sign in sign up
<<
Home , Cassowary, Ground sloth, Megafauna, Megalocnus, Meiolania, Stegomastodon

PERSPECTIVES

ECOLOGY An expanded concept elucidates how of the largest in The Forgotten Megafauna any ecosystem have similar effects.

Dennis M. Hansen1 and Mauro Galetti1,2

arge terrestrial vertebrates— Giant ground called megafauna—play key Lroles in ecosystem dynam- Giant ics by feeding on plants and by maintaining habitat hetero- geneity (1). A global wave of megafauna extinctions oc- curred 50,000 to 10,000 ago, when many large conti- nental were lost (2–5). Classical definitions of mega- ICA are based on such conti- MER TH A nental mammals and are vari- SOU ously given as larger than 44 kg (6) or above 1000 kg (7). Here, we argue that the mega- fauna concept should be extended beyond an SCAR MADAGA on May 10, 2009 absolute size to be context-dependent. In any given ecosystem, the largest vertebrates URITIUS have ecosystem impacts that are similar on a MA relative scale to those of the largest vertebrates Scaling the megafauna. The magnitude of loss of frugivorous megafauna is currently most dramatic on in another ecosystem: One ecosystem’s meso- islands, as illustrated by the smaller drawn sizes of the giant and the gomphothere from South fauna is another ecosystem’s megafauna. America, compared with the in and the of Mauritius. However, many An ecosystem function that clearly illus- continental regions are poised to catch up. trates this argument is animal-mediated seed

dispersal. Here, the link between animal body To illustrate our point, we have examined in relative terms, led to a greater megafaunal www.sciencemag.org mass and ecosystem function is straight- tropical and subtropical from three downsizing than the of even the forward: The larger the fruit-eating animal kinds of ecosystems: continental, continen- largest in (), the larger the fruits it can tal islands, and oceanic islands. For each (see the figure). consume. Thus, extinction-mediated “ecolog- fauna, we compared the body masses of the Moreover, in the relatively species-poor ical shrinkage”—the loss of species inter- largest extant frugivorous — and simple island ecosystems, cascading actions—in community-level seed dispersal , bird, or —to the largest that effects of megafaunal loss may manifest roughly scales with frugivore body mass. has gone extinct since the late . themselves faster and with more devastating Downloaded from Scientists have argued that in continental On continents, the body masses of extant results than in more complex continental Central and South America, the extinction of are an of magnitude lower ecosystems. For example, lost megafaunal the classic mammalian megafauna—such as than those of extinct frugivores; in contrast, seed-dispersal interactions on islands will not giant ground and gomphotheres— in some continental and oceanic islands, be compensated for by surviving frugivores, caused disruption of seed dispersal for large body masses of extant animals are two or because they are too small. In continental fruits (4, 5). However, on islands, the extinc- even three orders of magnitude lower than ecosystems, there is a higher functional tion of large and in the past two those of their extinct predecessors (see the redundancy, with medium- and even small- or three millennia has led to similar disrup- supporting online material). sized species capable of filling at least part of tions (8, 9). In both locations the demo- For instance, the largest frugivores in the megafaunal niche (12). graphic and genetic consequences of large- South America were gomphoteres (7580 Anthropogenic impacts are set to cause vertebrate extinctions for plants are likely kg), whereas the largest living frugivores are further extinctions among large vertebrates, similar—for example, disruption of long- the (300 kg). On the continental island with dramatic consequences for ecosystem distance gene flow or changes in species of Madagascar, the role of largest frugivore dynamics (11). If all currently threatened ver- composition (10, 11). Yet, by the classic def- has passed from the elephant bird (450 kg) to tebrate frugivores were to go extinct, the rela- initions, large insular vertebrates would not the radiated tortoise (10 kg). On Mauritius, tive ecological shrinkage in many continental be considered megafauna. giant weighing up to 100 kg were ecosystems would equal that of islands. For the largest native frugivores; today, the title instance, if all threatened frugivores in South goes to a fruitbat weighing only 0.54 kg. America were to go extinct, the largest 1Department of Biology, 371 Serra Mall, Stanford Uni- Thus, the loss of the island giants—tortoises, remaining frugivore would be the howler versity, Stanford, CA 94305, USA. 2UNESP-São Paulo State University, Department of Ecology, Rio Claro, São Paulo, , and flightless birds such as the monkey, weighing 9 kg—a factor of 700 less

