In This Issue: Infectious Three Mammals and Diseases and Two Plants Listed Endangered Species Management Agriculture, Wetlands and Endangered U.S

U PDATEEndangeredIncluding a Reprint Species of theTechnical latest USFWS Bulletin

THE UNIVERSITY OF MIC'HI(;,Zlri July 1989 Vol . 6 No. 9 School of Natural Resources

In this issue: Infectious Three Mammals and Diseases and Two Plants Listed Endangered Species Management Agriculture, Wetlands and Endangered U.S. Bans Ivory Species: The Food Imports Security Act Infectious Diseases and Endangered Species Management by Andrew Dobson and Dan Miller

Disease can play a pivotal role in standing of the types of pathogens that macroparasites (Anderson and May the population dynamics of endangered pose a threat to endangered popula- 1979). Microparasites include viruses, species. The mountain gorillas of tions. Several important general prin- bacteria, fungi, and many protozoans; Virunga recently suffered an epidemic ciples have emerged from the resur- these pathogens are characterized by of respiratory illness which infected an gence of interest in the ecology of para- their small size (invisible to the naked estimated ten percent of the population sites and disease of the last decade eye), their ability to reproduce directly and led to the death of six gorillas (Anderson and May 1979, May 1988, in infected hosts, and their ability to (Sholley 1989). The causative patho- Scott and Dobson 1989). The greatest induce sustained immunity to reinfec- logical agent was identified as measles, level of sophistication and detail is tion in their hosts. Macroparasites, presumably introduced through contact available for models of infectious dis- which include the parasitic helminths, with an infected human. Luckily an eases in human populations, where ticks, and fleas, are characterized by inoculation campaign was quickly mathematical models have become in- their larger size (visible to the naked mounted, and large scale vaccination of creasingly prominent as tools for the eye), and their relative inability to re- potentially susceptible individuals pre- study of disease dynamics. Many of the produce directly within infected hosts. vented a full epidemic. In a less fortu- important epidemiological features of This classification makes sense nate incident, North Sea populations of these models can be applied to systems when the characteristic patterns of the harbour seals (Phoca vitulinu) were where less comprehensive data are two types of infection are considered. struck by a viral epidemic last summer available. Essentially, hosts acquire macropara- which led to the deaths of more than sites as discrete numbers of infective 14,000 animals (Anon 1989). Such larvae which give rise to an infrapopu- epidemics are not confined to verte- Types of Pathogens lation of adult parasites in each infected brates. Crayfish plague, a disease host; these parasites induce a limited caused by a fungal pathogen, threatens Parasites and pathogens can be immune response in the host that may the endangered whiteclawed crayfish conveniently classified on epidemiol- diminish the rate of acquisition of sub- (Austropotamobius pallipes) in its last ogical grounds as microparasites and sequent challenge infections. In con- stronahold in the lakes of central trast, microparasites tend to infect Ireland (Reynolds 1988). These examples suggest that an understanding of the population dynamics of pathogens, and the diseases they cause, is essential to the management of populations of both endangered and apparently healthy populations. In this short review we outline some important features of the population biology of infectious diseases and discuss their implications for the conservation of endangered populations, The dynamics of parasites and pathogens are intrinsically diffi- cult to study in endangered species because of small population sizes and the concern for protecting the remaining populations of a species. However, studies on non- endangered species can be used as a means of gaining a basic under- black-footed ferret (Mustella nigripcs) M.Wa through the lifetime of a mature

