The behavioural ecology and population dynamics of a cryptic ground‐dwelling mammal in an urban Australian landscape

Urbanization results in widespread habitat loss and fragmentation and generally has a negative impact upon native wildlife, in particular ground‐dwelling mammals. The northern brown bandicoot (Isoodon macrourus; Marsupialia: Peramelidae) is one of relatively few native Australian ground‐dwelling mammals that is able to survive within urbanized landscapes. As a consequence of extensive clearing and urban development within the city of Brisbane, bandicoots are now restricted to the mostly small (<10 ha) bushland fragments scattered across the city landscape. Our study examined the behavioural ecology of northern brown bandicoots within habitat fragments located on a major creek‐line, using mark‐recapture population monitoring and radio telemetry. Bandicoots at monitored sites were found to occur at high densities (typically one individual ha^?1), although one‐third of the populations were transient. Radio tracking revealed that bandicoots had relatively small home ranges (mean 1.5 ± 0.2 ha) comprised largely of bushland/grassland with dense, often weed‐infested ground cover. Bandicoots sheltered by day in these densely covered areas and also spent most time foraging there at night, although they occasionally ventured small distances to forage in adjacent maintained parklands and residential lawns. We suggest that introduced tall grasses and other weeds contribute to high habitat quality within riparian habitat fragments and facilitate the persistence of high density populations, comprised of individuals with small home ranges. The generalized dietary and habitat requirements of northern brown bandicoots, as well as a high reproductive output, undoubtedly facilitate the survival of the species in urban habitat fragments. Further research is required on other native mammal species in urbanized landscapes to gain a greater understanding of how best to conserve wildlife in these heavily modified environments.     

Interactions between European rabbits and native marsupials in the absence of terrestrial predators

European rabbits (Oryctolagus cuniculus) are ubiquitous across Australia and have the ability to influence native species directly and indirectly. Despite this, limited research focuses on interspecific interactions between rabbits and native mammals. We aimed to determine how site occupancy, detection probability, and temporal activity periods of native mammals changed in response to the presence or absence of rabbits. We monitored three native mammal species using 85 camera traps in a systematic grid at Mt Rothwell Conservation and Research Reserve (Victoria, Australia), a predator-barrier fenced reserve with two distinct sections—an area with rabbits and an area without. Bettongs (rufous Aepyprymnus rufescens and eastern Bettongia gaimardi), eastern barred bandicoots (Perameles gunni), and southern brown bandicoots (Isoodon obesulus) had a naïve site occupancy of 71%, 42%, and 24%, respectively. Site occupancy for both bandicoot species decreased in areas with more clumping grass with eastern barred bandicoot occupancy increasing with leaf litter cover, and southern brown bandicoots with vegetation height. Rabbit presence did not influence site occupancy of any species. Species detection probabilities were generally positively associated with open vegetation and rabbit presence, except for southern brown bandicoots which were more detectable without rabbits. Both bandicoot species shifted their peak activity periods in the absence of rabbits having an earlier, and more defined activity peak. Our results demonstrate that the presence of rabbits in the absence of invasive predators may not influence the site occupancy of co-occurring native mammals, however, could influence the behaviour of smaller co-occurring mammals, either directly or indirectly.     

Habitat selection by vulnerable golden bandicoots in the arid zone

In 2010, vulnerable golden bandicoots (Isoodon auratus) were translocated from Barrow Island, Western Australia, to a mainland predator‐free enclosure on the Matuwa Indigenous Protected Area. Golden bandicoots were once widespread throughout a variety of arid and semiarid habitats of central and northern Australia. Like many small‐to‐medium‐sized marsupials, the species has severely declined since colonization and has been reduced to only four remnant natural populations. Between 2010 and 2020, the reintroduced population of golden bandicoots on Matuwa was monitored via capture–mark–recapture data collection, which was used in spatially explicit capture–recapture analysis to monitor their abundance over time. In 2014, we used VHF transmitters to examine the home range and habitat selection of 20 golden bandicoots in the enclosure over a six‐week period. We used compositional analysis to compare the use of four habitat types. Golden bandicoot abundance in the enclosure slowly increased between 2010 and 2014 and has since plateaued at approximately one quarter of the density observed in the founding population on Barrow Island. The population may have plateaued because some bandicoots escape through the fence. Golden bandicoots used habitats dominated by scattered shrubland with spinifex grass more than expected given the habitat''s availability. Nocturnal foraging range was influenced by sex and trapping location, whereas diurnal refuge habitat, which was typically under a spinifex hummock with minimal overstory vegetation, was consistent across sex and trapping location. Our work suggests that diurnal refuge habitat may be an important factor for the success of proposed translocations of golden bandicoots.     

