Les hommes ont oublié cette vérité. Mais tu ne dois pas l'oublier, dit le renard. Tu deviens responsable pour toujours de ce que tu as apprivoisé.
Le Petit Prince, chap. 21

Saturday, 27 August 2016

Feral cats overlap with prey and competitors in primary and altered habitats

Bogdan, V., Jůnek, T., & Vymyslická, P. J. (2016). Temporal overlaps of feral cats with prey and competitors in primary and human-altered habitats on Bohol Island, Philippines. PeerJ, 4, e2288.

The vertebrate fauna of the Philippines, known for its diversity and high proportion of endemic species, comprises mainly small- to medium-sized forms with a few large exceptions. As with other tropical ecosystems, the major threats to wildlife are habitat loss, hunting and invasive species, of which the feral cat (Felis catus) is considered the most damaging. Our camera-trapping study focused on a terrestrial vertebrate species inventory on Bohol Island and tempo-spatial co-occurrences of feral cats with their prey and competitors. The survey took place in the Rajah Sikatuna Protected Landscape, and we examined the primary rainforest, its border with agricultural land, and rural areas in the vicinity of villages. Altogether, over 2,885 trap days we captured 30 species of vertebrates–10 mammals (including Sus philippensis), 19 birds and one reptile, Varanus cumingi. We trapped 81.8% of expected vertebrates. Based on the number of events, the most frequent native species was the barred rail (Gallirallus torquatus). The highest overlap in diel activity between cats and potential prey was recorded with rodents in rural areas (Δ = 0.62); the lowest was in the same habitat with ground-dwelling birds (Δ = 0.40). Cat activity was not recorded inside the rainforest; in other habitats their diel activity pattern differed. The cats’ activity declined in daylight in the proximity of humans, while it peaked at the transition zone between rainforest and fields. Both rodents and ground-dwelling birds exhibited a shift in activity levels between sites where cats were present or absent. Rodents tend to become active by day in cat-free habitats. No cats’ temporal response to co-occurrences of civets (Paradoxurus hermaphroditus and Viverra tangalunga) was found but cats in diel activity avoided domestic dogs (Canis lupus familiaris). Our first insight into the ecology of this invasive predator in the Philippines revealed an avoidance of homogeneous primary rainforest and a tendency to forage close to human settlements in heterogeneous habitats. A detailed further investigation of the composition of the cat’s diet, as well as ranging pattern, is still needed.

Stray dogs den close to humans

Majumder, S. S., Paul, M., Sau, S., & Bhadra, A. (2016). Denning habits of free-ranging dogs reveal preference for human proximity. Scientific Reports, 6, 32014.

Dens are crucial in the early development of many mammals, making den site selection an important component of parental care in such species. Resource availability and shelter from predators primarily govern den selection. Species inhabiting human-dominated landscapes typically den away from human disturbance, often shifting dens to avoid humans during the early life of their young. Domesticated dogs have evolved in human proximity over centuries, being bred and reared in human homes for generations. While pets rely on their owners for shelter and care, free-ranging dogs roam uncared, and typically whelp in dens. We conducted a study on 148 free-ranging dog dens in India to understand their denning habits. Distance from resources influenced den choice, but anthropogenic disturbance did not. Dens were found in areas of high human activity, and begging from humans was preferred over scavenging. A study on 15 pregnant females revealed that females actively searched for denning sites, rejecting several intermediate ones before selecting the final den. We propose that the obvious preference of dogs for denning close to humans is a behavioural adaptation that helps them to survive in the urban landscape, in spite of the high human induced mortality during the early life of pups.

Saturday, 20 August 2016

Importance of CDV infection in free-ranging Iberian lynxes

Meli, M. L., Simmler, P., Cattori, V., Martínez, F., Vargas, A., Palomares, F., ... & Hofmann-Lehmann, R. (2010). Importance of canine distemper virus (CDV) infection in free-ranging Iberian lynxes (Lynx pardinus). Veterinary microbiology, 146(1), 132-137.

