SHORT COMMUNICATIONS 317

Wilson Bulletin 117(3):317±319, 2005

Divorce in the Stonechat ( dacotiae)

Juan Carlos Illera1

ABSTRACT.ÐI report the ®rst case of divorce for stricted to slopes of stony ®elds and ravines the Canary Islands Stonechat (Saxicola dacotiae), an covered by medium to large shrubs and large endemic of the semiarid island of Fuer- boulders; they avoid lava and sandy teventura (Canary Islands, Spain). I studied 72 pairs during three breeding seasons (2000±2001, 2001± (Illera 2001). Stonechats are thought to be 2002, and 2002±2003). In 2001±2002, a female di- monogamous, sedentary, and territorial (Mar- vorced after a successful ®rst nesting. This female set- tõÂn and Lorenzo 2001, Urquhart 2002, Illera tled in a neighboring territory where the owner was 2004b). Territory boundaries usually abut unpaired, built a new nest, and laid four eggs. The low those of neighboring pairs (JCI pers. obs.). rate of divorce (1.4%) suggests that unforced mate After settling, individual are extremely changes by Canary Islands Stonechats are rare. Re- ceived 22 July 2004, accepted 31 May 2005. faithful to their sites all year long, i.e., they do not move after the breeding season, al- though territory boundaries may shift between successive breeding seasons (Illera 2004b; JCI The Canary Islands Stonechat (Saxicola unpubl. data). The breeding period extends dacotiae) is an island endemic restricted to from December to April (MartõÂn and Lorenzo Island, Canary Islands, Spain. Bibby and Hill (1987) estimated the popula- 2001, Urquhart 2002, Illera 2004b; JCI un- tion size of the species as 650±850 pairs. Al- publ. data). Reproductive effort and the du- though there are no recent estimates of the ration of the breeding period depend proxi- abundance of the species, its status has been mately on rainfall and ultimately on food (ar- modi®ed recently by the Spanish Ornitholog- thropod) availability (Illera 2004b; JCI un- ical Society (SEO/BirdLife) from ``Near publ. data). Pairs breed over more extended Threatened'' to ``Endangered'' on the basis of periods and lay two clutches in wet years, increasing destruction and alteration of its whereas in dry years they breed only once or habitats (Illera 2004a). In spite of its critical not at all. Clutches are also larger in wet than status, very little is known about the biology in dry years (Illera 2004b; JCI unpubl. data). of this species. Here, I describe the ®rst case Results presented here were obtained dur- of divorce in the Canary Islands Stonechat. ing studies of stonechat breeding success and The Canary Islands archipelago is located site ®delity. Birds were trapped, banded, and in the Atlantic Ocean, 100±460 km off the monitored (Illera 2004b; JCI unpubl. data). I northwest coast of Africa, and comprises sev- monitored 72 pairs over three breeding sea- en main volcanic islands and several islets. sons (2000±2001, 2001±2002, and 2002± Fuerteventura is the easternmost (28Њ 46Ј N, 2003) at 12 study sites in which 1±10 pairs 14Њ 31Ј W), second largest (1,660 km2), and bred at least once in the 3 years. Of 72 pairs, oldest island (approximately 22 million years I color-banded 114 individuals: 32/42 (2000± old; Carracedo and Day 2002). The climate is 2001), 39/47 (2001±2002), and 43/54 (2002± semiarid with dry summers and scarce rainfall 2003). Due to sexual dimorphism and individ- in autumn and winter (mean annual precipi- ual variability in phenotypic traits, I was also tation ϭ 117 mm; Marzol-JaeÂn 1984). The able to identify unbanded individuals. How- vegetation is mainly sparse xerophytic shrub- ever, because sexual dimorphism traits shift land. Canary Islands Stonechats are largely re- after the molt period (Illera and Atienza 2002), they were only used to identify indi- viduals within each breeding season. The 1 Depto. de BiologõÂa (ZoologõÂa), Facultad de BiologõÂa, Univ. de La Laguna, E-38206 La Laguna number of banded pairs studied during two or (Tenerife), Canary Islands, Spain; e-mail: jcillera@ more breeding periods was seven. I monitored ull.es pairs at least weekly from late October to ear- 318 THE WILSON BULLETIN • Vol. 117, No. 3, September 2005 ly November (pre-breeding period) until a mate loss while feeding ¯edglings. Bibby and month after the last ¯edgling left the nest Hill (1987) reported that some unpaired male (post-breeding period). The time spent moni- Canary Islands Stonechats assist