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The Enigmatic Rallidae

Exploring the diverse morphology, unique behaviors, and evolutionary paradoxes of this cosmopolitan avian family.

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Overview

A Cosmopolitan Avian Family

The Rallidae family, commonly known as rails, encompasses a large and globally distributed group of small- to medium-sized birds. This diverse family includes well-known species such as crakes, coots, and gallinules, alongside many rarer or endangered forms. Their presence spans nearly every terrestrial habitat, excluding only arid deserts, polar or freezing zones, and high alpine regions above the snow line. Members of Rallidae are found on every continent except Antarctica, with numerous unique island species highlighting their remarkable adaptability.[2]

Preferred Habitats

Many rail species are intrinsically linked to wetland environments, with some exhibiting semi-aquatic tendencies akin to waterfowl, such as the coots. However, a significant number are wading birds or shorebirds. Their ideal habitats typically consist of marsh areas, including rice paddies, flooded fields, or open forests. These birds show a particular preference for dense vegetation, which provides crucial cover for nesting and protection.[2]

Etymological Roots

The common name "rail" is an anglicized rendition of the Old French word "râle," which itself originates from the Vulgar Latin "*rascula." This term is derived from the Latin "rādere," meaning "to scrape," a reference to the characteristic harsh cry emitted by these birds.[3]

Morphology

Size and Form Variation

Rails are characterized as ground-dwelling birds, exhibiting a considerable range in size. Their lengths can vary from a diminutive 12 cm (5 inches) to a substantial 63 cm (25 inches), with weights spanning from 20 grams (0.7 oz) to an impressive 3,000 grams (6 lb 10 oz). Many species possess elongated necks and are often noted for their laterally compressed bodies, a feature that aids their movement through dense vegetation.[4]

Bill Diversity

The bill is arguably the most variable morphological trait within the Rallidae family. Some species, such as the clapper rail of the Americas, display bills longer than their heads. In contrast, coots feature short and broad bills, while purple gallinules possess massive ones.[5] A distinctive feature in some coots and gallinules is a frontal shield, a fleshy extension of the upper bill that extends rearward. The horned coot, for instance, exhibits the most intricate form of this frontal shield.[6]

Sexual Dimorphism

Generally, rails exhibit minimal sexual dimorphism, meaning there are few discernible differences between males and females in terms of plumage or size. However, notable exceptions to this pattern include the watercock (Gallicrex cinerea) and the little crake (Zapornia parva), where sexual differences are more pronounced.[7]

Flight & Flightlessness

Avian Locomotion

The wings of all rail species are characteristically short and rounded. While the flight of volant (flying) Rallidae members may not be exceptionally powerful, it can be sustained over extended periods, enabling many species to undertake annual migrations. However, the inherent weakness of their flight makes them susceptible to being blown off course, a factor that has paradoxically facilitated their colonization of numerous isolated oceanic islands. Furthermore, these birds frequently opt to run rather than fly, particularly within dense habitats. Some species also experience periods of temporary flightlessness during their moult cycles.[8]

Parallel Evolution of Flightlessness

Flightlessness within the Rallidae family stands as a prime example of parallel evolution in the animal kingdom. Out of approximately 150 historically recognized rail species, 31 extant or recently extinct species independently evolved flightlessness from flying ancestors.[9] This evolutionary trajectory has given rise to the unique populations of flightless rails observed on Pacific islands today.

  • Absence of Predators: Small island ecosystems often lack mammalian predators, diminishing the selective pressure for flight as an escape mechanism.[9]
  • Energy Conservation: Flight is metabolically demanding, with the keel and flight muscles constituting up to 40% of a bird's body weight. Reducing these muscles and their associated metabolic requirements allows flightless rails to conserve energy, which is advantageous in resource-limited island environments.[9][10]
  • Morphological Predisposition: Rails inherently possess relatively small flight muscles and wings, which typically account for only 12–17% of their total body mass. This, combined with their terrestrial habits and behavioral reluctance to fly, contributes significantly to their remarkably rapid loss of flight.[9][13]
  • Climate Stability: Flightlessness is more prevalent on tropical islands, where a stable climate obviates the need for seasonal long-distance migrations, further reducing the selective pressure for flight.[16]

