A note on wildlife poisoning cases from Kerala, South India
Wildlife poisoning is an important conservation threat for endangered species in India. There are no publications in the scientific literature that identify the specific poisons or chemicals involved in wildlife poisoning cases from the state of Kerala. In this report, all cases of wildlife mortality recorded between 2011 and 2013 at the office of the Assistant Forest Veterinary Officer, Periyar Tiger Reserve in Kerala were reviewed and cases where poisoning was considered as a differential diagnosis were identified. Specific poisons or chemicals were identified in three cases, while in a fourth, poisoning was determined to have occurred based on physical traces of the poison in gut contents. The poisons identified include carbofuran (a carbamate pesticide) in a bonnet macaque (), warfarin (a rodenticide) in a mortality event involving four wild boars (), endosulfan (an organochlorine pesticide) toxicity in a gaur () and imidacloprid (a neonicotinoid pesticide) toxicity in a wild adult Asian elephant (). This communication thus reports for the first time on the specific chemical compounds identified in wildlife poisoning cases from Kerala state and argues for greater regulation of the sale and use of such toxic compounds in India.
Renewed coexistence: learning from steering group stakeholders on a beaver reintroduction project in England
Species reintroductions are growing in popularity, and example motivations include supporting species populations or the restoration of ecosystem function. Interactions between humans and the reintroduced species are likely to occur post-reintroduction. Coexistence between humans and wildlife is adaptive and dynamic, in part requiring management of conflicts between humans and wildlife, or of conflicts between humans over wildlife management. We seek to learn from the experiences of steering group members in a Eurasian beaver () project in England and identify how governance of coexistence with reintroduced species may differ from the governance of coexistence with species that are already present in the landscape. Using a qualitative thematic analysis of an online survey, we identify a series of lessons in six key areas: (1) project governance, (2) stakeholder engagement, (3) research and monitoring programme, (4) strategy to manage arising conflicts, (5) public engagement, and (6) broad perspectives on reintroduction trials. We advocate for reflective evaluation as an essential component of reintroduction projects to enable knowledge-sharing from experiences, leading to improved practices in the future. Reflecting on our analysis, we identify and define 'Renewed Coexistence'-a new term that draws on pre-existing coexistence knowledge but identifies the unique elements that relate to governing coexistence with reintroduced species.
Severe feather deformation in greater white-fronted goose () goslings during hot summer period on Kolguev Island 2016
In summer 2016, we observed premature feather malformation among goslings of greater white-fronted goose (), between 7 and 10 weeks of age on family gathering areas on Kolguev Island, Russia, the most important breeding island in the Western Palearctic. Rarely reported in wild birds, to our knowledge, this phenomenon has not been recorded in wild geese of this species, despite continuous ringing and marking of thousands of wild geese across Northern Europe and Arctic Siberia. This feather malformations were documented in 36 unfledged goslings showing weak feather basis, deformed or unevenly grown wing feathers or even dead feather buds. Approximately about one-third of all chicks were affected. Feather malformations like this, causing flightless chicks as a result, have never been noticed in any other of our 12 study years since 2006. The lesion was characterised by soft feather buds, weak or incomplete wing feathers and lack of feather development. No other abnormalities were observed in the goslings, so goslings did not differ in weight or body sizes. Affected fledglings never became airworthy and were killed in large numbers by predators or at latest perished during the Arctic winter.
Molecular surveillance revealed no SARS-CoV-2 spillovers to raccoons () in four German federal states
Raccoons (), which are closely related to the family Mustelidae, might be susceptible to natural infection by SARS-CoV-2. This assumption is based on experimental evidence that confirmed the vulnerability of farmed fur-carnivore species, including to SARS-CoV-2. To date, there are no reports of natural SARS-CoV-2 infections of raccoons in Germany. Here, we use RT-PCR to analyze 820 samples from raccoons hunted in Germany with a focus on 4 German federal states (Saxony-Anhalt, Thuringia, Hesse, North Rhine-Westphalia). Lung tissues were homogenized and processed for RNA extraction and RT-qPCR for detecting SARS-CoV-2 was performed. No viral RNA was detected in any samples (0/820). Next, we compared raccoons and human ACE-2 residues that are known to serve for binding with SARS-CoV-2 receptor binding domain (RBD). Interestingly, we found only 60% identity on amino acid level, which may have contributed to the absence of SARS-CoV-2 infections in raccoons. In conclusion, the chance of raccoons being intermediate reservoir hosts for SARS-CoV-2 seems to be very low.
