Biology of the California Gnatcatcher: Filling in the Gaps
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WESTERN BIRDS Volume 29, Number 4, 1998 BIOLOGY OF THE CALIFORNIA GNATCATCHER: FILLING IN THE GAPS JOHN T. ROTENBERRY, Natural Reserve System and Department of Biology, Universityof California, Riverside,California 92521 THOMAS A. SCOTT, Departmentof EnvironmentalPolicy, Science,and Manage- ment, Universityof California, Berkeley, California 94720 Interest in the coastal populations of the Black-tailed Gnatcatcher (Polioptila melanura) first diffusedoutside of the ornithologicalcommunity in 1978. Notable for its odd vocalizations, localized distribution, and co- occurrencewith a vegetationthen known as "inland sage scrub"(Thorne 1976), the coastal subspeciesof the Black-tailed Gnatcatcher (P.m. californica)was judgedby environmentalplanners to be sensitivebecause of its small range and vulnerabilityto habitat conversion(M. U. Evans pers. comm.). Therefore, it warranted discussionin reviews of development projectswritten in responseto the CaliforniaEnvironmental Quality Act. Discussionsof the bird's sensitivity,however, rapidly intensified as it became apparentthat the subspeciesinhabited some of the most developablelands of coastal California. Conservationof the subspeciessoon became synony- mous with broader goal of conservingcoastal scrub vegetation,which by 1980 was consideredone of the most rapidly disappearingvegetation types in California(Westman 1981). In the late 1980s, Phillips(1986:xxvi and 75, 1991:25-26) and Atwood (1988) elevatedthe subspecies,with its relatives in Baja California,to the level of a species,the CaliforniaGnatcatcher (P. californica), restrictedin the U.S. to the coastalsage scrub vegetationtype and its variations(Westman 1981, O'Leary 1990, White and Padley 1997). Althoughthe CaliforniaGnatcatcher was originallydescribed as a speciesby William Brewsterin 1881, its reappreciationas such a centurylater almost guaranteed that southern California would experience an endangered- species conflict involving billions of dollars of real-estatedevelopment. Becauseof extensivehabitat loss (due to conversionto urban, suburban,and agriculturaldevelopment; Westman 1981, O'Leary 1990) and associated declinein the species'population (e.g., Atwood 1993), the U.S. Fish and Western Birds 29:237-241, 1998 237 BIOLOGY OF THE CALIFORNIA GNATCATCHER: FILLING IN THE GAPS Wildlife Service listed the California Gnatcatcheras a "threatened"species under the EndangeredSpecies Act (ESA) in March 1993 (USFWS 1993). The USFWS went on to state that the speciesprobably warranted "endan- gered" status but that ongoing conservationplanning ("Natural Communi- ties ConservationPlan"; Calif. Dept. Fish & Game 1993) reduced the level of threat. In anticipationof (and subsequentto) the federal listingof the gnatcatcher under the ESA, local, state and federal agencies,as well as conservationists and developers,began collectinginformation and data to support policy decisionsand permits requiredunder the act. Unlike previousconflicts over endangered birds, which occurred primarily on federal lands and were investigatedby teams of agency and resource-industryscientists [e.g., the Northern Spotted Owl ($trix occidentalis caurina); Murphy and Noon 1992, Guti6rrez et al. 1996), that over the gnatcatcherrepresented a new model of conflictover private land, where scientistswould be contractedfor researchby a bewilderingarray of publicagencies, corporations, and private individuals. Most of this information-gatheringactivity focused on the fine-scale details of gnatcatcherdistribution, particularly the identificationof occupied and unoccupiedhabitat. Ancillary to these distributionalstudies, data were also gathered on local movements,breeding biology (particularlyreproduc- tive success),habitat association,and territory sizes. Much of the informa- tion amassedfor managementand conservationdecisions, however, was recordedonly in unpublishedreports, or even simplyburied in an individual's field notebook. Mechanismsfor objectivereview of the information used in policydecisions did not exist.Although thousands of hours of researcheffort had been expended on the California Gnatcatcher, few peer-reviewed papers describingthis work were published.Administrative functions, such as issuanceof permits, continuedin absenceof publishedinformation, but conservation science suffered. Researchers had few avenues of communica- tion, no one could build off the establishedwork of others, and mechanisms for identifyingreliable data went unused.Thus the need for a symposiumon the biology of the California Gnatcatcher,and perhaps a new manner of looking at endangered-speciesresearch, arose from the chaos createdwhen so many researchersplunged into so many studiesin such a short period. We organized