The Laysan rail, for example, is estimated to have lost the power of flight and developed reduced, stubby wings (useful only for balance during rapid running) in as little as 125,000 years.[15] Some researchers even suggest that the evolution of flightlessness in rails could be measured in generations rather than millennia.[11]

Dispersal Paradox and Social Evolution

Despite their apparent aversion to flight, the evolution of flightless rails on isolated islands necessitates a high capacity for long-distance dispersal by their volant ancestors.[13] Certain small-massed rail species, such as Gallirallus philippensis, Porphyrio porphyrio, and Porzana tabuensis, demonstrate a consistently high ability to disperse across vast distances among tropical Pacific islands. Interestingly, only the latter two gave rise to endemic flightless species throughout the Pacific Basin.[17] Research into the phylogeny of G. philippensis indicates that the flightless condition evolved in rails even before speciation was complete, highlighting a complex evolutionary pathway.[17]

A consequence of the reduced energy expenditure in flightless island rails has been linked to the evolution of increased "tolerance" and "approachability" in these species.[16] This shift in behavior, potentially leading to kin-selecting altruistic phenomena, reallocates resources to produce fewer, more competitive offspring, benefiting the overall population. However, with human colonization of most islands over the past 5,000 to 35,000 years, this tolerance has unfortunately reversed into a heightened wariness of humans and introduced predators, rendering many species vulnerable to extinction.[16]

Behavior & Ecology

Diet and Foraging

Members of the Rallidae family are generally omnivorous generalists, displaying a flexible diet. Many species consume a variety of invertebrates, alongside fruits and seedlings. A smaller subset of species are primarily herbivorous, adapting their diets to the available plant matter in their specific habitats.[2]

Vocalizations

The calls of Rallidae species are highly varied and frequently quite loud. Some vocalizations are whistle-like or squeak-like, while others can sound distinctly unbirdlike.[20] These loud calls serve a crucial function in dense vegetation or during nocturnal activity, where visual contact with other members of the species is challenging. Certain calls are also employed for territorial defense and communication.[5]

Terrestrial Habits and Habitat Preference

The most characteristic members of this family inhabit dense vegetation within damp environments, typically near lakes, swamps, or rivers. Reed beds are a particularly favored habitat. Species that undertake migration do so under the cover of night. Most rails construct their nests within dense vegetation, contributing to their generally shy, secretive nature, which makes them challenging to observe in the wild. They are adept at walking and running vigorously on their strong legs, which are equipped with long toes well-suited for navigating soft, uneven surfaces.[8]

Size and Naming Conventions

Reedbed species are often secretive (aside from their loud calls), crepuscular, and possess laterally flattened bodies. In the Old World, long-billed species are typically referred to as rails, while short-billed species are called crakes. In North America, however, species are generally termed rails irrespective of their bill length. The smallest known rail is Swinhoe's rail, measuring 13 cm (5.1 inches) and weighing 25 grams. Larger species may also be known by other names, such as coots (which are more adapted to open water) and gallinules or swamphens. The largest member of this group is the takahē, reaching 65 cm (26 inches) in length and weighing 2.7 kg (6.0 lb).

Reproduction

Breeding Systems

The breeding behaviors of many Rallidae species remain incompletely understood or entirely unknown. While most are presumed to be monogamous, instances of polygyny (one male mating with multiple females) and polyandry (one female mating with multiple males) have also been documented within the family.[24]

Clutch Size and Development

Typically, rails lay clutches consisting of five to ten eggs. However, clutch sizes can vary significantly, with records ranging from as few as one egg to as many as fifteen.[24] It is not uncommon for egg clutches to hatch asynchronously. Chicks achieve mobility within a few days of hatching, but generally remain dependent on their parents until they fledge, a developmental stage usually reached around one month of age.[6]

Rallidae & Humans

Human Interaction and Exploitation

Some of the larger and more abundant rail species have historically been hunted for food, and their eggs collected. A tragic example is the Wake Island rail, which was hunted to extinction by the starving Japanese garrison after the island's supply lines were severed during World War II.[26] Furthermore, certain species, such as the common moorhen and the American purple gallinule, have occasionally been regarded as agricultural pests.[25]

