Chromosomal homologies of the genera Vespertilio,Plecotus and Barbastella (Chiroptera: Vespertilionidae)

Q-banding (and partly G-banding) patterns of chromosomes were studied in three species of vespertilionid bats: Vespertilio murinus, Plecotus auritus and Barbastella barbastellus. The chromosomes of these species were identified and compared on the basis of the G-band diagram of Bickham (Cytologia 44, 1979) for the genus Myotis. The additional metacentric chromosomes which Vespertilio shows as compared to Myotis, differ in arm arrangements from those found in Plecotus and Barbastella. Banding patterns of P. auritus and B. barbastellus are nearly identical. It is remarkable that the smallest metacentric chromosome of P. auritus differs clearly in its banding pattern from that of the American P. townsendi. The significance of the results obtained for the systematic position of these species is discussed.     

Post-natal growth, mother-infant interactions and development of vocalizations in the vespertilionid bat Plecotus auritus

Female brown long-eared bats Plecotus auritus nurse their own young selectively. Choice experiments suggested that females recognized infants by using both olfactory and acoustic cues. Infant isolation calls (i-calls) showed individual variation, and vocal signatures allowed the females to recognize and suckle their own pup. As the pups grew, the i-calls increased in frequency and decreased in duration, and the i-calls appeared to change into orientation cries. Mothers were able to recognize recent calls of their own babies in preference to older calls. Echolocation sounds were distinguishable statistically between lactating females, and the behaviour of the babies during choice experiments suggested that in this species a mutual acoustic recognition occurs. The growth and development of wing shape in P. auritus are described.     

Genic differentiation and phylogenetic relationships within two New World bat genera

Phylogenetic placement of taxa within the two New World leaf-nosed bat genera, Tonatia and Micronycteris was carried out using allozymic characters. Of particular interest were the relationships between the karyotypically megaevolved species and their proposed congeners. Neither of the two karyotypically megaevolved Tonatia species shared synapomorphic allozymes with the other Tonatia taxa. In contrast, the three Micronycteris species identified as having undergone this unusual mode of karyotypic evolution fall within a common evolutionary assemblage with all but one of their congeners. Measures of genic similarity among the Tonatia and Micronycteris species yield relatively low Nei identity values. No correlation is found when the number of karyotypic and allozymic changes since a common ancestor are compared.     

Echolocation by the long-eared bat,Plecotus phyllotis

Summary The echolocating bat,Plecotus phyllotis (Vespertilionidae), uses long-CF/FM and FM sonar sounds in different situations. The CF component in long-CF/FM sounds occurs at 27 kHz and has a duration of 20 to 200 ms. The FM component sweeps down from 24 to 12 kHz, with a prominent second harmonic from 40 to 22 kHz. This second harmonic sweep is interrupted at 28 to 25 kHz, providing a notch in the spectrum of the FM component at the CF frequency. This notch probably permits isolation of CF and FM components in echoes for separate processing, thus avoiding mutual interference with the different kinds of target information the two components convey. The FM component is also used without the CF component as a sonar sound. Two other FM orientation sounds are used when the bat is in a confined space such as a room. One contains only the second and fourth harmonics of the 24 to 12 kHz fundamental sweep, while the other contains only the fifth harmonic. This bat's repertoire of sonar sounds closely matches the hearing capacities of the genus.We thank P.H. Dolkart and W.A. Lavender, of Washington University, and the Nevada State Parks Department for their assistance. This research was supported by Grant # BMS-72-02351-A01 from the National Science Foundation.     

On the phylogenetic relationships among tetraphyllidean,lecanicephalidean and diphyllidean tapeworm genera

