Research Article
The proboscis of tapirs (Mammalia: Perissodactyla): a case study in novel narial anatomy
- Lawrence M. Witmer, Scott D. Sampson, Nikos Solounias
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- Published online by Cambridge University Press:
- 27 February 2001, pp. 249-267
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The trunk-like proboscis of tapirs provides a prime case study in the evolution of anatomical novelty. Morphological study of this unique structure was undertaken employing several specimens and a combination of analytical techniques: gross anatomical dissection, radiographic imaging and histological sectioning. Evolution of the proboscis of tapirs entailed wholesale transformation of the narial apparatus and facial architecture relative to perissodactyl outgroups. This transformation involved retraction and reduction of the bony and cartilaginous facial skeleton, such that several structures present in outgroups are completely absent in tapirs, including cartilages surrounding the nasal vestibule (e.g. alar and medial accessory cartilages, rostral portion of the nasal septum) and associated musculature (dilatator naris apicalis, lateralis nasi pars ventralis). At the same time, soft tissues surrounding the upper lip and nose became elaborated to form a mobile, fleshy proboscis. Several key facial muscles (e.g. levator labii superioris, levator nasolabialis, caninus, lateralis nasi) have been co-opted to function in movement of the proboscis. The nasal vestibule is expanded and occupies approximately 75% of the nasal cavity. Vestibular expansion has compressed and simplified caudal components of the nasal cavity (e.g. reduction of dorsal and middle nasal conchae, loss of plica recta and plica basalis). The airway has become dorsally arched causing the ventral conchal complex to become inclined relative to the long axis of the skull. A few anatomical enigmas remain, such as the complicated maxilloturbinate that rostrally contacts the nasal septum and vomeronasal organ. Similarly, the meatal diverticulum, despite being both ancient and anatomically complex, has no obvious functional significance; it is clear that it is not homologous to the nasal diverticulum of horses and other equids. The reduction of the osseocartilaginous portion of the proboscis, coupled with expansion of the muscular and connective tissue components, has resulted in an organ that is best interpreted as a muscular hydrostat.
Ecological traits and conservation biology of five fossorial ‘sand-swimming’ snake species (Simoselaps: Elapidae) in south-western Australia
- Richard A. How, Richard Shine
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- 27 February 2001, pp. 269-282
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Although small, nocturnal, fossorial snakes are a significant component of the reptile fauna in many parts of the world, their biology is poorly known. An 11-year pit-trapping study in urban bushland remnants near the city of Perth, Western Australia, provided data from >500 captures of small fossorial snakes of the genus Simoselaps. The five species differed in relative abundances and in distribution, both among localities and among habitats within a single locality. For example, three saurophagous taxa (Simoselaps bertholdi, S. bimaculatus, S. calonotos) were most abundant in Banksia woodland, whereas two species that feed on reptile eggs (S. semifasciatus, S. fasciolatus) were most abundant in coastal heath. Capture rates for most species were low (for three of the five species, <one specimen captured per 1000 trapdays), and these taxa may be genuinely rare in most of the habitats that we surveyed. Activity patterns were highly seasonal, with little activity in winter or in midsummer. The two oophagous species showed a more restricted activity period (late spring–early summer) than did species with broader dietary habits. In the most abundant taxon (Simoselaps bertholdi), males were active mainly during spring (the mating season) and females during autumn, after oviposition. Capture rates and body condition of the captured snakes varied substantially among seasons and across years. Low capture rates mean that very prolonged surveys are needed to determine reliably whether or not a taxon occurs on any given site.
Group dynamics of humpback dolphins (Sousa chinensis) in the Algoa Bay region, South Africa
- Leszek Karczmarski
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- Published online by Cambridge University Press:
- 27 February 2001, pp. 283-293
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Group dynamics of humpback dolphins Sousa chinensis inhabiting the Algoa Bay region on the south Eastern Cape coast of South Africa, were investigated by means of boat-based photo-identification surveys undertaken between May 1991 and May 1994. Groups of humpback dolphins varied in size from three to 24 animals (
x = seven), with adults representing almost two-thirds of the group members. Births occurred predominantly in summer. Some females, however, may also cycle outside of the apparent summer breeding season, perhaps indicating a secondary winter season. Circumstantial evidence suggests a minimum of a 3-year calving interval. Maternal care lasts at least 3–4 years, but female–calf separation is seemingly not related to the female's next pregnancy. Humpback dolphins displayed varying degrees of residence/fidelity to Algoa Bay. Although a few individuals may possibly be classified as ‘resident’, most dolphins were infrequent visitors in the Bay and seem to be transient. The social system of humpback dolphins appears to be fluid with only casual and short-lasting affiliations. Strong bonds between individuals other than mothers and calves are uncommon. Lack of consistency in the group membership appears to be the general pattern. The weak site fidelity and possibly extensive long-range movement of the majority of dolphins may contribute to the dynamic nature of humpback dolphin groups. There is probably some form of segregation between sex and/or age classes among humpback dolphins in Eastern Cape waters. The nature and extent of this segregation, however, is not yet sufficiently understood. It is likely that the degree of site fidelity displayed by female humpback dolphins is related to their reproductive stage and increases during the nursing period. Mate-searching behaviour of male humpback dolphins is the most likely reproductive strategy of the species.
