skip to main content


Title: Madagascar’s extraordinary biodiversity: Evolution, distribution, and use
BACKGROUND The Republic of Madagascar is home to a unique assemblage of taxa and a diverse set of ecosystems. These high levels of diversity have arisen over millions of years through complex processes of speciation and extinction. Understanding this extraordinary diversity is crucial for highlighting its global importance and guiding urgent conservation efforts. However, despite the detailed knowledge that exists on some taxonomic groups, there are large knowledge gaps that remain to be filled. ADVANCES Our comprehensive analysis of major taxonomic groups in Madagascar summarizes information on the origin and evolution of terrestrial and freshwater biota, current species richness and endemism, and the utilization of this biodiversity by humans. The depth and breadth of Madagascar’s biodiversity—the product of millions of years of evolution in relative isolation —is still being uncovered. We report a recent acceleration in the scientific description of species but many remain relatively unknown, particularly fungi and most invertebrates. DIGITIZATION Digitization efforts are already increasing the resolution of species richness patterns and we highlight the crucial role of field- and collections-based research for advancing biodiversity knowledge in Madagascar. Phylogenetic diversity patterns mirror that of species richness and endemism in most of the analyzed groups. Among the new data presented, our update on plant numbers estimates 11,516 described vascular plant species native to Madagascar, of which 82% are endemic, in addition to 1215 bryophyte species, of which 28% are endemic. Humid forests are highlighted as centers of diversity because of their role as refugia and centers of recent and rapid radiations, but the distinct endemism of other areas such as the grassland-woodland mosaic of the Central Highlands and the spiny forest of the southwest is also important despite lower species richness. Endemism in Malagasy fungi remains poorly known given the lack of data on the total diversity and global distribution of species. However, our analysis has shown that ~75% of the fungal species detected by environmental sequencing have not been reported as occurring outside of Madagascar. Among the 1314 species of native terrestrial and freshwater vertebrates, levels of endemism are extremely high (90% overall)—all native nonflying terrestrial mammals and native amphibians are found nowhere else on Earth; further, 56% of the island’s birds, 81% of freshwater fishes, 95% of mammals, and 98% of reptile species are endemic. Little is known about endemism in insects, but data from the few well-studied groups on the island suggest that it is similarly high. The uses of Malagasy species are many, with much potential for the uncovering of useful traits for food, medicine, and climate mitigation. OUTLOOK Considerable work remains to be done to fully characterize Madagascar’s biodiversity and evolutionary history. The multitudes of known and potential uses of Malagasy species reported here, in conjunction with the inherent value of this unique and biodiverse region, reinforce the importance of conserving this unique biota in the face of major threats such as habitat loss and overexploitation. The gathering and analysis of data on Madagascar’s remarkable biota must continue and accelerate if we are to safeguard this unique and highly threatened subset of Earth’s biodiversity. Emergence and composition of Madagascar’s extraordinary biodiversity. Madagascar’s biota is the result of over 160 million years of evolution, mostly in geographic isolation, combined with sporadic long distance immigration events and local extinctions. (Left) We show the age of the oldest endemic Malagasy clade for major groups (from bottom to top): arthropods, bony fishes, reptiles, flatworms, birds, amphibians, flowering plants, mammals, non-flowering vascular plants, and mollusks). Humans arrived recently, some 10,000 to 2000 years (top right) and have directly or indirectly caused multiple extinctions (including hippopotamus, elephant birds, giant tortoises, and giant lemurs) and introduced many new species (such as dogs, zebu, rats, African bushpigs, goats, sheep, rice). Endemism is extremely high and unevenly distributed across the island (the heat map depicts Malagasy palm diversity, a group characteristic of the diverse humid forest). Human use of biodiversity is widespread, including 1916 plant species with reported uses. The scientific description of Malagasy biodiversity has accelerated greatly in recent years (bottom right), yet the diversity and evolution of many groups remain practically unknown, and many discoveries await.  more » « less
Award ID(s):
1655076
NSF-PAR ID:
10436279
Author(s) / Creator(s):
; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; more » ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; ; « less
Date Published:
Journal Name:
Science
Volume:
378
Issue:
6623
ISSN:
0036-8075
Format(s):
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. BACKGROUND Madagascar is one of the world’s foremost biodiversity hotspots. Its unique assemblage of plants, animals, and fungi—the majority of which evolved on the island and occur nowhere else—is both diverse and threatened. After human arrival, the island’s entire megafauna became extinct, and large portions of the current flora and fauna may be on track for a similar fate. Conditions for the long-term survival of many Malagasy species are not currently met because of multiple anthropogenic threats. ADVANCES We review the extinction risk and threats to biodiversity in Madagascar, using available international assessment data as well as a machine learning analysis to predict the extinction risks and threats to plant species lacking assessments. Our compilation of global International Union for Conservation of Nature (IUCN) Red List assessments shows that overexploitation alongside unsustainable agricultural practices affect 62.1 and 56.8% of vertebrate species, respectively, and each affects nearly 90% of all plant species. Other threats have a relatively minor effect today but are expected to increase in coming decades. Because only one-third (4652) of all Malagasy plant species have been formally assessed, we carried out a neural network analysis to predict the putative status and threats for 5887 unassessed species and to evaluate biases in current assessments. The percentage of plant species currently assessed as under threat is probably representative of actual numbers, except in the case of the ferns and lycophytes, where significantly more species are estimated to be threatened. We find that Madagascar is home to a disproportionately high number of Evolutionarily Distinct and Globally Endangered (EDGE) species. This further highlights the urgency for evidence-based and effective in situ and ex situ conservation. Despite these alarming statistics and trends, we find that 10.4% of Madagascar’s land area is protected and that the network of protected