Prey-handling Behaviour of Newly Hatched Snakes in Two Species of the Genus Elaphe with Comparison to Adult Behaviour

The prey-handling behaviour of two species of Asian ratsnakes was studied in the laboratory. In experiment 1, effects of prey size and type (mouse, lizard and frog) on capture position, direction of ingestion, condition of prey at ingestion, and prey-handling method were investigated using newly hatched Elaphe quadrivirgata. Prey size did not affect these variables except prey-handling method in frog trials: large frogs were more frequently constricted after a delay of more than 1 s than small frogs, and small frogs were usually simply seized. On the other hand, prey type affected prey-handling method: mice and lizards were more frequently constricted or pinioned than frogs. When hatchlings of E. quadrivirgata tried to coil around a prey animal immediately after striking, they frequently failed or released their coils within 10 s after striking. In experiment 2, prey-handling behaviour of adult E. climacophora was examined with mice of various sizes. Adult E. climacophora tended to constrict large mice immediately after striking, whereas they simply seized small mice. Only one out of 38 mice which were constricted immediately after striking was released from the coils within the subsequent 10 s. Large mice were killed prior to ingestion, whereas small mice were swallowed alive. Large mice tended to be swallowed head first. Experiment 3 was conducted to investigate effects of mouse size on prey-handling behaviour of newly hatched E. climacophora. Mouse size affected condition of prey at ingestion and prey-handling method but not capture position and direction of ingestion. Possible relationships between feeding ecology and the differences of prey-handling behaviour among Elaphe are discussed.     

Prey-handling and the evolutionary ecology of sand-swimming lizards (Lerista : Scincidae)

Fossorial lizards differ in morphology from their surface-dwelling relatives. The Australian sphenomorphine skink genus Ctenotus consists of surface-dwelling species, and is closely related to the genus Lerista, which includes both surface-dwelling and fossorial species. Sand-swimming represents the derived condition and has evolved independently in several lineages of Lerista. The heads of lizards in the two genera differ in shape (blunt snout for Ctenotus versus wedge-shaped for Lerista) and in length relative to the body (approximately 20% of snout-vent length for Ctenotus versus 12% for sand-swimming Lerista). Do these specializations affect the sizes or types of prey that can be consumed by Lerista? We compared prey-handling by Ctenotus and Lerista to correlate morphological differences with differences in prey-handling ability, and to distinguish the effects of snout shape and head length. Feeding trials included three categories of insect prey that the lizards normally eat: soft-bodied larvae (Lepidoptera), hard-bodied larvae (Coleoptera), and roaches (Blatoidea). In comparisons based on the mass of a prey item relative to the mass of a lizard, Lerista had longer handling times for all prey categories and were limited to smaller prey than were Ctenotus. However, when comparisons were based on the length of prey relative to the length of a lizard's head, Lerista ate some elongate prey as fast or faster than did Ctenotus, and both genera successfully swallowed prey more than twice the length of their own head. Thus, the differences in prey-handling performance of Ctenotus and Lerista probably result from the fact that Lerista have a relatively shorter head than Ctenotus. All Lerista species, surface-dwelling and fossorial, have short heads compared to primitive sphenomorphine lizards. Fossorial species of Lerista have elongate trunks, and consequently their heads are shorter in proportion to trunk length than those of surface-dwelling Lerista. However, most fossorial species of Lerista are longer and heavier than any of their surface-dwelling congeners, and the heads of these fossorial species are large relative to the prey they encounter. As a consequence, the diets of large fossorial species of Lerista do not appear to be limited by their morphological specialization for sand-swimming.     

ECOLOGICAL SPECIATION IN STICKLEBACKS: ENVIRONMENT-DEPENDENT HYBRID FITNESS

“Ecological” speciation occurs when reproductive isolation evolves as a consequence of divergent selection between populations exploiting different resources or environments. We tested this hypothesis of speciation in a young stickleback species pair by measuring the direct contribution of ecological selection pressures to hybrid fitness. The two species (limnetic and benthic) are strongly differentiated morphologically and ecologically, whereas hybrids are intermediate. Fitness of hybrids is high in the laboratory, especially F₁ and F₂ hybrids (backcrosses may show some breakdown). We transplanted F₁ hybrids to enclosures in the two main habitats in the wild to test whether the distribution of resources available in the environment generates a hybrid disadvantage not detectable in the laboratory. Hybrids grew more slowly than limnetics in the open water habitat and more slowly than benthics in the littoral zone. Growth of F₁ hybrids was inferior to the average of the parent species across both habitats, albeit not significantly. The contrast between laboratory and field results supports the hypothesis that mechanisms of F₁ hybrid fitness in the wild are primarily ecological and do not result from intrinsic genetic incompatibilities. Direct selection on hybrids contributes to the maintenance of sympatric stickleback species and may have played an important role in their origin.     