1 Endangered Species UPDATE Vob 6 No. 9 female in the absence of density de- population is disturbed or stressed; pendent consmints (Anderson and May these outbreaks may reduce the popula- 1982b). In both cases, R, must be equal tion to sizes where demographic sto- to or greater than one in order for the in- chasticity or AUee effects prevent re- fection to persist. NT is the threshold covery, Such effects may be com- Endangered Species density of susceptibles required for a pounded by increased levels UPDATE disease to establish in a population. R, of aggression in high density popula- and N, generally have an inverse rela- tions, which in turn reduce levels of A forum for information exchange on immunocompetence. example, in a endangered species issues tionship; if R, is large, then a smaller NT o or July 1989 is required for establishment of a pathu- captive population of macaques Vd. 6 No. 9 gen. Thus, particularly virulent species (Macaca mulatta), clinically confirmed require large populations to sustain tetanus accounted for approximately 25 percent of the 230 deaths which oc- Rob Bbir.,...... Editor them; when a pathogen species is highly Suzanne Jones ...... Assistant Editor specific towards its host species, it is curred during a five-year study (Rawl- unlikely to be the ultimate cause of ins and Kessler 1982). The tetanus extinction in an endangered population, organism, which is found in the feces of Instructions for Authors: because the pathogen itself is likely to infected animals, enters the body The Endongend Species UPDATE become extinct when host population through puncture wounds, postpartum welcomes articles related to species density falls below N, through host contamination of the uterus, or umbili- protection in a wide range of areas in- mortality or reductions in fecundity cal infections. Peak levels of annual cluding but not limited to: research and management activities for endangexd (Dobson and May 1986,1989). mortality occurred during the mating species, theoretical approaches to season when levels of aggression were species conservation, and habitat protec- at a peak and infection could occur tion and preserve design. Book reviews, Pathogens As A Threat to Endan- through septic wounds. editorial comments, and announcements of current events and publications are gered Species also welcome. One characteristic of most endan- Diseases That infect Multiple Readers include a broad range of gered species is small population sizes. Host Species professionals in both scientific and policy fields. Anicles should be written These small populations are often found in an'easily understandable style for a at artificiallyhigh densities in reserves, Pathogens that can utilize a range knowledgeable audience. thereby often facilitating the rapid of host species present the greatest Manuscripts should be 10-12 double spread of pathogens that may be intro- threat to endangered species. The pres- spaced typed pages. For further information please contact Rob Blair at duced into these populations, even if the ence of reservoir host species allows the number listed below. population density is below the thresh- generalist pathogens to establish and old density required to maintain an maintain infections in populations oth- Subscription Informafion: endemic infection. A classic example erwise too small to sustain them (Dob- The Endangered Species UPDATE is published approximately ten times per of this effect may be the outbreak of son and May 1989). Where the pathu- year by the School of Natural Resources canine distemper that nearly eradicated gen is more virulent towards rarer spe- at 'Ihe University of Michigan. Annual the black-footed ferret (Mustella ni- cies, extinction of a susceptible popula- subscriptions are $15 each ($18 outside gripes) shortly after its rediscovery in the U.S.). Send check or money order tion may be facilitated by the presence (made payable to 'Ihe University of 1981 near Meeteetse in Wyoming of a more common coexisting host. In Michigan) to: (Thome and Williams 1988). The free- many cases the impact of the pathogen. ranging ferret population, which results from differences in the lengths Endangered Species UPDATE School of Natural Resources peaked at 129 individuals in 1984, was of time for which the various host spe- The University of Michigan reduced to six individuals by the end of cies have had to develop a resistance to Ann Arbor, MI 48109-11 15 1985 by an epizootic of the disease the pathogen. The establishment of (3 13)763-3243 (Forrest et al. 1988). pathogens of this type is often the result Where small populations have re- of the induction of a non-native host duced genetic variability due to past species which acts as a reservoir for the Cover: bottleneck effects and inbreeding, the disease while remaining comparatively Elk (Cervus elaphus catdensir) loss of genetic diversity at such loci as immune to it. A classic example of this the major histocompatibility complex type of introduction has occurred in Photo: Colorado Department of Natural and other immune system loci may in- Hawaii where the extinction of several Resources, Wildlife Division crease population susceptibility to a species of endemic landbirds was facili- The views expressed in the Endangered disease epidemic (Allendorf 1986, tated by the induction of a suitable Species UPDATE are those of the author O'Brien and Evermann 1988). Simi- vector for transmissions of avian ma- and may not necessarily reflect those of the U.S. Fish and Wildlife Service or larly, epidemics often break out when a laria The pathogen had little impact on The University of Michigan.

