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17

Biodiversity Inventories in Costa Rica and Their Application to Conservation

Paul Hanson

many basic scientific disciplines as well as many applied sciences such as biotechnology, agriculture, fisheries, and conservation. Most inventorying and monitoring have involved organisms that are relatively well known taxonomically for example, vertebrates and vascular plants. Yet the poorly known groups of organisms, such as invertebrates and fungi, constitute the majority of the species. Conservation decisions based o n data for a limited range of organisms having relatively few species can be misleading (PrendergastetaL 1993). Moreover, poorly known groups of organisms tend to be smaller i n size and to shorter generation times, which means they respond more rapidly to environmenchanges (Brown 1991). Thus, taxonomically ilt organisms are often good indicators o f inmental change and can help us to realthat a problem exists before the plants and

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vertebrates are affected. However, basic taxonomic research is needed before these poorly known groups o f organisms can be used i n monitoring and other conservation activities. The problem is that there are too few taxonomists working on the most species-rich groups of organisms (Gaston and May 1992; Hawksworth and Ritchie 1993; Hammond 1995). Worse yet, taxonomists who have a lifetime's experience are retiring, and the knowledge they possess (much of it unpublished) is not being recovered through firsthand transfer to young recruits (Cotterill 1995). I n recent years developed countries have begun reducing government expenditures, and thus funding o f national museums has diminished markedly i n real terms. Increasingly such institutions are expected to become more self-sufficient. This means that traditional taxonomic research now has to compete with more attractive proposals i n evolutionary biology, the latter often addressing species/population level

toring provide essential information used by

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questions i n taxa that are already well known. Not surprisingly, job opportunities for taxonomists have fallen drastically, a n d graduate students choose research. Because the majority o f species do not yet have scientific names and the process o f describing new species is generally slow, an attractive alternative is a rapid biodiversity assessment o f taxonomicalfy well-known groups (Oliver and Beattie 1996). W h e n such rapid assessments do include poorly k n o w n taxa, they must rely on the separation o f morphospecies. A l t h o u g h t i m e is saved by not having to obtain names for the species, i f the sorting o f morphospecies is done by a nonspecialist, it is prone to serious errors (for example, many insects can be separated only by dissecting male genitalia). A l t h o u g h rapid biodiversity assessments are relatively inexpensive, they are no substitute for the more costly systematic inventories. Moreover, species names are required for determining which sites harbor phylogenetically isolated species and for serving a wider community o f users interested i n environmental education, bioprospecting, and the like. I n recent years a considerable amount o f taxonomic inventory w o r k has been carried out i n Costa Rica i n general and i n dry forests i n particular. Because the w o r k done i n the dry forests has occurred i n a national context, and because the problems facing inventories do n o t differ fundamentally between terrestrial ecosystems, this chapter treats the broader national process. I t thus differs f r o m other chapters i n this book by not focusing specifically o n dry forests. This chapter briefly evaluates inventory work i n Costa Rica by considering what some o f the problems have been and h o w future inventories m i g h t be improved. Three aspects o f the inventory process are discussed here: (1) the field operation o f collecting specimens, (2) the resource base for biodiversity assessment, especially the biological collections and h u m a n resources available i n the country, and (3) the information that is derived f r o m the inventory, especially as i t applies to conservation. financially more p r o m i s i n g fields o f

