Mixed-species foraging flocks

A mixed-species feeding flock, also termed a mixed-species foraging flock, mixed hunting party or informally bird wave, is a flock of usually insectivorous birds of different species, that join each other and move together while foraging. These are different from feeding aggregations, which are congregations of several species of bird at areas of locally high food availability.

A mixed-species foraging flock typically has "nuclear" species that appear to be central to its formation and movement. Species that trail them are termed "attendants". Attendants tend to join the foraging flock only when the flock enters their territory.

How such flocks are initiated is under investigation. But in Sri Lanka for example, vocal mimicry by the Greater Racket-tailed Drongo (Dicrurus paradiseus) might have a key role in the initiation of mixed-species foraging flocks, while in parts of the American tropics noisy packs of foraging Golden-crowned Warblers (Basileuterus culicivorus) might play the same role. Forest structure is also believed to be an important factor deciding the propensity to form flocks. In tropical forests, birds that glean food from foliage were the most abundant species in mixed-species flocks.

A typical Neotropic mixed feeding flock moves through the forest at about 0.3 kilometers per hour (0.19 miles per hour), with different species foraging in their preferred niches (on the ground, on trunks, in high or low foliage, etc.). Some species follow the flock all day, while others – such as the Long-billed Gnatwren (Ramphocaenus melanurus) – join it only as long as it crosses their own territories.

Costs and benefits
Several evolutionary mechanisms have been proposed to explain the formation of mixed-species flocks. These are usually described in terms of the costs and benefits to individuals. The key benefits that have been suggested are a reduction in predation risk through increased vigilance, that is, more eyes that can spot predators and raise an alarm and increased foraging efficiency. Costs could include the risk of kleptoparasitism.

In the Holarctic
In the North Temperate Zone, they are typically led by Paridae (tits and chickadees), often joined by nuthatches (Sitta), treecreepers (Certhia), woodpeckers (Picidae, such as the Downy Woodpecker "Picoides" pubescens and Lesser Spotted Woodpecker "P." minor), kinglets (Regulus), and in North America Parulidae (New World "warblers") – all insect-eating birds. This behavior is particularly common outside the breeding season.

The advantages of this behavior are not certain, but evidence suggests that it confers some safety from predators, especially for the less watchful birds such as Vireonidae (vireos) and woodpeckers, and also improves feeding efficiency, perhaps because arthropod prey that flee one bird may be caught by another.

In the Neotropics
Insectivorous feeding flocks reach their fullest development in tropical forests, where they are a typical feature of bird life. In the Neotropics the leaders or "core" members may be Black-throated Shrike-tanagers (Lanio aurantius) in southern Mexico, or Three-striped Warblers (Basileuterus tristriatus) elsewhere in Central America. In South America, core species may include antbirds (Thamnophilidae) such as Thamnomanes, antshrikes, Furnariidae (ovenbirds and woodcreepers) like the Buff-fronted Foliage-gleaner (Philydor rufum) or the Olivaceous Woodcreeper (Sittasomus griseicapillus), or Parulidae (New World "warblers") like the Golden-crowned Warblers (Basileuterus culicivorus). In open cerrado habitat, it may be White-rumped (Cypsnagra hirundinacea) or White-banded Tanagers (Neothraupis fasciata). Core species often have striking plumage and calls that attract other birds; they are often also known to be very active sentinels, providing warning of would-be predators.

But while such easy-to-locate bird species serve as a focal point for flock members, they do not necessarily initiate the flock. In one Neotropic mixed flock feeding on swarming termites, it was observed that Red-rumped Warbling-finches (Poospiza lateralis) were most conspicuous. As this species is not an aerial insectivore, it is unlikely to have actually initiated the flock rather than happening across it and joining in. And while Basileuterus species are initiators as well as core species, mixed flocks of Tangara species – in particular Red-necked (T. cyanocephala), Brassy-breasted (T. desmaresti) and Green-headed Tanager (T. seledon) – often initiate formation of a larger and more diverse feeding flock, of which they are then only a less significant component.

Nine-primaried oscines make up much of almost every Neotropical mixed-species feeding flock. Namely, these birds are from families such as the Cardinalidae (cardinals), Parulidae (New World "warblers"), and in particular Emberizidae (American "sparrows") and Thraupidae (tanagers). Other members of a Neotropic mixed feeding flock may come from most of the local families of smaller diurnal insectivorous birds, and can also include woodpeckers (Picidae), toucans (Ramphastidae) and trogons (Trogonidae). Most Furnariidae do not participate in mixed flocks very often, though there are exceptions such as Synallaxis spinetails and some species of the woodcreeper subfamily (Dendrocolaptinae) – e.g. those mentioned above or the Lesser Woodcreeper (Xiphorhynchus fuscus) – are common or even "core" members. Among the tyrant flycatchers (Tyrannidae) there are also some species joining mixed flocks on a somewhat regular basis, including the Sepia-capped Flycatcher (Leptopogon amaurocephalus), Eared Pygmy-tyrant (Myiornis auricularis), White-throated Spadebill (Platyrinchus mystaceus) and Oustalet's Tyrannulet (Phylloscartes oustaleti).