Vol. 6 No. 9 Endangered Species UPDA TE Table 1. Tactics taken in pathogen control may depend on the primary host More complex problems arise when elk feed alongside cattle during the period that abortions are occurring, Wild Host Domestic Host thus providing a potential source of infection for brucellosis-free cattle herds. Circumstantial evidence has implicated Reservoir Controlled culling of Control is likely if elk in two cases of brucellosis arising in Host wild host often seen economic losses in cattle in northwestern Wyoming as cheapest viable domestic animals (Boyce 1989, Tessaro 1986). In the means of control. are important. spring of 1989, an outbreak of bovine brucellosis which occurred near Dubois, Wyoming, was likely transmit- Path~l~gica 1 May exclude domestic Mortality in wild ted by infected elk and bison that were Host livestock from species may go feeding with this particular herd (E.T. "wild" areas. unnoticed. Thorne pers. comm.). A vaccination program at several feedgrounds was begun to control brucellosis in the elk. This control program the large variety of exotic bird species Brucellosis will help ease the conflicts between previously imported to the island, yet wildlife and livestock managers, and, if endemic Hawaiian species with no pre- Brucellosis is a major disease of successful, will set precedents for simi- vious exposure to the pathogen suc- domestic livestock that results in eco- lar programs involving other species. cumbed rapidly to infection (Warner nomic loss through abortions, infertil- The bison of Yellowstone National 1968, van Riper et al. 1986). ity, and loss of domestic animal prod- Park are also a reservoir of brucellosis Pathogens that infect both wild and ucts. It also poses a health risk to those (Tessaro 1986, McCorquodale and domestic species of hosts are of particu- who work with infected animals DiGiacomo 1985, Meagher 1973), as lar concern for conservation biologists (Thimm 1982). In the prairie regions of are the hybrid woods-plain bison (Bi- and managers of wild habitats. The northwestern Wyoming, elk and bison son bison athabascae X Bison bison control of these pathogens can lead to are reservoirs of the disease, with trans- bison) in Wood Buffalo National Park, conflicts between interest groups, pri- mission between elk occurring primar- Alberta (Broughton 1987). Like elk, marily livestock and wildlife managers. ily through contact with aborted fe- the presence of brucellosis in bison of- The problems that arise are often de- tuses. Infected elk have been present in fers several management problems. pendent upon whether domestic or wild Yellowstone National Park since at The fear of transmission of brucellosis animals are the primary host of the pa- least 1917 (Tunnicliff and Marsh 1935), from bison to cattle is such that all bison thogen (Table l). In essence, if the and infected animals are found in at migrating north across the Yellowstone pathogen is predominantly present in least 15 of the 23 herds that visit the Park border are shot (Meagher 1989). domestic animals, it is in everyone's winter feeding grounds in northwestern best interest to control it. However, if Wyoming (Thorne et al. 1979). The infections are maintained in wild ani- disease is most likely maintained be- Rinderpest mals, then it may be easier to control cause of the unnaturally crowded condi- interactions between wild and domestic tions resulting from winter feeding pro- In many ways rinderpest presents animals by reducing the size of the wild grams; non-feedground elk that have an obverse example to brucellosis. With host population. been tested have shown no evidence of rinderpest, domestic livestock act as the As examples of these two different disease (Thorne et al. 198 1). reservoir from which the pathogen can scenarios, we can compare and contrast Brucellosis in free-ranging elk has invade wild animals (Scott 1970, Plow- brucellosis with rinderpest - two mi- several implications for the manage- right 1982). The pathogen caused a croparasitic diseases which infect wild ment of elk. It causes abortion of the major pandemic in both cattle and wild and domestic ungulate populations in first pregnancy following infection, and game when first introduced into East the savanna grasslands of United States occasionally causes abortion of the sec- and Southern Africa at the end of the and East Africa, respectively. The ond. The net result is a 7 to 12 percent 19th century. causative agent of brucellosis is the bac- reduction in herd productivity, and a The development of a relatively terium Brucella abortus, while rinder- reduction in the potential sustainable effective vaccine in the 1940s led to pest is caused by a Morbillivirus. Both yield harvested by hunters (Boyce large scale inoculation schemes for pathogens cause problems around the 1989, Thorne, et al. 1979). Also, in- domestic livestock. However, these borders of nature reserves where do- fected elk pose a health problem for vaccines were only effective in animals mestic livestock come into contact with hunters handling infected animals that were devoid of colostral immunity wild ungulates. (Thorneet al. 1981, Kistneretal. 1982). (partial immunity conferred on newly-