with feeding toring each pair during each visit varied be- young of established pairs. However, during tween 30 min and 2 hr. Stonechats nested once 238.5 hr of observation of feeding (23 pairs), or not at all in the dry (27.3 mm rainfall) I did not record males or females assisting breeding season of 2000±2001, and laid two with feeding young. clutches in the two wet years (124.5 and 125.1 The low rate of within-season divorce mm rainfall in 2002 and 2003, respectively; (1.4%, n ϭ 72 pairs; three breeding seasons) Illera 2004b; JCI unpubl. data). and the lack of between-season divorce (0/7) Of 72 pairs monitored, only one divorced suggests that unforced mate changes by Ca- within a breeding season (2001±2002). Di- nary Islands Stonechats are rare. Strong site vorce was not recorded for pairs monitored (n ®delity reported for this species (Illera 2004b; ϭ 7) in successive breeding seasons. The case JCI unpubl. data.) also suggests that occur- of divorce reported here was the result of de- rences of between-season divorce are rare. Di- sertion by a female that subsequently settled vorce did not appear to have been triggered in a neighboring territory where the owner by poor breeding success, as mean fecundity was unpaired. Desertion occurred after the of all pairs monitored over 3 years was 2.25 original pair successfully reared a brood (three Ϯ 0.15 SE (n ϭ 68). The biotic and abiotic ¯edglings). In the ®rst brood (about 15 days homogeneity of Fuerteventura and the likely previous to divorce), during 90 min of obser- costs associated with territory switching (e.g., vation, 9 and 10 feeding visits were performed increased probability of predation after a by the female and the male, respectively. In move, aggression of adjacent territory holders, six visits to this territory, I did not observe and less ef®cient foraging in unfamiliar terri- the second male assisting with feeding young tories; Jakob et al. 2001, Yoder et al. 2004) or encounters between the female and the sec- probably makes divorce maladaptive in this ond male. All three adults were color-banded. species. Fifteen days after the ®rst brood ¯edged, the female built a new nest and laid four eggs. ACKNOWLEDGMENTS Both members of the new pair fed nestlings The Regional Government of the Canary Islands of the second brood, with no assistance from gave the of®cial permit to band this threatened bird the divorced male. The nest was depredated, species. D. Santana and N. HernaÂndez provided worms but the new pair did not attempt to breed used in trapping birds. A. Moreno and J. C. Atienza helped during banding sessions. M. Nogales reviewed again, and 2 weeks later the female disap- an early draft. M. DõÂaz encouraged me to write this peared from the territory. The female's origi- note and provided useful suggestions that improved nal mate continued singing and feeding its this paper a great deal. K. Koivula, E. D. Urquhart, three chicks alone during the 4 weeks after and an anonymous referee provided useful comments ¯edging. The divorced female never returned during revision. to her former territory even though the terri- LITERATURE CITED tories were adjacent. The deserted male did not mate with another female during the re- BIBBY,C.J.AND D. A. HILL. 1987. Status of the Fuer- mainder of the 2001±2002 breeding period, teventura Stonechat Saxicola dacotiae. Ibis 129: 491±498. and he ®nally moved to a new territory 5 CARRACEDO,J.C.AND S. DAY. 2002. Canary Islands. months later. Classic geology in Europe series. Terra Publish- Limited information is available on divorce ing, Hertfordshire, London, United Kingdom. in the genus Saxicola (Urquhart 2002). John- ILLERA, J. C. 2001. selection by the Canary son (1961, 1971) recorded several cases of Islands Stonechat (Saxicola dacotiae) (Meade- mate exchanges, polygyny, and polyandry in Waldo, 1889) in Fuerteventura Island: a two-tier a population of Common Stonechat (S. tor- habitat approach with implications for its conser- vation. Biological Conservation 97:339±345. quata), both within a given breeding season ILLERA, J. C. 2004a. Tarabilla Canaria (Saxicola daco- and between successive breeding seasons. H. tiae). Pages 327±328 in Libro rojo de las aves de Flinks (in Urquhart 2002) reported that male EspanÄa (A. MadronÄo, C. GonzaÂlez, and J. C. Common Stonechats might be vulnerable to Atienza, Eds.). SEO/BirdLife-DireccioÂn General SHORT COMMUNICATIONS 319