Detection of SARS-CoV-2 in a free ranging leopard () in India
We report an incidence of natural infection of SARS-CoV-2 in free-ranging Indian leopard (). The case was detected during routine screening. Post-mortem and laboratory examination suggested virus-induced interstitial pneumonia. Viral genome could be detected in various organs including brain, lung, spleen, and lymph nodes by real-time PCR. Whole-genome sequence analysis confirmed infection of Pango lineage B.1.617.2 of SARS-CoV-2. Till now, only Asiatic lions have been reported to be infected by SARS-CoV-2 in India. Infections in animals were detected during peak phase of pandemic and all the cases were captive with close contacts with humans, whereas the present case was observed when human cases were significantly low. No tangible evidence linked to widespread infection in the wild population and the incidence seems to be isolated case. High nucleotide sequence homology with prevailing viruses in humans suggested spillover infection to the animal. This report underlines the need for intensive screening of wild animals for keeping track of the virus evolution and development of carrier status of SARS-CoV-2 among wildlife species.
Diseases of Iberian ibex ()
Iberian ibex ( ) is an ecologically and economically relevant medium-sized emblematic mountain ungulate. Diseases participate in the population dynamics of the species as a regulating agent, but can also threaten the conservation and viability of vulnerable population units. Moreover, Iberian ibex can also be a carrier or even a reservoir of pathogens shared with domestic animals and/or humans, being therefore a concern for livestock and public health. The objective of this review is to compile the currently available knowledge on (1) diseases of Iberian ibex, presented according to their relevance on the health and demography of free-ranging populations; (2) diseases subjected to heath surveillance plans; (3) other diseases reported in the species; and (4) diseases with particular relevance in captive Iberian ibex populations. The systematic review of all the information on diseases affecting the species unveils unpublished reports, scientific communications in meetings, and scientific articles, allowing the first comprehensive compilation of Iberian ibex diseases. This review identifies the gaps in knowledge regarding pathogenesis, immune response, diagnostic methods, treatment, and management of diseases in Iberian ibex, providing a base for future research. Moreover, this challenges wildlife and livestock disease and wildlife population managers to assess the priorities and policies currently implemented in Iberian ibex health surveillance and monitoring and disease management.
Modelling Eurasian beaver foraging habitat and dam suitability, for predicting the location and number of dams throughout catchments in Great Britain
Eurasian beaver () populations are expanding across Europe. Depending on location, beaver dams bring multiple benefits and/or require management. Using nationally available data, we developed: a Beaver Forage Index (BFI), identifying beaver foraging habitat, and a Beaver Dam Capacity (BDC) model, classifying suitability of river reaches for dam construction, to estimate location and number of dams at catchment scales. Models were executed across three catchments, in Great Britain (GB), containing beaver. An area of 6747 km was analysed for BFI and 16,739 km of stream for BDC. Field surveys identified 258 km of channel containing beaver activity and 89 dams, providing data to test predictions. Models were evaluated using a categorical binomial Bayesian framework to calculate probability of foraging and dam construction. BFI and BDC models successfully categorised the use of reaches for foraging and damming, with higher scoring reaches being preferred. Highest scoring categories were ca. 31 and 79 times more likely to be used than the lowest for foraging and damming respectively. Zero-inflated negative binomial regression showed that modelled dam capacity was significantly related ( = 0.01) to observed damming and was used to predict numbers of dams that may occur. Estimated densities of dams, averaged across each catchment, ranged from 0.4 to 1.6 dams/km, though local densities may be up to 30 dams/km. These models provide fundamental information describing the distribution of beaver foraging habitat, where dams may be constructed and how many may occur. This supports the development of policy and management concerning the reintroduction and recolonisation of beaver.