Canine distemper virus (CDV) is a morbillivirus that is the etiological agent of one of the most important viral diseases affecting canids and an expanding range of other carnivores. Using real-time RT-PCR, CDV RNA was detected in organs of an Iberian lynx (Lynx pardinus) found dead in the Doñana National Park, Southwestern Andalusia, Spain. This finding may be of great importance for the conservation of the species; at present the Iberian lynx is the most critically endangered wild felid. The aim of the present study was to elucidate the significance of CDV for the Iberian lynx population. High viral loads were evident in the dead lynx, suggesting an etiological involvement of CDV in its death. When carnivores from the same region were analyzed by CDV RT-PCR, a stone marten (Martes foina) was positive. Phylogenetic analyses demonstrated high identity of the two detected CDVs and a close relationship to the European dog lineage of CDV. Antibodies to CDV were detected in 14.8% of 88 tested free-ranging Iberian lynxes. The sample seroprevalence was significantly higher in lynxes from the Doñana Natural Space (22.9%) than Sierra Morena (5%). The stone marten and a red fox (Vulpes vulpes) also tested seropositive. In conclusion, CDV is present in the Iberian lynx population, especially in the Doñana region, with sporadic cases of disease. To reduce the infectious pressure of CDV on this endangered population, a mass dog vaccination should be considered.

Human-cat relationship in an oceanic biosphere reserve

Medina, F. M., Nogales, M., Farnworth, M. J., & Bonnaud, E. (2016). Human-cat relationship in an oceanic biosphere reserve: The case of La Palma Island, Canary archipelago. Journal for Nature Conservation.

Removal of feral cats from island environments is a useful mechanism by which their ecological impact on endangered species can be reduced or ended. Nevertheless, because cats are anthropogenic in their origins, social perceptions of management practices play a large role in their implementation. Four-hundred questionnaires were delivered (386 were returned) with 100 going to each of the following: local residents; environmental workers; tourists; and, hunters. Questions explored respondents’ knowledge about island biodiversity and invasive species as well as attitudes towards cat population management methods. Habitat destruction and introduction of invasive species were considered the main threats for the conservation of island biodiversity. Most respondents considered cats to have a negative impact on biodiversity and sterilization campaigns were considered most appropriate for cat population control. Several free sterilization campaigns have been conducted in La Palma Island Biosphere Reserve in order to reduce free-ranging cats and were well received by local people. This research, which combined concepts of management, ecology and social sciences, provides valuable insights which may to be applicable on several other islands where cats and people are present and in conflict with conservation priorities.

Thursday, 18 August 2016

Preliminary data on the distribution of free-ranging dogs in a Bulgarian National Park

Doykin, N., Popova, E., Zlatanov, V., Petrov, P., & Zlatanova, D. 2016. Preliminary data on the distribution of free-ranging dogs (Canis familiaris L.) in NP Vitosha, Bulgaria. Annuaire de l’Université de Sofia “St. Kliment Ohridski” Faculte de Biologie 2016, volume 101, livre 4, pp. 11-22. Youth Scientific Conference “Kliment’s Days”, Sofia 2015

Free-ranging dogs often leave the urbanized areas and stray into nearby mountainous habitats, even entering protected areas. This causes problems for the wildlife due to either direct predation or disturbance. Our camera trap survey (July 2013 - November 2014) in NP Vitosha, Bulgaria resulted in a total of 199 independent registrations of free-ranging dogs in 81 locations. In this preliminary study, we present the distribution, habitat selection, and distance from settlements and activity of free ranging dogs in Vitosha NP. The free-ranging dogs in Vitosha are predominantly diurnal, and show preference towards coniferous and mixed forests, mostly closed. Their distribution and activity patterns are influenced by human presence, which is due to the fact that they at least partially rely on human-sourced food. Some data for observations of wild animals influenced by dogs is also discussed. 

Wednesday, 17 August 2016

Analysis of six Latin American cat populations through coat genes and molecular microsatellite markers

Ruiz-García, M., & Alvarez, D. (2003). Análisis de seis poblaciones latinoamericanas de gatos mediante genes del pelaje y marcadores microsatélites. Acta zoológica mexicana, (89), 261-286. (Analysis of six Latin American cat populations through coat genes and molecular microsatellite markers)