Phylogenetic and taxonomical relationships of the eight species in the melanogaster subgroup of the genus Drosophila (Sophophora) based on the electrophoretic mobility of the major chorion proteins and the eggshell ultrastructure
- Maria C. Kalantzi-Makri, Ioannis P. Trougakos, Triantafyllos P. Tafas, John Sourdis, Lukas H. Margaritis
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- 27 February 2001, pp. 295-306
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The eight species of the melanogaster species subgroup of the genus Drosophila were compared to each other with respect to: (a) the electrophoretic mobility of the major chorion proteins under SDS–PAGE; (b) the overall eggshell ultrastructure. The phylogenetic tree obtained using the first criterion showed that the eight species are divided into three clusters that contain the species of the melanogaster complex (D. melanogaster, D. sechellia, D. mauritiana and D. simulans), the two species of the yakuba complex (D. yakuba and D. teissieri) and the erecta complex species (D. erecta and D. orena). This clustering is in agreement to the generally accepted phylogeny of the subgroup based on exclusive criteria. The eggshell ultrastructure of the species at the main body of the egg revealed that the phylogenetically closely related species exhibit similar eggshell structure. We conclude that the criteria applied in this study could be eventually used in groups with unknown phylogeny to trace the phylogenetic relationships of the species.
Developmental and territorial aspects of suckling behaviour in the domestic pig (Sus scrofa f. domestica)
- B. Puppe, A. Tuchscherer
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- 27 February 2001, pp. 307-313
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The suckling behaviour of 327 piglets from 39 litters of primiparous domestic sows Sus scrofa f. domestica was analysed regarding the development of territorial suckling consistency during a 5-week suckling period. Additionally, the consistent use of a certain teat pair (1–7) during the whole suckling period was related to weight gain within the suckling period and to social dominance after weaning and mixing of the piglets. The piglets increased their suckling stability consistent with a three-parametric exponential model delivering a time-dependency that is typical for learning curves and which may be a useful methodological approach for studies on behavioural ontogeny. Thus, the development of suckling stability can be regarded as a continuous learning process. After extremely rapid learning within the first 4 days of life a moderate increase of suckling consistency with final stabilization occurred within the second week. After this time, the piglets maintained a high suckling consistency of about 95% up to the end of the observed suckling period. The anterior teats were preferred, whereas piglets suckling at the posterior teats showed a tendency to have lower weight gains and poorer dominance values. However, the lowest suckling stability (consistency) was found with piglets occupying the middle teats. The study shows that individual differences (e.g. weight gain, position in a social hierarchy) have their roots in early ontogenetic characteristics of the neonates (e.g. suckling behaviour, physical strength). The results are discussed with regard to findings and interpretations of behavioural ontogeny in pigs.
Life histories of the Insectivora: the role of phylogeny, metabolism and sex differences
- Matthew R. E. Symonds
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- Published online by Cambridge University Press:
- 27 February 2001, pp. 315-337
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Life-history data for 63 species from the mammalian order Insectivora have been collated from the literature. These data were analysed for covariation and for correlations with body mass, brain mass and mass-specific resting metabolic rate. An independent contrasts method has been used to remove the effect of phylogeny. Due to uncertainties surrounding their evolutionary relationships, 22 different phylogenies of insectivores have been used as a basis for comparative analysis. The results show that several key correlations between life-history variables are only significant when certain phylogenies are used, highlighting the problems of such analyses when the phylogeny used is inaccurate. After removing the effect of phylogeny, relatively few significant correlations remain. Insectivores that have a high body mass have relatively lower metabolic rates, longer lifespans and longer gestation lengths. There is some support for a fast–slow continuum in insectivore life-history evolution: there are some significant positive correlations between measures of growth rates (e.g. gestation length and age at weaning) and lifespan, and some negative correlations between growth rates and measures of reproductive output. It is suggested that the seasonality of life of many insectivores may have played an influential role in the evolution of the group, in particular in delaying the onset of sexual maturity. There is little indication that brain size influences life-history evolution in this order, but metabolism may play an important role. The energetic requirements of maintaining high metabolic rates in small mammals such as insectivores may be constraining life histories to a greater extent than occurs in larger mammals. This effect may have obscured the relationship between metabolic rate and life histories in wider inter-order analyses. Finally, there is considerable evidence that sex differences play a large role in shaping insectivore evolution, and it is suggested that this factor must be considered more often in future studies of mammalian life histories in general.