This study had two main objectives: (1) to construct an extensive, explicit list of characters and character states that might serve as a starting point, and perhaps even a model, for the compilation of a more complete list of characters for all cestode taxa; and (2) to use this character list to generate a hypothesis of the phylogenetic relationships among species representing most of the tetraphyllidean, lecanicephalidean and diphyllidean genera. Specimens of one species in each of 48 genera of tetraphyllideans, eight genera of lecanicephalideans, the three genera of diphyllideans, two genera of proteocephalideans and two genera of trypanorhynchs, were examined as whole-mounts and sections, with light and scanning electron microscopy. A list of 120 morphological characters was compiled. Four phylogenetic analyses were conducted using PAUP* and/or NONA. The first was a comprehensive analysis with the 56 tetraphyllidean and lecanicephalidean species as ingroups and the remaining seven species as outgroups. The second was an analysis of the three diphyllidean species as ingroups and the two proteocephalidean and the two trypanorhynch species as outgroups. The third was an analysis of the eight lecanicephalidean species and the tetraphyllideans Echeneibothrium sp. and Pseudanthobothrium n. sp. as ingroups and an outgroup consisting of the seven species used as outgroups in the first analysis. In the fourth analysis, the ingroup consisted of the 14 hooked tetraphyllideans (onchobothriids), and the outgroup consisted of the seven species used as outgroups in the first analysis. The results of these analyses support the following phylogenetic hypotheses: The diphyllideans are monophyletic and Echinobothrium n. sp. and Macrobothridium sp. are more closely related to one another than either is to Ditrachybothridium macrocephalum. The tetraphyllideans, lecanicephalideans and proteocephalideans are more closely related to each other than they are to the diphyllideans or the trypanorhynchs. The ordinal status of the lecanicephalideans is dubious. The lecanicephalidean species are more closely related to some of the tetraphyllidean taxa than these tetraphyllidean taxa are to the remainder of the tetraphyllidean taxa. The proteocephalideans appear to belong within the tetraphyllidean clade. The tetraphyllidean species Echeneibothrium sp. and Pseudanthobothrium n. sp. are members of the lecanicephalidean clade. The position of Discobothrium n. sp. within the lecanicephalideans is dubious. Within the tetraphyllideans, the non-acetabulate species Litobothrium daileyi, Disculiceps galapagoensis and Cathetocephalus sp. are the most basal members of the group. The family Onchobothriidae is monophyletic, as it is currently defined. Within the onchobothriids, the uniloculate species are basal to the multiloculate species; the species with unipronged hooks are basal to the species with multipronged hooks. Although relationships among the phyllobothriids, as they are currently defined, remain poorly resolved, the family Phyllobothriidae is not monophyletic. These results suggest that some aspects of the classification of the lecanicephalidean and tetraphyllidean taxa require revision. However, such revision should be based on further analyses including a broader representation of the genera and species in these groups.     

Evaluating the monophyly and phylogenetic relationships of Brassolini genera (Lepidoptera, Nymphalidae)

Abstract.  This study uses 80 morphological characters and cladistic analysis to evaluate the monophyly and phylogenetic relationships of 18 genera that constitute the butterfly tribe Brassolini. Most characters derive from genitalia, confirming previous generic definitions based mainly on wing characters, and showing that 16 of 18 genera are monophyletic. Mimoblepia Casagrande, syn.n. was subsumed within Opoptera Aurivillius to address the paraphyly of the latter, but resolution of the status of Aponarope Casagrande requires further study. The results suggest that the Brassolini includes six suprageneric groups/clades. Although this study verifies some genus-level relationships put forward over 100 years ago, some new hypotheses of relationships are proposed. Tracing larval host plant use onto the Brassolini phylogeny indicates that species in this tribe retain the use of Arecaceae and Poaceae from their 'satyroid' ancestors.     

Phylogenetic relationships of vespertilionid genera (Mammalia: Chiroptera) as revealed by karyological analysis

Banded karyotypes of 50 species belonging to 23 genera were analyzed. The diploid chromosome numbers ranged from 26 to 50. For karyotypic comparison we used a 44 chromosomes karyotype, consisting of 4 metacentric and 17 acrocentric autosomes, as “basic karyotype”. Almost all of its 25 autosomal arms could be identified in each individual karyotype. In 8 chromosomes, i. e. 1/2, 7, 11, 12, 13, 15, 23 and X, small inversions were detected. As a result, each of the chromosomes mentioned occurs in two states which differ slightly in their banding patterns. These were used as character states in the cladistic analysis together with other chromosomal rearrangements. The implications drawn from the cladogram obtained are: The Miniopterinae clearly belong to the Vespertilionidae but are the first to branch off from the common stem. The tribe Myotini should be raised to the rank of a subfamily. Within the largest subfamily Vespertilioninae, one autapomorphic chromosomal character was found for each of the tribes Vespertilionini and Pipistrellini. In addition, both tribes are distinguished from the other Vespertilioninae tribes by two synapomorphic features. These results allow for the first time an unequivocal classification of the systematically difficult “pipistrelloid” species. The species of the genus Pipistrellus (sensu Hill and Harrison 1987) are spread over the Pipistrelllni and Vespertilionini. We therefore suggest the splitting of this heterogenous genus into at least four genera. Only the members of the previous subgenus Pipistrellus constitute the genus Pipistrellus and belong to the Pipistrellini. The previous subgenera Hypsugo, Vespadelus and Falsistrellus, given generic rank in some recent studies, belong to the tribe Vespertilionini and are not closely related to Pipistrellus. For the genera Eptesicus and Hesperoptenus, which belong neither to the Vespertilionini nor to the Pipistrellini, the tribe Eptesicini was established. The phylogenetic relations of this tribe and the status of the presumably polyphyletic tribe “Nycticeiini” could not be solved.     