Beware of dogs! Domestic animals as a threat for wildlife conservation in Alpine protected areas
Diseases are natural regulating factors of wildlife populations, but some pathogens may become an important threat in wildlife conservation, especially for endangered species. The presence of domestic animals may foster the spread of diseases in natural population, although their role in the dynamic of infections in wildlife is not clear. In this study, we investigated the presence and prevalence of a range of multi-host pathogens in wild species (red fox, Eurasian badger, beech marten, pine marten, stoat for a total of 89 carcasses analysed) and domestic animals ( = 52 shepherd and = 25 companion dogs) living in a protected area of the Alps (the Gran Paradiso National Park) and discussed the role of domestic dogs as possible source of infection for wild species. Our results showed that domestic dogs are potential shedder of three important pathogens: Canine distemper virus, sp. and . In particular, shepherd dogs seem to represent a threat for wildlife as they are exposed to multiple pathogens because of free-roaming, scavenging lifestyles and close proximity to livestock. However, also companion dogs more subject to veterinary care may foster the spread of pathogens. Our results highlight the importance of regulating the access of domestic dogs to protected areas that aim at preserving biodiversity and enhancing the conservation of endangered species.
Fifty years of spring censuses in black grouse () in the High Fens (Belgium): did the rabies vaccination has a negative impact on a fox prey population?
Since 2017, a reinforcement programme was developed to save the last, endangered, Belgian population of black grouse (), in the High Fens Natural Park. To improve the success of this programme, an analysis of past data of this population was undertaken to understand the causes of its past decline. A time series analysis was applied, using annual spring male census data recorded between 1967 and 2016. In the period 1967-1993, there was a fluctuation around an equilibrium of a population of ca. 40-45 males. The peak of 85 males observed in 1971 was probably due to a succession of several favourable years in terms of environmental conditions, albeit without an exceptional annual growth rate. It seems that fox density, by using the occurrence of rabies as a proxy, has an impact on the black grouse population. After 1993, the population dynamic changed drastically, decreasing continuously until finally reaching quasi-extinction. On average, the population lost 15.4% of its size each year. Climate models, applied in previous studies to explain these population trends in the High Fens, failed to describe this major modification in this population's dynamic and its recent decline. We suggest that this negative effect was mainly induced by a significant increase in predation by red fox (), whose abundance has increased considerably since the 1990s, in particular, as a consequence of the eradication of fox rabies. We also discuss alternative hypotheses, such as the impact of other predator species, modification of the natural environment and climatic modifications.
Economic valuation of wildlife conservation
This paper reviews concepts and methods for the economic valuation of nature in the context of wildlife conservation and questions them in light of alternative approaches based on deliberation. Economic valuations have been used to set priorities, consider opportunity costs, assess co-benefits of conservation, support the case for conservation in public awareness and advocacy, and drive novel schemes to change incentives. We discuss the foundational principles of mainstream economic valuation in terms of its assumptions about values, markets, and human behaviour; propose a list of valuation studies in relation to wildlife protection; and explain the methods used. We then review critiques of these approaches focusing on the narrow way in which economics conceives of values, and institutional, power, and equity concerns. Finally, we complement conventional approaches commonly used for wildlife valuation with two forms of deliberative valuation: deliberated preferences and deliberative democratic monetary valuation. These are discussed in terms of their potential to address the drawbacks of mainstream economics and to realise the potential of valuation in bridging conservation of nature for its own sake and its important contributions to human well-being.
Lack of SARS-CoV-2 RNA evidence in the lungs from wild European polecats () from Spain
Data on SARS-CoV-2 infection in wildlife species is limited. The high prevalences found in mustelid species such as free-ranging American minks () and domestic ferrets () justify the study of this virus in the closely related autochthonous free-ranging European polecat (). We analysed lung samples from 48 roadkilled polecats collected when the human infection reached its highest levels in Spain (2020-2021). We did not detect infections by SARS-CoV-2; however, surveillance in wild carnivores and particularly in mustelids is still warranted, due to their susceptibility to this virus.
Your place or mine? Exploring birdwatching tourists' behaviour disturbing birds in a nature reserve
To manage for more sustainable wildlife viewing tourism, a better understanding of tourists' behaviour that might disturb and negatively affect wildlife such as birds is needed. We conducted a qualitative case study of visitors to Hornøya, a protected bird cliff in Northern Norway. Behaviours with the potential to disturb seabirds at the site were explored using the theory of planned behaviour as a guiding framework. In-depth interviews and observations were used to explore why some visitors perform illegal or unwanted, potentially harmful behaviours, and to understand attitudes, social norms, and perceived behaviour control as influencing factors. The tourists visited Hornøya to experience seabirds in their natural surroundings, without causing harm. They were emotionally affected by the bird encounters, which may lead to thoughts and reflections about environmental challenges and nature protection. Visitors generally did not intend to disturb birds during these encounters. However, many visitors interpreted the birds' behaviour as meaning that they were not easily disturbed. Poorly developed social norms among visitors, in combination with limited presence of surveillance/guide personnel, and strong behavioural control among some visitors (willingness and ability to engage in illegal behaviour) led to occasional breaching of formal rules as well as incidents of inappropriate, potentially disturbing behaviour towards birds. Experienced wildlife photographers tended to show willingness and ability to engage in illegal behaviour more often than the two other segments.