Feeding Wild Animals: The Urge,the Interaction,and the Consequences

Abstract

Endangered species recovery requires a rapid, concerted effort on several fronts, including an understanding of sociological considerations such as the values and attitudes of local people. We examined the values and attitudes of residents living in and around the City of Hamilton, Victoria, toward the endangered, mainland Australia population of eastern barred bandicoots (Perameles gunnii) and bandicoot conservation efforts using a sample telephone survey. Over 90% of those surveyed indicated that they favored efforts to save the bandicoot and nearly 100% believed that bandicoots have a right to exist. Support was broad and varied little with respect to gender, age, level of education, or income. Attitude scale scores supported these results, with most people scoring relatively high on moralistic and naturalistic/ecologistic scales and relatively low on utilitarian and negativistic scales. Of people surveyed, 45.5% owned cats. Cats pose a major threat to bandicoot survival because of predation and disease transmission. Although cat owners and nonowners displayed similar levels of support for bandicoot conservation, 70% of cat owners allowed their cats out at night, when bandicoots are most active. The results suggest that the local public would support new conservation measures invoked to enhance bandicoot survival and represents a potentially valuable source of volunteer labor. A public affairs program should educate the public about the role of domestic cats in bandicoot decline and explicitly point out the contradiction between strong support for bandicoot conservation and the behavior of permitting cats to roam free at night. To increase chances of success, managers and conservationists should recognize the importance of nonbiological aspects, such as values and attitudes of local people, to endangered species restoration efforts.     

Factors influencing the distribution of medium‐sized ground‐dwelling mammals in southeastern mainland Australia

Abstract The forage diggings of medium‐sized ground‐dwelling mammals (bandicoots and potoroos) were recorded over two seasons across 136 forested study sites, representing a stratified sample of the climatic, geological and topographic features of far south‐eastern mainland Australia. Diggings, presumably left by bandicoots, were recorded at a total of 42 sites, and those of potoroos at 27 sites. Statistical models were developed for the occurrence of these diggings, based on environmental attributes measured for each site. At a landscape scale, mean minimum temperature of the coldest month was an important explanatory variable for both groups of animals, with likelihood of their occurrence increasing with increasing temperature during that period. More locally, soil fertility and time since last fire also influenced the occurrence of bandicoot and potoroo diggings, while density of ground cover was additionally important in explaining the occurrence of bandicoot diggings. Bandicoot and potoroo diggings were more likely to occur with decreasing soil fertility and increasing time since fire, while occurrence of bandicoot diggings also increased with increasing density of ground cover. The possible management implications of our findings for conservation of medium‐sized ground‐dwelling mammals in multiple‐use forests are discussed.     

The phylogenetic relationships of the bilby, Macrotis lagotis (Peramelimorphia: Thylacomyidae), to the bandicoots- DNA sequence evidence

Partial sequencing of the 12S ribosomal RNA gene was used to test two competing hypotheses concerning the phylogenetic relationship of the bilby (Macrotis lagotis) to the Australian and New Guinean species of bandicoot. The first hypothesis proposes that the Australian and New Guinean bandicoots are in a monophyletic clade to the exclusion of the bilbies, whereas the second hypothesis proposes that the bilby is monophyletic with the Australian bandicoots to the exclusion of the New Guinean bandicoots. Phylogenies determined by both maximum-likelihood and neighbour-joining approaches supported the first hypothesis in which the bilby is excluded from the clade represented by the Australian and New Guinean bandicoots. Monophyly of the Australian and New Guinean bandicoots is consistent with the biogeographical scenario in which Australia and Papua New Guinea have undergone repeated connection and disconnection over the last 20 million years.     

