Leszek Rychlik

Overlap of temporal niches among four sympatric species of shrews

Hypotheses about the dependence of circadian activity from metabolic rate and the segregation of temporal niches among competing species were verified by the study of activity patterns in a shrew community of two semiaquatic species,Neomys anomalus Cabrera, 1907 andN. fodiens (Pennant, 1771), and two terrestrial species,Sorex araneus Linnaeus, 1758 andS. minutus Linnaeus, 1766, co-existing in wet habitats of Białowieża Forest (E Poland). In ten trapping sessions, performed in early summer between 1991 and 2000, traps were open 24 hours continuously and patrolled at 1:00, 5:00, 10:00, 15:00, and 20:00. All the shrew species were most active between 20:00 and 1:00, and least active around mid-day (10:00–15:00). However, activity of the twoSorex species was lower than that of the twoNeomys species in the period 20:00–1:00, but higher in the period 15:00–20:00. BothNeomys species displayed clearly nocturnal, unimodal patterns of activity. In contrast, activity of bothSorex species was relatively evenly distributed over 24 hours and they increased their activity earlier (ie after 15:00) than bothNeomys species (after 20:00). These results confirm the idea that small shrew species with higher metabolic rate have more frequent and more equally distributed activity bouts than large species. Overlap of temporal niches was the highest within genera (99.29% between bothNeomys species and 98.36% between bothSorex species), the lowest betweenN. fodiens andS. araneus (88.26%) andS. minutus (89.34%), and intermediate betweenN. anomalus and bothSorex species (91.78 and 93.34%, respectively). Such high interspecific overlaps in activity suggest a joint-action of other mechanisms that separate ecological niches of these species also in other dimensions (eg food, microhabitat).

Natural infections of small mammals with blood parasites on the borderland of boreal and temperate forest zones

Blood parasites of small mammals living in Białowieża Forest (eastern Poland) were investigated between 1996 and 2002. The following haemoparasite species were found:Trypanosoma (Herpetosoma) evotomys in bank voleClethrionomys glareolus; T. (H.) microti in root voleMicrotus oeconomus; Babesia microti in root vole;Hepatozoon erhardovae in bank vole andHepatozoon sp. in root vole. Some non-identifiedBartonella species were found in bank vole, root vole, field voleMicrotus agrestis, yellow-necked mouseApodemus flavicollis, common shrewSorex araneus, Eurasian water shrewNeomys fodiens, and Mediterranean water shrewN. anomalus. The prevalence and diversity of blood parasites were lower in shrews than small rodents. Totally, 52.0% of bank voles, 50.0% of root voles, 32.5% of common shrews, and 41.2% of Eurasian water shrews were infected with any of the blood parasites. Mixed infections were seldom observed in bank vole (17.3% of investigated individuals) and root vole (14.7%). No animals were infected with three or four parasites simultaneously. Infection of Białowieża small mammals with haemoparasites seemed to be similar to those described in other temperate forest regions rather than boreal ones. Infection rates of rodent species seem to be higher in their typical habitats: for bank vole it was the highest in mixed forest, whereas for root vole in sedge swamp. The results suggest that Arvicolidae play a greater role than Muridae or Soricidae in maintenance ofBabesia andHepatozoon foci in natural environments of central Europe.

Intra- and interspecific antagonistic behaviour in two sympatric species of water shrews:Neomys fodiens andN. anomalus

Ethological mechanisms which diminish the aggression and competition among sympatricNeomys fodiens (Pennant, 1771) [FF] andN. anomalus Cabrera, 1907 [AA] were studied in enclosure conditions. The animals were kept and subsequently tested (11 FF and 10 AA) in 1- or 2-species groups, in enclosures measuring 275×135 cm or 135×135 cm. During 240 h of direct observation, social behaviours and patterns of hiding-place usage by water shrews were recorded. Mutual avoidance (individual occupation of the nest boxes, territoriality, and spatial dispersion in the enclosure), active antagonism (threats, combats, chases) and absence of typical social hierarchy were the most characteristic intraspecific relations of FF. A high level of tolerance and group-occupation of the nest boxes were the most characteristic intraspecific relations of AA in socially stabilized communities. It seems that a group model of life without hierarchic relations is appropriate for this species. In mixed groups, FF dominated over AA. Over the course of time, the stabilization of interspecific relations and decrease of interspecific conflicts were observed. This may be the result of the habituation of one species to the presence of another, and of active avoidance of predominant species by submissive species (assemblage in 1 nest box, maintenance of distance), in which the same, or very similar, communication systems could be involved.

Behavioural mechanisms of conflict avoidance among shrews

Stable co-existence of similar species should be facilitated by mechanisms impairing, besides exploitative, interference competition. We investigated avoidance of intra- and interspecific conflicts in a four-species community of shrews [Sorex minutus Linnaeus, 1766,S. araneus Linnaeus, 1758,Neomys anomalus Cabrera, 1907, andN. fodiens (Pennant, 1771)], using the method of dyadic encounters in a neutral arena. We tested whether the use of passive (habituation, reduction of mobility, increase of inter-individual distance, and stillness) and active (‘to-and-fro’ and ‘keeping distance’ behaviours) forms of conflict avoidance depends on species, size or domination rank. The duration of conflicts was positively correlated with mobility and negatively with inter-individual distance, whereas it was unrelated to time of stillness and the active forms. The repertoire of conflict avoidance mechanisms was not species-specific and the display of these mechanisms depended rather on the size and domination rank of animals participating in a given interaction. In contrast to rodents, shrews did not avoid conflicts by the most passive forms: freeze and stillness reactions. All other forms were used with a higher or lower efficiency by all species. However, consistent with our predictions, large shrews (asN. fodiens) used mainly the passive mechanisms of conflicts avoidance (‘wait-and-see’ strategy), whereas small shrews (asS. minutus) invest proportionally more time in active forms (‘escape’ strategy).

