produce defenses against leaf-eating animals, the extensive use of spines by plants in the spiny forests suggest that they evolved to cope with leaf-eating lemurs, large and small. Seed dispersal Giant subfossil lemurs are thought to have also played a significant role in seed dispersal, possibly targeting species that did not attract the seed dispersal services of the extinct elephant birds. Biogeochemistry studies have shown that they may have been the primary seed dispersers for the endemic and native C3 trees in the spiny forests. Terrestrial species may have dispersed seeds for small bushes as well as tall trees. Seed dispersal can involve passing seeds through the gut (endozoochory) or attaching the seeds to the animal's body (epizoochory), and both processes probably occurred with subfossil lemurs. Seeds from Uncarina species embed themselves in lemur fur, and likely did the same with subfossil lemurs. Seed dispersal biology is known for very few species in the spiny forest, including genera of plants suspected of depending on giant lemurs, such as Adansonia, Cedrelopsis, Commiphora, Delonix, Diospyros, Grewia, Pachypodium, Salvadora, Strychnos, and Tamarindus. For example, Delonix has edible pods that are rich in protein, and Adansonia fruits have a nutritious pulp and large seeds that may have been dispersed by Archaeolemur majori or Pachylemur insignis. Seed size may be a limiting factor for some plant species, since their seeds are too large for living (extant) lemurs. The common brown lemur (Eulemur fulvus) can swallow seeds in diameter, while the black-and-white ruffed lemur (Varecia variegata) is capable of swallowing seeds up to in diameter. A large lemur, such as Pachylemur, which was more than twice the size of today's ruffed lemurs, could probably swallow even larger seeds. Seed dispersal limitations tied to megafaunal extinction are exhibited by Commiphora guillaminii. At present, this tree species has a short dispersal distance, but its genetics indicate higher levels of regional gene flow in the past, based on comparisons with a closely related species in Africa whose seeds are still dispersed by large animals. Discovery and research The writings of French colonial governor Étienne de Flacourt in the mid-17th century introduced the existence of giant Malagasy mammals to Western science with recorded eye-witness accounts from the local people of dangerous animals, hornless "water cows", and a large lemur-like creature referred to locally as the or . Today, the latter is thought to have been a species of Palaeopropithecus or