Home range area in the tortoise Testudo hermanni in relation to habitat complexity: implications for conservation of biodiversity

The tortoise Testudo hermanni is endangered by habitat fragmentation and loss in western Europe, where its high public profile and specific conservation projects make this a flagship species. Studies of movement in the peak reproductive season (June) showed that home ranges were substantially larger in France than in Greece. This difference was due to the intensity of use of the home range, not to the distance moved which was remarkably similar in the two areas. There was no sexual difference in home range area. The home range was therefore not large in France because of movement to nesting sites, but rather for utilization of the greater habitat complexity there. The need for reserves to include different vegetation types makes conservation of T. hermanni in France more difficult. Conversely, the need for large reserves increases its value as an umbrella species for conservation of biodiversity.     

Identification and characterization of microsatellite markers in Hermann's tortoise (Testudo hermanni,Testudinidae)

The isolation and characterization of six polymorphic loci from a Testudo hermanni genomic library is reported. Enrichment was performed for AC but four of the characterized microsatellites present also an additional motif. Variability was tested on populations of the two recognized subspecies, Testudo hermanni hermanni and Testudo hermanni boettgeri. For one locus, a size range specific for the subspecies T. h. hermanni was observed. These are the first primers identified directly in the genome of T. hermanni     

Daily movements and home range of Eastern Hermann's Tortoise,Testudo hermanni boettgeri (Reptilia: Testudines)

Understanding the movements of animals in the wild is critical for providing important information regarding their conservation and management. With this aim in mind, we studied a population of Testudo hermanni in their natural habitat using micro GPS devices. Additionally, using VHF transmitters we tracked six individuals (four females and two males) from May to August 2015. The overall home range size varied from 813 and 123,567 m². Home range size did not differ significantly between males and females, and was also independent of the size of the individual (expressed as straight carapace length). Individual movements were confined to short distances (males 25.7 m; females 18.5 m). Activity levels were highest in July and August for males and May and July for females.     

A rangewide phylogeography of Hermann's tortoise, Testudo hermanni (Reptilia: Testudines: Testudinidae): implications for taxonomy

Hermann's tortoise (Testudo hermanni), the best-known western Palaearctic tortoise species, has a rare natural distribution pattern comprising the Mediterranean areas of the Iberian, Apennine, and Balkan Peninsulas, as well as Sicily, Corsica and Sardinia. The western part of this range is traditionally considered habitat for T. h. hermanni, while T. h. boettgeri occurs in the Balkans. Taxonomy of this tortoise has been challenged in recent years, with the two subspecies being considered full species and the central Dalmatian populations of T. h. boettgeri being considered a third species, T. hercegovinensis. Using an mtDNA fragment approximately 1150 bp long (cytochrome b gene and adjacent portion of tRNA-Thr gene), we investigated mtDNA diversity with regard to contrasting concepts of two subspecies or three species. Seven closely related haplotypes were identified from the western Mediterranean and 15 different, in part much-differentiated, haplotypes from the Balkans. Western Mediterranean haplotypes differ from Balkan haplotypes in 16–42 mutation steps. One to seven mutation steps occur within western Mediterranean populations. Balkan haplotypes, differing in 1−37 nucleotides, group in parsimony network analysis into three major assemblages that display, in part, a similar degree of differentiation to that of western Mediterranean haplotypes relative to Balkan haplotypes. Rates of sequence evolution are different in both regions, and low divergence, palaeogeography and the fossil record suggest a slower molecular clock in the western Mediterranean. While monophyly in western Mediterranean haplotypes is well-supported, conflicting evidence is obtained for Balkan haplotypes; maximum parsimony supports monophyly of Balkan haplotypes, but other phylogenetic analyses (Bayesian, ML, ME) indicate Balkan haplotypes could be paraphyletic with respect to the western Mediterranean clade. These results imply a process of differentiation not yet complete despite allopatry in the western Mediterranean and the Balkans, and suggest all populations of T. hermanni are conspecific. In the western Mediterranean no clear geographical pattern in haplotype distribution is found. Distribution of Balkan haplotypes is more structured. One group of similar haplotypes occurs in the eastern Balkans (Bulgaria, Republic of Macedonia, Romania and the Greek regions Evvia, Macedonia, Peloponnese, Thessaly and Thrace). Two distinct haplotypes, differing in eight to nine mutation steps from the most common haplotype of the first group, are confined to the western slope of the Taygetos Mts. in the Peloponnese. Yet another group, connected over between four and 23 mutation steps with haplotypes of the eastern Balkan group, occurs along the western slope of the Dinarid and Pindos Mts. (Istria, Dalmatia, western Greece). Taygetos haplotypes are nested within other haplotypes in all phylogenetic analyses and support for monophyly of the other Balkan groups is at best weak. We conclude that using the traditional two subspecies model should be continued for T. hermanni. Phylogeographies of T. hermanni and Emys orbicularis, another codistributed chelonian, are markedly different, but share a few similarities. Both were forced to retreat to southern refuges during Pleistocene glaciations. With the advent of Holocene warming, E. orbicularis underwent rapid range expansion and temperate regions of Europe and adjacent Asia were recolonized from refuges in the Balkans and the northern Black Sea Region. By contrast, T. hermanni remained more or less confined to refuges and nearby regions, resulting in a much smaller range, and allopatric and parapatric distribution of haplotype groups and clades. MtDNA lineages are more diverse in E. orbicularis than they are in T. hermanni on southern European peninsulas, indicating several distinct glacial refuges in close proximity and extensive intergradation during Holocene range expansion for E. orbicularis. In T. hermanni it is likely that only on the Balkan Peninsula was more than one refuge located, corresponding to the parapatric ranges of haplotype groups currently there. On the old western Mediterranean islands Corsica and Sardinia no differentiated (E. orbicularis) or only weakly differentiated haplotypes (T. hermanni) occur, even though there is evidence for the presence of both species on Corsica since at least the Middle Pleistocene. High mountain chains constitute major barriers separating distinct mtDNA clades or groups in each species.     