An anomalous hybrid zone in Drosophila

Abstract Despite the genetic tractability of many of Drosophila species, the genus has few examples of the “classic” type of hybrid zone, in which the ranges of two species overlap with a gradual transition from one species to another through an area where hybrids are produced. Here we describe a classic hybrid zone in Drosophila that involves two sister species, Drosophila yakuba and D. santomea, on the island of SaTo Tomé. Our transect of this zone has yielded several surprising and anomalous findings. First, we detected the presence of an additional hybrid zone largely outside the range of both parental species. This phenomenon is, to our knowledge, unique among animals. Second, the genetic analysis using diagnostic molecular markers of the flies collected in this anomalous hybrid zone indicates that nearly all hybrid males are F₁s that carry the D. santomea X chromosome. This F₁ genotype is much more difficult to produce in the laboratory compared to the genotype from the reciprocal cross, showing that sexual isolation as seen in the laboratory is insufficient to explain the genotypes of hybrids found in the wild. Third, there is a puzzling absence of hybrid females. We suggest several tentative explanations for the anomalies associated with this hybrid zone, but for the present they remain a mystery.     

SEXUAL SELECTION AGAINST HYBRIDS BETWEEN SYMPATRIC STICKLEBACK SPECIES: EVIDENCE FROM A FIELD EXPERIMENT

Sexual selection against viable, fertile hybrids may contribute to reproductive isolation between recently diverged species. If so, then sexual selection may be implicated in the speciation process. Laboratory measures of the mating success of hybrids may underestimate the amount of sexual selection against them if selection pressures are habitat specific. Male F₁ hybrids between sympatric benthic and limnetic sticklebacks (Gasterosteus aculeatus complex) do not suffer a mating disadvantage when tested in the laboratory. However, in the wild males choose different microhabitats and parental females tend to be found in the same habitats as conspecific males. This sets up the opportunity for sexual selection against male hybrids because they must compete with parental males for access to parental females. To test for sexual selection against adult F₁ hybrid males, we examined their mating success in enclosures in their preferred habitat (open, unvegetated substrate) where limnetic males and females also predominate. We found significantly reduced mating success in F₁ hybrid males compared with limnetic males. Thus, sexual selection, like other mechanisms of postzygotic isolation between young sister species, may be stronger in a wild setting than in the laboratory because of habitat-specific selection pressures. Our results are consistent with, but do not confirm, a role for sexual selection in stickleback speciation.     

TWO MORPHOLOGICALLY SIMILAR BIOLOGICAL SPECIES: CHONDRUS PINNULATUS AND C. ARMATUS (GIGARTINACEAE,RHODOPHYTA)1

The nature of the relationship between Chondrus pinnulatus (Harvey) Okamura f. pinnulatus and C. pinnulatus f. armatus (Harvey) Yamada et Mikami (Gigartinaceae, Rhodophyta) was investigated by comparative analysis of their biogeography, phenologies, life histories, gross and vegetative morphology, crossability, and upper thermal tolerance. The forma pinnulatus has a more northerly distribution in Japan and adjacent waters, exhibiting adaptation to the cooler regions, whereas the forma armatus has a more southerly range. The latter may be the result of a higher thermal tolerance. Both formae have a Polysiphonia-type life history and are similar in their internal vegetative morphology. They can, however, be distinguished by gross morphology: forma pinnulatus has wide, flattened axes, compressed to flattened ultimate segments and proliferations, while forma armatus has narrow, compressed to subterete axes and subterete to terete ultimate segments and proliferations. These differences persist in laboratory culture. All intraformae crosses were positive, with carpospores from the cross developing into fertile F₁ tetrasporophytes releasing tetraspores that developed into dioecious F₁ gametophytes, the female gametophytes of which formed normal cystocarps. This suggests that members of populations of each forma freely interbreed. Among interformae crosses, only some offspring derived from geographically distant strains bore normal cystocarps in the F₁ female gametophytes. Other crosses showed that interbreeding between populations of these two formae was blocked by various isolating mechanisms: incompatibility, hybrid inviability, and hybrid sterility. Reproductive isolation between f. pinnulatus and f. armatus is virtually complete in wild populations, because hybrid populations have not been found in the wild. In addition, these two entities can be considered biological species that are also referred to the taxonomic species, C. armatus and C. pinnulatus, because they do not overlap with regard to the morphology of the ultimate segments and proliferations. Subtle (but significant) gross morphological differences, partial interfertility between the two species, and deleterious hybridization in the area in which they occur sympatrically suggest that their evolutionary divergence was relatively recent.     

