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Snowy Plover Reproductive Success in Beach and River Habitats

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Snowy Plover Reproductive Success in Beach and River Habitats J. Field Ornithol. 76(4):373±382, 2005 Snowy Plover reproductive success in beach and river habitats M. A. Colwell,1,3 C. B. Millett,1 J. J. Meyer,1 J. N. Hall,1 S. J. Hurley,1 S. E. McAllister,2 A. N. Transou,2 and R. R. LeValley2 1 Wildlife Department, Humboldt State University, Arcata, California 95521 USA 2 Mad River Biologists, 920 Samoa Boulevard, Suite 210, Arcata, California 95521 USA Received 25 October 2004; accepted 16 March 2005 ABSTRACT. Poor reproductive success has contributed to the decline and low population size of the federally listed Western Snowy Plover (Charadrius alexandrinus nivosus), especially where it breeds on coastal beaches used by humans for recreation. From 2001±2004, we compared reproductive success of color-marked plovers breeding on ocean beaches with those on gravel bars of the lower Eel River in coastal northern California, one of six recovery units as identi®ed by the species' recovery plan. In three of four years, more plovers (54±64%) nested in river than beach habitats, but this pattern was reversed in the last year of the study when 62% of plovers used beaches. Each year, a higher proportion of clutches hatched and more chicks ¯edged from river than beach habitats, producing a disproportionate number of yearlings recruited into the local population from the river. On average, river-nesting males tended signi®cantly fewer eggs, hatched similar numbers of chicks, and ¯edged signi®cantly more young compared with males breeding on beaches. Corvids were more prevalent in river habitats in two of four years, but beaches consistently had signi®cantly greater human activity. These habitat differences in reproductive success exist despite efforts to manage predators (e.g., exclosures around nests) and humans (e.g., signs, fencing, and vehicle restrictions) on beaches and almost no management of river habitats. SINOPSIS. EÂxito reproductico de Charadrius alexandrinus en habitats de playas y rios Un eÂxito reproductivo pobre ha contribuido a la merma del playero (Charadrius alexandrinus) particularmente cuando este anida en haÂbitats de playas utilizadas por humanos para la recreacioÂn. De 2001±2004, comparamos el eÂxito reproductivo de playeros que fueron anillados en playas y en graveros del rõÂo Eel que se encuentra en la parte norte de California. Esta es una de las seis localidades mencionadas en el Plan de RecuperacioÂn de esta especie que ha sido listada. En tres de los cuatro anÄos, una mayor cantidad de playeros (54±65%) anidaron en el haÂbitat de rõÂo que en el de playa, aunque este patroÂn se invirtio en el ultimo anÄo de estudio (64%). Cada anÄo una mayor proporcioÂn de camadas eclosionaron y un nuÂmero mayor de polluelos dejaron el nido en haÂbitats de rõÂos que en los de playa, produciendo una desproporcioÂn numeÂrica de volantones de un anÄo, reclutados en la poblacioÂn del rõÂo. En promedio, los machos que atendieron polluelos en rõÂos tuvieron que incubar menos huevos, ayudaron a eclosionar un numero similar de polluelos y produjeron maÂs volantones que los machos que utilizaron playas. Los coÂrvidos, que son depredadores, pasaron mas tiempo en haÂbitat de rõÂos en dos de los cuatro anÄos, aunque las playas consistentemente tuvieron mayor actividad de humanos, signi®cativamente. Las diferencias en el eÂxito reproductivo en los dos tipos de haÂbitats, existen a pesar del esfuerzo de manejar depredadores (e.j., sistemas que evitan que estos lleguen a los nidos), humanos (e.j., restriccioÂn al uso de vehõÂculos y roÂtulos) en las playas y virtualmente ninguÂn manejo en los rõÂos. Key words: Charadrius alexandrinus nivosus, ¯edging success, habitat quality, nesting success, reproductive success, site ®delity, Snowy Plover, threatened species For threatened and endangered species, it is population decline. This population decline essential that quanti®ed variation in habitat stemmed partly from poor reproduction asso- quality guide management efforts to increase ciated with predation of eggs and young by cor- reproduction or survival. In 1993, the U.S. Fish vids (Corvus corax, C. brachyrhynchos), gulls (La- and Wildlife Service (USFWS) listed the coastal rus spp.), red foxes (Vulpes vulpes), raccoons population segment of the Western Snowy Plo- (Procyon lotor), and striped skunks (Mephitis ver (Charadrius alexandrinus nivosus) as threat- mephitis) (USFWS 2001). In coastal central ened under the federal Endangered Species Act California, mammalian predator control suc- (ESA; USFWS 1993) based on a signi®cant cessfully increased hatching but not ¯edging success of Snowy Plovers (Neuman et al. 2004). 3 Corresponding author. Email: mac3@humboldt. Increased human recreation in the species' edu breeding habitats has also been cited as a cause 373 J. Field Ornithol. 