Hyalomma aegyptium as dominant tick in tortoises of the genus Testudo in Balkan countries, with notes on its host preferences

Collection of 1327 ticks sampled throughout Greece, Bulgaria, Romania and Croatia, from 211 tortoises belonging to three species, Testudo marginata Schoepff, T. graeca Linnaeus, and T. hermanni Gmelin, revealed the presence of four species of ixodid ticks, namely Hyalomma aegyptium (Linnaeus), Haemaphysalis sulcata Canestrini and Fanzago, H. inermis Birula and Rhipicephalus sanguineus (Latreille). Study confirmed the strong dominance of all life stages of H. aegyptium among ticks parasitizing west Palaearctic tortoises of genus Testudo Linnaeus. Furthermore, a considerable portion of ticks collected from tortoises in southwestern Bulgaria represent larvae and nymphs of H. sulcata. At the same area we collected as exception one larva and one nymph of H. inermis from a single specimen of T. hermanni. Our findings of four adults of R. sanguineus is the first record of this species from reptilian host. According to our results achieved on localities with syntopic occurrence of two tortoise species, T. marginata and T. graeca represent in the Balkans the principal hosts of H. aegyptium, whereas T. hermanni serves only as an alternative host in the areas close to range of either T. marginata or T. graeca.     

Home ranges in red fox: territoriality diminishes with increasing area

Red foxVulpes vulpes (Linnaeus, 1758) are generally regarded as strictly territorial animals, inhabiting distinct and well-separated ranges. The home ranges of these predators can vary in size from a few dozen hectares to as much as 20–30 km². Because it seemed impossible that foxes could defend areas so different in size with the same intensity, the published data were analysed in order to see how changes in home range size could affect the following parameters: home range overlap, the area visited daily and the distance travelled by animals during 24 h. It was found that the overlap between fox families (groups) occupying large ranges was much greater than between foxes living in small ranges. The ratio of the area visited daily to the seasonal home range size was greater in small ranges than in large ones. No significant correlation was found between the mean distance moved daily and home range size (r = −0.118,p = 0.75). These results suggest different patterns of the use of space and differences in territorial defence by red foxes inhabiting large and small ranges. The implications of home range overlap and intensity of contacts between foxes occupying neighbouring ranges for the spread of disease are discussed. The significant overlap between large ranges found in this paper is discussed in the light of Andrzejewski’s (2002) home range concept.

     

The effect of supplementary food on home range of the southern brown bandicoot,Isoodon obesulus (Marsupialia: Peramelidae)

The study investigated the relationship between home range and food abundance in a population of the southern brown bandicoot. Isoodon obesulus, in Western Australia. Home range areas were estimated seven times between 1986 and 1988 by live-trapping, spool-and-line devices and fluorescent pigment tracking. The abundance of invertebrate food was measured simultaneously by placing pitfall traps within the home ranges of individual animals, and by sampling invertebrates in topsoil and litter. Home range areas tended to be negatively correlated with food abundance, especially in the autumn and winter of 1986 and 1987. The influence of food on home range was investigated further in September 1988 by providing eight individual I. obesulus with a supplementary food mixture. Contrary to expectation, the added food caused an increase in home range area, home range overlap and displacement, as well as an influx of new individuals to food stations. In contrast, home range parameters in control (non-fed) individuals changed little during the experiment. The shuffling of home ranges due to feeding suggests that the home range system of I. obesulus is relatively flexible, with individuals monitoring and exploiting resources in an opportunistic manner. In contrast to previous studies, we found no evidence that I. obesulus was territorial. We speculate that individuals may be territorial at low population density if resources are defendable and intruder pressure is low, but occupy overlapping ranges if population density is high.     