areas (PAs) covers at least part of the range of 97.1% of terrestrial and freshwater vertebrates with known distributions (amphibians, freshwater fishes, reptiles, birds, and mammal species combined) and 67.7% of plant species (for threatened species, the percentages are 97.7% for vertebrates and 79.6% for plants). Complementary to this, ex situ collections hold 18% of vertebrate species and 23% of plant species. Nonetheless, there are still many threatened species that do not occur within PAs and are absent from ex situ collections, including one amphibian, three mammals, and seven reptiles, as well as 559 plants and more yet to be assessed. Based on our updated vegetation map, we find that the current PA network provides good coverage of the major habitats, particularly mangroves, spiny forest, humid forest, and tapia, but subhumid forest and grassland-woodland mosaic have very low areas under protection (5.7 and 1.8% respectively). OUTLOOK Madagascar is among the world’s poorest countries, and its biodiversity is a key resource for the sustainable future and well-being of its citizens. Current threats to Madagascar’s biodiversity are deeply rooted in historical and present social contexts, including widespread inequalities. We therefore propose five opportunities for action to further conservation in a just and equitable way. First, investment in conservation and restoration must be based on evidence and effectiveness and be tailored to meet future challenges through inclusive solutions. Second, expanded biodiversity monitoring, including increased dataset production and availability, is key. Third, improving the effectiveness of existing PAs—for example through community engagement, training, and income opportunities—is more important than creating new ones. Fourth, conservation and restoration should not focus solely on the PA network but should also include the surrounding landscapes and communities. And finally, conservation actions must address the root causes of biodiversity loss, including poverty and food insecurity. In the eyes of much of the world, Madagascar’s biodiversity is a unique global asset that needs saving; in the daily lives of many of the Malagasy people, it is a rapidly diminishing source of the most basic needs for subsistence. Protecting Madagascar’s biodiversity while promoting social development for its people is a matter of the utmost urgency Visual representation of five key opportunities for conserving and restoring Madagascar’s rapidly declining biodiversity identified in this Review. The dashed lines point to representative vegetation types where these recommendations could have tangible effects, but the opportunities are applicable across Madagascar. ILLUSTRATION: INESSA VOET 
    more » « less
  2. 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 (Sambaina 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. 
    more » « less
  3. 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 fosa 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. 
    more » « less
  4. null (Ed.)
    Background Madagascar is famous for its extremely rich biodiversity; the island harbors predominantly endemic and threatened communities meriting special attention from biodiversity scientists. Continuing ongoing efforts to inventory the Malagasy ant fauna, we revise the species currently placed in the myrmicine genus Aphaenogaster Mayr. One species described from Madagascar, Aphaenogaster friederichsi Forel, is synonymized with the Palearctic A. subterranea Latreille syn. nov. This species is considered neither native to Madagascar nor established in the region. This revision focuses on the balance of species in the A. swammerdami group which are all endemic to Madagascar. Methods The diversity of the Malagasy Aphaenogaster fauna was assessed via application of multiple lines of evidence involving quantitative morphometric, qualitative morphological, and DNA sequence data. (1) Morphometric investigation was based on hypothesis-free Nest Centroid clustering (NC-clustering) combined with PArtitioning based on Recursive Thresholding (PART) to estimate the number of morphological clusters and determine the most probable boundaries between them. This protocol provides a repeatable and testable approach to find patterns in continuous morphometric data. Species boundaries and the reliability of morphological clusters recognized by these exploratory analyses were tested via confirmatory Linear Discriminant Analysis (LDA). (2) Qualitative, external morphological characteristics (e.g., shape, coloration patterns, setae number) were subjectively evaluated in order to create a priori grouping hypotheses, and confirm and improve species delimitation. (3) Species delimitation analyses based on mitochondrial DNA sequences from cytochrome oxidase I (COI) gene fragments were carried out to test the putative species previously delimited by morphological and morphometric analyses. Results Five species can be inferred based on the integrated evaluation of multiple lines of evidence; of these, three are new to science: Aphaenogaster bressleri sp. n ., A. gonacantha (Emery, 1899), A. makay sp. n. , A. sahafina sp. n. , and A. swammerdami Forel, 1886. In addition, three new synonymies were found for A. swammerdami Forel, 1886 ( A. swammerdami clara Santschi, 1915 syn. n. , A. swammerdami curta Forel, 1891 syn. n. and A. swammerdami spinipes Santschi, 1911 syn. n. ). Descriptions and redefinitions for each taxon and an identification key for their worker castes using qualitative traits and morphometric data are given. Geographic maps depicting species distributions and biological information regarding nesting habits for the species are also provided. 
    more » « less
  5. Rarely have studies assessed Odonata diversity for the entire Nearctic realm by including Canada, the United States, and Mexico. For the first time, we explored Odonata diversity in this region according to a definition of natural community assemblages and generated species distribution models (SDMs). Species occurrence data were assembled by reviewing databases of specimens held by significant Odonata repositories and through an extensive search of literature references. Species were categorized as forest-dependent or non-forest-dependent, as lentic or lotic-dependent, and according to conservation status. Predicted distributions were stacked for all species across their entire ranges, including areas outside of the Nearctic. Species richness and corrected weighted endemism (CWE) were then calculated for each grid cell. We found a pattern of greater species richness in the eastern portion of the Nearctic, which can be explained by the higher aquatic habitat diversity at micro and macroscales east of the Rocky Mountains, promoting niche partitioning and specialization. In the Nearctic region, the southeastern US has the highest number of endemic species of dragonflies and damselflies; this degree of endemism is likely due to glacial refuges providing a foundation for the evolution of a rich and unique biota. 
    more » « less