Laughingthrush
Red-tailed laughingthrush
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Order:	Passeriformes
Superfamily:	Sylvioidea
Family:	Leiothrichidae Swainson, 1832
Genera
17; see article text

The laughingthrushes and their allies are a family, Leiothrichidae, of Old World passerine birds. The entire family used to be included in the Old World babbler family Timaliidae. The family contains 143 species and is divided into 17 genera. The species are diverse in size and coloration. These are birds of tropical areas, with the greatest variety in Southeast Asia and the Indian subcontinent.

They are small to medium-sized birds. They have strong legs, and many are quite terrestrial. They typically have generalised bills, similar to those of a thrush. Most have predominantly brown plumage, with minimal difference between the sexes, but many more brightly coloured species also exist.^[1]

This group is not strongly migratory, and most species have short rounded wings, and a weak flight. They live in lightly wooded or scrubland environments, ranging from swamp to near-desert. They are primarily insectivorous, although many will also take berries, and the larger species will even eat small lizards and other vertebrates.^[1]

The family Leiothrichidae was introduced (as a subfamily Leiotrichanae) by the English naturalist William Swainson in 1832.^[2] However, for much of the twentiest century, the genera that now comprise the family were all placed in the large Old World babbler family Timaliidae. The family was a notorious wastebasket taxon into which Old World insectivores were placed when they could not be placed anywhere else. The French ornithologist Jean Théodore Delacour developed the most influential revision of the family, published in 1946 and 1950. His review, based on plumage colouration and bill shape, identified five tribes with the Timaliidae, of which Turdoidini contained many of the genera now assigned to Leiothrichidae, including Garrulax, Liocichla, Turdoides.^[3] Other genera assigned to the tribe are now assigned to other babbler families, such as Yuhina, now with the Zosteropidae, or even less closely related, such as Pteruthius, now known to be a vireo. The next major revision was the Sibley–Ahlquist taxonomy of birds developed using DNA–DNA hybridization. This study removed some groups from the wider Timaliidae, and found the laughingthrushes in the genus Garrulax to be the sister taxon to the rest of the family. This work only examined 11 species and 9 genera.^[3]

A comprehensive molecular phylogenetic study of the family published in 2018 led to substantial revision of the taxonomic classification.^[4] The laughingthrushes in the genus Garrulax were found to belong to three separate clades that had diverged in the Miocene 7–9 million year ago. The genus was therefore split with Garrulax restricted to one clade and the genera Pterorhinus and Ianthocincla resurrected for the other two clades. The genus Turdoides was also split and species moved into the resurrected genus Argya.^[4][5]

In a separate change, the crocias were moved to the genus Laniellus Swainson, 1832 which has priority over Crocias Temminck, 1836.^[5][6][7]

The cladogram below showing the phylogenetic relationship of the Leiothrichidae to other families is based on a study of the babblers by Tianlong Cai and collaborators published in 2019.^[8][9]

The cladogram below shows the phylogenetic relationships between the genera in the family Leiothrichidae based on a study by Alice Cibois and collaborators published in 2018.^[4]

The laughingthrushes and allies are found in the tropics of Africa, the Middle East, through to South Asia, Southeast Asia and Eastern Asia, with some species reaching temperate climates in East Asia. They are predominantly woodland and forest birds, but some species are found in scrub and savanna, particularly birds in the genus Argya and Turdoides.^[10] They reach their greatest diversity in the Himalayas and South East Asia.^[4]

Like the other families of birds that are treated as Old World babblers, the Leiothrichidae do not undertake long distance migration, although some species do undertake shorter or irregular migrations. Some African species in the genus Turdoides will move in response to rains or drought. Several montane species display altitudinal migration, including the red-tailed milna, blue-winged minla, streaked barwing and red-billed leiothrix. The latter will migrates down from higher forest in the Himalayas in November before returning in late April.^[11]

Some species of the family have been introduced outside their native range. For example, the Chinese hwamei, a popular cagebird, has been accidentally introduced to several countries and states including Japan, where it is expanding its range. It was was even deliberately introduced to California, although that introduction failed.^[12] The red-billed leiothrix has been widely introduced across Europe, Hawaii, Reunion Island and Japan, and is considered a pest due to its tendency to spread non-native plants, compete with local species and acting as a reservoir for avian malaria.^[13]

