Palaelodidae

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Palaelodidae
Temporal range: Early Oligocene–Pleistocene
Palaelodus ambiguus skull and neck.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Phoenicopteriformes
Family:† Palaelodidae
Stejneger, 1885
Type species
Palaelodus ambiguus
Genera

Palaelodidae is a family of extinct birds in the group Phoenicopteriformes, which today is represented only by the flamingos. They were widespread during the Neogene, with fossil remains found on all continents other than Antarctica. The oldest remains referred to this group appeared in the fossil record during the Oligocene in Egypt and Belgium, before palaelodids reached their peak diversity during the Miocene. Following this the group declined in the early Pliocene before going extinct on most continents. However, remains found near Cooper Creek in the Lake Eyre Basin indicate that palaelodids managed to survive in Australia until the Pleistocene. Currently three genera are recognized by scientists: Adelalopus , Palaelodus and Megapaloelodus . Most fossil remains stem from Europe and have been assigned to the type species, Palaelodus ambiguus. Due to the fragmentary nature of most of these species, little is known about their ecology. They appear to have preferred brackish lakes and lagoons. Palaelodus has previously been thought to be a wader or diver, but recent research indicates that they were better suited for swimming and possibly fed on insect larvae and other aquatic invertebrates. At least Megapaloelodus appears to have adaptations for "locking" their legs in a standing position.

Contents

History of discovery

The first palaelodid remains were discovered during the middle of the 19th century in the region around Saint-Gérand-le-Puy in France. These remains were described by French scientist Alphonse Milne-Edwards who recognized several, in his mind distinct, forms of birds that he included in the new genus Palaelodus , a name he derived from the Ancient Greek words "palaios" (ancient) and "elodus" (inhabitant of marshes). [1] Milne-Edwards initially named three species; P. ambiguus, P. gracilipes and P. crassipes; before returning later and establishing two more species, P. minutus and P. goliath. [2] However most of these species would eventually come to be recognized as likely having belonged to a single form displaying variation in size among individuals. Approximately 20 years after the first description of Palaelodus, Leonhard Stejneger established the family Palaelodontidae after having misinterpreted the original etymology. This would eventually be amended by Howard, who changed it to Paloelodidae, and even later by Brotkorb who introduced the current spelling, Palaelodidae. [1] During the 1940s and 1950s remains similar to the ones from Europe would come to be discovered in the western United States, leading to the establishing of the genus Megapaloelodus by Alden H. Miller. [3] Although only known from a few fragmentary bones, Miller remarked that the American material appeared significantly larger than that of the European Palaelodus goliath and subsequently chose to reflect this in the genus name (although Miller misspelled Palaelodus in the process). An additional form, Megapaloelodus opsigonus, also known from fragmentary remains was named around 20 years later by Pierce Brodkorb. [1] The 1980s saw a major revision of the European species, with Jacques Cheneval working of older work and synonymizing much of the French material into a single species while also transferring P. goliath into Miller's Megapaloelodus. [4] The range of this family received a significant boost when fossil material of these birds was discovered in the early Oligocene of Africa and the Oligocene to Miocene of Australia. [5] Another significant find was the discovery of unambiguous skull remains, first described by Cheneval and Escuillié in 1992. [6] Another potential revision was suggested in 2002 by Jiří Mlíkovský, who proposed that Palaelodus and Megapaloelodus should be lumped into a single genus. This was however met with criticism and not widely accepted. [7] The same year as Mlíkovský's suggested revision, Mayr and Smith described Adelalopus (an anagram of Palaelodus), an entirely new genus of palaelodid from the early Oligocene. [8] The 2000s and 2010s also saw the description of multiple new species in the established genera. [9]

Species

GenusSpeciesAgeLocationNotes
Adelalopus [8] Adelalopus hoogbutseliensis Early Oligocene Flag of Belgium (civil).svg  Belgium The oldest known palaelodid.
Megapaloelodus Megapaloelodus connectens [3] Miocene Flag of the United States.svg  USA The type species of Megapaloelodus.
Megapaloelodus goliath [4] Oligocene-Miocene Flag of France.svg  France

Flag of Germany.svg  Germany

M. goliath was originally described as a species of Palaelodus. Its referral to Megapaloelodus is uncertain.
Megapaloelodus opsigonus [1] Early Pliocene Flag of the United States.svg  USA

Flag of Mexico.svg  Mexico

The youngest species of Megapaloelodus.
Megapaloelodus peiranoi [9] Miocene Flag of Argentina.svg  Argentina M. peiranoi is thought to be the basalmost species of Megapaloelodus.
Palaelodus Palaelodus ambiguus [4] Oligocene-Miocene Flag of France.svg  France

Flag of Germany.svg  Germany

Flag of Brazil.svg  Brazil?