Evaporative water loss in two sympatric species of vespertilionid bat, Plecotus auritus and Myotis daubentoni: relation to foraging mode and implications for roost site selection

Simultaneous measures of oxygen consumption and evaporative water loss (EWL) were made in two species of temperate-zone vespertilionid bat ( Plecotus auritus and Myotis daubentoni ; mean body mass 9.12 and 10.12g, respectively) at ambient temperatures (T_a) of 5, 15 and 25°C and variable vapour pressure deficit. EWL was directly dependent on vapour pressure deficit and oxygen consumption and inversely dependent on T_a. EWL was significantly greater in P. auritus than in M. daubentoni. A model for EWL in P. auritus under a variety of environmental conditions (5–25°C and 20–80% relative humidity) suggested that EWL from bats in shallow summer torpor will be lowest at low T_a, and that, except at low (> 50%) relative humidity, EWL from euthermic bats will be lowest at high T_a. At low relative humidity (< 20%), resting bats could lose over 30% of body mass per day (24 h) through evaporation. At high T_a (> 25°C), EWL from euthermic bats could be over 65% lower at high (> 80%) compared to low (< 20%) relative humidity. In bats in shallow summer torpor at low (5°C) T_a the equivalent saving was > 96%. At low relative humidity predicted EWL from bats in shallow summer torpor was 34 to 81% of that from euthermic bats, and at low T_a and high relative humidity was only 2%. In the wild, M. daubentoni has freer access to drinking water than does P. auritus and yet EWL at rest was higher in the latter species. We suggest that post-prandial dumping of urinary water by M. daubentoni leads to a limit in the amount of body water available to this species to cover evaporative losses once within the day roost, which in turn has led to an adaptation of physiology towards the minimization of EWL when at rest.     

Population genetic structure and gene flow in a gleaning bat,Plecotus auritus

During summer the brown long-eared bat Plecotus auritus (Vespertilionidae) forms stable colonies, comprised of both adult females and males and young of the year. A long-term ringing study conducted in north-east Scotland has established that little movement occurs among colonies and that both sexes are recruited into their natal colony. The aim of the present study was to investigate, using microsatellite DNA markers, if genetic structure within the population reflects the spatial structure indicated by ringing. Inter-colony F_ST estimates obtained for all colony members, and for females and males separately, were low (0.019, 0.026 and 0.011, respectively), but all values differed significantly from zero. These data indicate high gene flow between colonies, although some coancestry among colony members is evident in both sexes. On combining the ringing and genetic data, it is concluded that gene flow occurs via extra-colony copulation, rather than natal dispersal, and that each colony behaves as a distinct subpopulation. Microgeographical genetic isolation by distance was demonstrated for, to our knowledge, the first time in a bat species, and found to be apparent both across the entire study area and along one river valley. The results suggest that extensive macrogeographical population genetic structure may be evident across the species'' range.     

Induction of ovulation by clomiphene citrate in the Indian vespertilionid bat, Scotophilus heathi

The ovulation induction property of clomiphene citrate (CC) and human chorionic gonadotropin (hCG) was studied in Scotophilius heathi, an Indian tropical vespertilionid bat, during the period of delayed ovulation between December to early January. The results of the study showed that 10 microg of CC alone was ineffective to induce ovulation, whereas 100 microg CC and 10 IU hCG alone induced ovulation. A significant (P < 0.01) increase in the ovulation rate was observed when 10 microg CC followed by 10 IU hCG, compared to 10 IU hCG and 100 microg CC alone groups. Finally, CC at a 100 microg dose, followed by 10 IU hCG, produced superovulation (14.00 +/- 0. 70), which is significantly different in comparison to all other groups. This is the first report of ovulation induced by CC in the Indian tropical bat as well as in any animal model that exhibits temporary anovulation similar to polycystic ovary syndrome (PCOD) during the normal physiology of reproduction.     