Antipredator tactics: a kin-selection benefit for defensive spines in coral catfish?

Morphological features that impair a predator's ability to consume a prey item may benefit individual prey; but what of features that prolong prey-handling but do not enhance prey survival? For example, a striped eel catfish Plotosus lineatus will be fatally envenomated if struck by its specialist predator, the greater sea snake Hydrophis major. Nonetheless, the catfish typically erects long, toxic pectoral and dorsal spines that increase prey-handling times for the snake by around eightfold. Because the catfish travel in swarms of closely-related individuals, the delay enforced by spines may enable the victim's swarm-mates to disperse before the snake is able to search for another meal. In keeping with that hypothesis, defensive spines tend to be longer in catfish from regions where the greater sea snake occurs, than from areas where the snake does not occur.     

Identification of a short rDNA spacer sequence highly specific of a tomato line containing Tm-1 gene introgressed from Lycopersicon hirsutum

Summary We studied rDNA restriction fragment length polymorphism between two tomato lines used for F₁ hybrid seed production: line A, containing the Tm-1 gene responsible for tobacco mosaic virus tolerance introgressed from the wild species Lycopersicon hirsutum, and line B, a tobacco mosaic virus sensitive line. Hybridization patterns led to distinct rDNA maps with two size classes, 10.4 and 10.7 kb, in line A and a single, 8.9-kb class in line B. Size differences were located in the intergenie sequence (IGS). A highly specific 54-bp TaqI fragment was cloned from the line A IGS and used in dot blot experiments to probe total DNA from line A, line B, and their F₁ hybrid. It proved capable of discriminating B from A and the hybrid. This probe could thus serve to screen inbreds in commercial seed lots where line A is used as male. This fragment showed 80–90% sequence homology with the 53-bp subrepeats previously characterized in a region of the tomato IGS close to the 25S rRNA gene. Preliminary comparison of rDNA in line A and several wild related species indicated that the L. hirsutum H₂ genotype was the closest to line A. rDNA variations between line A and this wild genotype could be explained by recombination during the introgression process involving numerous backcrosses or by an important intraspecific polymorphism. Our results strongly suggest that Tm-1 and the rDNA were introgressed together into tomato from L. hirsutum through linkage drag.     

NUMERICAL TAXONOMY APPLIED TO CUCURBITA RELATIONSHIPS

In a numerical study of taxonomic relationships in Cucurbita, the inclusion of F₁ interspecific hybrids did not materially alter the species grouping in the phenogram. The F₁ interspecific hybrids tend to cluster with one of the parent species or species groups unless the parents are widely divergent. Also, the F₁ interspecific hybrids between wild and domesticated species cluster with the wild parent Both parent species or species groups of an F₁ interspecific hybrid can usually be determined from Q-correlation values. It is essential that compatibility relations be determined between species; i.e., species groups must be established before Q-correlation values can be successfully used in determining the parentage of hybrids. Undescribed collections can usually be correctly placed in the proper species group.     

Hybridization and sequential components of reproductive isolation between parapatric walnut‐infesting sister species Rhagoletis completa and Rhagoletis zoqui