COSTA RICAN INVENTORIES: THE FIELD OPERATION

As i n most countries, the early inventories i n Costa Rica were quite sporadic (see Gomez and Savage 1983 for a b r i e f history). Subsequent inventories, primarily by staff o f the National M u s e u m and the University o f Costa Rica, have been p r i m a r i l y museum-based and driven by the m o t i v a t i o n o f a few individuals. W i t h the creation o f the National Biodiversity Institute (INBio) i n 1989 there was for the first t i m e a coordinated and institutionalized effort to survey the species present i n the country. INBio is a private, nonprofit organization whose mission is to promote a greater consciousness o f the value o f biodiversity i n order to conserve i t and hence improve the quality o f h u m a n life. INBio is not intended to be an institution dedicated to taxonomy as an end i n itself, and the t e r m museum has been carefully shunned. From the beginning i t was felt that neither the t i m e nor the economic resources were available to train Costa Rican taxonomists i n foreign universities (Gamez 1999). Instead, foreign taxonomists have been encouraged to visit INBio i n order to help identify specimens collected and curated by INBio staff and to train the collectors and curators. The primary emphasis has been o n vascular plants and i n sects, although fungi, mollusks, and nematodes have been added. At the front end o f I N Bio's inventory are the "parataxonomists"rural people w h o grew u p i n areas surrounding protected w i l d lands and are trained to collect a n d m o u n t specimens. After being m o u n t e d at the field stations, specimens are brought to INBio, where they are labeled, bar-coded, and classified by technicians and curators (Janzen et al. 1993). H o w well have the parataxonomists performed w i t h respect to the biodiversity inventory? O n the positive side they have amassed the largest collection o f i n sects i n Central America i n a very short period o f time. Because insects were collected primarily by hand-netting and at lights, the inventory has been most successful i n those taxa that are

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readily collected by these methods. Many species are represented by l o n g series f r o m various localities, thus providing i n f o r m a t i o n o n geographical distribution and intraspecific variation w i t h i n the country. I n general the parataxonomists who have been most motivated have been the most successful. For example, the team o f parataxonomists i n the Guanacaste Conservation Area have produced more specimen-based information on host ranges o f parasitoids than was previously k n o w n f r o m all o f tropical America (I. Gauld pers. comm.). A l t h o u g h the use o f parataxonomists has proved to be a very effective means to jump-start a collection, i n the l o n g r u n i t becomes ever more challenging to employ t h e m efficiently. Without sufficient feedback f r o m curators or taxonomists, overzealous collecting eventually yields vast quantities o f the c o m m o n species, a problem that INBio is now beginning to address. Meanwhile, the m i n u t e insects are generally missed by hand-collecting, and the few that are encountered are often poorly mounted, since the m o u n t i n g is done i n the field. To be fair, no h u m a n being could possibly do a first-rate job o f collecting and m o u n t i n g all types o f insects; that is one o f the reasons that entomologists specialize i n particular taxa. Recently INBio has trained parataxonomists w h o specialize i n just one order o f insects, b u t there are still certain taza that are most efficiently collected by the taxonomists themselves. I n order to improve the collection o f smallersized taxa, I N B i o has begun placing more emphasis on trapping methods. However, there are still not enough laboratory technicians to handle these mass samples, perhaps because o f the institute's legacy o f being predominantly fieldoriented. For example, a parataxonomist needs only about an hour per m o n t h to m a i n t a i n a Malaise trap, but a laboratory technician w i l l require a m i n i m u m o f 20 to 30 hours to separate the focal taxa f r o m this same sample. Thus, when it comes to processing b u l k trap samples, INBio continues to invest a disproportionate amount o f h u m a n resources i n a task that requires only

a small fraction o f the total time. Parataxonomists can, o f course, be refocused i n their collecting methodsfor example, to obtain more biological i n f o r m a t i o n (which is being done), b u t this still does not address the shortage o f h u m a n resources for separating b u l k samples. To decide o n a set o f methods for a particular taxon, we need to establish criteria for measuri n g the performance o f different collecting strategies. The n u m b e r o f specimens collected and processed, although widely cited i n annual reports, is a poor indicator o f performance because it reveals n o t h i n g about h o w many species were collected or the quality o f the processed specimens. I n cases i n w h i c h a specialist is actively w o r k i n g w i t h the collection, a m u c h better i n dicator o f performance is the number o f new species collected and h o w m u c h i t cost to obtain them.