Six Latin American cat populations (La Havana, San Jose, Bogotá, Asunción, Buenos Aires and Santiago) have been studied from a population genetics standpoint by using different morphological coat and molecular microsatellite markers (FCA43, FCA45, FCA96 and FCA126). The main aims of the current work are as follows: (1) To determine whether the type and intensity of the genetic differences found for diverse morphological loci among the current British cat populations and those from the British oversea colonies (USA, Canada and Australia) agree with the differences among the current Spanish cat populations and those from Latin America and (2) to determine if the genetic relationships among some of these Latin American cat populations are in agreement by using independently morphological and molecular microsatellite markers. The different results obtained were as follows: (A) All populations analyzed were in Hardy-Weinberg equilibrium at the O, S and at the four microsatellite loci studied with the exception of La Havana at the S locus. (B) The trees obtained showed that the relationships of the six cat populations studied regard to the Spanish populations, in particular, and with the European populations, in general, were extremely heterogeneous. Therefore, for instance, Asuncion was genetically identical to some Catalonian populations meanwhile Santiago (Chile) revealed more resemblance with the cat populations of presumed British origin in the Eastern Coast of the United States by means of the coat color genes. The striking genetic heterogeneity among some of these Latin American cat populations could be explained by the existence of different geographic, or temporal, migrations from Spain and/or that diverse gene drift degrees were present in the foundation of the diverse populations studied. Finally, the molecular results were similar to those obtained with the gross morphological genes. Therefore, the overall evolution of these morphological markers is controlled more probably by neutral stochastic forces than by selective ones.

Seis poblaciones latinoamericanas de gatos (La Habana, San José, Bogotá, Asunción, Buenos Aires y Santiago) han sido estudiadas desde una perspectiva genético poblacional con marcadores que codifican características morfológicas del pelaje y marcadores moleculares nucleares microsatélites (FCA43, FCA45, FCA96, FCA126). A partir de las frecuencias alélicas de ambos tipos de marcadores genéticos se investigó: (1) si el tipo y la intensidad de las diferencias genéticas encontradas para diversos loci morfológicos entre las poblaciones de gatos en Gran Bretaña y en sus ex-colonias transmarítimas (EU, Canadá, Australia) se dio también entre las poblaciones de gatos actuales en España y en Latinoamérica y (2) si las relaciones genéticas de esos caracteres morfológicos entre algunas de esas poblaciones latinoamericanas de gatos fue paralela a las relaciones encontradas con marcadores moleculares microsatélites. Los resultados obtenidos fueron: (A) Todas las poblaciones analizadas estuvieron en equilibrio Hardy-Weinberg para los loci O, S y para los cuatro loci microsatélites estudiados, con la excepción de la población de La Habana para el locus S. (B) Los fenogramas obtenidos mostraron que las relaciones de las seis poblaciones latinoamericanas de gatos respecto a las poblaciones españolas y europeas fueron muy heterogéneas. Por ejemplo, la población de Asunción (Paraguay) fue genéticamente indistinguible de algunas poblaciones de gatos analizadas en Cataluña, tanto con los genes morfológicos como con los microsatélites, mientras que Santiago presentó más semejanzas con las poblaciones de gatos de presunto origen británico en la costa Este de los Estados Unidos cuando se utilizaron los genes del pelaje. La fuerte heterogeneidad genética entre algunas de las poblaciones latinoamericanas estudiadas hace pensar en que diversas migraciones geográficas, o temporales, se dieron desde España, o que diversos grados de deriva genética se dieron en la fundación de las diferentes poblaciones latinoamericanas estudiadas. Finalmente, los resultados moleculares son similares a los obtenidos con los genes de codificación morfológica por lo que la evolución global de éstos parece más modulada por fuerzas neutrales que selectivas.

Saturday, 13 August 2016

Spatial ecology and population genetics of cats living in or near conservation-sensitive areas

Cross, C. (2016). Spatial ecology and population genetics of cats (Felis catus) living in or near conservation-sensitive areas (Doctoral dissertation, University of Otago).

Human-mediated dispersal of organisms across the world has resulted in species introductions into many vulnerable ecosystems. Invasive mammalian predators have had detrimental impacts on native island biota, leading to declines and extinctions of many endemic prey species. Humans have transported cats (Felis catus) across the world as mousers on ships and as companion animals. The role cats (especially feral) have played in the decline and extinction of several island species is clear; however, different types of cats classified by their associations with humans has an influence on the public perception of cat impacts on wildlife and acceptance of appropriate management strategies.
I studied the spatial ecology of two different types of cats in two different conservation-sensitive areas (Te Anau Basin and Canterbury/North Otago) in the South Island of New Zealand. I conducted this research to gain an insight into companion cat spatial ecology and feral cat population genetics. Specifically, to investigate individual cat movement patterns and population level movements to discover putative geographic barriers to movement. Additionally, I intended to aid formulation and reinforcement of appropriate and current management strategies with respect to conservation-sensitive areas that support high levels of native biodiversity.
Cat capture rates in the Tasman Valley from March 2005 toFebruary 2013 (included as 2012).