Mating patterns, relatedness and the basis of natal philopatry in the brown long-eared bat, Plecotus auritus

The brown long-eared bat, Plecotus auritus, is unusual among temperate zone bats in that summer maternity colonies are composed of adult males and females, with both sexes displaying natal philopatry and long-term association with a colony. Here, we describe the use of microsatellite analysis to investigate colony relatedness and mating patterns, with the aim of identifying the evolutionary determinants of social organization in P. auritus. Mean colony relatedness was found to be low (R=0.033 +/- 0.002), with pairwise estimates of R within colonies ranging from -0.4 to 0.9. The proportion of young fathered by males in their own colony was investigated using a Bayesian approach, incorporating parameters detailing the number of untyped individuals. This analysis revealed that most offspring were fathered by males originating from a different colony to their own. In addition, we determined that the number of paternal half-sibs among cohorts of young was low, inferring little or no skew in male reproductive success. The results of this study suggest that kin selection cannot account for colony stability and natal philopatry in P. auritus, which may instead be explained by advantages accrued through the use of familiar and successful roost sites, and through long-term associations with conspecifics. Moreover, because the underlying causes of male natal dispersal in mammals, such as risk of inbreeding or competition for mates, appear to be avoided via extra-colony copulation and low male reproductive skew, both P. auritus males and females are able to benefit from long-term association with the natal colony.     

The phylogenetic relationships of the physaloid genera (Solanaceae) based on morphological data

A group of genera, e.g., Chamaesaracha, Leucophysalis, Physaliastrum, Margaranthus, and Withania, in the subfamily Solanoideae (Solanaceae) is centered around the genus Physalis and has been named the physaloid group. It comprises a number of small and often poorly known genera, sometimes seen as united with Physalis and/or each other. A hypothesis of the phylogenetic relationships within this group, based on parsimony analyses of morphological data, is here presented for the first time. The result is discussed in relation to prevailing generic circumsciptions and taxonomic consequences. It is also compared with hypotheses of relationships based on cpDNA data.     

Frequency sensitivity and directional hearing in the gleaning bat,Plecotus auritus (Linnaeus 1758)

1. The neural audiogram of the common long-eared bat, Plecotus auritus was recorded from the inferior colliculus (IC). The most sensitive best frequency (BF) thresholds for single neurones are below 0 dB SPL between 7-20 kHz, reaching a best value of -20 dB SPL between 12-20 kHz. The lower and upper limits of hearing occur at 3 kHz and 63 kHz, respectively, based on BF thresholds at 80 dB SPL. BF threshold sensitivities are about 10 dB SPL between 25-50 kHz, corresponding to the energy band of the sonar pulse (26-78 kHz). The tonotopic organization of the central nucleus of the IC (ICC) reveals that neurones with BFs below 20 kHz are disproportionately represented, occupying about 30% of ICC volume, occurring in the more rostral and lateral regions of the nucleus. 2. The acoustical gain of the external ear reaches a peak of about 20 dB between 8-20 kHz. The gain of the pinna increases rapidly above 4 kHz, to a peak of about 15 dB at 7-12 kHz. The pinna gain curve is similar to that of a simple, finite length acoustic horn; expected horn gain is calculated from the average dimensions of the pinna. 3. The directional properties of the external ear are based on sound diffraction by the pinna mouth, which, to a first approximation, is equivalent to an elliptical opening due to the elongated shape of the pinna. The spatial receptive field properties for IC neurones are related to the directional properties of the pinna. The position of the acoustic axis of the pinna and the best position (BP) of spatial receptive fields are both about 25 degrees from the midline between 8-30 kHz but approach the midline to 8 degrees at 45 kHz. In elevation, the acoustic axis and the BP of receptive fields move upwards by 20 degrees between 9-25 kHz, remaining stationary for frequencies up to 60 kHz. 4. The extremely high auditory sensitivity shown by the audiogram and the directionality of hearing are discussed in terms of the adaptation of the auditory system to low frequencies and the role of a large pinna in P. auritus. The functional significance of low frequency hearing in P. auritus is discussed in relation to hunting for prey by listening and is compared to other gleaning species.     