this symposiumfor the purpose of bringing gnatcatcher researcherstogether and lobbyingfor the publicationof the large body of data generated on the species. The California GnatcatcherSymposium, held 15-16 September1995 on the campusof the Universityof California, Riverside, drew 160 participants from academia, biological consulting businesses,and governmentagencies. We asked the participantsto present existing data and analyses and to help us identify the current status of scientificallybased knowledgeon the species.Our hope for the symposium proceedingswas to move as much of this knowledgeas possiblefrom the "gray" literatureinto the peer-reviewed(and more widely accessible)main- stream. The papers containedin this issue of Western Birds represent a major portion of the realizationof that goal. In further support of that goal, we were successfulin making a collectionof previouslyunpublished reports availablethrough the auspicesof the Van Tyne Memorial Library, curatedby 238 BIOLOGY OF THE CALIFORNIA GNATCATCHER: FILLING IN THE GAPS the Wilson Ornithological Society, at the Museum of Zoology at the University of Michigan. Although these reports have not undergonepeer review,they containvaluable data; each is cited in one or more of the papers in this volume as appropriate. At the symposium45 oral and poster papers were presented,represent- ing most aspects of California Gnatcatcherbiology. Although many were explicitly conservation or management oriented, that focus was not a criterion for inclusion on the program. From these 45, 22 appear in this collection;we are aware of at least sevenother presentations,portions or all of which have appeared or are currently in review in other professional journals. Publication of these proceedings would not have been possible without the generous support of Ogden Environmental Services, LSA Associates, the Metropolitan Water District of Southern California, the Riverside County Habitat Conservation Agency, and the University of California. From these papers we can distill several facts about the California Gnatcatcher.For example, at a regionalor landscapescale, the distribution of the speciesappears largely constrained by the distributionof coastalsage scrub vegetation;however, at more local scalesbirds may be found in (and perhaps depend on) additionalvegetation types, dependingon the particular local mosaic. Because individualsand pairs can be found in patches of suitable habitat quite isolated from the nearest population of any conse- quence, the power of dispersal of this nonmigratory species may be substantial.It is quite clear that gnatcatchersin southern California occupy a fragmented landscape and are governed by metapopulation dynamics. Territory sizes are highly variable, increasing significantly as one moves inland from the coast,and in generalare much larger on averagethan those of other passerinesof comparablebody mass. They are persistentnesters, constructingas many ten nests within a breedingseason. However, nests are frequentlyabandoned before eggs are laid, so that the number of clutchesis usuallyless than the number of nests built by a pair. Clutch size is variable, and much of the variation seemsrelated to variationin weather,particularly precipitation.Although California Gnatcatchersserve as frequent hosts to the Brown-headedCowbird (Molothrus ater), they also suffer a high rate of nest predation,which seems to exert the greatestcontrol over reproductive success. Nevertheless,numerous gaps in our knowledge of gnatcatcherbiology remain. For example,we lack certainty about the birds' mating systemand criteria for mate choice; although they appear sociallymonogamous, they also demonstratebehaviors that clearly suggesta potential for extra-pair copulationsfor both sexes.While mating systemsare of academicinterest to behavioral ecologists, they are also relevant to conservation biologists because of the implications they may have for genetic structuring of populationsand effective populationsize. Likewise, although there have been severalinvestigations of gnatcatchergenetics with a goal of elucidating taxonomicrelationships, we are currentlyunaware of any studiesexamining the geneticsof populationsor demes. In addition to providinginformation on effective population size, properly done molecularstudies also assess gene flow, genetictraces of the connectionsamong subpopulationsdistrib- 239 BIOLOGY OF THE CALIFORNIA GNATCATCHER: FILLING IN THE GAPS uted over a fragmentedlandscape. Because gnatcatchers nest persistently, their reproductiveeffort can be high; althoughthere is a rich body of life- history theory that deals with trade-offs among aspects of reproduction, none has been systematicallyapplied to the California Gnatcatcher. And althoughclutch size varies from year to year, it also varies from pair to pair