In the Te Anau Basin, the township of Te Anau lies on the edge of Lake Te Anau, directly adjacent to Fiordland National Park. The Kepler Mire conservation area, also situated in the Te Anau Basin, is a nearby wetland that supports a diverse range of fauna. I GPS tracked 32 local companion cats (11F:21M) for a maximum of 10 to 14 days over the austral spring/summer. I recorded a total of 19,157 locations prior to filtering data for erroneous locations. Home range and habitat analysis were performed on a filtered dataset of 13,241 locations using 100% minimum convex polygons (MCP) and Objective-Restricted-Edge Polygons (OREP). Dispersal barriers might be acting to prevent movement of tracked cats into Fiordland National Park, but not the Kepler Mire conservation area. I found males (mean MCP: 22.13 ha, OREP: 1.05 ha) exhibited larger movements (home range and distance travelled from home) than females (mean MCP: 8.83 ha, OREP: 0.45 ha) and rural-living cats (mean MCP: 32.54 ha, OREP 1.33 ha) exhibited larger movements than urban-living cats (mean MCP: 5.90 ha, OREP: 0.46 ha).  Cats showed a tendency to preferentially select Built, Cover and Sealed habitat features.  Although there was great individual variation in the ranging behaviour, there was no sex or age-related difference observed in the cats’ resource selection.
To infer population movements, I used 10 microsatellite loci and a sexidentification marker, in a multiplex framework, to infer population structure of 157 feral cats in the upper Waitaki Basin (Tasman Valley, Ohau River and Ahuriri Valley) and Macraes Flat. I found some evidence of population connectivity between the sites based on migration rates and low FST values, indicating features in the landscape that act to facilitate dispersal. Bayesian clustering analysis noted the presence of three separate clusters; however, assignment rates were low for the Ohau River, Tasman Valley and Macraes Flat sites. Spatial autocorrelation and Mantel tests indicated rough terrain (i.e. mountain ranges) might limit dispersal. Macraes Flat and Ohau River might function as man-made sinks due to lower relatedness scores. Lower relatedness, genetic differentiation scores, and proximity to human habituation suggested there might be genetic input from nearby stray and companion cat populations. Due to large movements exhibited by feral cats in these areas, reinvasion into trapped areas seems likely; however, the Tasman Valley might be able to be managed as an eradication unit, if movement out of the Ohau River and surrounding area is reduced. Continued genetic monitoring of
these sites and sampling of local stray and companion cats might help to identify if there is connectivity between different types of cats (i.e. companion, stray and feral). Additionally, continued genetic monitoring might be able to determine if genetic differentiation increases between each site in response to trapping operations.
Tighter regulations regarding companion cat management might aid New Zealand conservation efforts by reducing and restricting movement and cat interactions with native wildlife. Stricter companion and stray cat regulations might also benefit feral cat control efforts; however, this aspect requires further analysis. 

Sunday, 7 August 2016

High-impact conservation: invasive mammal eradications from the islands of western Mexico

Aguirre-Muñoz, A., Croll, D. A., Donlan, C. J., Henry III, R. W., Hermosillo, M. A., Howald, G. R., ... & Samaniego-Herrera, A. (2008). High-impact conservation: invasive mammal eradications from the islands of western Mexico. AMBIO: A Journal of the Human Environment, 37(2), 101-107.