THE RESOURCE BASE FOR BIODIVERSITY ASSESSMENT

Most inventories result i n a collection o f specimens, w h i c h usually consist o f both identified (voucher) specimens and unidentified specimens o f taxa for w h i c h no specialists are currently available. Voucher collections are essential for the verification o f field data and to provide a permanent historical record. Biological reference collections i n general, w i t h their associated l i braries and staff, are a vital source o f baseline data for future inventorying and monitoring (Cotterill 1995). Biological collections are n o t mere warehouses for specimens. A t the very m i n i m u m they require constant vigilance to prevent the ravages o f fungal and arthropod pests, w h i c h can rapidly destroy specimens. However, a collection whose staff does n o t h i n g more t h a n ensure its physical well-being is effectively a dead collection. A n active and useful collection requires a constant i n p u t o f labor for sending and receiving loans o f specimens, organizing (curating) specimens so that individual taxa are readily accessible t o taxonomists, and similar

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tasks. Because the c o n t i n u i n g cost o f maintaini n g such collections is frequently viewed w i t h suspicion by auditors, i t is essential to publicize the importance o f biological collections as research facilities, every b i t as valuable as their more dazzling counterparts i n nuclear physics facilities. I n Costa Rica the oldest biological collection is that o f the National M u s e u m . This collection was mismanaged i n the late 1940s and d i d not recover u n t i l the 1970s. Despite some unexplained disappearances o f type specimens i n the past, the National M u s e u m currently houses one o f the best vascular plant collections i n Central America. Its insect collection is p r i m a r i l y restricted to the butterflies. At the University o f Costa Rica the H e r b a r i u m was initiated i n the 1940s and the Zoology M u s e u m and Insect M u seum i n the 1960s. U n l i k e other large herbaria i n Central America, that o f the university i n cludes a considerable n u m b e r o f nonvascular plants and fungi. The Zoology M u s e u m has substantial collections o f marine invertebrates, arachnids, aquatic insects, and especially vertebrates. The Insect M u s e u m has focused on bees, certain families o f beetles, and parasitic wasps. The university's biological collections w o u l d be impressive i f their contents were better documented. The preservation and growth o f these biological collections have been assured primarily by motivated individuals w o r k i n g f r o m shoestring budgets, although recently the university has taken a greater interest i n t h e m . The collections at INBio have been an institutional priority since the inception o f the i n stitute. The combination o f a large, well-curated collection and enthusiastic staff has acted as a magnet for many foreign taxonomists. Perhaps for the first t i m e i n the history o f Costa Rica, at least some insect taxonomists are c o m i n g not primarily to collect but rather to work w i t h the existing collection i n INBio. As w i t h the other collections i n the country, the staff o f INBio has been quite enlightened about loaning specimens to the international taxonomic community. Besides collections, two other vital resources for carrying out biodiversity assessment are ac-

cess to scientific literature and people w i t h experience i n identifying particular taxa. Libraries i n Costa Rica cannot afford subscriptions to most o f the necessary scientific journals, w h i c h is not surprising because even libraries i n developed countries are reducing their subscriptions to these journals. Moreover, despite m u c h recent pontificating, the nuts and bolts o f taxonomic literature are still unavailable on the I n ternet and show no sign o f becoming available anytime soon. Thus, b u d d i n g taxonomists i n Costa Rica, like those i n other developing countries, have to obtain the necessary literature t h r o u g h their o w n efforts and those o f foreign collaborators. W i t h respect to h u m a n resources, most o f the curators at INBio have received firsthand t r a i n i n g under the guidance o f foreign taxonomists, and some are u t i l i z i n g this training to publish descriptions o f new species. O n the other hand, only a few o f the curators have had graduate-level courses i n systematics, genetics, or evolution, w h i c h w o u l d greatly facilitate separating the species, understanding species distributions, and providing advice on conservationrelated issues. This deficiency is especially ironic because courses i n these subjects are taught at the University o f Costa Rica.