Threats and Conservation Challenges

Rails have disproportionately suffered from human-induced environmental changes. Their inherent tendency towards flightlessness has rendered many island species particularly vulnerable to introduced predators. The most severe human-caused extinctions occurred in the Pacific Ocean as people colonized the islands of Melanesia, Polynesia, and Micronesia. During this period, an estimated 750–1800 bird species became extinct, with approximately half of these being rails.[27]

While many island rail species remain endangered, conservation organizations and governments are actively working to prevent further extinctions. Notable examples include:

  • The Lord Howe woodhen and the takahē, which have shown modest recoveries due to dedicated conservation efforts.
  • The Guam rail faced imminent extinction following the introduction of brown tree snakes to Guam. However, some of the last remaining individuals were brought into captivity and have successfully bred. Although reintroduction attempts have yielded mixed results, these programs offer a glimmer of hope for the species.[28][29][30]
Critical Concern: The introduction of terrestrial predators such as cats, foxes, weasels, mongooses, rats, and pigs has been a primary driver of extinction for many flightless island rail species.

Systematics & Evolution

Taxonomic History

The Rallidae family was formally introduced (as Rallia) by the French polymath Constantine Samuel Rafinesque in 1815.[31][32] Traditionally, this family has been grouped within the order Gruiformes, alongside larger birds like cranes and bustards, as well as several smaller families of typically "primitive" midsized amphibious birds. The alternative Sibley-Ahlquist taxonomy, widely accepted in America, elevates the family to an ordinal level as the Ralliformes. Given the ongoing uncertainties regarding gruiform monophyly, this reclassification may be justified, potentially including the Heliornithidae (finfoots and sungrebes), a tropical group exhibiting convergent evolution with grebes, often united with rails in the Ralli.[7]

Phylogenetic Relationships

Recent phylogenomic reconstructions, based on ultra-conserved elements, have significantly clarified the evolutionary relationships and timescale of rails and their allies (Ralloidea). These studies have provided the foundation for a revised classification of the Rallidae family.[1] The family is broadly divided into several subfamilies and tribes, reflecting distinct evolutionary lineages. For instance, the Rallinae subfamily includes tribes such as Pardirallini and Rallini, while the Himantornithinae subfamily encompasses Himantornithini, Fulicini, Porphyrionini, Laterallini, Zapornini, and Amaurornithini.[1][7][33]

The Rallidae family is broadly structured into two main subfamilies, each containing several tribes:

Rallinae Subfamily

  • Pardirallini: Includes genera like Anurolimnas (chestnut-headed crake), Mustelirallus (crakes), Pardirallus (rails), Amaurolimnas (uniform crake), and Aramides (rails).
  • Rallini: Comprises genera such as Rallus (typical rails), Crecopsis (African crake), Rougetius (Rouget's rail), Dryolimnas (rails), Crex (corn crake), Lewinia (rails), Aptenorallus (Calayan rail), Habroptila (invisible rail), Gallirallus (weka), Eulabeornis (chestnut rail), and Hypotaenidia (Austropacific rails and woodhen).

Himantornithinae Subfamily

  • Himantornithini: Contains the genus Himantornis (Nkulengu rail).
  • Fulicini: Includes genera like Porzana (crakes and sora), Tribonyx (nativehens), Paragallinula (lesser moorhen), Gallinula (moorhens), and Fulica (coots).
  • Porphyrionini: Features the genus Porphyrio (swamphens, gallinules, and takahes).
  • Laterallini: Encompasses genera such as Rufirallus (crakes), Coturnicops (rails), and Laterallus (crakes and rails).
  • Zapornini: Includes Zapornia (rails and crakes) and Rallina (crakes).
  • Amaurornithini: Contains genera like Megacrex (New Guinea flightless rail), Poliolimnas (white-browed crake), Aenigmatolimnas (striped crake), Gallicrex (watercock), and Amaurornis (waterhen and bush-hens).

Extant Genera

Living Diversity

The International Ornithological Committee (IOC) currently recognizes 153 extant species distributed across 43 genera within the Rallidae family.[34] This remarkable diversity highlights the family's evolutionary success and adaptability to a wide array of ecological niches.

