Frequency of interspecific mating in salamanders of the plethodontid genus Desmognathus: different experimental designs may yield different results

Heterosexual courtship trials were staged between dusky salamanders of two species ( Desmognathus fuscus and D. santeetlah ) from two allopatric populations in the southern Appalachian Mountains of eastern North America. Two experiments were conducted in order to test for effects of experimental design on estimates of the strength of sexual isolation between these species. In Experiment 1, frequencies of interspecific mating were determined when: (1) a male was presented with a conspecific and a heterotypic female successively; and (2) both types of females were presented simultaneously to a male. The frequency of interspecific mating by male D. fuscus was greatest when females were presented successively. The frequency of interspecific mating by male D. santeetlah was not influenced by method of female presentation. An index of sexual isolation of +0.29 was obtained between D. fuscus and D. santeetlah when females were presented successively, and of +0.63 when females were presented simultaneously. In Experiment 2, interspecific mating frequencies were determined in courtship arenas of two different sizes. No significant effect of arena size on frequency of interspecific mating was found for D. fuscus. In D. santeetlah , a trend towards greater mating frequency in small arenas was almost significant. These results suggest that interspecific mating frequencies (and thus estimates of the strength of sexual isolation between populations or species) may be sensitive to the experimental design employed. The validity and generality of this conclusion for other animals is an urgent, empirical issue.     

Evolution of Courtship Behaviour Patterns and Reproductive Isolation in the Desmognathus ochrophaeus Complex

The extent to which differences in courtship behaviour patterns act as mechanisms of reproductive isolation is critical to understanding both speciation and the evolution of these behaviour patterns. While numerous studies have investigated intraspecific and interspecific differences in courtship, fewer interpret results in a phylogenetic framework. We describe and analyse geographic variation in the courtship behaviour patterns of the Allegheny Dusky salamander ( Desmognathus ochrophaeus ). We then examine courtship among closely related species in the D. ochrophaeus complex in a phylogenetic context. We found that populations of D. ochrophaeus separated by extensive geographic distances show little variation in courtship behaviour patterns and are sexually compatible. This contrasts with significant levels of sexual isolation between D. ochrophaeus and other species in the complex. Mapping behaviour patterns onto a phylogeny that we generated from cytochrome b sequences indicates that two behaviour patterns present in the courtship sequence of other members in the complex have either been lost in D. ochrophaeus or gained independently in other species in the complex. Loss of these behaviour patterns may result in reproductive isolation between D. ochrophaeus and its sister taxon, D. orestes .     

Systematics of dusky salamanders, Desmognathus (Caudata: Plethodontidae), in the mountain and Piedmont regions of Virginia and North Carolina, USA

We analysed mitochondrial (cytochrome  b ) nucleotide sequences, nuclear allozyme markers, and morphometric characters to investigate species boundaries and phylogenetic relationships among dusky salamanders ( Desmognathus ) in the southern Blue Ridge and adjacent Piedmont Physiographic Provinces of Virginia and North Carolina. Our results revealed four distinct mitochondrial DNA clades that are also characterized by distinct allozyme markers. One clade consists of sequences derived from populations distributed from New England to south-western Virginia that are referable to Desmognathus fuscus Rafinesque, 1820, although there is considerable sequence and allozyme divergence within this clade. A second clade consists of sequences derived from populations referable to Desmognathus planiceps Newman, 1955, a form that we resurrect from its long synonymy under D. fuscus . Desmognathus planiceps and D. fuscus also differ in mandibular tooth shape. Two other cytochrome  b sequences recovered from populations along the Blue Ridge escarpment in southern Virginia are quite divergent from those of the previous two clades, and these populations may represent yet another distinct species. Sequences from a population in the Brushy Mountains in the Piedmont of northern North Carolina are similar to those of Desmognathus carolinensis . Population groupings indicated by allozyme data generally correspond to the cytochrome  b clades. Cryptic diversity in Appalachian desmognathan salamanders clearly requires further study. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 115–130.     