( Continued on UPDATE page 4)

3 Endangered Species UPDATE Vo1. 6 No. 9 born offspring by immune mothers). during the dry season, which was re- lowers herd immunity. Re-exposure - Because vaccination campaigns were quired because fencing around the park which is inevitable given a sufficiently carried out only once a year, substantial boundary prevented the game popula- long time horizon - will ultimately populations of immature susceptible tions from making their normal migra- resort in higher levels of mortality. It is animals built up, and these were large tion. The presence of large populations therefore important to keep popula- enough to sustain rinderpest infections of ungulates throughout the year and the tions of endangered species in condi- in many areas (Plowright 1982). The alkaline conditions of the pools proved tions where they lose natural immunity development of a culture-attenuated ideal for the survival of anthrax spores; to common infections. Indeed, in the virus vaccine in the late 1950s produced the mortality from anthrax accounts for very long term, selection by parasites a vaccine that was effective in cattle of an estimated 60 to 70 percent of diag- may help to maintain population ge- all ages, and this facilitated the interna- nosed wildebeest deaths. The presence netic variation. Thus, the threat of ex- tional campaign which attempted to of large numbers of readily available posure to pathogens is one of the strong- completely control rinderpest in Afri- corpses has led to an increase in the est arguments that can be made for sub can livestock. In areas of Afiica where density of scavenging predators in the dividing populations of endangered this campaign was successfully park, which in turn has led to increased species into different isolated nature mounted it had the additional side-ef- predation and disturbance of uninfected reserves. fect of removing the source of infection prey species (Berry 1981). Ratios of for wild animals, and many populations predator to prey in the area are thus consequently increased in numbers. In significantly higher than they are in the Serengeti region of Tanzania, num- other comparable habitats. "In many cases the ef- bers of buffalo and wildebeest have ficacy of vaccination increased by almost a factor of five and other disease since the control of rinderpest (Sinclair Controlling Pathogens In Wild control schemes is 1977). Populations In areas of Africa where wars or determined by who economic problems make administra- Once it is realized that artificially foots the bill..." tion of an effective vaccination scheme high population densities and associa- impossible, rinderpest continues to af- tion with potential reservoir hosts are fect both wild and domestic ungulate major causes of disease outbreaks, it is populations (Rossiter et al. 1987). possible to make recommendations In situations where pathogens are These economically troubled areas about the steps that should be taken to naturally present in wild and domestic serve as primary loci of infection from prevent outbreaks. populations, there will always be inher- which rinderpest can occasionally It is important to differentiate be- ent difficulties in controlling them; bru- spread with the movement of infected tween pathogens introduced from inter- cellosis, for example, may be controlled cattle or game, and cause outbreaks in actions with other species and those that fairly readily in elk, but not in bison due areas free from infection. These out- are normally present in host popula- to differences in feeding habits, locale, breaks are potentially devastating be- tions. If pathogens are introduced from and the effects of the vaccine on the cause the majority of populations of other species it is important to minimize animals. Similarly, although it is pos- game animals in rinderpest-control ar- the opportunity for chance introduc- sible to vaccinate several thousand eas contain few immune individuals tions, which may be done isolating the cattle a day with rinderpest vaccine, it is once the pathogen has been controlled endangered population from contact impossible to vaccinate more than 20 to for several years. with potential sources of infection. For. 30 wild buffalo (Plowright 1982); even example, in cases where pathogens af- vaccinating this many buffalo would fect both wild and domestic species, involve considerable risk of physical Anthrax fencing can be used to control diseases injury to the workers doing the vaccina- such as trypanosomiasis and foot-and- tion. Under these circumstances it is Pathogens can also have effects on mouth disease in Africa (Taylor and highly unlikely that sufficient levels of the structure of ecological communities Martin 1987). However, the cost of herd immunity would be attained in the that lead to changes in the numbers of fences can be expensive and may dis- wild population to come anywhere near species that are not infected by the dis- rupt the normal migration routes of wild controlling the disease. ease. A long term study of predator- species, as in Etosha. prey relationships in Etosha National Where pathogens directly threaten Park has shown that providing water by the welfare of endangered species, iso- Economics of Disease Control digging waterholes and gravel pits has lation may seem an attractive option. led to an epidemic of anthrax in several However, it may ultimately prove detri- In many cases the efficacy of vacci- ungulate species (Berry 1981; Ebedes mental because the absence of exposure nation and other disease control 1976). The waterholes were dug to to other common pathogens relaxes se- schemes is determined by who foots the provide a source of water for ungulates lection for resistance to pathogens, and bill for control (May 1988). The eco-

