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1. Description of the subfossil crocodylians from a new Late Pleistocene subfossil site (Tsaramody, Sambaina Basin) in central Madagascar

   Rakotozandry, R. ; Ranivoharimanana, L. ; Ranaivosoa, V. ; Rasolofomanana, N. ; Hekkala, E. ; Samonds, K.E. ( December 2021 , Malagasy nature)

   Madagascar is famous today not only for its unique biodiversity, but also for the high levels of endemism of plants and animals. Less appreciated is the fact that, in the recent past, the island had even greater biodiversity with many other endemic animals such as giant lemurs, elephant birds, pygmy hippopotami, tortoises, and crocodiles that have gone extinct within the past 2000 years. The extinction of many of these groups is thought to be the result of both human activities and environmental change. Most research has focused on the lemurs, hippopotami, and elephant birds. Other recently extinct animals, including the Malagasy horned crocodile (Voay robustus), are relatively poorly known. Madagascar’s subfossil crocodylians include two taxa: the extinct V. robustus (the Malagasy horned crocodile) and the extant Crocodylus niloticus. The latter arrived on Madagascar relatively recently and we know little about the habitat preferences, distributions and ecological interactions (if any) of either species during the Holocene. In order to better understand the recent history of crocodylian extinction in Madagascar, we must first identify which species were present and where they were found. We present here a description of subfossil crocodylian material collected from the newly discovered subfossil site of Tsaramody (Sambainamore »Basin), a high-elevation wetlandenvironment. At 1655 m, it represents the highest elevation subfossil site on the island. Here we describe both cranial (e.g., premaxillary, jugal, and squamosal “horns”) and postcranial elements (e.g., osteoderms). Our research indicates that crocodile material from Tsaramody appears morphologically to belong to V. robustus, the extinct species. However, oval tuberosities on the frontal bone and a triangular extension of the squamosal bone suggest previously unrecognized variation.« less
2. Subfossil lemur discoveries from the Beanka Protected Area in western Madagascar

   https://doi.org/10.1017/qua.2019.54  

   Burney, David A. ; Andriamialison, Haingoson ; Andrianaivoarivelo, Radosoa A. ; Bourne, Steven ; Crowley, Brooke E. ; de Boer, Erik J. ; Godfrey, Laurie R. ; Goodman, Steven M. ; Griffiths, Christine ; Griffiths, Owen ; et al ( January 2020 , Quaternary Research)

   A new fossil site in a previously unexplored part of western Madagascar (the Beanka Protected Area) has yielded remains of many recently extinct vertebrates, including giant lemurs (Babakotia radofilai, Palaeopropithecus kelyus, Pachylemur sp., and Archaeolemur edwardsi), carnivores (Cryptoprocta spelea), the aardvark-like Plesiorycteropus sp., and giant ground cuckoos (Coua). Many of these represent considerable range extensions. Extant species that were extirpated from the region (e.g., Prolemur simus) are also present. Calibrated radiocarbon ages for 10 bones from extinct primates span the last three millennia. The largely undisturbed taphonomy of bone deposits supports the interpretation that many specimens fell in from a rock ledge above the entrance. Some primates and other mammals may have been prey items of avian predators, but human predation is also evident. Strontium isotope ratios (87Sr/86Sr) suggest that fossils were local to the area. Pottery sherds and bones of extinct and extant vertebrates with cut and chop marks indicate human activity in previous centuries. Scarcity of charcoal and human artifacts suggests only occasional visitation to the site by humans. The fossil assemblage from this site is unusual in that, while it contains many sloth lemurs, it lacks ratites, hippopotami, and crocodiles typical of nearly all other Holocene subfossilmore »sites on Madagascar.« less
3. The stories people tell, and how they can contribute to our understanding of megafaunal decline and extinction in Madagascar

   Nomenjanahary, E.S. ; Freed, B.Z. ; Dollar, L.J. ; Randrianasy, J. ; Godfrey, L.R. ( December 2021 , Malagasy nature)

   Most researchers believe that Madagascar’s megafauna went extinct between 2000 and 1000 years ago. Across Madagascar, fossil specimens of the island’s endemic (and now extinct) pygmy hippopotamuses, elephant birds, giant lemurs, horned crocodiles, and other vertebrates larger in body size than 10 kg commonly date to the first millennium of the Common Era (CE) or earlier; few records date to the second millennium CE. Whereas megafaunal populations appear to have crashed almost simultaneously near the end of the first millennium CE, small populations can survive in remote pockets for centuries after precipitous species decline, perhaps longer. Examining the differences in the population dynamics of declining species and other factors can help to better identify the ultimate timing of extinction. Ever since Etienne de Flacourt traveled to Madagascar in the late 1600s, Malagasy stories of large-bodied wild animals have been recorded. Many include fantastic, clearly mythical creatures, but some provide anatomical or behavioral details which are consistent with legends or even direct observations of real, albeit potentially already extinct, species (including elephant birds, hippopotamuses, and some giant lemurs). In December 1989, at 06:00 hours, one of us (BZF) witnessed a large euplerid carnivoran locally known as fosabe (big fosa) or fosamore »jobijoby (blackish fosa) who had entered his field tent at Montagne d’Ambre. The animal was “twice the size and much darker than the common fossa” (Freed, 1996, p. 34). The individual was black and weighed approximately 20-25 kg. Freed wrote that the animal was well known to the local people and that “many local people also reported seeing it”. The animal fits paleontologists’ expectations for Cryptoprocta spelea, a large carnivoran known from the fossil record, believed to have been extinct for at least 1000 years. In June 2020, we recorded modern accounts of the big fosa. One of us (ESN) visited villages in four different sectors (Northwest, Northeast, East, and West) of Montagne d’Ambre National Park and the Forêt d’Ambre Special Reserve to examine potential regional differences and/or similarities in the stories of this animal, and whether such accounts include mythical elements, relevant anatomical information, and/or credible recent sightings. We also recorded stories of an Endangered extant animal, the aye-aye of the genus Daubentonia (also known locally as the kakahely). Ultimately, we believe this folklore provides clues that may help elucidate the geography of decline and possible late survival of an “extinct” megafaunal animal on Madagascar.« less
4. Evolutionary and phylogenetic insights from a nuclear genome sequence of the extinct, giant, “subfossil” koala lemur Megaladapis edwardsi