Insights into the primate trade into the European Union and the United Kingdom
Illegal and/or unsustainable trade is a major obstacle to effective primate conservation. The wildlife trade in the European Union (EU) and the United Kingdom (UK) is significant, but for many species, such as primates, the trade is poorly understood and sparsely reported. All EU countries are Party to the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES); all primates are listed on Appendix I or II of CITES and are included on Annex A or B of Regulation (EC) No 338/97. We here combine data from several databases (CITES, UN Comtrade, TRAFFIC WiTIS) and seizure reports, to provide a narrative of the trade in primates into and within Europe. The legal import of live primates (2002-2021) amounted to 218,000-238,000 individuals (valued at US$ 869 million), with France, the UK, and Spain as the main importers and Mauritius, Vietnam, and China as the main exporters. Over 21,000 primate parts (trophies, skulls, bodies) were imported mainly from African countries, and UN Comtrade data suggests that ~ 600 tonnes of primate meat was imported mainly from Asia. The vast majority of live primates are either captive-born or captive-bred, and this proportion has increased over time. Reports of the illegal primate trade are far from complete, but the illegal trade of specific species or primate meat can have negative impacts of wild populations of already imperiled species. Stronger policies and more effective enforcement in consumer countries, such as the EU, would also aid in, and garner support for, better protecting primates in primate range states.
The evolutionary dynamics history of canine distemper virus through analysis of the hemagglutinin gene during 1930-2020
Canine distemper virus (CDV) is a lethal viral disease of carnivores which is considered to be a serious threat to domestic and wild species. Despite the widespread use of vaccines, CDV still occurs in vaccinated animals and current vaccines does not guarantee complete protection. In this study, a total of 286 hemagglutinin (H) gene sequences of the virus isolated in 25 countries during 90 years (1930-2020) were analyzed by Bayesian maximum likelihood analysis to estimate the population dynamics. We identified the most recent common ancestor (TMRCA) of the virus in 1868 in the USA which arrived in continental Europe in 1948, and from there, the virus spread rapidly to other continents. The family was identified as the original host as well as a source of the subsequent spread. We identified 11 lineages of geographic co-circulating strains globally. The effective population size experienced a two-phase-exponential growth between 2000-2005 and 2010-2012. Our findings provide a novel insight into the epidemic history of canine distemper virus which may facilitate more effective disease management. This study uses a large set of sequencing data on the H gene of CDV to identify distinct lineages of the virus, track its geographic spread over time, analyze its likelihood of transmission within and between animal families, and provide suggestions for improved strategies to combat the virus.
Hunting techniques and their harvest as indicators of mammal diversity and threat in Northern Angola
Over-exploitation of wildlife especially bushmeat trade is the second most important threat to animal biodiversity. This also applies to Northern Angola but data on bushmeat and hunting techniques for this region are rare. Therefore, we study the most common hunting techniques, frequently captured species, and their economic value, and discuss the local resource use in relation to Angolan law and urgent global crises like the loss of biodiversity, the food supply in South African countries, and the risk of zoonoses. We recorded bushmeat hunting in 27 localities in the province of Uíge, accompanied hunters along their snare lines and interviewed additional 20 locals. Seven main types of snares and traps and their characteristics were defined. Hunters own on average 92 ± 128.7 snares and traps and capture about 25.3 ± 23.6 animals monthly. In total, respondents recognized 28 species of mammals of which one is considered as extinct and two as very rare. The majority of recorded species are hunted regularly. Rodents are most commonly caught followed by primates and duikers. Harvesting rates decrease with species' body size, leading to high economic value of and achievable prices for rare, large animals. Overall, our results document the hunting pressure on mammals and the persisting popularity of bushmeat in Northern Angola which poses an imminent threat to remaining mammal populations. Moreover, it endangers ecosystem integrity, rural livelihoods, and human health through the risk of new zoonoses. Our findings underscore the urgent need for sustainable solutions. The Angolan government should play a more active role in enforcing existing hunting legislation to reduce illegal bushmeat trade.