Hybridization can provide a window into how populations diverge to form new species. Here, we confirm hybridization between Rhagoletis completa Cresson, 1929 and Rhagoletis zoqui Bush, 1966, two species of walnut husk‐infesting flies that geographically overlap in a narrow area of parapatry in Northeastern Mexico. Rhagoletis completa and R. zoqui are members of a species group (Rhagoletis suavis) that has been hypothesized to speciate in allopatry. Sexual selection has been argued to be a potentially important factor for generating pre‐mating isolation among walnut husk flies, because of the differences in wing morphology and body coloration, and the existence of sexual dimorphism within species. However, there was no evidence for pre‐mating isolation between R. completa and R. zoqui, based on choice and no‐choice mating experiments conducted on adults of fly populations outside the contact zone. There was also no support for reduced fertility of hybrid matings or for F₁ inviability; however, F₁ hybrids appeared to have lower fertility and F₂ offspring have reduced survivorship. Postzygotic isolation in later generation hybrids of mixed ancestry therefore appears to be the first intrinsic barrier to gene flow evolving between R. completa and R. zoqui. We discuss the implications of our results for allopatric speciation in walnut flies and the potential evolutionary fate of R. zoqui and R. completa if they were to come into broad geographic contact in the future. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

ESTIMATING THE MATING BEHAVIOR OF A PAIR OF HYBRIDIZING HELICONIUS SPECIES IN THE WILD

Premating isolation between incipient species is rarely studied in nature, even though mating tests in captivity may give an inaccurate picture of natural hybridization. We studied premating barriers between the warningly colored butterflies Heliconius erato and H. himera (Lepidoptera) in a narrow contact zone in Ecuador, where hybrids are found at low frequency. Eggs obtained from wild-mated females, supplemented with eggs and young larvae collected from the wild, were reared to adulthood. Adult color patterns of these progeny were then used to infer how their parents must have mated. Likelihood was used to estimate both the frequencies of potential parental genotypes from adult phenotypes collected in the wild, and the degree of assortative mating from the inferred parents. The frequencies of parental genotypes varied across the hybrid zone, but our statistical method allowed estimates of hybrid deficit and assortative mating to be integrated across all sites sampled. The best estimate of the frequency of F₁ and backcross hybrid adults in the center of the hybrid zone was 10%, with support limits (7.1%, 13.0%; support limits are asymptotically equivalent to 95% confidence limits). Mating was highly assortative: in the center of the hybrid zone the cross-mating probability between H. erato and H. himera was only 5% (0.3%, 21.4%). Wild hybrids themselves mated with both pure forms, and the probabilities that they mated in any direction were not significantly lower than those among conspecifics. These results are consistent with earlier laboratory studies on mate choice, and suggest that selection against hybrids must be strong to prevent formation of a hybrid swarm. Unfortunately, the wide support limits on mating behavior precluded a measure of the strength of selection from these data alone. Our statistical approach provides a useful general method for estimating mate choice in the wild.     

Fitness of reciprocal F1 hybrids between Rhinanthus minor and Rhinanthus major under controlled conditions and in the field

The performance of first‐generation hybrids determines to a large extent the long‐term outcome of hybridization in natural populations. F₁ hybrids can facilitate further gene flow between the two parental species, especially in animal‐pollinated flowering plants. We studied the performance of reciprocal F₁ hybrids between Rhinanthus minor and R. major, two hemiparasitic, annual, self‐compatible plant species, from seed germination to seed production under controlled conditions and in the field. We sowed seeds with known ancestry outdoors before winter and followed the complete life cycle until plant death in July the following season. Germination under laboratory conditions was much lower for the F₁ hybrid formed on R. major compared with the reciprocal hybrid formed on R. minor, and this confirmed previous results from similar experiments. However, this difference was not found under field conditions, which seems to indicate that the experimental conditions used for germination in the laboratory are not representative for the germination behaviour of the hybrids under more natural conditions. The earlier interpretation that F₁ hybrid seeds formed on R. major face intrinsic genetic incompatibilities therefore appears to be incorrect. Both F₁ hybrids performed at least as well as and sometimes better than R. minor, which had a higher fitness than R. major in one of the two years in the greenhouse and in the field transplant experiment. The high fitness of the F₁ hybrids confirms findings from naturally mixed populations, where F₁ hybrids appear in the first year after the two species meet, which leads to extensive advanced‐hybrid formation and introgression in subsequent generations.     