Brazil 13506-900. E-mail: [email protected] that were once found on many islands—has, than the giant ground sloth. Even some islands BY CARL BUELL ARTWORK MEGAFAUNA CREDIT:

42 3 APRIL 2009 VOL 324 SCIENCE www.sciencemag.org Published by AAAS PERSPECTIVES

stand to suffer further losses; in Mauritius, the References and Notes 10. R. G. Colevatti et al., Mol. Ecol. 12, 105 (2003). largest nonthreatened native frugivore is the 1. R. Dirzo, A. Miranda, in Plant-Animal Interactions: 11. S. J. Wright et al., Biotropica 39, 289 (2007). Evolutionary Ecology in Tropical and Temperate Regions, 12. W. Hallwachs, in Frugivores and Seed Dispersal, A. Estrada, gray white-, a bird weighing a mere 0.009 P. W. Price, T. M. Lewinshon, G. W. Fernandes, W. W. T. H. Fleming, Eds. (Kluwer, Boston, 1986), pp. 285–304. kg (see the supporting online material). Benson, Eds. (Wiley, New York, 1991), pp. 273–287. 13. A. R. Wallace, The Geographical Distribution of Animals An extended megafauna concept has the 2. P. L. Koch, A. D. Barnosky, Ann. Rev. Ecol. Evol. Syst. 37, (Harper, New York, 1876). 215 (2006). 14. Supported by the Velux Foundation (D.M.H.), Fundação potential to promote synergy between other- 3. A. D. Barnosky et al., Science 306, 70 (2004). de Amparo à Pesquisa do Estado de São Paulo (M.G.), wise disparate research and conservation foci, 4. D. H. Janzen, P. S. Martin, Science 215, 19 (1982). and Conselho Nacional de Desenvolvimento Científico e and to facilitate broader syntheses of eco- 5. P. R. Guimaraes Jr., et al., PLoS ONE 3, e1745 (2008). Tecnológico (M.G.). system-level effects of extinctions of the 6. P. S. Martin, R. G. Klein, Quaternary Extinctions: A Prehistoric Revolution (Univ. of Arizona Press, Tucson, Supporting Online Material largest vertebrates and the resulting ecological 1995). www.sciencemag.org/cgi/content/full/324/5923/42/DC1 shrinkage. It is high time to more fully under- 7. R. N. Owen-Smith, Megaherbivores: The Influence of Very Fig. S1 Table S1 stand and ameliorate the recent and ongoing Large Body Size on Ecology (Cambridge Univ. Press, New York, 1988). References losses of all “the hugest, and fiercest, and 8. H. J. Meehan et al., J. Biogeogr. 29, 695 (2002). strangest forms” (13). 9. D. M. Hansen et al., PLoS ONE 3, e2111 (2008). 10.1126/science.1172393

COMPUTER SCIENCE Computers with intelligence can design and run experiments, but learning from the results Automating Science to generate subsequent experiments requires even more intelligence. David Waltz1 and Bruce G. Buchanan2

he idea of automating as- motion; today, computer-controlled on May 10, 2009 pects of scientific activity data collection is commonplace Tdates back to the roots of and necessary for both experi- computer science, if not to Francis mental and observational science. . Some of the earliest pro- Automating many activities be- grams automated the processes of yond data collection offers even creating ballistic tables, cracking more benefits. cyphers, collecting laboratory In the near term, a useful meta- data, etc., by carrying out a set of phor is to consider computers as