The type species of Palaelodus. Two other species, P. gracilipes and P. crassipes are thought to simply be smaller and larger individuals of this species respectively. It is the best understood species of palaelodid, known from thousands of individual bones. It is also the only species known from skull material.
Palaelodus aotearoa [7] Early Miocene Flag of New Zealand.svg  New Zealand A medium-sized species of Palaelodus.
Palaelodus kurochkini [10] Middle Miocene Flag of Mongolia.svg  Mongolia P. kurochkini shows several features that might indicate that it should be placed in its own, distinct genus.
Palaelodus pledgei [5] Oligocene-Miocene Flag of Australia (converted).svg  Australia The smallest species of palaelodid.
Palaelodus wilsoni [5] Oligocene-Pleistocene?Flag of Australia (converted).svg  Australia Fragmentary remains from Cooper Creek might indicate that this species survived until the Pleistocene.

The genera Probalearica as well as the species "Grus" miocenicus and Pliogrus germanicus were all initially described as cranes, but were later found to represent members of the Palaelodidae. [11] [12]

Range

Palaeolodus ambiguus skeleton Palaeolodus ambiguus skeleton.JPG
Palaeolodus ambiguus skeleton

Palaelodids were a successful group of bird found across all continents except for Antarctica, with their diversity being the greatest during the Miocene period.

The most extensive record of the family can be found in Europe, beginning with Adelalopus in the early Oligocene of Hoogbutsel, Belgium. [8] While this genus is currently only found in one locality, the younger Palaelodus appear to have been much more widespread, being found in the form of thousands of bones in Saint-Gérand-le-Puy, [4] [13] France, and multiple localities in Germany including the Mainz Basin, [13] Nördlinger Ries [7] and the Steinheimer Basin. [7] Megapaloelodus remains were also found in Saint-Gérand-le-Puy, [4] the Steinheimer Basin and Nördlinger Ries. [7] Other fossils have been discovered in Bohemia in the Czech Republic [7] and Romania. [11] Records in Asia are more sparse, with only a single described species known from the Middle Miocene Ooshin Formation in Mongolia. [10]
America primarily yielded remains of the large genus Megapaloelodus, which is chiefly known from formations of the west coast. This includes fossils from Oregon's Juntura Formation [1] and California's Barstow Formation [14] and the Sharktooth Hill Bonebed. [15] Further inland fossils were found in the Rosebud Formation of South Dakota. [3] These fossils mostly date to the Miocene, with the exception of the material from Oregon which suggests an early Pliocene age. Further south Megapaloelodus was found in the Miocene to Pliocene Almejas Formation of Cedros Island (Baja California). [16] In South America Megapaloelodus fossils were found in the Andalhualá [9] and Ituzaingó Formation [17] [18] of Argentina while material assigned to Palaelodus cf. ambiguus have been identified from Brazil's Taubuté Basin. [19]
While there are no described species from Africa, remains are still known from multiple localities. Two differently sized types have been recovered from the early Oligocene Jebel Qatrani Formation of Fayum, Egypt. [20] A possible species of Megapaloelodus was found in the early Miocene Northern Sperrgebiet of Namibia. [21]
Three species of Palaelodus are currently accepted from Australasia. Two of these were present in the Oligocene to Miocene Etadunna and Namba Formation within the Lake Eyre Basin. [5] While the third was found in New Zealand's Bannockburn Formation, corresponding to the early Miocene in age. [7] While palaelodid remains are primarily known from these Neogene localities, isolated remains were also discovered in the much younger Kutjitara Formation, indicating that the group may have survived until 0.2 mya. [5]

Description

Broadly speaking palaelodids resemble their modern relatives, the flamingos, in proportions. They had long legs and elongated necks. Despite these similarities, the two groups still differed in how these features were expressed or in how well they were developed. The neck for example lacks the pronounced kink where the vertebrae of the neck connect to those of the notarium and the tarsometatarsus is shorter than the humerus unlike in flamingos. Additionally, the limb bones are laterally flattened which is closer to the condition seen in grebes. The toes share this flattening and are limited in how much they can flex. The anatomical features responsible for the later are additionally associated with the presence of webbed feet, lending credence to the hypothesis that they were divers or swimmers. However, while grebes feature similar adaptations, the details of the flattening suggests that both groups evolved these traits independently from one another. Skull material is only known from a single species, Palaelodus ambiguus, and features a straight, conical bill very much unlike the curved bills of their modern relatives. [22] Although it is uncertain if other palaelodids would have the same head shape, at least P. ambiguus appears to mix traits of grebes with those of flamingos. Matching with their preference for brackish water, P. ambiguus confirms that at least some species within the family had salt glands. [13]