DATA AND INFORMATION MANAGEMENT AND COMMUNICATION

A general problem among many institutions worldwide is launching a database without clearly defining its intended use, thus forgetting that data management is a tool rather t h a n an end i n itself. A l t h o u g h developing one's o w n software for data management may be superficially attractive, i t is usually difficult for a local instit u t i o n do this i n a cost-effective way (Olivieri et al. 1995). INBio has invested heavily i n developi n g its own software, and i t w o u l d be interesting to analyze whether this effort has been costeffective. Nonetheless, there has been an institutional c o m m i t m e n t to data management (see Zeledn 2 0 0 0 : 100), w h i c h allows u p g r a d i n g as new technology arrives. I n contrast, data

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management i n other collections i n Costa Rica has generally relied on individual c o m m i t m e n t , w h i c h often results i n disconnected and potentially incompatible databases. Very little o f the inventory effort i n Costa Rica has entailed quantitative sampling, w h i c h can provide valuable data on the relative abundance o f different species and what proportion o f the biota remains to be collected. The notable except i o n is the Arthropods o f La Selva project (Longino and Colwell 1997). Yet i t is difficult to fault the rest o f the inventory w o r k for being nonquantitative because most tropical organisms do not even have names and merely addressing this essential first step is an enormous task. Different taxa require different sampling methods (New 1998), and for a particular taxon there is often uncertainty as to how efficient one method is i n comparison w i t h others ( H a m m o n d 1994; Samways 1994). Not surprisingly, we must often rely on the taxonomists' i n t u i t i o n . A l t h o u g h further biological data are usually beyond the scope o f most inventories, various field biologists have been carrying out research i n Costa Rica for many years, and this informat i o n is being compiled by the Organization for Tropical Studies (BINABITROP, or National Bibliography o n Tropical Biology, currently comprising 12,000 references). I n the future more emphasis needs to be placed on gathering ecological information d u r i n g the inventory, as is currently being done w i t h macrolepidopteran caterpillars i n the Guanacaste Conservation Area (http://Janzen.sas.uperm.edu:59i/rearingdb.htm) and the ants o f Costa Rica project (www.evergreen.edu/ants). The information obtained f r o m inventories has to be made available for a variety o f uses, notably scientific advancement, environmental education, and conservation management. Toward these ends the National M u s e u m , the public universities, and INBio have all been actively producing a variety o f publications (both scientific and popular), although a perennial problem i n Latin America i n general is the distribution o f these publications. Both INBio and the Arthropods o f La Selva project are actively producing

species Web pages, w h i c h has the potential to disseminate natural history information m u c h more widely than traditional publications. Finally, how is the inventory information bei n g used i n m a k i n g decisions about conserving biodiversity? I n general the staff o f the conservation areas has been concerned w i t h how to protect rather than how to manage. The most frequent requests coming f r o m both the conservation areas and the private reserves are for lists o f species and field guides, w h i c h are useful for environmental education and ecotourism, b u t less so for management and decision making. Information on rare and endangered plant species was used i n p l a n n i n g the Osa and "Paso de la Danta" corridors and i n planning a nature trail i n Corcovado National Park, b u t such examples are rare (R. Garcia pers. comm.). Conservation decisions often rely more o n ecological information (e.g., m a m m a l migrations) than o n r a w taxonomic i n f o r m a t i o n , yet the latter provides the foundation for the former. Therefore, the sequence o f information transfer that needs to occur is systematics > ecology conservation, although systematics can occasionally provide information directly to conservation (e.g., phylogenetically isolated taxa).