Patterns of trombiculid mite (Hannemania dunni) parasitism among plethodontid salamanders in the western Piedmont of North Carolina

Trombiculid mites are known to parasitize a variety of amphibian species, yet few comparisons of mite parasitism among amphibian species have been made. In this study, we investigated patterns of trombiculid mite parasitism among 3 plethodontid salamanders (Desmognathus fuscus, Eurycea cirrigera, and Plethodon cylindraceus) in the western Piedmont of North Carolina. All 3 salamander species were parasitized by a single species, Hannemania dunni. Desmognathus fuscus harbored mites more frequently (60.4% of individuals) than E. cirrigera (11.1%) or P. cylindraceus (14.6%). Desmognathus fuscus also had higher parasite loads than E. cirrigera or P. cylindraceus (P < 0.001). Mites on D. fuscus were found more frequently on the limbs than other body locations (P < 0.001). We found no correlation between salamander size and mite abundance (P = 0.689), but salamander collection sites influenced the abundance of mites on D. fuscus (P = 0.002). We found no effect of season on mite abundance in D. fuscus (P = 0.952). Salamander habitat preferences and edaphic or climatic differences among study sites may influence patterns of Hannemania sp. parasitism of salamanders.     

Diffusion-limited gas exchange across amphibian skin

Cutaneous gas exchange function of amphibians is conveniently analyzed in lungless salamanders in which all gas exchange is through the skin. Measurements in Desmognathus fuscus (Plethodontidae) have yielded the following results: 1) Cutaneous transfer of O2 and CO2 is limited predominantly by diffusion. 2) Accordingly, in environmental hypoxia, O2 uptake decreases. 3) Anaerobiosis contributes to energy balance during the first few hours of hypoxia only. Thereafter unknown biochemical adjustments take place that allow the animal to restore metabolite levels characteristic of normoxia regardless of the continued reduced oxygen uptake.     

Male—male aggression in the plethodontid salamander Desmognathus ochrophaeus

Aggressive encounters between pairs of male mountain dusky salamanders (Desmognathus ochrophaeus) were staged in the laboratory in the absence of any obvious limited resource. We observed aggressive interactions which sometimes escalated to biting. Physical injuries were inflicted during some aggressive encounters; however, the consequences of injury for future fitness are uncertain. Several behaviour patterns used in contests apparently functioned as threat displays, and these sometimes ended contests before they escalated to biting. We could detect no significant influences of three types of asymmetries among males in determining the winners and losers of contests (body size, recent mating experience and familiarity with the testing arena). Three cases of 'homosexual' courtship were observed. Both intermale aggression and intermale courtship may be interpreted as forms of competition for mates.     

Antennal courtship and functional morphology of tyloids in the parasitoid wasp Syrphoctonus tarsatorius (Hymenoptera: Ichneumonidae: Diplazontinae)

Courtship behaviour and associated morphological characters are believed to evolve under diversifying sexual selection. In Hymenoptera, sexually dimorphic antennal structures, the ‘tyloids’, show a large variability. Although crucial for functional interpretation, the link between tyloid morphology and courtship behaviour has gained only limited attention. Here, we investigate antennal morphology and antennal courtship in the parasitoid wasp Syrphoctonus tarsatorius (Hymenoptera: Ichneumonidae: Diplazontinae). We confirm the glandular nature of the tyloids by light and scanning electron microscopy. Moreover, we report a new form of antennation during courtship, antennal double-coiling, which links morphology and behaviour by bringing the tyloids in direct contact with the antennae of the female, thus probably facilitating the transfer of a contact pheromone. We show that a change in haemolymph pressure is the activator of the antennal movement and that it can be reproduced in the laboratory using amputated antennae. Investigations of antennal structure and movement in three additional hymenopteran species suggest that the number and location of tyloids coincide with the modality of antennal coiling. Our method for simulating antennal movement will enable retrieving information about courtship behaviour from museum specimens, thus leading to a better understanding of the evolution of courtship behaviour in Hymenoptera.     

It Takes Two to Tango: Relative Influence of Male and Female Identity and Morphology on Complex Courtship Display in a Newt Species

Consistency in behaviour is currently receiving a renewed interest. Although courtship display is generally consistent in terms of behavioural sequence and structure, there is also commonly important variation in the intensity of courtship display between and within males of a given species. Indeed, not all males have the same ability to perform courtship display (variation between males), and each male can potentially adjust his courtship effort in response to the environment (variation within a male). Although the study of male courtship display has received considerable attention in recent years, it is still unclear which part of the variation can be explained by male ability or motivation. We investigated this issue on two phases of the complex courtship display of the palmate newt Lissotriton helveticus. Overall, we found that both male and female identities affected courtship behaviour, but the relative influence of each sex depended on the courtship phase. Male identity explained variation in fan and creep‐quiver display, whereas female identity explained variation in creep‐quiver only. Interestingly, we did not find any link between the expression of courtship display and male or female morphological traits. Our study showed consistency of male courtship display in newts and successfully dissects the different sources of variation that can affect behavioural repeatability/consistency of courtship display.     