The laughingthrushes and their allies are small to medium sized songbirds that are generally larger and longer-tailed than species in the other babbler families (Pellorneidae and Timaliidae). They range in size from the largest, the greater necklaced laughingthrush, which can be up to 34 cm (13 in) in length and weigh up to 170 g (6.0 oz),^[14] to the fulvettas in the genus Alcippe like the Nepal fulvetta, which is only 12.5 cm (4.9 in) in length and weighs 13–18 g (0.46–0.63 oz).^[15]

Plumage varies widely across the family, with forest species ranging from black and white species to extremely colourful species to drabber species. Drabness is the rule for scrub or savanna species, particularly those in Africa.^[10] Many of the laughingthrushes in the genus Trochalopteron, Garrulax, and Ianthocincla are known for bright markings on their plumage which are counterintuitively difficult to see in the dark overgrown habitats they frequent unless at close range.^[16] Mimicry has been observed in the family, with lesser necklaced laughingthrushes mimicing the plumage of the greater necklaced laughingthrush, which is in a different genus, in order to allow them to forage closely together.^[17]

The laughingthrushes and allies, like other babbler families, do not generally exhibit much sexual dimorphism beyond males sometimes averaging slightly larger than females in some species. A few genera do exhibit differences in plumage between the sexes, such as Leiothrix, Minla and Cutia.^[18] However, some of the differences are subtle. For example a study examining the reflectance spectrometry of two laughingthrush species with no apparent differences in plumage between the sexes, Elliot's laughingthrush and plain laughingthrush, found that in both species the males had brighter plumage and the females, possibly due to sexual selection or to make the female less conspicuous to predators or brood parasites.^[19]

The diet of the laughingthrushes and their allies is dominated by invertebrates, mostly insects and other arthropods. Some species take fruit, nectar and seeds, and larger species will take small vertebrates.^[10] Among the insectivores there are several guilds that specialise in certain feeding in certain ways. Smaller arboreal species glean insects from leaves, twigs and branches, including Alcippe, Leiothrix, Minla and the crocias in Laniellus. The barwings in Actinodura and the two cutias also glean insects from epiphytes. Other guilds represented by the laughingthrushes include terrestrial hunters, like Ianthocincla and Trochalopteron, which hunt on the ground and in the undergrowth, and generalists that take prey via gleaning from leaves and branches and the ground.^[20]

The laughingthrushes and allies are highly social species, so it is no surprised that much feeding is undertaken in family groups or small flocks. Many species associate with mixed-species feeding flocks.^[21] Members of the family can even act as the nucleus species for such flocks, such as the Nepal fulvetta.^[22] These associations are not without cost; greater racket-tailed drongos that participate in flocks with greater and lesser necklaced laughingthrushes in Myanmar will pursue or attack the laughingthrushes in order to steal prey items they have located. In spite of this the laughingthrushes still participate in the feeding-flocks, presumably because they receive a benefit from the drongo also acting as a sentinel for the flock, alerting them to predators.^[23]

The family contains 143 species in 17 genera:^[9]

• Alcippe – fulvettas (10 species)
• Grammatoptila – striated laughingthrush
• Cutia – cutias (2 species)
• Laniellus – crocias (2 species)
• Trochalopteron – laughingthrushes (19 species)
• Actinodura – barwings and minlas (9 species)
• Montecincla – laughingthrushes (4 species)
• Minla – red-tailed minla
• Leioptila – rufous-backed sibia
• Leiothrix – (2 species)
• Liocichla – liocichlas (5 species)
• Heterophasia – sibias (7 species)
• Argya – mainly babblers (16 species); previous placed in Turdoides
• Turdoides – babblers (19 species)
• Garrulax – laughingthrushes (14 species)
• Ianthocincla – laughingthrushes (8 species); previously placed in Garrulax
• Pterorhinus – laughingthrushes and babaxes (23 species); previously placed in Garrulax

• Collar, Nigel; Robson, Craig (2007), "Family Timaliidae (Babblers)", in del Hoyo, Josep; Elliott, Andrew; Christie, David (eds.), Handbook of the Birds of the World. Volume 12, Picathartes to Tits and Chickadees, Barcelona: Lynx Edicions, pp. 70–140, ISBN 978-84-96553-42-2