Vol. 6 No. 9 Endangered Species UPDA TE 4 nomic problems involved here are fur- Acknowledgements Meagher, M. 1973. The Department of Agricul- ther compounded by the rate at which ture, the Department of the Interior, bllo- different interest groups discount the sis, and the fanner. BioScience 23:311-312. Andrew Dobson is an assistant Plowright, W. 1982. The effects of rinderpest future. Many farmers, wildstock man- professor, and Dan Miller is a graduate and ~&lpeStcontrol on wildlife in Africa. agers, and even politicians are as con- student, at the Department of Biology, Symp. Zool. Soc. hd.50:l-28. cerned for the future of different game University of Rochester, in Rochester, O'Brien, S. J., and J.F. Evennann. 1988. Inter- active influence of infectious disease and ge- species as are conservation biologists NY. Their work on wildlife diseases is netic diversity in natural populations. Trends and wildlife managers. However, the supported by a grant from Wildlife Ecd. Evd. 3:254:259. financial incentive determining how Conservation In tema tional . Rossiter, P. B., W. P. Taylor, B. Bwangamoi, A. much each group should pay towards R. H. Ngereza, P. D. S. Moorhouse, J. M. Haresnap, J. S. Wafula, J.. F. C. Nyange, and the control of a specific disease is highly Literature Cited I. D. Gumm. 1987. Continuing presence of dependent upon both the benefits that rindexpest virus as a threat in East Africa, each receives from the control and the Mendorf, F. W. 1986. Genetic drift and the loss 1983-1985. Veterinary Record 120:5942. length of time those benefits last. Un- of alleles versus hetemzygosity. Zoo Biology Scott, G. R. 1970. Rinderpest. Pages 20-35 in J. 5181-190. W. Davis, L H. Karstad and D. 0. Trainer, fortunately, there is often an inverse Anderson, R. M. and R. M. May. 1979. Popula- eds. Infectiovr Diseases of Wiki Mammals. tion biology of infectious diseases: Pan I. Iowa State University Press, Iowa. Nature 280:361-367. Scot!, M. E., and A. Dobson. 1989. The role of "Perhaps the present Anderson, R. M. andR. M. May. 1982a. Directly parasites in regulating host abundance. Para- epidemic of AIDS and transmitted infectious diseases: mtrol by sitology Today 5: 176-183. vaccination. Science 215: 1053-1060. Sinclair, A. R. E. 1977. The African BIcffalo: A the resurgence of Anderson, R. M. and R. M. May. 1982b. Popu- Study of Resource Limitation of Populations. malaria will alter the lation dynamics of human helminth infec- University of Chicago Press, Chicago, EL. ... tions: control by chemotherapy. Nature Sholley, C. R. 1989. Mountain gorilla update. perspectives of politi- 297:557-563. Oryx 2357-58. Anon. 1989. Death in the North Sea. Oryx Rawlins, B. G. and M. J. Kessler. 1982. A five- cians from their current 23:s-6. year study of tetanus in the Cayo Santiago short-term focus ..." Beny, H. H. 1981. Abnormal levels of disease Rhesus monkey colony: behavioral descrip- and predation as limiting factors for wilde- tion and epizootiology. Am. J. Primatology beest in the Etosha National Park. Madoqua 3:23-29. relationship between the finance base of 12:242-253. Reynolds, J. D. 1988. Crayfish extinctions and each of these organizations and the Broughton, E. 1987. Diseases affecting bison. crayfish plague in central Ireland. Biol. Cons. length of time for which they plan. The Pages 34-38 in H. W. Reynolds and A. W. L. 45:279-285. economic decisions made by the finan- Hawley, eds. Bison ecology in relation to Taylor, R. D., and R. B. Martin. 1987. Effects of agricultural development in the Slave River vete~aryfences on wildlife conservation in cially more vociferous agricultural and lowlands,NWT. Occ. Paper No. 63. Canadian Zimbabwe. Environ. Manage. 