374 M. A. Colwell et al. Autumn 2005 of low reproductive success (USFWS 2001). At docino counties (Colwell et al. 2004). Two sites Point Reyes National Seashore, the timing of (Clam Beach and South Spit/Eel River Wildlife chick mortality for beach nesting plovers oc- Area) are ocean-fronting beaches with vegeta- curred disproportionately on weekends, which tion dominated by European beach grass. At Ruhlen et al. (2003) suggested correlated with these beaches, most plovers nested on relatively high human disturbance of breeding plovers. homogeneous sandy substrates, often amidst European beach grass (Ammophila arenaria), an shells, driftwood, and other debris. Plovers (and introduced species that colonizes and alters the especially their tracks) were conspicuous in open sandy habitats preferred by plovers, has these ®ne substrates, making nests and broods also degraded breeding habitat (USFWS 2001). easy to ®nd and vulnerable to predation (Col- Along the Paci®c coast, Snowy Plovers breed well et al. 2004). After hatch, adults moved in sparsely vegetated habitats of salt pans and chicks to the upper reaches of ocean-fronting levees, dredge spoil islands, river gravel bars, beach, where debris offered food and cover. At and ocean beaches from central Washington both beach sites, but especially Clam Beach, south to Baja California, Mexico; within the humans used plover breeding habitats for rec- U.S., most individuals breed in California reation such as clamming, jogging, horseback (USFWS 2001). In northern California, plovers riding, dog walking and driving vehicles. winter and breed along ocean beaches (Page Plovers bred at gravel bars along the lower and Stenzel 1981) and river bars of the lower Eel River, from its con¯uence with the Paci®c Eel River (Tuttle et al. 1997), which led the Ocean upriver approximately 14 km (Colwell USFWS (2001) to designate coastal northern et al. 2004). River-breeding plovers nested in California as a discrete management unit (Re- coarse, heterogeneous substrates varying in size covery Unit 2). Within this area, the number from sand to pea-sized gravel and large stones, of breeding plovers has probably never been which were sparsely vegetated by willow (Salix high. In 1977, Page and Stenzel (1981) report- spp.) and white sweet clover (Melilotus alba). ed 80 plovers (26 nests) at 10 locations and Compared to beaches, plovers are more dif®cult estimated that this represented 6% of the pop- to observe along the river, nests are more cryp- ulation in coastal California. In the early 1990s, tic, and plover tracks are rarely seen in these this area supported 22±50 breeding plovers an- coarse substrates. Adults tended broods in the nually (USFWS 2001). Until recently, these same areas where they nested. Humans fre- plovers were not known to nest in habitats oth- quented the river much less than beach sites. er than coastal beaches. In 1996, however, Tut- tle et al. (1997) ®rst observed breeding plovers METHODS in a unique habitat, river bars of the lower Eel River; it remains unknown how long plovers Field observations. Intensive monitoring have bred in this habitat. Currently, the popu- of color-marked plovers began in 2001. Each lation size (for coastal northern California) rep- year, we captured plovers at nests using noose resents , 5% of the estimate for the listed pop- mats and circular nest traps, and banded nearly ulation and 40% of the recovery objective for all breeding plovers. Each individual received a the recovery unit (USFWS 2001). unique combination of colored leg bands. From From 2001±2004, we intensively monitored mid March to early September, observers sur- a marked population of plovers in Recovery veyed breeding plovers almost daily within each Unit 2. Here, we compare annual reproductive habitat. During surveys, we recorded the iden- success of plovers breeding in river and beach tity of banded plovers, found nests, and mon- habitats, and provide evidence that corvids and itored broods. Upon ®nding a nest, we recorded humans compromise the quality of breeding the number of eggs and estimated hatch date habitat for beach-breeding plovers. (for complete clutches) using egg ¯otation methods (Alberico 1995) or by the timing of STUDY AREA clutch completion for nests found with an in- complete clutch. On beaches, we erected pred- We monitored plovers at three principal sites ator exclosures at most (61%) nests; we never in Humboldt County, California (Fig. 1) and exclosed river nests. several secondary sites in Humboldt and Men- At hatching, we marked chicks with a brood- Vol. 76, No. 4 Plover Habitat Quality 375 Fig. 1. Location of beach (Clam Beach; South Spit and Eel River Wildlife Area) and river (Eel River gravel bars) habitats where Snowy Plovers bred in Humboldt County, California,
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