instructions from start to finish. intelligent assistants. Some assis- www.sciencemag.org Starting with DENDRAL in the tants gather data and attend to 1960s (1), artificial intelligence such tasks as noise filtering, data programs such as Prospector (2), smoothing, outlier rejection, and Bacon (3), and Fahrenheit (4) data storage. Other assistants are automated some of the planning, specialists at statistical analysis, Semiautomated. Scientists at Stanford’s Instrumentation Research Laboratory analysis, and discovery portions still others at bench work. This (circa 1970) linked a gas chromatograph and high-resolution mass spectrome- of the scientific enterprise. How- metaphor has driven many re- ter to computers to automate studies of biological fluids, meteorites, and other Downloaded from ever, most of these programs were materials. Stanford’s DENDRAL Project experimented with automated interpreta- search projects over the past sev- still designed to run a calculation tion of the data and experiment planning to specify nuclear magnetic resonance eral decades and has led to many to completion, produce an answer, or infrared data that would resolve ambiguities in the mass spectral data. of the most successful applica- and then stop. They did not fully tions of computers. “close the loop” in the sense of examining the systems, derived from visual observation of An early articulation of this metaphor results of their actions, deciding what to try such systems. As these reports show, it is pos- is Joshua Lederberg’s effort at Stanford next, potentially cycling forever. sible for one computer program to step University School of Medicine to develop an Two reports on pages 85 and 81 of this through the activities needed to conduct a con- automated biomedical laboratory to examine issue push the boundaries of automatic scien- tinuously looping procedure that starts with a the soil of Mars for traces of life, as part of tific experimentation and discovery. King question, carries out experiments to answer the 1975 Viking mission deployed by the et al. (5) describe a robotic system for running the question, evaluates the results, and refor- U.S. National Aeronautics and Space Ad- biological experiments, evaluating their mulates new questions. ministration. The robot assistant Lederberg results, and deciding what experiments to try The main goals of automation in science designed, with engineer Elliott Levinthal, next. Schmidt and Lipson (6) describe their have been to increase productivity by increas- consisted of a conveyor belt that scooped up work on discovering compact equations that ing efficiency (e.g., with rapid throughput), to samples of Martian soil and deposited them characterize complex nonlinear dynamical improve quality (e.g., by reducing error), and within a computer-controlled mass spectrom- to cope with scale, allowing scientific treat- eter. Each soil sample was bombarded with 1Center for Computational Learning Systems, Columbia ment of topics that were previously impossi- electrons, producing a fragmentation pattern 2 University, New York, NY 10115, USA. Computer Science ble to address. Tycho Brahe spent a lifetime that sorted the charged particles (ions) accord- Department, University of Pittsburgh, Pittsburgh, PA 15260, USA. E-mail: [email protected]; waltz@ recording observations that allowed Johannes ing to their mass. This pattern was transmitted

CREDIT: ROBERT K. LINDSAY ROBERT CREDIT: ccls.columbia.edu Kepler to formulate Kepler’s laws of planetary to Earth, where scientists could analyze it for

www.sciencemag.org SCIENCE VOL 324 3 APRIL 2009 43 Published by AAAS

www.sciencemag.org/cgi/content/full/324/5923/42/DC1

Supporting Online Material for

The Forgotten Megafauna Dennis M. Hansen* and Mauro Galetti

*To whom correspondence should be addressed. E-mail: [email protected]

Published 3 April 2009, Science 324, 42 (2009) DOI: 10.1126/science.1172393

This PDF file includes:

Fig. S1 Table S1 References Supplementary online material

Figure S1. Current and projected ecological shrinkage in seed dispersal interactions on continents, continental islands, and oceanic islands, illustrated by declines in frugivore body mass. Arrows span declines from the largest recently extinct frugivore to the largest extant frugivore (red arrows), as well as maximum projected future decline if all currently threatened frugivores went extinct (IUCN status of VU, EN or CR; yellow arrows). Regions: [AF], South America [SA], [AU], Southeast [AS]; [CU], Madagascar [MA], [NC]; Tonga [TO], Mauritius [MR], [FI].

Table S1. Largest extinct and extant frugivores in selected continents, continental islands and oceanic islands. Body mass in kg. IUCN status, from least concern to critically endangered: LC = Least Concern, NT = Near Threatened, VU = Vulnerable, EN = Endangered, CR = Critically Endangered.