Enterobacteriaceae and contamination of wild boar () carcasses: comparison between different sampling strategies
During 2020, a total of 64 wild boar carcasses were tested for Enterobacteriaceae count (EBC), and in the abdominal region (i) within 5 h after hunting in the game collection point and (ii) before dressing and processing in the game-handling establishment (GHE) (49 carcasses-average time interval between (i) and (ii): 4.3 days). Because of COVID-19 restrictions, 15 carcasses were transported to a near slaughterhouse (average time interval between (i) and (ii): 2.3 days). Mesenteric lymph nodes (MLNs) were collected and tested for and . Results are shown in relation to sampling A (49 carcasses-GHE) and sampling B (15 carcasses-slaughterhouse). Sampling A: EBC median values were (i) 2.51 log CFU/cm and (ii) 2.79 log CFU/cm. EBC increase between (i) and (ii) was statistically significant ( = 0.001). prevalence on carcasses varied from (i) 2.0 to (ii) 6.1%. Sampling B: EBC median values were (i) 3.1 log CFU/cm and (ii) 3.32 log CFU/cm. EBC increase between (i) and (ii) was not statistically significant ( = 0.191). prevalence on carcasses varied from (i) 6.7 to (ii) 0.0%. The prevalence (sampling A + B) of lymphatic carriers was 7.8% (5/64). From carcasses and/or MNLs, the serovars Enteritidis, Typhimurium, Agama, Zaiman and Diarizonae O:50 (z) were detected. was never isolated. Long chilling periods prior to wild game processing should be avoided, and carcasses should be tested at GHE rather than after shooting to proper reflect the microbial load of wild boar meat entering the food chain.
A study of the incidence of bovine tuberculosis in the wild red deer herd of Exmoor
Bovine tuberculosis (bTB) is an infectious disease which thrives at the wildlife-livestock interface. Exmoor has the largest herd of wild red deer () in England, and also a large number of dairy and beef farms. The population, health and well-being of the herd are managed by a combination of hunting with hounds and by stalking. This study used a serological assay to determine the incidence of bTB in the population of 106 wild red deer of Exmoor, the relationship between regional deer densities and the presence of bTB in deer, and domestic cattle. The overall number of bTB positive deer was 28.3% (30/106). Stalked deer had a slightly higher incidence of bTB (19 out of 55, 34.55%) than hunted deer (11 out of 51, 21.57%). There was no clear pattern of distribution except for one region which showed an incidence of 42.22% compared with 16.4% in the remainder of the moor. There was little difference in the incidence of bTB between male and female animals. The age of animals in the study ranged from < 1 year to over 10 years. There was no clear difference in the incidence across the age range (< 1 year- > 10 years) with the exception of a particularly high incidence in those animals aged 1 year or less. There was a significant correlation between the presence of deer with bTB and the number of farms reporting bTB positive cattle, but not between the regional population of red deer and bTB in deer or cattle.
Seasonal pattern in browsing trees and shrubs by reintroduced Cuvier's gazelles: a study in Morocco with management implications
Cuvier's gazelle () was reintroduced in Tergou reserve (TR) in 2018-2019 to help restore its population at the Moroccan Central Plateau. A study on browsing trees and shrubs was conducted in this reserve to identify (i) the seasonal pattern of this activity and (ii) the most solicited tree and shrub species. Here, we used six random 500-m transects to collect, each month and at each transect, the number of fresh browsing traces of each encountered tree and shrub species. We used principal component analysis (PCA) and linear regression model (LM) to test seasonal variation of the composition of trees and shrubs browsed. Our results showed that, in this reserve, , , , and are mostly browsed during winter, whereas and are mainly used during spring. This information is useful to the managers of the TR not only to consider the importance of the aforementioned trees and shrubs for reintroduced gazelles but also to take the necessary measures so that the forest stand of this reserve is maintained at an appropriate level. Thus, the pursuit of monitoring is necessary to understand the bio-ecology of this semi-captive population more deeply. For this reason, the TR remains a useful and appropriate reference framework offering the opportunity for consolidating and improving the knowledge on this vulnerable mammal species in the view of a future successful release in the wild.