de Biodiversidad/Ministerio de Medio Ambiente, JOHNSON, E. D. H. 1971. Observations on a resident Madrid, Spain. population of Stonechats in Jersey. British Birds ILLERA, J. C. 2004b. BiogeografõÂa y ecologõÂa de la Ta- 64:201±213, 267±279. rabilla Canaria (Saxicola dacotiae) con implica- MARTÂõN,A.AND J. A. LORENZO. 2001. Aves del Ar- ciones para su conservacioÂn. Ph.D. dissertation, chipieÂlago Canario. Francisco Lemus Editor, La University of La Laguna, Tenerife, Spain. Laguna, Tenerife, Spain. ILLERA,J.C.AND J. C. ATIENZA. 2002. DeterminacioÂn MARZOL-JAEÂ N, M. V. 1984. El clima. Pages 157±202 del sexo y edad en la Tarabilla Canaria Saxicola in GeografõÂa de Canarias (L. Afonso, Dir.). Inter- dacotiae mediante el estudio de la muda. Ardeola insular Canaria, Santa Cruz de Tenerife, Spain. 49:273±281. URQUHART, E. D. 2002. Stonechats: guide to the genus JAKOB, E. M., A. H. PORTER, AND G. W. UETZ. 2001. Saxicola. Christopher Helm, London, United Site ®delity and the costs of movement among Kingdom. territories: an example from colonial web-building YODER, J. M., E. A. MARSCHALL, AND D. A. SWANSON. spiders. Canadian Journal of Zoology 79:2094± 2004. The cost of dispersal: predation as a func- 2100. tion of movement and site familiarity in Ruffed JOHNSON, E. D. H. 1961. The pair relationship and po- Grouse. Behavioral Ecology 15:469±476. lygyny in the Stonechat. British Birds 54:213± 225.

Wilson Bulletin 117(3):319±321, 2005

Regurgitated Mistletoe Seeds in the Nest of the Yellow-crowned Tyrannulet (Tyrannulus elatus)

Peter A. Hosner1

ABSTRACT.ÐI describe a Yellow-crowned Tyran- cies (Tyrannidae: Traylor 1977; Lanyon 1984, nulet (Tyrannulus elatus) nest built largely of mistletoe 1988; Mobley 2002; Fitzpatrick 2004). A sin- seeds, which differs from the cup of plant matter typ- gle atypical nest of Fork-tailed Flycatcher ically constructed by this species. Mistletoe seeds have been observed in the nests of at least two other bird (Tyrannus savana) in Brazil contained mistle- species, but this observation is the ®rst where the nest toe seeds regurgitated by an adult into the nest appeared to be purposely constructed from seeds, pos- cup during incubation, apparently to help keep sibly to take advantage of their adhesive properties. the loosely structured nest from falling apart Received 20 September 2004, accepted 31 May 2005. (Sick 1985). Other descriptions of T. savana nests make no mention of mistletoe seeds (Hilty and Brown 1986, Stiles and Skutch Adhesive substances, such as the saliva of 1989, Howell and Webb 1995, Hilty 2003, Aerodramus and Collocalia swiftlets (Smy- Fitzpatrick 2004). In Venezuela, a single glob- thies 1999, Hansell 2000), and regurgitated ular nest of the Great Kiskadee (Pitangus sul- materials, such as fruits and seeds ingested by phuratus) was observed to contain a few re- Oilbirds (Steatornis caripensis; Hilty and gurgitated mistletoe seeds about the lower rim Brown 1986), are regularly used for nest con- struction by these and a few other bird spe- and below the side entrance to the nest (J. A. cies. However, the use of adhesive mistletoe Mobley pers. obs.). No published descriptions fruit seeds in nest construction is almost un- of P. sulphuratus nests mention mistletoe known. There have been only two other re- seeds (Hilty and Brown 1986, Stiles and ports of mistletoe seeds used in nest construc- Skutch 1989, Howell and Webb 1995, Hilty tion, both in nests of unrelated ¯ycatcher spe- 2003, Fitzpatrick 2004). Here, I provide the ®rst report of a nest covered almost entirely 1 Dept. of Ecology and Evolutionary Biology, Cor- with mistletoe seeds, by the Yellow-crowned nell Univ., Ithaca, NY 14853, USA; e-mail: Tyrannulet (Tyrannulus elatus). [email protected] Birds, including T. elatus (Hilty and Brown 320 THE WILSON BULLETIN • Vol. 117, No. 3, September 2005