A NEW SPECIES OF BANDICOOT FROM THE OLIGOCENE OF NORTHERN AUSTRALIA AND IMPLICATIONS OF BANDICOOTS FOR CORRELATING AUSTRALIAN TERTIARY MAMMAL FAUNAS

Abstract:  A new species of primitive bandicoot, Yarala kida sp. nov., is described from Kangaroo Well, a site in the Northern Territory of Australia. This species is possibly ancestral to Yarala burchfieldi , the type species of the Yaraloidea, and supports a late Oligocene age for the Kangaroo Well Local Fauna. The yaraloid bandicoots are likely to become important biochronological tools for Australian faunas of late Oligocene to early Miocene age, as they are widespread and diverse. Developing morphoclines for this group is therefore essential, as is publication of the mostly undescribed bandicoot material known from other sites of similar age.     

Molecular Relationships of the Extinct Pig-Footed Bandicoot Chaeropus ecaudatus (Marsupialia: Perameloidea) Using 12S rRNA Sequences

The pig-footed bandicoot, Chaeropus ecaudatus, is presumed to be extinct as no specimens have been collected or seen since early this century. Usually classified as a specialized member of the family Peramelidae, there is nevertheless still some doubt as to its taxonomic affinities, because this animal is highly specialized and shows several uniquely derived characters. We report here the first attempt to determine the molecular relationships of this animal using mitochondrial 12S rRNA sequences derived from spirit-preserved museum specimens. Phylogenetic analysis shows that the sequence derived from the Chaeropus sample is clearly that of a bandicoot. Within the bandicoot clade, the pig-footed bandicoot is quite distinct from all other taxa. Divergence-time estimates from the 12S rRNA sequences suggest that Chaeropus diverged from the other bandicoot genera in the late Oligocene or early Miocene and that bandicoots diverged from other Australian families in the late Paleocene–early Eocene.     

Overcoming prey naiveté: Free‐living marsupials develop recognition and effective behavioral responses to alien predators in Australia

Naiveté in prey arises from novel ecological mismatches in cue recognition systems and antipredator responses following the arrival of alien predators. The multilevel naiveté framework suggests that animals can progress through levels of naiveté toward predator awareness. Alternatively, native prey may be preadapted to recognize novel predators via common constituents in predator odors or familiar predator archetypes. We tested predictions of these competing hypotheses on the mechanisms driving behavioral responses of native species to alien predators by measuring responses of native free‐living northern brown bandicoots (Isoodon macrourus) to alien red fox (Vulpes vulpes) odor. We compared multiple bandicoot populations either sympatric or allopatric with foxes. Bandicoots sympatric with foxes showed recognition and appropriate antipredator behavior toward fox odor via avoidance. On the few occasions bandicoots did visit, their vigilance significantly increased, and their foraging decreased. In contrast, bandicoots allopatric with foxes showed no recognition of this predator cue. Our results suggest that vulnerable Australian mammals were likely naïve to foxes when they first arrived, which explains why so many native mammals declined soon after fox arrival. Our results also suggest such naiveté can be overcome within a relatively short time frame, driven by experience with predators, thus supporting the multilevel naiveté framework.     

Common names of species, the curious case of Capra pyrenaica and the concomitant steps towards the ‘wild-to-domestic’ transformation of a flagship species and its vernacular names

Common names (CN) add to linguistic richness and ultimately derive from how a majority of people refer to a species. CN have a biological and–above all—practical importance given that they are essential for connecting specialists and lay people. To illustrate the diversity of CN between and within species, we made an overview of common name in Caprinae species—flagship species in mountain ecosystems. Then, using Capra pyrenaica as a study case, we highlighted that the choice of CN is inextricably linked to current debates and trends in wildlife management that should never be ignored, given their importance in the fields of ethics, zoology, systematics, conservation and current management. We underline the need to investigate further the probable relationships linking common names, human perception and wildlife management. Researchers, citizens and policy-makers will have to be watchful that clumsy common names, such as ‘wild-to-domestic’ transformed ones, will not hamper the conservation of wild species as a ‘Common Heritage’.     