   https://doi.org/10.1073/pnas.2022117118  

   Marciniak, Stephanie ; Mughal, Mehreen R. ; Godfrey, Laurie R. ; Bankoff, Richard J. ; Randrianatoandro, Heritiana ; Crowley, Brooke E. ; Bergey, Christina M. ; Muldoon, Kathleen M. ; Randrianasy, Jeannot ; Raharivololona, Brigitte M. ; et al ( June 2021 , Proceedings of the National Academy of Sciences)

   No endemic Madagascar animal with body mass >10 kg survived a relatively recent wave of extinction on the island. From morphological and isotopic analyses of skeletal “subfossil” remains we can reconstruct some of the biology and behavioral ecology of giant lemurs (primates; up to ∼160 kg) and other extraordinary Malagasy megafauna that survived into the past millennium. Yet, much about the evolutionary biology of these now-extinct species remains unknown, along with persistent phylogenetic uncertainty in some cases. Thankfully, despite the challenges of DNA preservation in tropical and subtropical environments, technical advances have enabled the recovery of ancient DNA from some Malagasy subfossil specimens. Here, we present a nuclear genome sequence (∼2× coverage) for one of the largest extinct lemurs, the koala lemur Megaladapis edwardsi (∼85 kg). To support the testing of key phylogenetic and evolutionary hypotheses, we also generated high-coverage nuclear genomes for two extant lemurs, Eulemur rufifrons and Lepilemur mustelinus , and we aligned these sequences with previously published genomes for three other extant lemurs and 47 nonlemur vertebrates. Our phylogenetic results confirm that Megaladapis is most closely related to the extant Lemuridae (typified in our analysis by E. rufifrons ) to the exclusion of L. mustelinus , whichmore »contradicts morphology-based phylogenies. Our evolutionary analyses identified significant convergent evolution between M. edwardsi and an extant folivore (a colobine monkey) and an herbivore (horse) in genes encoding proteins that function in plant toxin biodegradation and nutrient absorption. These results suggest that koala lemurs were highly adapted to a leaf-based diet, which may also explain their convergent craniodental morphology with the small-bodied folivore Lepilemur .« less
5. Ecological Consequences of a Millennium of Introduced Dogs on Madagascar

   https://doi.org/10.3389/fevo.2021.689559  

   Hixon, Sean W. ; Douglass, Kristina G. ; Godfrey, Laurie R. ; Eccles, Laurie ; Crowley, Brooke E. ; Rakotozafy, Lucien Marie ; Clark, Geoffrey ; Haberle, Simon ; Anderson, Atholl ; Wright, Henry T. ; et al ( June 2021 , Frontiers in Ecology and Evolution)

   Introduced predators currently threaten endemic animals on Madagascar through predation, facilitation of human-led hunts, competition, and disease transmission, but the antiquity and past consequences of these introductions are poorly known. We use directly radiocarbon dated bones of introduced dogs ( Canis familiaris ) to test whether dogs could have aided human-led hunts of the island’s extinct megafauna. We compare carbon and nitrogen isotope data from the bone collagen of dogs and endemic “fosa” ( Cryptoprocta spp.) in central and southwestern Madagascar to test for competition between introduced and endemic predators. The distinct isotopic niches of dogs and fosa suggest that any past antagonistic relationship between these predators did not follow from predation or competition for shared prey. Radiocarbon dates confirm that dogs have been present on Madagascar for over a millennium and suggest that they at least briefly co-occurred with the island’s extinct megafauna, which included giant lemurs, elephant birds, and pygmy hippopotamuses. Today, dogs share a mutualism with pastoralists who also occasionally hunt endemic vertebrates, and similar behavior is reflected in deposits at several Malagasy paleontological sites that contain dog and livestock bones along with butchered bones of extinct megafauna and extant lemurs. Dogs on Madagascar have had amore »wide range of diets during the past millennium, but relatively high stable carbon isotope values suggest few individuals relied primarily on forest bushmeat. Our newly generated data suggest that dogs were part of a suite of animal introductions beginning over a millennium ago that coincided with widespread landscape transformation and megafaunal extinction.« less