The order Anguilliformes forms a natural group of eel-like species. Moringua edwardsi (Moringuidae) is of special interest because of its peculiar fossorial lifestyle: this species burrows head-first. Externally pronounced morphological specializations for a fossorial lifestyle include: reduced eyes, lack of color, low or absent paired vertical fins, elongated, cylindrical body, reduced head pores of the lateral line system, etc. Many fossorial amphibians, reptiles, and even mammals have evolved similar external specializations related to burrowing. The present study focuses on osteological and myological features of M. edwardsi in order to evaluate the structural modifications that may have evolved as adaptations to burrowing. Convergent evolutionary structures and possible relations with head-first burrowing, miniaturization, feeding habits, etc., were investigated. Body elongation, reduction of the eyes, modified cranial lateral line system, and modified skull shape (pointed though firm) can be considered specializations for head-first burrowing. Hyperossification can probably be regarded more as a specialization to both head-first burrowing and feeding, even though an impact of miniaturization cannot be excluded. Hypertrophied adductor mandibulae muscles and the enlarged coronoid process can be associated with both feeding requirements (it enhances bite forces necessary for their predatory behavior) and with a burrowing lifestyle, as well as miniaturization.

SUMMARYFish swimming is probably one of the most studied and best understood locomotor behaviors in vertebrates. However, many fish also actively exploit sediments. Because of their elongate body shape, anguilliform fishes are not only efficient swimmers but also very maneuverable. Consequently, many species live in complexly structured environments near the bottom and many are known to burrow into the sediment. To better understand burrowing and subsurface locomotion in anguilliform fish we provide descriptive kinematic data on subsurface locomotion in a burrowing eel (Pisodonophis boro) using videofluoroscopy. We also measured the maximal forces that can be exerted by this species during head-first and tail-first burrowing, and explored the implications of head-first burrowing on mechanical stress distribution in the skull. Our data show that P. boro uses lateral undulation to penetrate and move in sandy sediments under water. The kinematics of subsurface locomotion are different from those observed during swimming and are characterized by a very high slip factor. These observations differ considerably from recently published data in terrestrial sand-swimming lizards, and suggest that the sediment behaves like a solid rather than a frictional fluid. Finally, our finite element models show that the cranial shape and structure in the head-first burrowing P. boro is mechanically more suited for head-first burrowing than that of an obligate tail-first burrowing species, Heteroconger hassi.

The existence of two morphotypes, broadheaded and narrowheaded, in European eels Anguilla anguilla is common knowledge among fishermen and eel biologists in Europe. To test whether European eels really are dimorphic in head shape, a total of 277 specimens from two locations in Belgium (Scheldt-Lippenbroek and Lake Weerde), in combination with a larger data set of 725 eels from river systems across Flanders (the northern part of Belgium) were examined. Our biometric data support the hypothesis that a head shape variation in 'Belgian' European eel is best described as having a bimodal distribution. Literature data suggest that this may be the result of phenotypic plasticity related to trophic segregation between morphs.

The remarkable lifestyle of heterocongrines has drawn the attention of many authors in the past, though no or little attention has been paid to the morphology of the tail and the head of these species. In order to examine the true nature of possible morphological specializations of the head and tail and their relation to their tail-first burrowing habit and/or feeding mode, a detailed myological and osteological study of Heteroconger hassi and Heteroconger longissimus was performed. The osteological similarities of the cranial skeleton between H. hassi and H. longissimus are striking. Most of the cranial muscles show no variation in presence, insertion or origin between these two species except for the adductor mandibulae complex, the adductor hyomandibulae and the intermandibularis. The adductor mandibulae complex is small, compared to that of other anguilliform species, and is probably related to their suction-dominated feeding mode and a diet, comprising mainly small, soft prey items. Heterocongrinae have undergone several morphological specializations in the tail for their tail-first burrowing lifestyle. The skeleton and musculature of the tail of H. hassi and H. longissimus are similar. In both species the caudal skeleton is highly reduced and fortified, forming a firm, pointed burrowing tool. Intrinsic caudal musculature is reduced and some muscles (interradials, supracarinalis) are even absent.

The rice paddy eel, Pisodonophis boro (P. boro), is of special interest because of its peculiar burrowing habits. P. boro penetrates the substrate tail-first, a technique common for ophichthids, but it is able to burrow head-first as well. P. boro exhibits three feeding modes: inertial feeding, grasping, and spinning. Rotational feeding is a highly specialized feeding mode, adopted by several elongate, aquatic vertebrates and it is likely that some morphological modifications are related to this feeding mode. The detailed morphology of the head and tail of P. boro is examined with the goal to apportion the anatomical specializations among head-first burrowing, tail-first burrowing, and rotational feeding. The reduced eyes, covered with thick corneas may be beneficial for protection during head-first burrowing, but at the same time decreased visual acuity may have an impact on other sensory systems (e.g. cephalic lateral line system). The elongated and pointed shape of the skull is beneficial for substrate penetration. The cranial bones and their joints, which are fortified, are advantageous for resisting high mechanical loads during head-first burrowing. The aponeurotic connection between epaxial and jaw muscles is considered beneficial for transferring these forces from the body to the head during rotational feeding. Hypertrophied jaw muscles facilitate a powerful bite, which is required to hold prey during spinning movements and variability in the fiber angles of subdivisions of jaw muscles may be beneficial for preventing the lower jaw from being dislodged or opened. Furthermore, firm upper (premaxillo-ethmovomerine complex) and lower jaws (with robust coronoid processes) and high neurocranial rigidity are advantageous for a solid grip to hold prey during rotational feeding. The pointed shape of the tail and the consolidated caudal skeleton are beneficial for their tail-first burrowing habits. It is quite likely that the reduction of the caudal musculature is related to the tail-first burrowing behavior because the subtle movements of the caudal fin rays are no longer required.