Wild boar visits to commercial pig farms in southwest England: implications for disease transmission
Contact between wild animals and farmed livestock may result in disease transmission with huge financial, welfare and ethical consequences. Conflicts between people and wildlife can also arise when species such as wild boar (Sus scrofa) consume crops or dig up pasture. This is a relatively recent problem in England where wild boar populations have become re-established in the last 20 years following a 500-year absence. The aim of this pilot study was to determine if and how often free-living wild boar visited two commercial pig farms near the Forest of Dean in southwest England. We placed 20 motion-sensitive camera traps at potential entry points to, and trails surrounding, the perimeter of two farmyards housing domestic pigs between August 2019 and February 2021, covering a total of 6030 trap nights. Forty wild boar detections were recorded on one farm spread across 27 nights, with a median (range) of 1 (0 to 7) night of wild boar activity per calendar month. Most of these wild boar detections occurred between ten and twenty metres of housed domestic pigs. No wild boar was detected at the other farm. These results confirm wild boar do visit commercial pig farms, and therefore, there is potential for contact and pathogen exchange between wild boar and domestic pigs. The visitation rates derived from this study could be used to parameterise disease transmission models of pathogens common to domestic pigs and wild boars, such as the African swine fever virus, and subsequently to develop mitigation strategies to reduce unwanted contacts.
Effects of dehorning on population productivity in four Namibia sub-populations of black rhinoceros ()
The black rhinoceros (rhino) () is critically endangered, with poaching being one of several threats to the species' survival. Many reserves across several countries, including Namibia, South Africa, and Zimbabwe, now dehorn their rhinos in an attempt to reduce poaching. Historical data collected by the Namibian Ministry of Environment, Forestry, and Tourism and Save the Rhino Trust were used to investigate whether dehorning has an effect on age at first reproduction (AFR), inter-calving interval (ICI), birth sex ratios, calf survival, cause of death, and lifespan. These were assessed in four Namibian sub-populations (hereafter referred to as 'populations') of black rhino (denoted A, B, C, and D) which have undergone varying levels of dehorning. No significant difference was found in any of the variables between dehorned and horned individuals. Population was a significant predictor of AFR (LRT = 7.433, = 0.024) and ICI (LRT = 13.281, = 0.004), although pairwise comparisons only found populations A and B to be significantly different (AFR: = -2.736, = 0.017, ICI: = -3.705, = 0.001). Additionally, a significantly higher number of males than females were born in population D ( = 0.021, CI = 0.544, 0.960). The main cause of death across all individuals was poaching, although there was no significant difference in the proportion of rhinos poached between dehorned and horned individuals ( = 0.638, = 0.424, = 265). No evidence was found to suggest that dehorning has any effect on AFR, ICI, birth sex ratios, calf survival, or lifespan in the black rhino, which is reassuring in the continued use of dehorning as an anti-poaching technique in this species.
Consequences of polyploidy and divergence as revealed by cytogenetic mapping of tandem repeats in African clawed frogs (, Pipidae)
Repetitive elements have been identified in several amphibian genomes using whole genome sequencing, but few studies have used cytogenetic mapping to visualize these elements in this vertebrate group. Here, we used fluorescence in situ hybridization and genomic data to map the U1 and U2 small nuclear RNAs and histone H3 in six species of African clawed frog (genus ), including, from subgenus , the diploid and its close allotetraploid relative and, from subgenus , the allotetraploid species , , , and . Results allowed us to qualitatively evaluate the relative roles of polyploidization and divergence in the evolution of repetitive elements because our focal species include allotetraploid species derived from two independent polyploidization events - one that is relatively young that gave rise to and another that is older that gave rise to the other (older) allotetraploids. Our results demonstrated conserved loci number and position of signals in the species from subgenus ; allotetraploid has twice as many signals as diploid . However, the content of repeats varied among the other allotetraploid species. We detected almost same number of signals in as in and same number of signals in , , as in the diploid . Overall, these results are consistent with the proposal that allopolyploidization duplicated these tandem repeats and that variation in their copy number was accumulated over time through reduction and expansion in a subset of the older allopolyploids.