CONCLUSIONS
We obviously cannot afford to delay conservat i o n efforts u n t i l all taxa have been fully inventoried. The urgency o f the current situation requires us to select areas that need protection o n the basis o f existing information and rapid biodiversity assessments o f taxonomically wellk n o w n organisms. At the same time, however, i t is prudent to continue the inventories o f lesserk n o w n taxa such as microorganisms, fungi, and invertebrates. These taxa constitute the majority o f the world's species, and many o f t h e m are critical to ecosystem functioning. Moreover, certain species-rich areas would probably be overlooked i n conservation planning i f we totally ignored the species-rich taxa. Inventories i n Costa Rica have been very patchy i n their coverage o f the country. Inventory

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w o r k by the universities has been done primarily outside the national parks, whereas INBio's inventory has been restricted to protected areas. Some areas (e.g., Guanacaste and the Osa Peninsula) have received more emphasis t h a n others (e.g., Braulio Carillo National Park). Very little inventory w o r k has been carried out i n the private reserves, w h i c h are potentially quite important i n conservation (Herzog and Vaughan 1998). A national inventory that includes both protected and nonprotected areas w o u l d provide informat i o n o n the identity and geographical distribut i o n o f the species occurring w i t h i n the country as a whole. This is essential for determining what proportion o f the species are not i n protected areas and consequently w h i c h nonprotected areas are i n greatest need o f protection. Inventories are also needed after lands have been acquired for conservation. These lands need to be managed and monitored, but typically only vascular plants and vertebrates have been i n ventoried. I n fact, Costa Rican law requires only that trees and vertebrates be included i n environmental impact studies. Yet inventory w o r k i n the country has advanced to the point where other taxa could and should be included. The laws are especially outdated i n the case o f freshwater ecosystems, where environmental impact studies are required only to measure physical and chemical parameters ( M . Springer pers. comm.), w h i c h recover f r o m d u m p i n g events (e.g., pulp f r o m coffee berries) more quickly than do populations o f aquatic invertebrates. National biodiversity inventories have several functions, three o f w h i c h have been highlighted here: (1) b u i l d i n g local taxonomic expertise and biological collections, w h i c h are the basis for future scientific studies and conservation efforts, (2) providing basic i n f o r m a t i o n that can be used i n environmental education, and (3) supplying the i n f o r m a t i o n needed i n developing management plans for conservation. I n general, inventories i n Costa Rica have succeeded m u c h better i n the first two than i n the t h i r d . There are many good references treating the types o f information needed i n conservation (e.g., Noss and Cooperrider 1994), and so the problem is not i n de-

ciding what type o f i n f o r m a t i o n is needed but rather i n providing i t i n an accessible format, disseminating it, and u t i l i z i n g it. A great deal o f the information already exists, some o f it readily available but m u c h o f it requiring synthesis and dissemination. Perhaps the most serious probl e m is the final stepapplying the information. Too often inventory i n f o r m a t i o n is viewed as an end i n itself, and even w h e n the i n f o r m a t i o n reaches the decision makers, i t is often exiled to the file cabinet. Based o n the biodiversity inventory w o r k that has been done i n Costa Rica, we can draw on certain facts to make suggestions for future biological inventories: Fact: A l t h o u g h collecting is obviously an essential first step i n the taxonomic process and parataxonmists have been useful i n this process, the rate-limiting step i n inventory w o r k is the description o f new species. This can be done only by taxonomic experts, but they should not have to undertake all the tasks themselves as they have traditionally done. Suggestion: Develop more and varied h u m a n resources to support monographic taxonomy. For example, the concept o f a parataxonomist could be broadened to i n clude people who assist taxonomists i n describing new species. After the taxonomist has separated the species and written a prel i m i n a r y key, these individuals could do some o f the more routine tasks (e.g., recordi n g locality data f r o m labels and measuring specimens). Quality control should be m a i n tained by the taxonomist, and the usual peer review processes should be continued. Fact: There are not enough international taxonomic resources to support an INBio clone i n the majority o f tropical countries. INBio has benefited greatly f r o m being one o f the first such institutions to appear. For example, British taxpayers have invested approximately a m i l l i o n dollars i n the Costa Rican inventory t h r o u g h salaries paid to staff o f the Natural History M u s e u m who have participated i n this inventory (I. Gauld pers. comm.).