Geographic patterns of postzygotic isolation between two closely related widespread dung fly species (Sepsis cynipsea and Sepsis neocynipsea; Diptera: Sepsidae)

Identifying the contribution of pre‐ and postzygotic barriers to gene flow is a key goal of speciation research. The widespread dung fly species Sepsis cynipsea and Sepsis neocynipsea offer great potential for studying the speciation process over a range of opportunities for gene exchange within and across sister species (cross‐continental allopatry, continental parapatry and sympatry). We examined the role of postcopulatory isolating barriers by comparing female fecundity and egg‐to‐adult viability of F₁ and F₂ hybrids, as well as backcrosses of F₁ hybrids with the parental species, via replicated crosses of sym‐, para‐ and allopatric populations. Egg‐to‐adult viability was strongly but not totally suppressed in hybrids, and offspring production approached nil in the F₂ generation (hybrid breakdown), indicating yet unspecified intrinsic incompatibilities. Viable F₁ hybrid offspring showed almost absolute male (the heterogametic sex) sterility while females remained largely fertile, in accordance with Haldane's rule. Hybridization between the two species in European areas of sympatry (Swiss Alps) indicated only minor reinforcement based on fecundity traits. Crossing geographically isolated European and North American S. neocynipsea showed similar albeit weaker isolating barriers that are most easily explained by random genetic drift. We conclude that in this system with a biogeographic continuum of reproductive barriers, speciation is mediated primarily by genetic drift following dispersal of flies over a wide (allopatric) geographic range, with some role of natural or sexual selection in incidental or direct reinforcement of incompatibility mechanisms in areas of European sympatry. S(ubs)pecies status of continental S. neocynipsea appears warranted.     

SEX CHROMOSOME LINKAGE OF MATE PREFERENCE AND COLOR SIGNAL MAINTAINS ASSORTATIVE MATING BETWEEN INTERBREEDING FINCH MORPHS

Assortative mating is a key aspect in the speciation process because it is important for both initial divergence and maintenance of distinct species. However, it remains a challenge to explain how assortative mating evolves when diverging populations are undergoing gene flow (e.g., during hybridization). Here I experimentally test how assortative mating is maintained with frequent gene flow between diverged head‐color morphs of the Gouldian finch (Erythrura gouldiae). Contrary to the predominant view on the development of sexual preferences in birds, cross‐fostered offspring did not imprint on the phenotype of their conspecific (red or black morphs) or heterospecific (Bengalese finch) foster parents. Instead, the mating preferences of F₁ and F₂ intermorph‐hybrids are consistent with inheritance on the Z chromosomes, which are also the location for genes controlling color expression and the genes causing low fitness of intermorph‐hybrids. Genetic associations between color signal and preference loci on the sex chromosomes may prevent recombination from breaking down these associations when the morphs interbreed, helping to maintain assortative mating in the face of gene flow. Although sex linkage of reproductively isolating traits is theoretically expected to promote speciation, social and ecological constraints may enforce frequent interbreeding between the morphs, thus preventing complete reproductive isolation.     

Exploratory behaviour and novel predator recognition: behavioural correlations across contexts

It was hypothesized that the exploratory behaviour of an individual measured in a novel environment could predict its behaviour in response to a novel predator. This study examined novel predator recognition in the western mosquitofish Gambusia affinis, a species with individual differences in risk‐taking, activity and exploration in novel environments. Prey responded with characteristic shoaling and avoidance in response to native predators, but did not show characteristic antipredator behaviour towards novel predators. Furthermore, G. affinis exhibited individual‐level behavioural correlations across contexts but only when prey were tested with native predators. This could be the result of native predatory selection on behavioural correlations in the prey species.     

Identification of larch species (Larix decidua, Larix kaempferi and Larix X eurolepis) and estimation of hybrid fraction in seed lots by RAPD fingerprints

Species-specific RAPD markers were used to identify the different larch species (Larix decidua and Larix kaempferi) and their interspecific hybrid (Larix X eurolepis). Although morphological differences between pure species and the hybrids exist, differentiation is not always possible, especially at an early stage (seed or plantlet). Eleven RAPD markers differentiated the two larch species, and 4 species-specific markers were sufficient to estimate the F₁ hybrid fraction in a seed lot. The species-specific markers were tested on individual trees of European and Japanese larches of diverse geographic origins and on several seed lots of different origins (F₁, F₂ hybrids and pure species). The 4 specific markers found for the European larch and the Japanese larch were monomorphic and present in all provenances and in all F₁ hybrid trees tested. Polymorphic SCAR fragments were obtained for 3 of the 11 fragments originally selected for the RAPD screening phase. For 2 of them, the sequence had some homology with the mitochondrial genome of other organisms and is thus mitochondrial. The two mitochondrial fragments and the OPF-13₁₀₀₀ fragment exhibited one polymorphic band, thereby maintaining its species-specific identity: OPF-13₁₀₀₀ is specific to the European larch. The 4 RAPD primers selected in this study offer a reliable, quick and cheap tool for the identification of different larch species (Larix decidua and Larix kaempferi) and their interspecific hybrid (Larix X eurolepis). Received: 28 February 1999 / Accepted: 29 April 1999     