11:327-334. political lobbies are usually determined Wildlife Service. Tessaro, S. V. 1986. The existing and potential by events that occur on a timescale of Boyce, M. S. 1989. The Jackson Elk Herd. importance of brucellosis and tuberculosis in Cambiige University Press, New Rochelle. Canadian wildlife: a review. Can. Vet. J. months and years. In contrast, the rela- Dobson, A. P., and R. M. May. 1986. Disease 27:119-124. tively impoverished environmental or- and Conservation. Pages 345-365 in M. ?himm, B. M. 1982. Brucellosis:Dirtribidion in ganizations are concerned with times- SoulC, ed. ConservationBiology: The Science Man, Dcwnestic, and Wild Animals. Springer- cales that are measured in decades and ofScarclty and Diversity. Sinauer Associates, Verlag, Berlin. Sunderland, MA. Thome, E. T., and E. S. Williams. 1988. Disease centuries. Thus, financial aid is often Dobson, A. P. and R. M. May. 1989. Parasites, and endangered species: the black-footed fer- made available to control epidemic out- cuckoos and avian population dynamics. In ret as a recent example. Conserv. Biol. 266- breaks of disease where immediate C.M. Pemons, ed. Bird Population Studies: 74. profits are threatened, but little finan- Their Relevance lo Conrervation and Man- Thome, E. T., J. K. Morton, and W. C. Ray, 1979. agement. Oxford University Press (in pss). Brucellosis, its effect and impact on elk in cial incentive exists to continually Ebedes, H. 1976. Anthrax epizdcs in Etosha western Wyaning. Pages 212-220 in M. S. monitor the possibility of outbreaks in National Park. Madoqua 10:99-118. Boyce and L. D. Hayden-Wig, eds. North wild species of limited economic value. Forrest, S. C., D. E. Biggins, L. Richardson. T. American Elk: Ecology, Behavior, and Man- AIDS W. Clark, T. W. Campbell III, K. A. Fager- agement. The University of Wyoming. Lar- Perhaps the present epidemic of stone, and E. T. Thorne. 1988. Population amie, WY. and the resurgence of malaria - dis- attributes for the Mack-footed ferret (Mustela Thome, E. T., T. J. Walthall, and A. Tebaldi. eases whose economic and social ef- nigripes) at Meeteetse, Wyoming, 1981-1985. 1981. The incidence and imponance of bru- fects will be felt on the timescale of J. Mammal. 69:261-273. cellosis in elk in northwestern Wyoming. Job Kistner, T. P. 1982. Diseases and parasites. bp.Rep. Wyo. Game and Fish Canm., decades -will alter the perspectives of Pages 181-218in J. Thomas and D. E. Toweil, Lararnie, WY. politicians from their present short-term eds. Elk of North America: ecology and man- Tunnicliff, E. A., and H. Marsh. 1935. Bang's focus to one that promotes a longer term agement. Stackpde Books, Harrisburg. PA. disease in bison and elk in the Yellowstone approach to rational disease control. May. R. M. 1988. Conservation and disease. National Park and on the National Bison Cons. Bio. 2:28-30. Range. J. Am. Vet Med. Assoc. 86:745-752. This change in attitude is essential if the McCorquodale, S. M., and R. F. DiGiacxno. van Riper XU, C., S. G. van Riper, M. L Goff and major breakthroughs in epidemiologi- 1985. The role of wild North American mgu- M. Laird 1986. The epkmtidogy and eco- cal theory of the last 10 years are to be lates in the epidemiology of bovine brucello- logical signifkame of malaria in Hawaiian capitalized upon in the control of infec- sis. J. Wildl. Dis. 121:351-357. landbirds. Ecol. Monogr. 563327-344. Meagher, M. 1989. Evaluation of boundary Warner, R. E. 1%8. The role of introduced tious diseases in both wild and domestic control for bison of Yellowstone National diseases in the extinciion of the endemic populations. Park. Wildl. Soc. Bull. 17:15-19. Hawaiian avifauna. Condor 70:lOl-120.