Stifled sex in sympatry: patterns of sexual incompatibility among desmognathine salamanders

Sexual incompatibility due to differences in mate-recognition systems can be an important factor in preventing hybridization and gene flow between animal populations in sympatry. We tested in the laboratory for sexual incompatibility between populations of two species of desmognathine salamanders that occur in sympatry in south-eastern Kentucky, North America, Desmognathus monticola and D. welteri (Caudata: Plethodontidae: Desmognathinae). Spermatophore deposition and insemination (two related measures of sexual success) were frequent in intraspecific encounters, but interspecific encounters never resulted in the deposition of spermatophores by males (Experiment 1). The sexual behaviour patterns of these two species are similar qualitatively, but all encounters staged between them failed to progress beyond the most preliminary of courtship interactions (Experiment 2), perhaps due to species differences in chemosensory communication. Our data on sexual incompatibility between D. monticola and D. welteri are similar to those available for other desmognathine taxa that occur in sympatry. Review of a broader data-set on patterns of sexual incompatibility both within and between species, and in allopatry and in sympatry, provides evidence for both divergence of mate-recognition systems in allopatry and enhancement of sexual incompatibility in sympatry for desmognathine salamanders. We hypothesize that diversification in allopatry may be a consequence of natural and/or sexual selection acting to promote sexual success within populations. Enhancement in sympatry may be a consequence of subsequent selection for accurate species recognition under the threat of hybridization or gene flow between species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 367–375.     

The courtship behaviour of the pipefish Nerophis lumbriciformis: reflections of an adaptation to intertidal life

The courtship behaviour of Nerophis lumbriciformis (Pisces: Syngnathidae) consists of three distinct phases (initial courtship, spawning, and embrace) marked by prominent behavioural changes. The first courtship phase is characterised by female quivering. Courtship activity increases from low to high levels of quivering, causing undulatory head movements in the female. In the second phase, the female transfers her eggs onto the male incubating ventral surface. During the final phase, the male wraps his body around the female (embrace). Females actively initiate courtship indicating that these pipefishes are courtship role reversed. The complete lack of swimming and vertical movements in the courtship behaviour of N. lumbriciformis, unlike in the behaviour of other syngnathid species, suggests an adaptation to intertidal conditions.     

Mating behaviour and reproductive isolation of three species of reptile tick

The mating behaviour of three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum is similar and involves a female sex pheromone which activates males to detach from their hosts and search. After contacting females, a stereotyped six-phase courtship sequence occurs. There are qualitative differences between the species in courtship behaviour at phase 3 (reversal of position by dorsally mounted males) and phase 6 (the copulation position). On-host observations of non-conspecific mating show that females of the three species have species-specific activation pheromones, which is contrary to reports in other species of tick. Such specificity should result in reproductive isolation of the three species; however, under certain circumstances it may not prevent non-conspecific contacts between the sexes. Off-host observations of courtship behaviour show that once males of the three species contact non-conspecific females, they attempt courtship and are persistent with their courtship. Rarely did non-conspecific courtships proceed beyond phase 4 of the courtship sequence, as non-conspecific females did not lift their bodies to allow males venter contact. Differences between the species in leg orientation in the copulation position, together with body size differences, are responsible for a complete barrier to successful non-conspecific copulation. The observations illustrate the role that behavioural mechanisms play in reproductively isolating these three species of tick.     

Inhibitory effects of courtship in the wasp Nasonia vitripennis and a new interpretation of the biological significance of courtship in insects

Courtship has a cumulative inhibitory effect on the sexual behaviour of male Nasonia. The biological significance of this inhibition is considered. Courtship inhibits other activities, including the movements that normally follow courtship but precede insemination. If the female is receptive, this delay gives time for mutual recognition, and makes possible correct orientation and synchronization. Courtship also inhibits itself: if the female is non-receptive, courtship does not go on indefinitely. This termination of courtship allows the female to resume other activities and increases the male's chances of meeting other females, which it may inseminate.     