Winter reduction in body mass in a very small, nonhibernating mammal: consequences for heat loss and metabolic rates.

Low temperatures in northern winters are energetically challenging for mammals, and a special energetic burden is expected for diminutive species like shrews, which are among the smallest of mammals. Surprisingly, shrews shrink their body size in winter and reduce body and brain mass, an effect known as Dehnel’s phenomenon, which is suggested to lower absolute energy intake requirements and thereby enhance survival when food availability is low. Yet reduced body size coupled with higher body-surface-to-mass ratio in these tiny mammals may result in thermoregulatory heat production at a given temperature constituting a larger proportion of the total energy expenditure. To evaluate energetic consequences of reduced body size in winter, we investigated common shrews Sorex araneus in northeastern Poland. Average body mass decreased by 19.0% from summer to winter, and mean skull depth decreased by 13.1%. There was no difference in Dehnel’s phenomenon between years despite different weather conditions. The whole-animal thermal conductance (proportional to absolute heat loss) in shrews was 19% lower in winter than in summer; the difference between the two seasons remained significant after correcting for body mass and was caused by improved fur insulation in winter. Thermogenic capacity of shrews, although much enhanced in winter, did not reach its full potential of increase, and this corresponded with relatively mild subnivean temperatures. These findings indicate that, despite their small body size, shrews effectively decrease their costs of thermoregulation. The recorded decrease in body mass from summer to winter resulted in a reduction of overall resting metabolic rate (in thermoneutrality) by 18%. This, combined with the reduced heat loss, should translate to food requirements that are substantially lower than would be the case if shrews did not undergo seasonal decrease in body mass.

Allozyme differentiation in four sympatric species of european shrews (Soricidae: Mammalia)

Abstract Four species of shrews from the Bialowieza Primeval Forest, eastern Poland, were analyzed electrophoretically: Sorex araneus Linnaeus, 1758, Sorex minutus Linnaeus, 1766, Neomys anomalus Cabrera, 1907, and Neomys fodiens (Pennant, 1771). Two of the 25 loci studied were monomorphic for the same allele in all species, nine loci were monomorphic for the same allele in both Sorex species and an alternate allele was fixed in the two species of Neomys . Only one locus ( Pgm-3 ) was diagnostic with a different allele in all species. The mean heterozygosity values in S. araneus and N. fodiens (0.069 and 0.061) were significantly higher than those in S. minutus and N. anomalus (0.021 and 0.029). The Neis genetic distances were small between the two Neomys species (0.123) and between the two Sorex species (0.330). The highest Neis distance was found between S. minutus and N. anomalus (2.487).

The transstadial persistence of tick-borne encephalitis virus in Dermacentor reticulatus ticks in natural conditions

There are a number of reports regarding natural infection of Dermacentor reticulatus ticks with TBE virus; however, the transmission mode of TBE virus in this tick population has not been investigated. This study was conducted in Białowieża Primeval Forest, east Poland. Forty fully engorged nymphs of D. reticulatus were sampled from root voles (Microtus oeconomus). Ticks were kept until molting. All ticks were screened for the presence of TBE virus by nested RT-PCR. Three adult ticks were positive for infection with TBE virus. The present study for the first time demonstrates the possibility of transstadial mode of TBEV transmission in D. reticulatus ticks.

Apparent scarcity of glial fibrillary acidic protein expression in the brain of the pygmy shrew Sorex minutus as revealed by immunocytochemistry

We examined astroglial cells in the brain of the pygmy shrew Sorex minutus (Insectivora). For that purpose we labeled glial fibrillary acidic protein (GFAP) immunohistochemically in brain sections with a polyclonal antibody. Antigen retrieval experiments were performed to counteract formaldehyde fixation masking of GFAP epitopes. Our results showed remarkable paucity of GFAP-immunoreactive cells and fibers in the cerebral cortex and nuclei, as well as in the majority of the diencephalic and mesencephalic structures. In the forebrain, significant numbers of GFAP-containing astrocytes were found only in the ependyma and subventricular zones, superficial part of layer I of the cerebral cortex, and the majority of white matter structures. In the diencephalon, habenular nuclei were rich in GFAP-immunopositive astrocytes and labeled radial fibers were extended between median eminence and the third ventricle. A considerably higher density of labeled astrocytes was detected in the caudal brainstem and cerebellum. In contrast, in the mouse brain, immunoreactive astrocytes were present in large quantities in various structures. Staining of sections of the shrew brain against glutamine synthetase revealed abundance of immunofluorescent astrocytes in many areas, especially in the shrew cerebral cortex. It seems probable that in the shrew brain only a limited fraction of astroglia expresses GFAP, while other astroglial cells can be detected with different markers. It is possible that the rodent type of astroglial GFAP expression might not be common to insectivores and probably to some other mammalian orders.