5 Endangered Species UPDATE Vol. 6 No. 9 Invertebrate Subspecies and the Endangered Species Act by Dennis D. Murphy

Taxonomists' decisions can have new-found power of taxonomists to the pending publication of the paper enormous impact on our estimates of influence application of the Endangered alone was adequate to squelch the list- rates of vanishing biological diversity Species Act to invertebrates is well- ing. One of the remaining populations and our ability to employ the Endan- illustrated by the case of the callippe has become surrounded by housing de- gered Species Act. In the original Act, silverspot butterfly (Speyeria callippe velopments, thus the future of Speyeria Congress legislated "species" protec- callippe) . callippe callippe (sensu lato) is now in tion that could be extended to individ- Known from just three relict popu- additional jeopardy. ual populations. An amendment in the lations around San Francisco Bay, the Needless to say, this soft under- 1978 reauthorization of the Act re- silverspot was an inarguably strong belly of the Act has not been lost on the stricted protection for invertebrates to candidate for endangered status. As development interests that perceive subspecies or higher taxonomic entities time came for the USFWS to render its themselves threatened by specific in- (species, genera, etc.) that are threat- "biological opinion" on the proposal, a vertebrate listing proposals. Defense ened or endangered in their entirety. manuscript purporting to "revise" (reas- contractor United Technologies Corpo- With that amendment, Congress cre- sign) the subspeciesnames of Speyeria ration, for example, recently hired Ar- ated a source of unending controversy, callippe was circulated. Using princi- nold to make a similar taxonomic argu- and, certainly unintentionally, granted pal components analysis, Arnold (1985, ment to stop the listing of the bay check- taxonomists the power to determine Pan. Pac. Ent. 61: 1-23) drew the con- erspot butterfly, Euphydryas editha which among the world's millions of clusion that the species Speyeria cal- bayensis. That auempt ultimately invertebrates can be considered for fed- lippe consisted of not the 16 tradition- failed, but during the more than two eral protection. ally accepted, narrowly distributed sub- year delay that it caused, several popu- While the subspecies designation species, but just three broadly distrib- lations of the butterfly declined and one is generally recognized as a valuable uted subspecies. went extinct. tool for describing infraspecific geo- Principal components analysis is a Both of the above examples clearly graphic variation, it is also recognized valuable tool for assessing variation, indicate that irresponsible revisions that as an often arbitrary taxonomic con- but use of this technique does not guar- lump subspecies compromise the abil- struct that conveys little genetic or evo- antee unequivocal taxonomic assign- ity of the USFWS to fulfill the Congres- lutionary information. Wilson and ments. Biological insight is required in sional mandate to protect Earth's bio- Brown (1953, Sys. Zool. 2:97-111) choosing appropriate characters and in logical diversity, including diversity long ago noted that the use of different interpreting the results. In the case of within species. A better Endangered sets of physical characters can result in the silverspot, from an essentially infi- Species Act would confer federal pro- the division of species into different nite pool of phenetic, genetic, and eco- tection upon invertebrate popula- numbers of subspecies. Thus, all taxo- logical characters, Arnold selected for tions-for it is on that level that the nomic treatments below the species analysis but eight characters, all wing genetic diversity of invertebrate species level are particularly subject to dis- markings. Based on ventral hindwing is shaped and must be preserved. But, agreement . color, nine otherwise highly morpho- until the Act is improved, taxonomists The U.S. Fish and Wildlife Serv- logically and ecologically distinct sub- will continue to bear a heavy conserva- ice, short on staff and appropriations, species distributed from Mexico to tion responsibility-lumping of sub- and faced with thousands of candidates Oregon were lumped as one--Speyeria species as a by-product of studies of in- for endangered species protection, un- callippe callippe. Since this study was fraspecific variation could well mean derstandably attempts to avoid initiat- published, specialists have indicated the extinction of threatened or endan- ing the listing process for controversial that the characters chosen for the analy- gered populations. With that in mind candidates-exemplars of which are sis would not have separated full spe- taxonomists: do feel free to continue to organisms whose taxonomic status can- cies within the genus Speyeria, and that study infraspecific variation but, not be resolved with even the best avail- individuals of species other than Spey- please, lump subspecies only after all able scientific evidence. In this context, eria callippe appear to have been used factors are considered. otherwise innocuous infraspecific taxo- in the original analysis. Nonetheless, nomic decisions can have far-reaching following that treatment, the silverspot Dennis Murphy is director of The Center for Con- consequences. The implications of this would not qualify for protection, and servation Biology at Stanford University.