Region Largest extinct Largest extant Largest extant non-threatened

Body Body IUCN Body IUCN mass mass status (1) mass status (1) Africa Pleistocene elephant 6500 3940 NT African bush elephant 3940 NT iolensis (2) Loxodonta africana (2) Loxodonta africana (2) South Gomphothere 7580 Baird’s 300 EN Red howler monkey 9 LC America Stegomastodon superbus (2) Tapirus bairdii (2) Alouatta seniculus (2) Australia 2700 Southern 85 VU 60 LC Diprotodon australis (3) Casuarius casuarius (8) novaehollandiae (8) SE Asia 10000 3000 VU Tibetan macaque 45 NT Stegodon sp. (2) Elephas maximus (2) Macaca thibetana (2) Cuba Cuban ground sloth 200 Cuban rock iguana 4.1 VU Prehensile-tailed hutia 1.9 NT rodens (4) Cyclura nubila (9) Mysateles prehensilis (11) Madagascar Elephant bird 450 Radiated tortoise 10 VU Brown lemur 5.8 LC maximus (5) Astrochelys radiata (10) Eulemur fulvus (2) New Giant horned tortoise 500 Pacific flying fox 0.6 LC Pacific flying fox 0.6 LC Caledonia sp. (6) Pteropus tonganus (2) Pteropus tonganus (2) Tonga Giant iguana 3.3 Pacific flying fox 0.6 LC Pacific flying fox 0.6 LC Brachylophus gibbonsi (7) Pteropus tonganus (2) Pteropus tonganus (2) Mauritius Mauritian saddle-backed giant 100 Greater Mascarene 0.54 EN Mauritius grey white-eye 0.009 LC tortoise flying fox Zosterops mauritianus (8) Cylindraspis triserrata (6) Pteropus niger (2) Fiji Giant horned tortoise 300 Pacific flying fox 0.6 LC Pacific flying fox 0.6 LC Meiolania sp. (6) Pteropus tonganus (2) Pteropus tonganus (2)

Acknowledgements: Thanks to the following for providing information relating to Table S1: Richard Corlett, Anthony Cheke, Trevor Worthy, Christopher Johnson, Pedro Jordano.

References: 1. IUCN Redlist online: http://www.iucnredlist.org (accessed February 01, 2009). 2. Smith, F. A. et al. Ecological Archives E084-094, (2003) and personal communications (see above). 3. C. Johnson, Australia’s Mammal Extinctions: A 50,000 history. Cambridge University Press (2006) 4. D. W. Steadman et al., Proc. Natl. Acad. Sci. U.S.A. 102, 11763 (2005). 5. D. Amadon, 49, 159 (1947). 6. No published body mass estimates exist for these species; we estimated the body masses based on known carapace length/body mass relations for the giant Aldabran tortoise, Aldabrachelys gigantea, from M. J. Coe, D. Bourn, I. R. Swingland, Phil. Trans. R. Soc. Lond. B. 286, 163 (1979). 7. No published body mass estimate exists for this species; we estimated the body mass using the SVL reported in G. K. Pregill, D. W. Steadman, J. Herpetol. 38, 15 (2004), and the relationship between SVL and body mass for Iguana iguana reported in R. W. Van Devender, in Iguanas of the World, G. M. Burghardt, A. S. Rand, Eds. (Noyes Publications, Park Ridge, 1982), pp. 162–183. 8. J. B. Dunning, Ed., CRC Handbook of Avian Body Mass, 2nd edition (CRC Press, Cleveland, 2007). 9. K. Beovides–Casas, C. A. Mancina, Animal Conservation and 29, 1 (2006). 10. M. Pedrono, The Tortoises and of Madagascar (Natural History Publications, Borneo, 2008). 11. M. Hutchins, D. Kleiman, V. Geist, M. McDade, Eds. Grzimek's Animal Life Encyclopedia, Second Edition (Thomson-Gale, Farmington Hills, 2003).