Islands harbor a disproportionate amount of the earth's biodiversity, but a significant portion has been lost due in large part to the impacts of invasive mammals. Fortunately, invasive mammals can be routinely removed from islands, providing a powerful tool to prevent extinctions and restore ecosystems. Given that invasive mammals are still present on more than 80% of the world's major islands groups and remain a premier threat to the earth's biodiversity, it is important to disseminate replicable, scaleable models to eradicate invasive mammals from islands. We report on a successful model from western México during the past decade. A collaborative effort between nongovernmental organizations, academic biologists, Mexican government agencies, and local individuals has resulted in major restoration efforts in three island archipelagos. Forty-two populations of invasive mammals have been eradicated from 26 islands. For a cost of USD 21 615 per colony and USD 49 370 per taxon, 201 seabird colonies and 88 endemic terrestrial taxa have been protected, respectively. These conservation successes are a result of an operational model with three main components: i) a tri-national collaboration that integrates research, prioritization, financing, public education, policy work, capacity building, conservation action, monitoring, and evaluation; ii) proactive and dedicated natural resource management agencies; and iii) effective partnerships with academic researchers in México and the United States. What is now needed is a detailed plan to eradicate invasive mammals from the remaining islands in the region that integrates the needed additional financing, capacity, technical advances, and policy issues. Island conservation in western México provides an effective approach that can be readily applied to other archipelagos where conservation efforts have been limited.

Saturday, 6 August 2016

Ancient wolf genome reveals an early divergence of domestic dog ancestors and admixture into high-latitude breeds

Skoglund, P., Ersmark, E., Palkopoulou, E., & Dalén, L. (2015). Ancient wolf genome reveals an early divergence of domestic dog ancestors and admixture into high-latitude breeds. Current Biology, 25(11), 1515-1519.

The origin of domestic dogs is poorly understood, with suggested evidence of dog-like features in fossils that predate the Last Glacial Maximum  conflicting with genetic estimates of a more recent divergence between dogs and worldwide wolf populations. Here, we present a draft genome sequence from a 35,000-year-old wolf from the Taimyr Peninsula in northern Siberia. We find that this individual belonged to a population that diverged from the common ancestor of present-day wolves and dogs very close in time to the appearance of the domestic dog lineage. We use the directly dated ancient wolf genome to recalibrate the molecular timescale of wolves and dogs and find that the mutation rate is substantially slower than assumed by most previous studies, suggesting that the ancestors of dogs were separated from present-day wolves before the Last Glacial Maximum. We also find evidence of introgression from the archaic Taimyr wolf lineage into present-day dog breeds from northeast Siberia and Greenland, contributing between 1.4% and 27.3% of their ancestry. This demonstrates that the ancestry of present-day dogs is derived from multiple regional wolf populations.
A model of population history (admixture graph) fitted to the data

Thursday, 4 August 2016

Exposure of Free-Ranging Wild Carnivores and Domestic Dogs to CDV and Parvovirus in Central Brazil

Furtado, M. M., Hayashi, E. M. K., Allendorf, S. D., Coelho, C. J., de Almeida Jácomo, A. T., Megid, J., ... & Neto, J. S. F. Exposure of Free-Ranging Wild Carnivores and Domestic Dogs to Canine Distemper Virus and Parvovirus in the Cerrado of Central Brazil. EcoHealth, 1-9.

Human population growth around protected areas increases the contact between wild and domestic animals, promoting disease transmission between them. This study investigates the exposure of free-ranging wild carnivores and domestic dogs to canine distemper virus (CDV) and parvovirus in Emas National Park (ENP) in the Cerrado savanna of central Brazil. Serum samples were collected from 169 wild carnivores, including the maned wolf (Chrysocyon brachyurus), crab-eating fox (Cerdocyon thous), hoary fox (Pseudalopex vetulus), puma (Puma concolor), ocelot (Leopardus pardalis), pampas cat (Leopardus colocolo), jaguarundi (Herpailurus yagouaroundi), striped hog-nosed skunk (Conepatus semistriatus) and coati (Nasua nasua), and from 35 domestic dogs living on rural properties bordering ENP. Serological tests showed that 10.6% of wild carnivores (maned wolves, crab-eating foxes and ocelots) and 71.4% of domestic dogs were exposed to CDV, and 56.8% of wild carnivores, including all species sampled except coatis, and 57.1% of domestic dogs were exposed to parvovirus. This report is the first to indicate that the free-ranging pampas cat, jaguarundi and striped hog-nosed skunk are exposed to parvovirus. CDV and parvovirus deserve attention in ENP, and it is extremely important to monitor the health of carnivore populations and perform molecular diagnosis of the viruses to determine the possible involvement of the domestic dog in their transmission.

Home range of feral cats on Rota Is.

Leo, B. T., Anderson, J. J., Brand Phillips, R., & Ha, R. R. (2016). Home range estimates of feral cats (Felis catus) on Rota Island and determining asymptotic convergence 1. Pacific Science, 70(3), 323-331.