Vol. 6 No. 9 Endangered Species UPDATE 6 Bulletin Board

Restoration Conference North America north of Mexico was velopment (USAID) program for the recently begun by Mason E. Hale, Jr. Conservation of Biological Diversity. The second annual conference of and Sherry K. Pittam of the Smith- The program will promote sustainable the Society for Ecological Restoration sonian Institution, in cooperation with economic development in developing will be held in Chicago on April 29- The Nature Conservancy. The project's countries through better conservation May 3,1990. The program will include goal is to generate a list of rare and and use of biological resources. several special sessions and a full pro- endangered lichens, along with the in- The project is composed of five gram of contributed papers, lectures, formation necessary to provide protec- major parts: 1) technical assistance of- workshops, and field trips to facilitate tion for these species as needed. This fered to AID missions, host country communication among restorationists, information will then be made available institutions, local private volunteer or- decision makers, and thegeneral public. to The Nature Conservancy, as well as ganizations and the Peace Corps; 2) a Special session topics will include: other conservation organizations and small grants program for specific re- prairie and savanna restoration, restora- land management agencies. Those search issues relevant to AID'S world- tion and global climate change, setting interested are invited to submit the wide conservation activities; 3) train- standards for monitoring restoration names of potentially rare or endangered ing for local scientists in identifying projects, restoration and recovery of lichen species, along with any support- research priorities and preparing pro- endangered species, and restoration ing information. Please send to: Sherry posals; 4) an information and evalu- philosophy. The Society invites sub- K. Pittam, Rare Lichens Project, Smith- ation network on USAID conservation mission of paper abstracts on these top- sonian Institution, Botany/NHB 166, activities linked with other relevant ics in parbcular, as well as other aspects Washington, DC 20560; (202) 357- databases; and 5) pilot demonstration of ecological restoration. Deadline for 2545. projects funded largely by USAID mis- submission is January 15, 1990. Sub- sion and regional bureau buy-ins. For mission forms may be obtained from more information, contact: Dr. Gary S. the Society's offices: S.E.R., 1207 Conservation of Biological Diver- Hartshorn, Biodiversity Director, Seminole Highway, Madison, WI sity Program World Wildlife Fund, 1250 24th Street, 5371 1; (608) 263-7889. NW, Washington, DC 20037; (202) The World Wildlife Fund, the 778-9600. World Resources Institute, the Center Rare Lichens Project for International Development and Environment, and The Nature Conser- Bulletin bwrd information provided in part by A project to determine theconserva- vancy, are implementing a centrally- Jane Villa-Lobos,Smilhsonian Institution. tion status of the lichens of Hawaii and funded U.S. Aid for International De-

Non-Profit Organizat~on U.S. POSTAGE Endangered Species PAID Ann Arbor, MI UPDATE Permit No. 144

School of Natural Resources The Univers~tyof Michigan Ann Arbor, MI 48109-1115