carassius auratus. Sensory fibers of the trigeminal nerve distribute over the head via four branches. The ophthalmic branch distributes fibers to the region above the eye and naris. The maxillary and mandibular branches innervate the regions of the upper and lower lip, respectively. A fourth branch of the trigeminal nerve was demonstrated to be present in the hyomandibular trunk. Upon entering the medulla the trigeminal afferent fibers divide into a rostromedially directed bundle and a caudally directed bundle. The rostromedially directed bundle terminates in the sensory trigeminal nucleus (STN) located within the rostral medulla. The majority of fibers turn caudally, forming the descending trigeminal tract. Fibers of the descending trigeminal tract terminate within three medullary nuclei: the nucleus of the descending trigeminal tract (NDTV), the spinal trigeminal nucleus (Spv), and the medial funicular nucleus (MFn). All projections, except for those to the MFn, are ipsilateral. Contralateral projections were observed at the level of the MFn following the labeling of the ophthalmic and maxillomandibular branches. All branches of the trigeminal nerve project to all four of the trigeminal medullary nuclei. Projections to the STN and MFn were found to be topographically organized such that the afferents of the ophthalmic branch project onto the ventral portion of these nuclei, while the afferents of the maxillo- and hyomandibular branches project to the dorsal portion of these nuclei. Cells of the mesencephalic trigeminal nucleus were retrogradely labeled following HRP application to the ophthalmic, maxillary, and mandibular branches of the trigeminal nerve. In addition to demonstrating the ascending mesencephalic trigeminal root fibers, HRP application to the above-mentioned branches also revealed descending mesencephalic trigeminal fibers. The descending mesencephalic trigeminal fibers course caudally medial to the branchiomeric motor column and terminate in the ventromedial portion of the MFn.