However, even of commonly-participating families not all species join mixed flocks. There are genera such as Vireo in which some species do not join mixed flocks, while others (e.g. Red-eyed Vireo, V. olivaceus) will even do so in their winter quarters. Of the three subspecies groups of the Yellow-rumped Warbler (Dendroica coronata), only one (Audubon's Warbler, D. c. auduboni group) typically does. And while the importance of certain Thraupidae in initiating and keeping together mixed flocks has been mentioned already, for example the Black-goggled Tanager (Trichothraupis melanops) is an opportunistic feeder that will appear at but keep its distance from any disturbance – be it a mixed feeding flock, an army ant column or a group of monkeys – and pick off prey trying to flee.

Conopophagidae (gnateaters) are notable for their absence from these flocks, while Apodidae (swifts) and Hirundinidae (swallows) rarely join them, but will if there is for example an ant or termite swarm. Cotingidae (cotingas) are mainly opportunistic associates which rarely join flocks for long if they do so at all; the same holds true for most Muscicapoidea (mockingbirds and relatives), though some thrushes (Turdidae) may participate on more often. And though most Tityridae rarely join mixed flocks, becards (Pachyramphus) do so regularly. Tapaculos (Rhinocryptidae) are rarely seen with mixed flocks, though the Collared Crescentchest (Melanopareia torquata), doubtfully assigned to that family, may be a regular member. Icteridae (grackles and relatives) are also not too often seen to take part in these assemblages, though caciques like the Golden-winged (Cacicus chrysopterus) or Red-rumped Cacique (C. haemorrhous) join mixed flocks on a somewhat more regular basis. Cuculiformes (cuckoos and allies) are usually absent from mixed feeding flocks, but some – e.g. the Squirrel Cuckoo (Piaya cayana) can be encountered not infrequently.

Some species appear to prefer when certain others are present: Cyanolyca jays like to flock with Unicolored Jays (Aphelocoma unicolor) and the Emerald Toucanets (Aulacorhynchus prasinus species complex). Many Icteridae associate only with related species, but the western subspecies of the Yellow-backed Oriole (Icterus chrysater) associates with jays and the Band-backed Wren (Campylorhynchus zonatus).

Other species participate to varying extents depending on location or altitude – presumably, the different species composition of mixed flocks at varying locations allows these irregular members more or less opportunity to get food. Such species include the Grey-hooded Flycatcher (Mionectes rufiventris: Tyrannidae), or the Plain Antvireo (Dysithamnus mentalis: Thamnophilidae) and the Red-crowned Habia (Habia rubica: Cardinalidae ) which are often recorded in lowland flocks but rarely join them at least in some more montane regions.

In the Old World tropics
The flocks in the Old World are often much more loosely bonded than in the Neotropics, many being only casual associations lasting the time the flock of core species spends in the attendants' territory. The more stable flocks are observed in tropical Asia, and especially Sri Lanka. Flocks there may number several hundred birds spending the entire day together, and an observer in the rain forest may see virtually no birds except when encountering a flock. For example, as a flock approaches in the Sinharaja Forest Reserve in Sri Lanka, the typical daytime quiet of the jungle is broken by the noisy calls of the Orange-billed Babbler (Turdoides rufescens) and Greater Racket-tailed Drongo (Dicrurus paradiseus), joined by species such as the Ashy-headed Laughingthrush (Garrulax cinereifrons), Kashmir Flycatcher (Ficedula subrubra) and Velvet-fronted Nuthatch (Sitta frontalis).

A mixed flock in the Cordillera Central of Luzon in the Philippines was mainly composed of Bar-bellied Cuckoo-shrikes (Coracina striata: Campephagidae), Philippine Fairy-bluebirds (Irena cyanogaster: Irenidae) and Violaceous Crows (Corvus (enca) violaceus: Corvidae). Luzon Tarictic Hornbills (Penelopides manillae: Bucerotidae) were also recorded as present. With the crows only joining later and the large hornbills probably only opportunistic attendants rather than core species, it is likely that this flock was started by one of the former species – probably the bold and vocal cuckoo-shrikes rather than the more retiring fairy-bluebirds, which are known to seek out such opportunities to forage.

African rainforests also hold mixed-species flocks, the core species including Pycnonotidae (bulbuls) and Nectarinidae (sunbirds), and attendants being as diverse as the Red-billed Dwarf Hornbill (Tockus camurus) and the Tit-hylia (Pholidornis rushiae), the smallest bird of Africa. Dicruridae (drongos) and Monarchidae (paradise-flycatchers) are sometimes described as the sentinels of the flock, but they are also known to steal prey from other flock members. Acanthizidae are typical core members in New Guinea (Gerygone) and Australia (Acanthiza); in Australia, fairy-wrens (Malurus) are also significant. The core species are joined by birds of other families such as minivets (Pericrocotus).