Effects of Food Availability on Space and Refuge Use by a Neotropical Scatterhoarding Rodent

Animals that rely on refuges for safety can theoretically increase their foraging area without simultaneously increasing predation risk and travel costs by using more refuges. The key prediction of this theory, a negative correlation between food abundance, home range size and the number of refuges used, has never been empirically tested. We determined how home range size and refuge use by the Central American agouti (Dasyprocta punctata) varied across a gradient of abundance of the agoutis' principal food source: seeds and fruits of the palm Astrocaryum standleyanum. We used both manual and automated radio telemetry to measure space use of 11 agoutis during 2 mo of the Astrocaryum fruiting season, and of another set of 10 agoutis during 6 mo in which the animals largely relied on cached Astrocaryum seeds. We found that agoutis living in areas of lower food density had larger home ranges, and that all individuals used multiple refuges. The number of refuges, however, was not correlated with home range size. Consequently, agoutis that had larger home ranges roamed farther from their refuges. These results suggest that agoutis increase their home range size in response to food scarcity at the cost of their safety.     

First record of the family Hygrobiidae (Coleoptera: Adephaga) from Turkey

With the discovery of Hygrobia hermanni, the family Hygrobiidae is reported for the first time from Turkey. An illustration of the habitus and line drawings of diagnostic characters are provided.     

Home-range size and use and territorial behavior in the common marmoset,Callithrix jacchus jacchus,at the tapacura field station,recife, Brazil

Data were obtained both from observation and by radiolocation on the size and use of the home range of the common marmoset, Callithrix jacchus jacchus.Ranges were small compared to those of other callitrichids, varying between 0.72 to 1.62 ha, and a high proportion of the home range was used each day. The marmosets showed territorial behavior, defending an area almost equivalent to the home range. Despite this, mating between animals belonging to neighboring groups was seen. Correlations between the use of the home range and some environmental variables showed little consistency between two groups which occupied ranges of differing resource density, and their activity patterns, as measured by movement through their ranges, were also different. Possible reasons for these inconsistencies are discussed.     

Home range use and the exploitation of gum in the marmosetCallithrix jacchus jacchus

Three wild groups of common marmoset, Callithrix jacchus jacchus,in north-east Brazil, of approximately similar size, had home ranges between 2.5 and 6.5 ha. But their core areas were similar in size between 1.0 and 1.5 ha, with a monthly area of heavy use between 1.1 and 1.6 ha. The groups were selective in the use of their home ranges, even though they were small: they used some areas heavily and others lightly. The core areas had higher densities of trees that produced gum exudates than did other parts of the home ranges. Our data suggest that a group of marmosets in this habitat may require a minimum of about 50 gum trees in its home range at a minimum density of about 50 trees/ha. In addition, the animals require suitable trees in which to sleep. We suggest that patches of forest with these desirable properties remain relatively fixed in size and location over the years and that individual animals are constantly in flux between them.     

An ecological study of troop fissions of Japanese monkeys (Macaca fuscata yakui) on Yakushima Island,Japan

Wild, habituated, Japanese monkeys were observed from 1975 to 1979 on Yakushima Island, Southern Japan. The monkey troops had a continuous distribution in a warm temperate forest. Demographic data on local populations was collected. The population density was 33 animals/km². The growth rate of the studied troop was 3.0% per year. A significant correlation between home range areas (R) and troop size (P) was found (r=0.955,p<0.005), using anR-P equation,R=1.84P. One troop split into three troops through two successive fissions. Twenty-one intertroop encounters were observed. Five types of encounters were distinguished. The encounters were apparently territorial defence. Increases in birth rate and socionomic sex ratio after the fissions were prominent. The following four factors had a direct effect upon the dispersion of the troops after fission: (1) dominance relation between the fission troops; (2) social pressure of the neighbors; (3) troop's attachment to its home range; and (4) structure of the environment. The home range of Japanese monkeys is a territory, and territoriality is a population regulating mechanism which serves to reduce competition for food.     