1986, Stiles and Skutch 1989, Hilty 2003, nest did not appear large enough to accom- Fitzpatrick 2004), are well-known consumers modate a second egg. Nest-construction ma- and dispersers of mistletoes (Viscaceae, Lo- terials and the pattern of egg markings dif- ranthaceae) in the Neotropics (e.g., Calder and fered from those described in other published Bernhardt 1983, Sargent 1994, Restrepo et al. accounts (Penard and Penard 1910, Snethlage 2002). However, other uses of these plants by 1935, Hilty and Brown 1986, Stiles and birds are poorly known. The seeds of some Skutch 1989, Hilty 2003, Fitzpatrick 2004); mistletoe species contain an extremely adhe- these authors described the nest as a shallow sive layer of viscin tissue under the fruit ¯esh; cup built of small twigs, mosses, spiderwebs, once digested, viscin can sometimes elongate grasses, tree bark, feathers, and other ®ne ma- to form a thread 3±4 cm long that is attached terial, and containing a clutch of two un- to one end of the seed. During regurgitation marked white or cream-colored eggs. A nest or defecation of the seed, the loose end of this of T. elatus found in the same clearing on Bar- thread will stay in the stomach or cloaca until ro Colorado in 2003 (R. Moore pers. obs.) was it is wiped on a substrate, usually a branch. similar to the published descriptions; there- Leverage is then used to remove the strand fore, the nest built of mistletoe seeds was not entirely from the body (Kuijt 1969, Calder likely to have been a local adaptation. and Bernhardt 1983). This is an effective dis- On 20 March 2004, the bird attending the persal strategy for mistletoes, as the seeds are egg (distinguishable from its mate by awk- occasionally placed on a branch in a location wardly arranged feathers on the crown) left suitable for germination. for short foraging bouts that lasted ϳ5 min. The T. elatus nest was found on 20 March Upon returning to the nest, it would resume 2004, on Barro Colorado Island, Panama incubation, and then regurgitate three to four (9Њ 9Ј 49Љ N, 79Њ 50Ј 16Љ W; 80 m in elevation) additional seeds on the nest. The bird then in a clearing in humid, lowland evergreen for- used its bill to move the seeds to a location est. When found, the nest was complete and on the nest's exterior, or work them into the contained a single egg. The nest was placed plant material in the cup wall, thus continuing approximately 7 m above ground on a small to construct the nest after the egg was laid. horizontal branchÐabout 2 cm in diameterÐ Placement of each seed took a short period of in a 20-m broadleaf evergreen tree parasitized timeÐbetween 5 and 15 sec in 3 hr of obser- by several mistletoe plants, on which the ¯y- vation. I observed four such foraging bouts catcher pair fed. The cup-shaped nest was followed by seed regurgitation and placement. small, approximately 5 cm in diameter (out- Both the attending bird and its mate also side rim to outside rim), and was built of un- wiped regurgitated seeds on branches in the determined plant material. Regurgitated seeds nest tree. Both adults actively defended the covered the entire exterior of the nest, and territory against Palm Tanagers (Thraupis pal- there was a band of regurgitated seeds that marum) and Social Flycatchers (Myiozetetes wrapped beneath the branch on which the nest similis), two other species that may consume was built. Seeds were also worked into the mistletoe fruits. plant material in the nest's interior. Two hypotheses might explain wiping mis- The mistletoe seeds were probably of the tletoe seeds on nests: either the adhesive prop- genus Struthanthus or Oryctanthus, with Stru- erties of the seeds were used intentionally to thanthus more likely based on the round (as aid in nest construction, or the nest was sim- opposed to more ovoid) seed shape (S. Sar- ply the most convenient object on which the gent pers. comm.) Upon regurgitation, the incubating bird could wipe seeds. The careful seeds were a whitish color, but seeds already placement of regurgitated seeds by T. elatus placed on the nest had dried and turned a on the nest exterior, however, suggests that the crimson-red to orange-red color. The single seeds were being intentionally used for their egg was white with small, evenly dispersed adhesive properties to hold the small nest to- blotches of pale cinnamon to raw umber. The gether and to adhere it to the branch. Placing color appeared to be from pigment in the