Palaelodids generally fall within the size range of modern species of flamingo. Palaelodus is considered to be the smallest genus in the family, [3] [7] with the smallest species being Palaelodus pledgei. [5] P. kurochkini, which is described as medium-sized for its genus, has been said to be the size of an American flamingo. [10] Larger Palaelodus species overlap with small species of Megapaloelodus, such as M. goliath [10] and M. opsigonos. [16] M. connectens and Adelalopus were both noted to be larger than M. goliath, the former reaching a size similar to a large male greater flamingo [23] and the later being said to have been slightly larger than M. goliath. [8]

Phylogeny

Flickr - Rainbirder - Great Crested Grebe (Podiceps cristatus).jpg
Phoenicopterus roseus 237172900.jpg
Palaelodids are most closely related to modern flamingos, linking them to the diving grebes

The Palaelodidae is a family within the clade Phoenicopteriformes, which includes the modern flamingos of the Phoenicopteridae. Together with grebes they form the clade Mirandornithes. The relationship between these groups is well supported by morphological and molecular data respectively, with palaelodids being the sister family to flamingos and forming an important link between the two extant families. Based on Palaelodus, these birds displayed general proportions similar to flamingos but legs akin to those of grebes. [13] [24] [25] The precise internal relationships in this family are not well understood. Adelalopus appears to have been the oldest member of the group and shares similarities with the later Megapaloelodus. [8] Among the species of Megapaloelodus, M. peiranoi is thought to be the basalmost member due to its resemblance to some early mirandornithes, specifically Juncitarsus . [9] The following phylogenetic tree depicts Mirandornithes as recovered by Torres and colleagues in 2015. [22]

Mirandornites

† Juncitarsus

Podicipediformes

Phoenicopteriformes

† Palaelodidae

Phoenicopteridae

Paleobiology

Paleoenvironment

Many of the rock units that yielded palaelodid remains are thought to have represented shallow lakes or similar environments at the time the bird fossils were deposited. Furthermore, many of these are believed to have been either occasionally or permanently brackish or even saline, indicating that these animals had habitat preferences similar to modern flamingos. The genus Adelalopus for instance appears to have inhabited the shores of a lagoon in what is now Belgium. [8] In a 1989 study Jacques Cheneval analysed the bird species of the type locality of Saint-Gérand-le-Puy, finding that over half of the identified taxa show a preference for brackish conditions while freshwater exclusives appear to have been absent. [26] This was later supported by studies that proposed that the locality underwent cyclical dry and wet seasons, with the salinity of the lake changing through the evaporation of water. This appears to have been an attractive environment for early phoenicopteriforms, given the presence of vast numbers of Palaelodus and more rarely Megapaloelodus and Harrisonavis . [22] Additionally, the discovery of juvenile birds and eggs suggests that palaelodids also bred there. [27] Another similar environment was present in the Lake Eyre Basin, which is thought to have supported salt lakes throughout much of the Neogene and early Quaternary. These localities mirror those of Europe in preserving not just the remains of juvenile palaelodids but also the fossils of more derived flamingos that seemingly coexisted with their more basal relatives. [5] In the case of Australia specifically, this included Phoenicopterus novaehollandiae and Phoeniconotius eyerensis at Lake Palankarinna.

However, palaelodids show some additional range beyond brackish lakes and lagoons. The Jebel Qatrani Formation, where two indetermined palaelodids were discovered, is thought to represent a mix of tropical lowland swamp and marshland supported by a series of river channels that empty into the Tethys Ocean. [28] Palaelodus aotearoa from New Zealand inhabited paleo-Lake Manuherikia, which preserves a rich fauna of water fowl and even the remains of crocodiles. Worthy and colleagues describe the environment as a large lake fed by a river delta that was highly affected by wave activity and currents within the lake itself. [7] The Northern Sperrgebiet of Namibia also preserves the presence of freshwater without signs of saline lakes like those inhabited by the European and Australian taxa. [21] The Barstow Formation also likely represented a freshwater rather than brackish environment. [23]

Palaelodids may have fed on caddisfly larvae and other small invertebrates Freeliving caddisfly larva, member of the R. Brunnea-Vemna Group (9647705348).jpg
Palaelodids may have fed on caddisfly larvae and other small invertebrates