Metabolic and ventilatory physiology of the Barrow Island golden bandicoot (Isoodon auratus barrowensis) and the northern brown bandicoot (Isoodon macrourus)

Metabolic and ventilatory parameters were measured for the smallest and largest Isoodon bandicoots; the arid-adapted Barrow Island golden bandicoot (Isoodon auratus barrowensis) and the tropical northern brown bandicoot (Isoodon macrourus). I. a. barrowensis has a number of physiological characteristics that aid its tolerance of high T_a and survival in a hot and dry climate, including a low and labile body temperature, a very low basal metabolic rate, low total evaporative water loss, and an effective panting mechanism. I. macrourus generally has an “average” physiology for a bandicoot despite its size, although a number of its physiological characteristics aid survival in (sub)tropical conditions. These include a low body temperature, low total evaporative water loss and minute ventilation at high ambient temperatures, and an average thermal conductance. These data support the theory that phylogeny is a more important predictor of bandicoot physiology than habitat/distribution.     

Effects of long-term captivity on thermoregulation,metabolism and ventilation of the southern brown bandicoot (Marsupialia: Peramelidae)

Thermoneutral metabolic and ventilatory parameters were measured every 3 months over 2 years for southern brown bandicoots held in captivity, and from a nearby reserve. Captive bandicoots were 130 g (9.9%) heavier than wild bandicoots. Long-term captivity had no effect on body temperature, basal metabolic rate (oxygen consumption), thermal conductance or respiratory ventilation, but there was an effect on carbon dioxide production, respiratory exchange ratio and total evaporative water loss (values were between 15 and 25% higher for captive than for wild bandicoots). Diet may be influencing these aspects of captive bandicoot physiology; the diet of captive bandicoots would be considerably different to that of wild bandicoots. Water availability seems to have a minimal effect. This study has important implications regarding physiological measurement for captive and wild mammals. For bandicoots at least, captive animals are equivalent to wild animals for some physiological parameters at thermoneutrality (body temperature, resting metabolic rate and thermal conductance), but not others.     

Entrainment of Circadian Wheel-Running Rhythms of the Northern Brown Bandicoot, Isoodon macrourus, by Daily Restricted Feeding Schedules

Northern brown bandicoots (Isoodon macrourus) were subjected to restricted feeding for 3 h in the middle of the light period of a 14: 10 light/dark cycle and immediately following this in constant dark. When feeding was restricted to the middle of the light period of the light/dark cycle, all bandicoots maintained a nocturnal activity rhythm. In addition to the nocturnal rhythm, a few bandicoots showed meal-anticipatory activity during the light period. In bandicoots that did not show meal-anticipatory activity, diurnal activity was sometimes evident either during or shortly after the daily meal time. The observation of meal-anticipatory activity in some bandicoots suggests that this species may have a mechanism separate from the light-entrainable mechanism that allows the daily anticipation of periodically available food sources. In the next stage of the experiment, which was in constant dark, the meal was presented at the same time of day as it had been in the previous stage. In all bandicoots, the previously light-entrained component of activity free-ran and was eventually affected by the restricted feeding schedule to some degree. Bandicoots showed weak entrainment and relative coordination, suggesting that restricted feeding is a weak zeitgeber in this species. Evidence also suggesting that two separate but coupled pacemakers control the activity rhythms of the bandicoot was that (a) bandicoots simultaneously showed free-running light-entrainable rhythms and meal-entrained anticipatory rhythms; (b) in several bandicoots, the light-entrainable rhythm was phase advanced when it free-ran through the meal time; and (c) in one bandicoot, meal-entrained anticipatory activity was forced away from the meal time when the previously light-entrained component of activity free-ran through it.     

Relative influence of habitat structure,species interactions and rainfall on the post‐fire population dynamics of ground‐dwelling vertebrates

The long‐term impacts of wildfires on animal populations are largely unknown. We used time‐series data based on a tracking index, from coastal NSW spanning 28 years after a wildfire, to investigate the relative influence of habitat structure, species interactions and climate on post‐fire animal population dynamics. The fire had an immediate impact on habitat structure, reducing and simplifying vegetation cover, which then underwent post‐fire successional change including an increase and plateau in tree canopy cover; an increase, stabilization and then decline in shrub cover; and an increase in ground litter cover. Population changes of different animal species were influenced by different components of successional change, but there was also evidence that species interactions were important. For example, bandicoots (Isoodon obesulus and Perameles nasuta combined) increased concurrent with an increase in shrub cover then declined at a faster rate than a direct association with senescing shrub cover would suggest, while the feral cat (Felis catus) population changed with the bandicoot population, suggesting a link between these species. Potoroos (Potorous tridactylus) increased 10 years after the fire concurrent with the closing tree canopy, but there was also evidence of a negative association with feral foxes (Vulpes vulpes). Variation in rainfall did not have significant effects on the population dynamics of any species. Our results suggest that changes in habitat structure play a key role in the post‐fire dynamics of many ground‐dwelling animals and hence different fire regimes are likely to influence animal dynamics through their effects on habitat structure. However, the role of predator–prey interactions, particularly with feral predators, is less clear and further study will require manipulative experiments of predators in conjunction with fire treatments to determine whether feral predator control should be integrated with fire management to improve outcomes for some native species.     