Postzygotic isolation drives genomic speciation between highly cryptic Hypocnemis antbirds from Amazonia

How species evolve reproductive isolation in the species-rich Amazon basin is poorly understood in vertebrates. Here, we sequenced a reference genome and used a genome-wide sample of SNPs to analyze a hybrid zone between two highly cryptic species of Hypocnemis warbling-antbirds—the Rondonia warbling-antbird (H. ochrogyna) and Spix's warbling-antbird (H. striata)—in a headwater region of southern Amazonia. We found that both species commonly hybridize, producing F₁s and a variety of backcrosses with each species but we detected only one F₂-like hybrid. Patterns of heterozygosity, hybrid index, and interchromosomal linkage disequilibrium in hybrid populations closely match expectations under strong postzygotic isolation. Hybrid zone width (15.4 km) was much narrower than expected (211 km) indicating strong selection against hybrids. A remarkably high degree of concordance in cline centers and widths across loci, and a lack of reduced interspecific F_st between populations close to versus far from the contact zone, suggest that genetic incompatibilities have rendered most of the genome immune to introgression. These results support intrinsic postzygotic isolation as a driver of speciation in a moderately young cryptic species pair from the Amazon and suggest that species richness of the Amazon may be grossly underestimated.     

Hybridization and introgression across different ploidy levels in the Neotropical orchids Epidendrum fulgens and E. puniceoluteum (Orchidaceae)

The hypothesis of gene flow between species with large differences in chromosome numbers has rarely been tested in the wild, mainly because species of different ploidy are commonly assumed to be reproductively isolated from each other because of instantaneous and strong postzygotic barriers. In this study, a broad‐scale survey of molecular variation was carried out between two orchid species with different ploidy levels: Epidendrum fulgens (2n = 2x = 24 chromosomes) and Epidendrum puniceoluteum (2n = 4x = 52 chromosomes). To test the strength of their reproductive barriers, we investigated the distribution of genetic variation in sympatric and allopatric populations of these two species and conducted crossing experiments. Nuclear and plastid microsatellite loci were used to genotype 463 individuals from eight populations across the geographical range of both species along the Brazilian coastal plain. All six sympatric populations analysed presented hybrid zones, indicating that hybridization between E. fulgens and E. puniceoluteum is a common phenomenon. Bayesian assignment analysis detected the presence of F₁ and F₂ individuals and also signs of introgression, demonstrating a high potential for interspecific gene flow. Introgression occurs preferentially from E. fulgens to E. puniceoluteum. Pure parental individuals of both species display strong genotype–habitat associations, indicating that environment‐dependent selection could be acting in all hybrid zones. This study suggests that hybridization and introgression are evolutionary processes playing a role in the diversification of Epidendrum and indicates the importance of investigations of hybrid zones in understanding reproductive barriers and speciation processes in Neotropical orchid species.     

DEVELOPMENTAL STABILITY,FITNESS, AND TRAIT SIZE IN LABORATORY HYBRIDS BETWEEN EUROPEAN SUBSPECIES OF THE HOUSE MOUSE

The effects of hybridization on developmental stability and size of tooth characters were investigated in intersubspecific crosses between random-bred wild strains of the house mouse (Mus musculus domesticus and M. m. musculus). Fluctuating asymmetry (FA) and trait size were compared within and between parental, F₁, backcross, and F₂ hybrid groups. The relationship between FA and reproductive fitness within the F₁ hybrids was also studied. The results indicated that both FA and character size levels differed significantly between the two subspecies. The F₁ hybrids and the recombined groups (backcrosses and F₂ hybrids) showed heterosis for both parameters. No significant differences in the FA of fertile and sterile F₁ hybrid individuals were found. Comparison of the FA levels obtained in this study with those found in wild populations from the hybrid zone in Denmark showed that the levels of FA were lower in laboratory-bred samples than in the wild populations. This study provides further evidence that, in hybrids, the developmental processes underlying most of the morphological traits we studied benefit from a heterotic effect, despite the genomic incompatibilities between the two European house mice revealed by previous genetical and parasitological studies.