An open nest of a House Sparrow (Passer domesticus, Ploceidae) was located in the same position in the left corner of the church front, likewise protected by the column. Passer domesticus is a typical cavity nester, accordind to la Peña 1987, Scott et al. 1977, Sick 1985, Stoner and Stoner 1952, Summers-Smith 1988 and pers. obs. The nest was typically globular, probably closed (the rear was not visible) and occupied the whole width of the niche; some space was free at the top. The wide, round entrance was frontal. Some nest material hung down from the nest, and some had fallen to the ground. The materials used were grasses (stems, flowering branches and some leaves, mainly of Panicum maximum, Poaceae), very thin tree stems, kapok (of Chorisia speciosa, Bombacaceae), pieces of varied strings and shreds, and fragments of plastic tapes.

On later days two adults were seen feeding at least two nestlings at the entrance of the nest. On the ground under the nest, I found many fragments of insects (mainly Coleoptera), seeds (mostly of Murraya paniculata [Rutaceae] and some of Livistona chinensis [Arecaceae]) and pellets (containing chiefly insect remains and seeds of Murraya paniculata). On 25 September the nest was inactive, but on 8 October a pair of Passer domesticus had adopted the nest, and these later bred successfully. Five years later, the remains of this same nest were still visible.

On 7 December 1993 another nest, practically identical to the nest described above, was found in a niche 10.54 m high in the left wall of the façade and partially protected from lateral view by the large column. The entrance of the nest was about 9 cm in diameter. Some old typical Pitangus sulphuratus pellets were on the ground under the nest. Because of its construction, this nest may be safely ascribed to Pitangus sulphuratus. The nest was then occupied by a pair of Passer domesticus that was seen feeding two nestlings at the entrance. Another seven open nests built by Pitangus domesticus were placed in the niches, mainly at medium heights. A niche in the higher row contained a nest of Polybia sp. (Hymenoptera, Vespidae) attached to its roof. No nest of niche-nesting Columba livia (Columbidae), very common in Piracicaba, were found in the façade, probably because the niches favored by that species are more concealed and less visible. Pitangus sulphuratus is an occasional predator on bird nests (Sick 1985), but the species was not seen preying on the contents of these Passer nests.

On 27 November 1995 a third nest, practically identical to the previous nests, was found in the same niche, but no one individual of Pitangus sulphuratus was seen then. The 1993 nest had been removed by maintenance workers in 1994. Several references of nests of Pitangus sulphuratus built inside cavities (sensu lato, including niches, notches, crevices, rolled barks, domed nests of other birds, spaces under massive plant parts, hollow fallen logs, rock chaps, mammal burrows, cliff holes and other concealed spaces) exist in the literature. Zuberbhuler (1971) described an unfinished nest of Pitangus sulphuratus, discovered in Argentina, built into a nest of Anumbius annumbi (Furnariidae). Anumbius builds voluminous closed nests of large twigs provided with an entrance tunnel and a central chamber (Sick 1985).

Haverschmidt (1974) reported a nest of Pitangus sulphuratus built in an old woodpecker (Picidae) hole. Belton (1985) described a nest of Pitangus sulphuratus built in an abandoned oven nest of Furnarius rufus (Furnariidae), in which the side wall separating the nesting chamber from the entrance passage had been destroyed so the chamber opened directly to the exterior.

La Peña (1987) described nests of Pitangus sulphuratus placed in a cavity on the cement base of a bridge, and in a semi-destroyed oven nest of Furnarius rufus (Furnariidae). Smith (1962) and Sick (1985) asserted that Pitangus sulphuratus may build a cup nest when in a concealed situation, like under the broad leaf sheath of a palm. J. W. Fitzpatrick observed Pitangus sulphuratus building a cup nest in dense foliage over water, in southeastern Peru (pers. comm., in lit.).

Pitangus sulphuratus has been seen building a perfectly normal open cup, which it then gradually covered over during the first two weeks of incubation (Fitzpatrick, unpubl. obs., in Traylor and Fitzpatrick 1982). Therefore, the distinction between cup and ball nests appears not to be especially rigid in this species, suggesting an evolution pathway from one to the other.

The two species formerly ascribed to the genus Pitangus have nests of divergent structure. Pitangus lictor builds a cup nest (Smith 1962,, Willis 1962, Haverschmidt 1968) so utterly divergent from that of Pitangus sulphuratus that Haverschmidt (1957) wondered if these two species are really so closely related as to warrant their inclusion in the same genus. Due to this nest divergence and some syringeal traits, Lanyon (1984) created the genus Philohydor for Pitangus lictor.

Cavity nesting is refered in literature for some close relatives of Pitangus sulphuratus. Myiozetetes similis, a near relative of Pitangus, occasionally places its bulky grass nest (usually round and exposed) inside large holes, which is suggestive of an evolutionary pathway toward hole nesting (Traylor and Fitzpatrick 1982).

Mousley (1916) and Cottam (1938) related nesting by Tyrannus tyrannus in an old nest of Icterus galbula (Icteridae). H. L. Harlee (apud Bent 1942) observed that Tyrannus tyrannus built a nest and laid a set of eggs in a gourd that was suspended for Progne subis breeding. Brewster (1937) observed that the species nested mostly within hollow tree-trunk over water.