Courtship herding in the fiddler crab <Emphasis Type="Italic">Uca elegans</Emphasis>

Male and female animals are not always complicit during reproduction, giving rise to coercion. One example of a system that is assumed to involve sexual coercion is the mate herding behaviour of fiddler crabs: males push females towards the home burrow with the goal of forcing copulation at the burrow entrance. We recorded and analysed in detail the courtship behaviour of a North Australian species of fiddler crab Uca elegans. Courtship was composed of four main phases: broadcast waving, outward run, herding and at burrow display. During interactions males produced claw-waving displays which were directed posteriorly towards the female and which varied in timing and structure depending on the courtship phase. We suggest that courtship herding in U. elegans is driven primarily by mate choice for the following reasons, (1) females can evade herding, (2) no other reproductive strategies were observed, (3) males broadcast their presence and accompany courtship with conspicuous claw waves, and (4) the behaviour ends with the female leading the male into the home burrow. As an alternative function for herding in U. elegans we suggest that the behaviour represents a form of courtship guiding, in which males direct complicit females to the correct home burrow.     

MATING BEHAVIOUR OF DEROCERAS RODNAE GROSSU & LUPU, 1965 AND D. PRAECOX WIKTOR, 1966 (PULMONATA: AGRIOLIMACIDAE)

The mating behaviour of Deroceras rodnae Grossu & Lupu,1965 and D. praecox Wiktor, 1966 was observed under laboratoryconditions. Both species differ by the shape and functioningof the sarcobelum, the timing and duration of courtship andthe colour of the sperm mass. Particularly the different timingof the courtship behaviour seems to act as an effective prezygoticisolation mechanism preventing interspecific crosses. Usually,sperm masses are exchanged mutually. In three homospecific pairings,unilateral sperm transfer was observed followed by a characteristicbehaviour of the donor slug. This observation is discussed inlight of the hermaphrodite's dilemma and sperm trading models. (Received 23 May 1994; accepted 1 December 1994)     

Time courses of thermal acclimation for critical thermal minima in the salamanders Desmognathus quadramaculatus, Desmognathus monticola, Desmognathus ochrophaeus, and Plethodon jordani

1. We examined the time courses of thermal acclimation for critical thermal minima (CTMin) in the salamanders Desmognathus quadramaculatus, D. monticola, D. ochrophaeus, and Plethodon jordani. 2. D. quadramaculatus showed no annual differences in their time courses, but we noted seasonal differences in the CTMin with these salamanders having lower CTMin in the spring. 3. Gain in CTMin (25 degrees C to 5 degrees C transfer) was faster than its corresponding loss (5 degrees C to 25 degrees C transfer) in D. quadramaculatus. 4. Comparison of all three Desmognathus species revealed no differences in the CTMin for the 5 degrees C to 25 degrees C time courses, but P. jordani had lower CTMin than corresponding D. quadramaculatus. 5. Our results indicate that the pattern of CTMin acclimation correlates with the constraints acting on salamanders in their natural environment.     

Changes in male courtship intensity and androgen levels during brood cycling in the blenniid fish Rhabdoblennius nitidus

Androgens, for example testosterone, are major hormones that affect male courtship activity, territorial activity, sexual dimorphism, and reproductive tactics; they reach, and are maintained at, a sufficient level to express such sexual traits at the appropriate time for reproductive success. This study examined the effects of androgen levels (testosterone and 11-ketotestosterone) on male brood cycling with two distinct reproductive phases (i.e. courtship and parental phases) of a paternal brooding blenny Rhabdoblennius nitidus. Our study showed that time spent on courtship behaviour and androgen levels decreased with progress of brood cycling. In addition, time spent on courtship behaviour of males administered cyproterone acetate, an anti-androgen, was shorter than that of control males. These results indicate that the brood cycling is affected by the change in androgen levels. The study also showed that androgen levels decreased after acquisition of the eggs, irrespective of available spawning space in the nests. This result suggests that the presence of eggs themselves may be a trigger for the shift from the courtship phase to parental phase.