Phylogenetics,population structure and genetic diversity of the endangered southern brown bandicoot (<Emphasis Type="Italic">Isoodon obesulus</Emphasis>) in south-eastern Australia

The southern brown bandicoot (Isoodon obesulus) has undergone significant range contractions since European settlement, and it is now considered Endangered throughout south-eastern mainland Australia. This species currently has a highly fragmented distribution inhabiting a mosaic of habitats. This project uses mitochondrial DNA (mtDNA) and microsatellite data to determine levels of genetic diversity, population structure and evolutionary history, which can aid wildlife managers in setting priorities and determining management strategies. Analyses of genetic diversity revealed low levels of mtDNA variability (mean h=50.42%, =0.76%) and divergence (mean d_A=0.29%) across all regions investigated, and was among the lowest recorded for marsupials. These data indicate a relatively small female effective population size, which is most likely a consequence of a large-scale population contraction and subsequent expansion occurring in pre-history (mismatch distribution analysis, SSD P-value=0.12). Individuals from the Sydney region experienced significant reductions in microsatellite diversity (A=3.8, H_E=0.565), with the Garigal National Park (NP) population exhibiting genetic reduction signatures indicating a recent population bottleneck. Population differentiation analysis revealed significant genetic division amongst I. obesulus individuals from Sydney, East Gippsland and Mt Gambier regions (=0.176–0.271), but could not separate the two Sydney populations (Ku-ring-gai NP and Garigal NP). Based on these data and habitat type, translocations could readily be made between the two Sydney populations, but not between the others. Phylogenetic comparisons between I. obesulus and I. auratus show little support for current Isoodon taxonomy, consistent with the findings of Pope etal. 2001. We therefore recommend the recognition of only three I. obesulus sub-species and suggest that these comprise a single morphologically diverse species that once was widespread across Australia.     

Genetic dissection of grain yield in bread wheat. II. QTL-by-environment interaction

The grain yield of wheat is influenced by genotype, environment and genotype-by-environment interaction. A mapping population consisting of 182 doubled haploid progeny derived from a cross between the southern Australian varieties ‘Trident’ and ‘Molineux’, was used to characterise the interaction of previously mapped grain yield quantitative trait locus (QTL) with specific environmental covariables. Environments (17) used for grain yield assessment were characterised for latitude, rainfall, various temperature-based variables and stripe rust infection severity. The number of days in the growing season in which the maximum temperature exceeded 30°C was identified as the variable with the largest effect on site mean grain yield. However, the greatest QTL-by-environmental covariable interactions were observed with the severity of stripe rust infection. The rust resistance allele at the Lr37/Sr38/Yr17 locus had the greatest positive effect on grain yield when an environment experienced a combination of high-stripe rust infection and cool days. The grain yield QTL, QGyld.agt-4D, showed a very similar QTL-by-environment covariable interaction pattern to the Lr37/Sr38/Yr17 locus, suggesting a possible role in rust resistance or tolerance. Another putative grain yield per se QTL, QGyld.agt-1B, displayed interactions with the quantity of winter and spring rainfall, the number of days in which the maximum temperature exceeded 30°C, and the number of days with a minimum temperature below 10°C. However, no cross-over interaction effect was observed for this locus, and the ‘Molineux’ allele remained associated with higher grain yield in response to all environmental covariables. The results presented here confirm that QGyld.agt-1B may be a prime candidate for marker-assisted selection for improved grain yield and wide adaptation in wheat. The benefit of analysing the interaction of QTL and environmental covariables, such as employed here, is discussed.     