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Global taxonomic resources are simply not large enough to duplicate such support i n most other tropical countries. Suggestion: Promote greater regional interinstitutional collaboration and increase employment opportunities for taxonomists. I t is w o r t h rem e m b e r i n g that no country w i l l ever be self-sufficient w i t h respect to taxonomic expertise. Even the United States and Canada had to i m p o r t taxonomic expertise i n order to produce keys to the genera o f Chalcidoidea o f North America (Gibson et al. 1997). Fact: Globally, taxonomy is primarily funded through science research budgets and is subject to changes i n science priority. The user c o m m u n i t y wants taxonomists to produce the tools by w h i c h organisms can be identified, but the research required to produce such tools is not currently viewed by the research c o m m u n i t y as a priority undertaking. Suggestion: Potential user communities should identify specific taxonomic needs (e.g., an identification manual for the Neotropical genera o f Chalcidoidea), and funding agencies should invite tenders to produce these products rather than invest i n more nebulous "capacity b u i l d i n g . " Funding opportunities for specific products w o u l d stimulate organizations to collaborate i n formulating competitive bids, w h i c h i n t u r n w o u l d create jobs for young taxonomists, particularly i n developing countries w i t h comparatively low overheads. Costa Rica has received international accolades for its efforts i n conservation and inventory. H o w well it lives u p to this reputation i n the future depends on a continued c o m m i t m e n t to these interrelated activities. The often repeated phrase "learning occurs t h r o u g h doing" is appropriate, and for other tropical countries that have barely begun doing it, the Costa Rican experience can provide several take-home lessons.

several important ideas for this chapter. I also thank the following persons for providing useful comments on previous versions o f the chapter: Zaidett Barrantes, Wills Flowers, Gordon Frankie, Randall Garcia, Jorge Gomez Laurito, Luis Diego Gomez, Barry H a m m e l , Jack Longino, Julian Monge, and Rodrigo Zeledon.

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ACKNOWLEDGMENTS
I especially t h a n k I a n Gauld for n u m e r o u s thought-provoking discussions that provided

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Janzen, D. H . , W. Hallwachs, J. Jimenez, and R. Gamez. 1993. The role o f the parataxonomists, inventory managers, and taxonomists i n Costa Rica's national biodiversity inventory. I n Biodiversity prospecting: Using genetic resources for sustainable development, ed. W. V. Reid et al., 223-54. Baltimore: World Resources Institute. Longino, J. T , and R. K. Colwell. 1997. Biodiversity assessment using structured inventory: Capturing the ant fauna o f a tropical rain forest. Ecological Applications 7:1263-77. New, T. R. 1998. Invertebrate surveys for conservation. Oxford: Oxford University Press. 240 pp. Noss, R. F., and A. Y. Cooperrider. 1994. Savingnature's legacy: Protecting and restoring biodiversity. Washington, D.C.: Island Press. 416 pp. Oliver, I . , and A. J. Beattie. 1996. Designing a costeffective invertebrate survey: A test of methods for rapid assessment o f biodiversity. Ecological Applications 6:594-607.

Olivieri, S. T , J. Harrison, and J. R. Busby. 1995. Data and information management and communication. I n Global biodiversity assessment, ed. V. H . Heywood and R. T. Watson, 607-70. Cambridge: Cambridge University Press. Prendergast, J. R., R. M . Quinn, J. H . Lawton, B. C. Eversham, and D. W. Gibbons. 1993. Rare species, the coincidence o f diversity hotspots and conservation stategies. Nature 365:335-37. Rossman, A. Y., R. E. Tulloss, T. E. O'Dell, and R. G. Thorn. 1998. Protocols for an all taxa biodiversity inventory of fungi in a Costa Mean conservation area. Boone, N.C.: Parkway Publishers. 195 pp. Samways, M . J. 1994. Insect conservation biology. London: Chapman and Hall. 358 pp. Zeledn, R. 2000. 10 aos del INBio: De una utopia a una realidad. Santo Domingo de Heredia: Instituto Nacional de Biodiversidad. 144 pp.

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