egg- seeds underneath the supporting branch and shell rather than from stains from the mistle- working seeds into the nest cup takes more toe seeds, as the colors were different. The concentrated effort and care compared to wip- SHORT COMMUNICATIONS 321 ing them on the side of the nest or a nearby HANSELL, M. H. 2000. Bird nests and construction be- branch. Because mistletoe seeds are adhesive havior. Cambridge University Press, Cambridge, and viscin strands are strong, it seems possible United Kingdom. HILTY, S. L. 2003. Birds of Venezuela, 2nd ed. Prince- that the seeds were used to strengthen the nest, ton University Press, Princeton, New Jersey. perhaps allowing the overall size of the nest HILTY,S.L.AND W. L. BROWN. 1986. A guide to the to be smaller than previously described nests, birds of Colombia. Princeton University Press, which would decrease the effort needed to Princeton, New Jersey. build a nest. Because the bright, red-orange HOWELL,S.N.G.AND S. WEBB. 1995. A guide to the color of the seeds contrasted strongly with the birds of Mexico and northern Central America. Oxford University Press, New York. surroundings, it seems unlikely that the seeds KUIJT, J. 1969. The biology of parasitic ¯owering were used as camou¯age. plants. University of California Press, Berkeley. Conversely, Sick's (1985) description of the LANYON, W. E. 1984. A phylogeny of the kingbirds T. savana nest and J. A. Mobley's (pers. and their allies. American Museum Novitates comm.) observations of P. sulphuratus nests 2797:1±28. LANYON, W. E. 1988. A phylogeny of the thirty-two indicate that the addition of mistletoe seeds to genera in the Elaenia assemblage of tyrant ¯y- the nest may be only a convenient means of catchers. American Museum Novitates 2914:1± wiping seeds while incubating, with no intent 57. to alter nest structure. Sick (1985) suggested MOBLEY, J. A. 2002. Molecular phylogenetics and the the seeds were inadvertently voided in the evolution of nest building in kingbirds and their nest; neither he nor J. A. Mobley (pers. allies (Aves: Tyrannidae). Ph.D. dissertation, Uni- versity of California, Berkeley. comm.) observed active placement of, or nest PENARD,F.P.AND A. P. PENARD. 1910. De vogels van building with, mistletoe seeds, and the few Guyana (Suriname, Cayenne en Demerara), vol. seeds present were not enough to affect nest 2. Published by F. P. Penard, Paramaribo, Surina- construction. me. RESTREPO, C., S. SARGENT,D.J.LEVEY, AND D. M. ACKNOWLEDGMENTS WATSON. 2002. The role of vertebrates in the di- versi®cation of New World mistletoes. Pages 83± I would like to thank I. J. Lovette, Cornell Univer- 98 in Seed dispersal and frugivory: ecology, evo- sity, and the National Science Foundation (04222333) lution, and conservation (D. J. Levey, W. R. Silva, for funding ®eldwork. I also thank R. Moore for ad- and M. Galetti, Eds.). CABI Publishing, New ditional observations; J. A. Mobley for observations York. and suggestions; K. Zyskowski for translations of two SARGENT, S. 1994. Seed dispersal of mistletoes by nest descriptions into English, as well as comments birds in Monteverde, Costa Rica. Ph.D. disserta- and suggestions; W. Belton, J. Tobias, M. Hansell, and tion, Cornell University, Ithaca, New York. S. Sargent for their expertise in Neotropical birds, nest SICK, H. 1985. Birds in Brazil: a natural history. biology, and mistletoe biology; and D. W. Winkler and Princeton University Press, Princeton, New Jer- M. J. Andersen for comments. Last, I thank R. E. Ben- sey. netts and two anonymous reviewers for comments and SMYTHIES, B. E. 1999. Birds of Borneo, 4th ed. (re- suggestions that improved the manuscript. vised by G. W. H. Davison). Natural History Pub- lications, Kota Kinabalu, Borneo. LITERATURE CITED SNETHLAGE, E. 1935. BeitraÈge zur Fortp¯anzungs-biol- ogie brasilianischer VoÈgel. Journal fuÈr Ornitholo- CALDER,M.AND P. B ERNHARDT. 1983. The biology of gie 83:532±562. mistletoes. Academic Press, New York. STILES,F.G.AND A. F. SKUTCH. 1989. A guide to the FITZPATRICK, J. W. 2004. Family Tyrannidae (tyrant- birds of Costa Rica. Cornell University Press, Ith- ¯ycatchers). Pages 170±463 in Handbook of birds aca, New York. of the world, vol. 9: cotingas to pipits and wagtails TRAYLOR, M. A. 1977. A classi®cation of the tyrant (J. del Hoyo, A. Elliott, and D. A. Christie, Eds.). ¯ycatchers (Tyrannidae). Bulletin of the Museum Lynx Edicions, Barcelona, Spain. of Comparative Zoology 148:129±184.