  • Canirallus – grey-throated rail
  • Anurolimnas – chestnut-headed crake
  • Mustelirallus – (4 species)
  • Pardirallus (3 species)
  • Amaurolimnas – uniform crake
  • Aramides – wood rails (8 species)
  • Rallus – typical rails (14 species)
  • Crecopsis – African crake
  • Rougetius – Rouget's rail
  • Dryolimnas – (1 living species, 1 recently extinct)
  • Crex – corn crake
  • Aramidopsis – snoring rail
  • Lewinia – (4 species)
  • Aptenorallus – Calayan rail
  • Habroptila – invisible rail
  • Gallirallus – weka
  • Eulabeornis – chestnut rail
  • Cabalus – (1 possibly extinct species, 1 recently extinct)
  • Hypotaenidia – Austropacific rails (8 living species, 4 recently extinct)
  • Porphyriops – spot-flanked gallinule
  • Porzana – (3 species)
  • Tribonyx – nativehens (2 species)
  • Paragallinula – lesser moorhen
  • Gallinula – moorhens (5 living species, 2 recently extinct)
  • Fulica – coots (10 living species, one recently extinct)
  • Porphyrio – swamphens and purple gallinules (10 living species, 2 recently extinct)
  • Rufirallus – (5 species)
  • Coturnicops – (2 species)
  • Laterallus – (11 species)
  • Zapornia – (10 living species, 5 recently extinct)
  • Rallina – (4 species)
  • Gymnocrex – (3 species)
  • Himantornis – Nkulengu rail
  • Megacrex – New Guinea flightless rail
  • Poliolimnas – white-browed crake
  • Aenigmatolimnas – striped crake
  • Gallicrex – watercock
  • Amaurornis – bush-hens (5 species)

Recently Extinct Genera

Lost Species

The Rallidae family has unfortunately seen a significant number of extinctions, particularly among island species, largely due to human activities and the introduction of non-native predators. These recently extinct genera represent a poignant loss to global biodiversity.

  • Mundia – Ascension crake (flightless, single island, lost by early 1800s to introduced cats and rats)
  • Aphanocrex – Saint Helena rail (flightless, single island, lost by 1500s to introduced cats and rats)
  • Diaphorapteryx – Hawkins's rail (flightless, two islands, lost between 1500 and 1700 to overhunting)
  • Aphanapteryx – Red rail (flightless, single island, lost by 1700 to overhunting and introduced pigs, cats and rats)
  • Erythromachus – Rodrigues rail (flightless, single island, lost by 1760 to overhunting, destruction of habitat by tortoise hunters, and introduced cats)
  • Genus Cabalus – Chatham rail and New Caledonian rail (sometimes included in Gallirallus; extinct around 1900)
  • Genus Capellirallus – Snipe-rail (flightless, single island, lost by no later than 1400s to introduced rats)
  • Genus Vitirallus – Viti Levu rail (flightless, single island, lost by no later than early Holocene)
  • Genus Hovacrex – Hova gallinule (flight ability uncertain, single island, lost by no later than Late Pleistocene)

Additionally, the undescribed Fernando de Noronha rail, of undetermined genus and species, survived into historic times. The extinct genus Nesotrochis from the Greater Antilles was once considered a rail but is now recognized as an independent lineage of gruiforms, more closely related to Sarothruridae and adzebills.

Fossil Record

Ancient Lineages

The fossil record of long-extinct prehistoric rails is extensively documented, particularly from well-researched geological formations in Europe and North America, as well as from less comprehensively studied strata in other regions.[35] These fossil discoveries provide invaluable insights into the deep evolutionary history and diversification of the Rallidae family.