Genetic structure and effective population size of Sydney rock oysters in eastern Australia

Oyster reef habitats are critical to coastal biodiversity and their decline has prompted restoration efforts in Australia. Knowledge gaps exist regarding the population structure and diversity of key species in these habitats. This may be critical information for the design of effective restoration programs. Sydney rock oysters (Saccostrea glomerata) are the dominant reef-forming bivalve in eastern Australia. Wild populations of S. glomerata have declined due to overharvesting, disease outbreaks, coastal development and reduced water quality. Here, we use genetic markers identified by genome-wide sequencing to investigate the genetic structure and diversity of wild Sydney rock oysters throughout their distribution in eastern Australia. We examine evidence for past population bottlenecks and spatial genetic structure associated with the East Australian Current. Analysis of 3, 400 neutral single-nucleotide polymorphisms (SNPs) revealed a single population, and an overlap with two other Saccostrea sp. at the northernmost boundary of the distribution. We detected signals of asymmetric gene flow consistent with the direction of the East Australian Current, and spatial structure patterns of limited genetic isolation by distance and spatial autocorrelation in the northern region (which experiences stronger effects of the East Australian Current) but not in the southern region of the distribution. We found no evidence of significant recent bottlenecks, with high effective population size throughout the species’ range. This information will provide a baseline against which to assess the impact of restoration projects, and guide strategies for sourcing stock for the enhancement of wild oyster populations. Our results provide a positive outlook for the resilience and adaptive capacity of Sydney rock oysters, and highlight wild populations as valuable resources for aquaculture and restoration initiatives.

     

Plasma concentrations of progesterone and 13,14-dihydro-15-keto-prostaglandin F-2 alpha during regression of the corpora lutea of lactation in the bandicoot (Isoodon macrourus)

Plasma progesterone concentrations (mean +/- s.e.m.) declined from 7.5 +/- 1.2 ng/ml and 7.5 +/- 1.0 ng/ml to less than 1 ng/ml after removal of pouch young (RPY) from bandicoots at Days 24 and 30 of lactation respectively. In all 7 bandicoots, the corpora lutea of lactation showed signs of regression and, in 5 of these bandicoots, a premature ovulation had occurred 6-9 days after RPY. There was no change in the concentration of PGFM after RPY, and uterine prostaglandin F-2 alpha may not be involved in luteal regression in the bandicoot.     

Population genetic evidence for the east–west division of the narrow-barred Spanish mackerel (<Emphasis Type="Italic">Scomberomorus commerson,</Emphasis> Perciformes: Teleostei) along Wallace’s Line

Mitochondrial DNA D-loop (control) region (426-bp) was used to infer the genetic structure of Spanish mackerel (Scomberomorus commerson) from populations in Southeast Asia (Brunei, East and West Malaysia, Philippines, Thailand, Singapore, and China) and northern Australia (including western Timor). An east–west division along Wallace’s Line was strongly supported by a significant AMOVA, with 43% of the total sequence variation partitioned among groups of populations. Phylogenetic and network analyses supported two clades: clade A and clade B. Members of clade A were found in Southeast Asia and northern Australia, but not in locations to the west (Gulf of Thailand) or north (China). Clade B was found exclusively in Southeast Asia. Genetic division along Wallace’s Line suggests that co-management of S. commerson populations for future sustainability may not be necessary between Southeast Asian nations and Australia, however all countries should share the task of management of the species in Southeast Asia equally. More detailed genetic studies of S. commerson populations in the region are warranted.     

Microsatellite and mitochondrial DNA variation defines island genetic reservoirs for reintroductions of an endangered Australian marsupial, <Emphasis Type="Italic">Perameles bougainville</Emphasis>

Natural populations of the endangered western barred bandicoot (Perameles bougainville) now exist on only two islands in Shark Bay, Western Australia. Our aim was to investigate genetic diversity in natural, reintroduced, and captive populations of the bandicoots and to assess the extent of divergence between the populations. The contemporary isolation of the natural populations has resulted in heterogeneity of allele frequency between the islands, which has acted to maintain a higher combined diversity than would be expected from either population on its own. These findings highlight how remnant island populations can act as genetic reservoirs to maximize diversity for reintroductions into a species former range. Although diversity is high between island populations, diversity within populations, based on six microsatellite loci, are amongst the lowest ever recorded for populations of marsupials. The mtDNA sequence data indicate that the two remaining natural populations show only minor divergence from each other, with the five haplotypes separated by just single base pairs. The reintroduced population and captive colonies show evidence for the loss of diversity related to genetic drift operating on small isolated populations.