The fighting technique of male fallow deer (Dama dama): an analysis of agonistic interactions during the rut
- Valeria Mattiangeli, Silvana Mattiello, Marina Verga
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- Published online by Cambridge University Press:
- 27 February 2001, pp. 339-346
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A population of 560 individually recognizable European fallow deer (Dama dama) was studied in Phoenix Park, Dublin, Ireland, during the rut in order to examine buck fighting technique in relation to age and rank. The percent usage and association among behavioural elements used were analysed in 162 fights and 82 sparrings. During fight sequences, high-ranking bucks tend to use a higher percentage of low-risk behaviours (all behaviours which do not involve antler contact), especially when they interact with lower-ranking deer. Fights between combatants of the same rank class show a higher use of high-risk behaviours (characterized by antler contact), and cluster analysis of a preceding–following matrix revealed that high-risk behaviours are closely associated with each other in these interactions. During sparrings, the low intensity of the interaction allows the deer to test their ability and strength using a higher percentage of behaviours which are generally considered high-risk, without risking injury. The data suggest that the percent usage and the sequences of behavioural elements performed in agonistic interactions are affected by deer rank, probably in relation to the different risks and benefits that different hierarchical positions can offer.
Seasonal changes in the foraging behaviour of Leisler's bats (Nyctalus leisleri) in Ireland as revealed by radio-telemetry
- C. B. Shiel, R. E. Shiel, J. S. Fairley
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- Published online by Cambridge University Press:
- 27 February 2001, pp. 347-358
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Leisler's bats Nyctalus leisleri, from two nursery colonies were radio-tracked from April to August in 2 years. They commuted directly to foraging sites up to 13.4 km away at speeds often exceeding 40 km h-1. Except during lactation, individuals occasionally day-roosted away from the nursery in buildings or hollow trees. These were also sometimes used as night roosts, especially during rain, which also stimulated return to the nursery. On most nights the first flight lasted the longest. During preparturition, when distances from roost to foraging site were greatest, and time spent flying per night (tF) was least, there was often only one flight, with a second towards dawn on warmer nights, and occasionally a third. After parturition, in June, tF and the number of flights (NF) increased up to weaning, corresponding to rising energy demands. But distances to foraging sites fell, rising only slightly in postlactation, presumably because insects, and thus feeding patches, were more abundant than in preparturition. NF fell to preparturition levels after lactation. Exploratory behaviour by juveniles intensified with time – distances to feeding sites, tF and the total distance flown each night all increased; the distance flown each night was eventually greater than any recorded for adults. Roosting away from the nursery was also more common. Before dispersal, there were extended migrations by juveniles outside the summer range of the adults. Distances moved by two dispersed juveniles exceeded 34 km. Two-thirds of the recorded foraging time was over pasture or drainage canals. Foraging over other habitats, particularly lake and conifer forest, was greatest in preparturition. Other habitats foraged included lights, estuary, stream, beach and dunes.
Echolocation signal design in Kerivoulinae and Murininae (Chiroptera: Vespertilionidae) from Malaysia
- Tigga Kingston, Gareth Jones, Zubaid Akbar, Thomas H. Kunz
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- 27 February 2001, pp. 359-374
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Echolocation signal design in nine syntopic vespertilionid bats from the Malaysian rain forest understorey was studied. Four species of Kerivoula, two species of Phoniscus (Kerivoulinae) and three species of Murina (Murininae) all emitted calls that are typical of species that glean insects from surfaces: broadband, frequency-modulated (FM) calls of low intensity and short duration. However, calls were highly distinctive in the use of very large bandwidths (range: 89–123 kHz) and extremely high frequencies (start frequency 152–180 kHz; end frequency 43–86 kHz). Furthermore, calls were produced in groups of 2–15 at very high pulse repetition rates (37–105 Hz). The functional significance of these characteristics with respect to foraging strategy is discussed. Large signal bandwidths facilitate highly accurate target localization in terms of both range and angle estimation and can thus be interpreted as an adaptation to foraging in the highly cluttered environment of the forest understorey. The use of high frequencies so far in excess of those seen in other FM gleaning bats of the same size is less easily explained, but may represent a mechanism by which these species distinguish prey items using echolocation alone, without recourse to visual or auditory cues. Species exhibited differences in echolocation parameters, particularly the end frequency and the number of calls per group of calls. The two subfamilies differed from each other in multivariate space derived from echolocation parameters; calls of the Murininae were of lower frequency than the Kerivoulinae and were typically produced in smaller groups or singly. Within the subfamilies there was considerable overlap between species of Murina, but the Kerivoula spp. were clearly distinct from one another but not from the two Phoniscus spp.