野放普氏野马(Equus przewalskii)家域面积及其影响因素

采用MCP方法研究了2011年至2012年新疆卡拉麦里山有蹄类自然保护区野放普氏野马家域的变化。通过方差分析验证了年间、季节间不同群体家域及其两两重叠无差异。以家族群大小为协变量进行了野放野马家域协方差分析。利用野放野马家族大小为协变量的协方差分析分析检验了野放野马家族大小与家域关系。结果表明:(1)野马平均家域面积由2011年的(20±2)km~2/匹扩大到2012年的(30±2)km~2/匹。对部分野放群体中头马未发生更替的野马群的研究表明,随着野马群体增大,其家域面积显著增大(P0.05)。(2)单因素方差分析显示,不同野马群的家域面积在不同年份差异显著,且春季家域秋季家域夏季家域。(3)2011年不同群家域两两间相互重叠面积与群大小无显著相关(r=0.256,P=0.5800.05)。而2012年野马群家域两两之间重叠面积有显著差异(F=4.521,df=8,P0.001)。家域两两相互重叠面积与群大小显著相关(r=0.706,P=0.0330.05)。(4)不同季节间野马群家域重叠面积有显著差异(F=5.695,df=8,P0.001)。5号群、7号群和8号群的自身家域重叠面积(P0.05),3号群、6号群和9号群的家域重叠面积(P0.05)。(5)影响野放野马家域面积的生物因子有草本盖度、灌木盖度,非生物因子主要有温度、湿度、风速、最近水源地距离和最近居民点距离等。温度与草本盖度是影响野放野马家域面积大小的主要因素,两者与野放野马家域面积显著相关(P0.01)。     

Die ökologische Spezialisierung der EidechseCryptoblepharus boutoni cognatus (Boettger) auf das Leben in der Gezeitenzone (Reptilia,Skinkidae)

Zusammenfassung Auf Madagaskar läuft die EidechseCryptoblepharus boutoni cognatus einmal am Tag bei Einsetzen der Ebbe in die langsam trockenfallende Gezeitenzone, wo sie nach Insekten, Krebsen und Fischen (jungePeriophtalmus kohlreuteri) jagt. Die gezeitenanhängige Aktivität, definiert als die Anzahl der frei umherlaufenden Tiere, wurde gemessen.Die Eidechse ist ortstreu und wandert auf erlernten Wegen zwischen den Ruheplätzen an Land und den Jagdgründen in der Gezeitenzone. Bei Verfrachtung über ca. 200 m außerhalb ihres Aktionsraumes (home range) kehrt sie in ihr Heimatareal zurück. Ortstreue, gezeitenperiodische Wanderungen und Heimfinde-Verhalten werden als Anpassung für das Leben in der Gezeitenzone diskutiert.Die Eidechse ist ein Gezeitenzonen-Spezialist, die ohne Aufgabe ihrer terrestrischen Lebensweise in diese neue ökologische Nische eingedrungen ist.

---

Ecological adaptation of the lizzardCryptoblepharus boutoni cognatus (boettger) for life in the interditel zone (reptilia, skinkidae)

Summary The lizardCryptoblepharus boutoni cognatus was observed in Madagaskar. The animals descended daily into the intertidal zone to feed there on insects, crustaceans and fish (juvenilePeriophthalmus kohlreuteri). The pattern of activity of the animals is related to the movements of the tides.The lizards have a definite home range and migrate between intertidal feeding area and resting place on land along pathways established by experience. They return to their home even if released at about 200 m from their home range because they seem to acquire a knowledge of the surrounding by occasional exploratory visits. The tendency to establish a home range and homing behaviour are considered as adaptations for life in the intertidal zone, but the lizard remains a purely terrestial animal.

---

---

FrüherAblepharus boutoni, s. Fuhn, I. E.: Z. f. Systematik u. Evolutionsforschung7 (1), 67–76 (1969).     