Feral cats (Felis catus) have been shown to be a main contributor to species decline throughout the world and are especially threatening to insular species that lack appropriate defense characteristics. To mitigate the impact of feral cats on threatened species, space-use data are commonly used to design control strategies. In this article we report on the performance of GPS datalogging collars and provide baseline information on daily space use and home ranges of feral cats that threaten an endangered species on Rota Island in the Commonwealth of the Northern Mariana Islands. Using 100% Minimum Convex Polygon (MCP), average adult male home range was 1.32 km2(n = 2) and average adult female home range was 0.22 km2(n = 3). Home ranges were deemed fully revealed if asymptotes were approached using incremental analysis. Currently, there is no objective method for assessing where an asymptote is approached. Here, we describe a methodology to do so with the application of a Michaelis-Menten model to incremental data. We conclude that GPS datalogging collars are a viable tool for feral cat location data collection on Rota Island and that the Michaelis-Menten model is useful for determining asymptotic convergence of incremental location data.

Epidemiological survey of zoonotic helminths in feral cats in Gran Canaria

Rodríguez-Ponce, E., González, J. F., de Felipe, M. C., Hernández, J. N., & Raduan Jaber, J. (2016). Epidemiological survey of zoonotic helminths in feral cats in Gran Canaria island (Macaronesian archipelago-Spain). Acta Parasitologica, 61(3), 443-450.

The presence of zoonotic parasites in feral cats have been widely considered all over the world. In Gran Canaria (Macaronesian archipelago, Canary Islands, Spain) the number of feral cats has grown out of control in urban and rural areas. 48 of Felis catus captured in different Gran Canaria areas were studied. Animals were necropsied and several organs were systematically examined in order to collect and identify macroscopic parasites. In addition, coprological tests were done in 28 cats. There were no statistically significant differences in the prevalence rate among sex, age or capture area, showing an overall prevalence of helminths of 77.1%. The most common tapeworms were Dipylidium caninum (64.6%) and Taenia taeniaeformis (31.3%), followed by the nematodes Toxocara cati (20.8%), Ancylostoma tubaeforme (18.8%), Aelurostrongylus abstrusus (10.4%) and Trichuris vulpis (2.08%). We also find several eggs of Alaria alata in the small intestine of one cat (2.08%), being the first description of this trematode in cats in the Canary Islands. Aproximatelly, 40% of the studied cats harboured more than one parasite. High rates of zoonotic species found in these animals suggest the need of controling parasitic infections and preventive measures against them.

Wednesday, 3 August 2016

3D morphometric analysis of fossil canid skulls contradicts the suggested domestication of dogs during the late Paleolithic

Drake, A. G., Coquerelle, M., & Colombeau, G. (2015). 3D morphometric analysis of fossil canid skulls contradicts the suggested domestication of dogs during the late Paleolithic. Scientific reports, 5.

Whether dogs were domesticated during the Pleistocene, when humans were hunter-gatherers, or during the Neolithic, when humans began to form permanent settlements and engage in agriculture, remains controversial. Recently discovered Paleolithic fossil skulls, Goyet dated 31,680 +/− 250 YBP and Eliseevichi MAE 447/5298 dated 13,905 +/− 55 YBP, were previously identified as dogs. However, new genetic studies contradict the identification of these specimens as dogs, questioning the validity of traditional measurements used to morphologically identify canid fossil skulls. We employ 3D geometric morphometric analyses to compare the cranial morphology of Goyet and Eliseevichi MAE to that of ancient and modern dogs and wolves. We demonstrate that these Paleolithic canids are definitively wolves and not dogs. Compared to mesaticephalic (wolf-like breeds) dog skulls, Goyet and Eliseevichi MAE, do not have cranial flexion and the dorsal surface of their muzzles has no concavity near the orbits. Morphologically, these early fossil canids resemble wolves, and thus no longer support the establishment of dog domestication in the Paleolithic.

(a) 3D plot of PC1, PC2 and PC3, (b) 3D morphs of extremes along PC 1, PC 2 and PC 3. 95% Confidence interval ellipsoids of modern dogs, Alaskan wolves, and European wolves are outlined. Unclassified specimens are labelled separately in this and other figures. A 3D version of this figure is available as Supplementary Figure S1.

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