Utility of arginine kinase for resolution of phylogenetic relationships among Brachyuran genera and families

The molecular phylogenetics of decapod crustaceans has been based on sequence data from a limited number of genes. These have included rapidly evolving mitochondrial genes, which are most appropriate for studies of closely related species, and slowly evolving nuclear ribosomal RNA genes, which have been most useful for resolution of deep branches within the Decapoda. Here we examine the utility of the nuclear gene that encodes arginine kinase for phylogenetic reconstruction at intermediate levels (relationships among genera and families) within the decapod infraorder Brachyura (the true crabs). Analyses based on arginine kinase sequences were compared and combined with those for the mitochondrial cytochrome oxidase I gene. All of the genera in our taxon sample were resolved with high support with arginine kinase data alone. However, some of these genera were grouped into clades that are in conflict with recognized brachyuran families. A phylogeny based on cytochrome oxidase I was consistent with the arginine kinase phylogeny, but with weaker support. A recently proposed measure of phylogenetic informativeness indicated that arginine kinase was generally more informative than cytochrome oxidase I for relationships above the level of genus. Combined analysis of data from both genes provided strong support for clades that are in conflict with current assignments of genera to the families Epialtidae, Mithracidae, Pisidae, and Portunidae.     

Analysis of the diet of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus in the West of Ireland

The diets of Natterer's bat Myotis nattereri and the common long-eared bat Plecotus auritus were investigated at a nursery colony of each species by analysis of droppings collected monthly from May to September. Respectively, 68% and 42% of the diets comprised items presumed to have been gleaned from foliage or other surfaces: diurnal insects, insects which rarely fly, and non-flying arthropods. Such surfaces included the ground: centipedes were eaten by both bats, and the muscid, Scatophaga stercoraria , usually associated with cattle dung, was a common prey. Indeed, the prevalence of cattle-farming around both of the bat roosts almost certainly contributed indirectly, in the form of S. stercoraria , to a significant part of the Diptera consumed. The chief food of M. nattereri was the larger Diptera, but Trichoptera, Hymenoptera and Arachnida were also important. Lepidoptera, Coleoptera and Hemiptera were minor prey, with Dermaptera and Chilopoda taken occasionally. Diptera, closely followed by Lepidoptera, together accounted for nearly two-thirds of the diet of P. auritus. Also taken, in descending order of importance, were Trichoptera, Arachnida, Chilopoda, Coleoptera, Dermaptera, Hymenoptera and Hemiptera. Plecotus auritus , but not M. nattereri , was evidently able to take a few small insects such as aphids and lesser nematoceran Diptera.     

Copepod parasites of the fatheads (pisces, psychrolutidae) and their implication on the phylogenetic relationships of psychrolutid genera

Nine species of psychrolutids kept in the Marine Zoology of Hokkaido University were found to carry eight species of copepod parasites. The parasites and their hosts are: Bobkabata kabatabobbus Hogans and Benz, 1990 on Malacocottus zonurus Bean; Chondracanthus parvus n. sp. on Eurymen gyrinus Gilbert and Burke; Chondracanthus yabei n. sp. on Dasycottus setiger Bean and M. zonurus; Ch. yanezi Atria, 1980 on Psychrolutes phrictus Stein and Bond; Caligus similis n. sp. on Neophrynichthys latus (Hutton); Clavella adunca (Str?m, 1762) on M. zonurus; Neobrachiella amphipacifica Ho, 1982 on Ambophthalmos angustus (Nelson), Cottunculus sp., D. setiger, Ebinania brephocephala (Jordan and Starks), E. vermiculata Sakamoto, and P. phrictus; and Naobranchia occidentalis Wilson, 1915 on D. setiger and M. zonurus. Chondracanthus parvus is closest to Ch. deltoideus Fraser, 1920, but differs from it in having only one pair of small knobs on the head and carrying a pair of lateral processes on the second pediger. Chondracanthus yabei resembles Ch. yanezi Atria, 1980, but can be distinguished from it by the presence of three, low protrusions on the mid-dorsal surface of the trunk; besides, maxillule also shows difference. As the name indicates, C. similis resembles several species of Caligus that bear a short abdomen and a formula of I; IV on the exopod of leg 4. However, it can be separated from them by the combination of the following characters: a genital complex distinctly smaller than the cephalothoracic shield, a pair of relatively short caudal rami, a smooth dentiform process on maxillule, and a pair of truncate tines on sternal furca. Analysis of the occurrence of Chondracanthus on the psychrolutid fishes shows that the phylogeny of Chondracanthus is in congruence with that of the Psychrolutidae. It implies that parasitism of Chondracanthus occurred after the Dasycottus clade diverged off the main stock of the Psychrolutidae and the occurrence of Ch. yabei on D. setiger is resulted from a later colonization.