  • Genus Eocrex (Wasatch Early Eocene of Steamboat Springs, USA; Late Eocene – ?Oligocene of Isfara, Tadzhikistan)
  • Genus Palaeorallus (Wasatch Early Eocene of Wyoming, USA)
  • Genus Parvirallus (Early – Middle Eocene of England)
  • Genus Aletornis (Bridger Middle Eocene of Uinta County, USA) – includes Protogrus[36]
  • Genus Fulicaletornis (Bridger Middle Eocene of Henry's Fork, USA)
  • Genus Latipons (Middle Eocene of Lee-on-Solent, England)
  • Genus Ibidopsis (Hordwell Late Eocene of Hordwell, UK)
  • Genus Quercyrallus (Late Eocene -? Late Oligocene of France)
  • Genus Belgirallus (Early Oligocene of WC Europe)
  • Genus Rallicrex (Corbula Middle/Late Oligocene of Kolzsvár, Romania)
  • Rallidae gen. et sp. indet. (Late Oligocene of Billy-Créchy, France) – a small species of rail[37]
  • Genus Palaeoaramides (Late Oligocene/Early Miocene – Late Miocene of France)
  • Genus Rhenanorallus (Late Oligocene/Early Miocene of Mainz Basin, Germany)
  • Genus Paraortygometra (Late Oligocene/?Early Miocene -? Middle Miocene of France) – includes Microrallus
  • Genus Australlus (Late Oligocene – Middle Miocene of NW Queensland, Australia)
  • Genus Pararallus (Late Oligocene? – Late Miocene of C Europe) – possibly belongs in Palaeoaramides
  • Genus Litorallus (Early Miocene of New Zealand)
  • Rallidae gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) – dozens of mostly broken isolated skull and limb bones of a rail or crake[38]
  • Rallidae gen. et sp. indet. (Bathans Early/Middle Miocene of Otago, New Zealand) – quadrate and 2 femora of a large rail or crake[39]
  • Genus Miofulica (Anversian Black Sand Middle Miocene of Antwerp, Belgium)
  • Genus Miorallus (Middle Miocene of Sansan, France -? Late Miocene of Rudabánya, Hungary)
  • Genus Youngornis (Shanwang Middle Miocene of Linqu, China)
  • Rallidae gen. et sp. indet. (Sajóvölgyi Middle Miocene of Mátraszőlős, Hungary) – several limb bones of a smallish rail[40]
  • Rallidae gen. et sp. indet. (Middle Miocene of Grive-Saint-Alban, France) – partial hand of a common moorhen-sized rail[41]
  • Rallidae gen. et sp. indet. (Late Miocene of Lemoyne Quarry, USA)
  • Rallidae gen. et sp. indet. UMMP V55013-55014; UMMP V55012/V45750/V45746 (Rexroad Late Pliocene of Saw Rock Canyon, USA)
  • Rallidae gen. et sp. indet. UMMP V29080 (Rexroad Late Pliocene of Fox Canyon, USA)
  • Genus Creccoides (Blanco Late Pliocene/Early Pleistocene of Crosby County, USA)
  • Rallidae gen. et sp. indet. (Bermuda, West Atlantic)
  • Rallidae gen. et sp. indet. (formerly Fulica podagrica) (Late Pleistocene of Barbados)[42]
  • Genus Pleistorallus (mid-Pleistocene New Zealand). The holotype of Pleistorallus flemingi is housed in the Museum of New Zealand Te Papa Tongarewa.[43][44]

Doubtfully Placed Taxa

Some fossil taxa have been tentatively associated with the Rallidae family, but their precise phylogenetic placement remains uncertain. Further research and fossil discoveries are often required to definitively classify these enigmatic ancient birds.

  • Genus Ludiortyx (Late Eocene) – includes "Tringa" hoffmanni, "Palaeortyx" blanchardi, "P." hoffmanni
  • Genus Telecrex (Irdin Manha Late Eocene of Chimney Butte, China)
  • Genus Amitabha (Bridger middle Eocene of Forbidden City, USA) – possibly a phasianid?
  • Genus Palaeocrex (Early Oligocene of Trigonias Quarry, USA)
  • Genus Rupelrallus (Early Oligocene of Germany)
  • Neornithes incerta sedis (Late Oligocene of Riversleigh, Australia) – a left carpometacarpus piece of a bird about the size of Lewin's rail. Probably from a rail, but too damaged for precise affiliation.[45]
  • Genus Euryonotus (Pleistocene of Argentina)

The presumed scolopacid wader Limosa gypsorum (Montmartre Late Eocene of France) is occasionally considered a rail and subsequently placed in the genus Montirallus.[46]

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References

References

  1.  Partial hand of a common moorhen-sized rail: Ballmann (1969)
A full list of references for this article are available at the Rail (bird) Wikipedia page

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