Molecular systematics of Scaphirhynchinae: an assessment of North American and Central Asian Freshwater Sturgeon Species

The sturgeon subfamily Scaphirhynchinae contains two genera of obligate freshwater sturgeon: Scaphirhynchus and Pseudoscaphirhynchus, from North America and Central Asia, respectively. Both genera contain morphologically variable species. A novel data set containing multiple individuals representing four diagnosable morphological variants for two species of Pseudoscaphirhynchus, P. hermanni and P. kaufmanni, was generated. These data were used to test taxonomic hypotheses of monophyly for the subfamily Scaphirhynchinae, monophyly of both Scaphirhynchus and Pseudoscaphirhynchus, monophyly of P. hermanni and P. kaufmanni, and monophyly of the recognized morphological variants. Monophyly of the subfamily Scaphirhynchinae is consistently rejected by all phylogenetic reconstruction methodologies with the molecular character set while monophyly of both river sturgeon genera is robustly supported. The molecular data set also rejects hypotheses of monophyly for sampled species of Pseudoscaphirhynchus as well as monophyly for the recognized intraspecific morphological variants. Interestingly both Scaphirhynchus and Pseudoscaphirhynchus demonstrate the same general pattern in reconstructed topologies; a lack of phylogenetic structure in the clade with respect to recognized diversity. Despite rejection of monophyly for the subfamily Scaphirhynchinae with molecular data, reconstructed hypotheses from morphological character sets consistently support monophyly for this subfamily. Disparities among the data sets, as well as reasons for rejection of monophyly for Scaphirhynchinae and species of Scaphirhynchus and Pseudoscaphirhynchus with molecular characters are examined and a decreased rate of molecular evolution is found to be most consistent with the data.     

Ecological study of the saddle backed tamarin (Saguinus fuscicollis) in Northern Bolivia

The author carried out an ecological survey of the saddle backed tamarin (Saguinus fuscicollis) in the Pando, northern Bolivia, from July to December 1979. It was found thatS. fuscicollis formed a family group consisting of two to seven individuals and spent almost all the daytime in moving to forage insects, although it fed mainly on fruits of the family Moraceae. Primary forest was preferred for sleeping sites, whereas the animals spent more hours in secondary forest than in primary forest in their home range. Marking sites were concentrated in the center of the home range but encounter behaviors tended to take place near the borderlines of the home range. It was also found that infants began feeding on fruit at about 5 weeks of age, whereas they shared insects obtained by their mothers until about 10 weeks of age.     

Interactions between density,home range behaviors,and contact rates in the Channel Island fox (Urocyon littoralis)

Many of the mechanisms underlying density‐dependent regulation of populations, including contest competition and disease spread, depend on contact among neighboring animals. Understanding how variation in population density influences the frequency of contact among neighboring animals is therefore an important aspect to understanding the mechanisms underlying, and ecological consequences of, density‐dependent regulation. However, contact rates are difficult to measure in the field and may be influenced by density through multiple pathways. This study explored how local density affects contact rates among Channel Island foxes (Urocyon littoralis) through two pathways: changes in home range size and changes in home range overlap. We tracked 40 radio‐collared foxes at four sites on San Clemente Island, California. Fox densities at the four sites ranged from 2.8 ± 1.28 to 42.8 ± 9.43 foxes/km². Higher fox densities were correlated with smaller home ranges (R² = 0.526, F_1,38 = 42.19, P < 0.001). Thirty foxes wore collars that also contained proximity loggers, which recorded the time and duration of occasions when collared foxes were within 5 m of one another. Contact rates between neighboring fox dyads were positively correlated with home range overlap (R² = 0.341, P = 0.008), but not fox density (R² = 0.012, P = 0.976). Individuals at high densities had more collared neighbors with overlapping home ranges (R² = 0.123, P = 0.026) but not an increase in the amount of contact between individual neighbors. This study was the first time contact rates were directly measured and compared to density and home range overlap. Results suggest that foxes exhibit a threshold in their degree of tolerance for neighbors, overlap is a reliable index of the amount of direct contact between island foxes, and disease transmission rates will likely scale with fox density.     

Nutrition and husbandry conditions of Palearctic tortoises (Testudo spp.) in captivity

Mediterranean and Russian tortoises (Testudo spp.) are popular companion animals (pets), despite ongoing controversy concerning privately keeping reptiles. The arguments used during these controversial discussions have often been based on outdated facts. Therefore, a survey was developed to evaluate the current population structure, husbandry conditions, diet regime, and health status of Testudo species in captivity. More than 75% of the 1075 respondents housed their tortoises in an outdoor enclosure containing a greenhouse or cold frame, which is considered the most species-appropriate way of husbandry. Of the respondents, 67.7% fed their tortoises with the optimum diet of more than 80% grasses and weeds during the summer vegetation period. Only 8.2% of respondents owned a tortoise with a diagnosed disease. According to the results, the likelihood of tortoises developing pyramidal growth syndrome, which can be used as an indicator of the quality of tortoise husbandry, was high in tortoises kept in a terrarium and/or fed a diet of less than 80% grasses and weeds in summer. This likelihood varied among species, with a higher incidence in Hermann’s tortoises (Testudo hermanni).