Kennedy (1915) observed that Tyrannus verticalis nested inside old nests of Icterus galbula. Munro (1919) found these birds using frequently abandoned holes carved by Colaptes auratus (Picidae). Bent (1942) stated that where no suitable trees were available, Tyrannus verticalis selected nest boxes for nesting.

Hunt (1964) observed Tyrannus melancholicus in Barro Colorado Island, Panama, incubating two eggs in a partially domed nest, possibly built by Myiozetetes sp.

Several obligatory or facultative cavity nesters are found in the following genera of Tyrannidae: Tityra, Xolmis, Sayornis, Colonia, Fluvicola, Machetornis, Myiodynastes, Conopias, Myiozetetes, Tyrannus, Attila, Deltarhynchus, Pyrrhomyias, Ramphotrigon, Rhytipterna, Sirystes, Casiornis and Myiarchus (Euler 1900, von Ihering 1904, Bent 1942, Tyler 1942, Haverschmidt 1957, Skutch 1960, Traylor and Fitzpatrick 1982, Lanyon and Fitzpatrick 1983, Parker 1984, Lanyon 1985, Sick 1985, Hilty and Brown 1986, Tostain 1989). As summarized here, Pitangus is a documented member of this list. Sick (1985) also found cavity nesting tendencies (or behavior vestiges) in the genera Serpophaga, Knipolegus and Hirundinea.

Myiodynastes and Conopias are genera of obligatory cavity nesters in the Pitangus (or Tyrannus) assemblage. This assemblage comprises twelve genera and 33 recognized species, according to Traylor (1977), Traylor and Fitzpatrick (1982) and Lanyon (1984): Pitangus (1 sp.), Philohydor (1 sp.), Megarhynchus (1 sp.), Myiozetetes (4 spp.), Conopias (3 spp.), Phelpsia (1 sp.), Myiodinastes (5 spp.), Legatus (1 sp.), Empidonomus (1 spp.), Griseotyrannus (1 spp.), Tyrannopsis (1 sp.) and Tyrannus (13 spp.).

Lanyon (1985) suggested that hole-nesting behavior in the several subfamilies of Tyrannidae can be attributed to convergent or parallel evolution.

I hypothesize that the habit of nesting in cavities is a primitive character in certain branches of the Tyrannidae. Some assemblages within the family (notably the Tyrannus relatives) maintain cavity nesting behavior, also showing an evolution for building exposed, open nests and, as a refinement, domed nests.

Cavity nesting must have appeared early in the Tyrannus group, being a primitive (generalized) character because the cavity nesters need only build a mere cup nest inside the cavity chosen. The evolution for building exposed, domed nests would ensure that some advantages of a cavity concealment of eggs and young, stable attachment, protection from rain and winds, escape from predators) would be retained. The independence from having to use cavities would bring the advantage of not needing to search and defend the cavities, a resource normally rare and intensively exploited by many birds, mammals, arthropods, other animals (Collias and Collias 1984; McComb and Noble 1981) and epiphytic plants, as Ficus spp. (Moraceae), Clusia spp. (Guttiferae), Hippeastrum spp. (Amaryllidaceae), Gesneriaceae, Bromeliaceae and many others (pers. obs.). The evolution of aggressive habits among the Tyranninae might be related to building and maintaining nests in very exposed situations, as once suggested by von Ihering (1904) for the larger species of Tyranninae.

The fact that several species normally building closed nests (Pitangus sulphuratus and Myiozetetes spp.) occasionally return to nest in cavities suggests that the domed nest evolved as a substitute for a cavity. Legatus evolved parasitic use of closed nests (Skutch 1960, Sick 1985), and this may be a sign of earlier building of closed nests or of cavity nests.

Occasional or obligatory cavity nesting is found in several branches of the phylogenetic tree proposed by Lanyon (1984) for the Tyranninae; whether it be a derived character state, the habit would must concentrate in some branches having a common trunk. That ubiquity in distribution of the character state forces it to the basis of the tree, being useless for clustering genera in the assemblage as any generalized (therefore primitive) state.

The genetic background for cavity nesting may be still present and unexpressed in some groups of Tyrannidae, but ready to be expressed occasionally by some individuals or populations in species that normally build cup or domed nests. I hypothesize this to be the case for Pitangus sulphuratus and Myiozetetes similis.

Under this evolutionary scenario, the occasional use of cavities by these two species, including typical woodpecker holes, may be looked at not as an opportunistic behavior by an exposed-nester but as an expression of ancestral behavior by some individuals.

I propose the latin compound term arbocavicola (arbocavicolous, in English) as a technical and more concise substitute for “tree-cavity nesting”, “tree-hole nesting” or “tree-cavity using” animals, following the rules for scientific neologisms (Brown 1985). The complete neologism would be arbocavinidicola, regarding to the use of a tree cavity for breeding. General neologisms are cavicola, related to the use of any cavities and cavinidicola, regarding to the use of any cavities for breeding.


Acknowledgments. I wish to thank Keith S. Brown, Jr. and an anonymous reviewer, for manuscript revision and valuable comments, Edwin O. Willis, Luiz G. E. Lordello and Telma E. Candido for providing access to literature.



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