Locomotion and feeding

Little is known about the precise habits of most palaelodids given the lack of substantial fossil material outside of Palaelodus ambiguus. Subsequently, most of what has been proposed for this group is based on said species. Palaelodids lack several of the adaptations found in flamingos that help them in their filter feeding lifestyle, such as the curved jaw, attachment point for keratinous lamellae and the absence of the pronounced kink in the spine prior to the shoulder girdle. The absence of these features all point towards the fact that they had a notably different way of feeding despite their affinity for similar environments. One possibility that has been suggested is that palaelodids fed on various aquatic invertebrates, specifically snails and caddisflies. The later are known to have been especially abundant in the French deposits, where their casings make a substantial contribution to the local limestone. It is possible that the modern filter-feeding apparatus of flamingos evolved from this or a similar feeding method. [13]

Their precise way of life has also been the subject of much debate. Earlier research on P. ambiguus, primarily the works of Cheneval and Escuillié, proposed that they may have been leg-propelled divers. [4] [6] More recently this hypothesis has been called into question, partially due to the fact that no diving bird possesses limbs as elongated as what was present in palaelodids. Worthy and colleagues suggest a wading lifestyle, arguing that the compressed and elongated limbs may have aided in wading more easily through deep water. [7] Mayr on the other hand proposes a swimming lifestyle, pointing towards the reduced ability of the toes to flex being unsuited for wading. In his hypothesis the webbed feet and compressed toes would be more suited to work as propulsion while swimming. [25] [22] Given its more fragmentary nature, Megapaloelodus has been subject to less speculation regarding its lifestyle. Miller notes that the bones of the lower leg appear to have a mechanism that could "lock" them in place. This could help stabilize the bird while standing in a position without moving. [23]

Related Research Articles

<span class="mw-page-title-main">Gaviiformes</span> Order of birds

Gaviiformes is an order of aquatic birds containing the loons or divers and their closest extinct relatives. Modern gaviiformes are found in many parts of North America and northern Eurasia, though prehistoric species were more widespread.

<span class="mw-page-title-main">Grebe</span> Order of birds

Grebes are aquatic diving birds in the order Podicipediformes. Grebes are widely distributed freshwater birds, with some species also found in marine habitats during migration and winter. Most grebes fly, although some flightless species exist, most notably in stable lakes. The order contains a single family, the Podicipedidae, which includes 22 species in six extant genera.

<span class="mw-page-title-main">Flamingo</span> Family of birds

Flamingos or flamingoes are a type of wading bird in the family Phoenicopteridae, which is the only extant family in the order Phoenicopteriformes. There are four flamingo species distributed throughout the Americas, and two species native to Afro-Eurasia.

<span class="mw-page-title-main">Phoenicopteriformes</span> Order of birds

Phoenicopteriformes is a group of water birds which comprises flamingos and their extinct relatives. Flamingos (Phoenicopteriformes) and the closely related grebes (Podicipedidae) are contained in the parent clade Mirandornithes.

<i>Larus</i> Genus of birds

Larus is a large genus of gulls with worldwide distribution.

<span class="mw-page-title-main">Mousebird</span> Order of birds

The mousebirds are birds in the order Coliiformes. They are the sister group to the clade Cavitaves, which includes the Leptosomiformes, Trogoniformes (trogons), Bucerotiformes, Piciformes and Coraciformes. This group is now confined to sub-Saharan Africa, and it is the only bird order confined entirely to that continent, with the possible exception of turacos which are considered by some as the distinct order Musophagiformes, and the cuckoo roller, which is the only member of the order Leptosomiformes, and which is found in Madagascar but not mainland Africa. Mousebirds had a wider range in the Paleogene, with a widespread distribution in Europe and North America during the Paleocene.

<span class="mw-page-title-main">Sagittariidae</span> Family of birds

Sagittariidae is a family of raptor with one living species—the secretarybird native to Africa—and a few fossil taxa.

<i>Palaelodus</i> Extinct genus of birds

Palaelodus is an extinct genus of bird of the Palaelodidae family, distantly related to flamingos. They were slender birds with long, thin legs and a long neck resembling their modern relatives, but likely lived very different livestyles. They had straight, conical beaks not suited for filter feeding and legs showing some similarities to grebes. Their precise lifestyle is disputed, with researchers in the past suggesting they may have been divers, while more recent research suggests they may have used their stiff toes as paddles for swimming while feeding on insect larvae and snails. This behavior may have been key in later phoenicopteriforms developing filterfeeding bills. The genus includes between five and eight species and is found across Europe, Australia, New Zealand, Asia and possibly South America. However some argue that most of the taxa named from Europe simply represent differently sized individuals of one single species. Palaelodus was most abundant during the Late Oligocene to Middle Miocene periods, but isolated remains from Australia indicate that the genus, or at least a relative, survived until the Pleistocene.

Megapaloelodus is an extinct genus of stem flamingo of the family Palaelodidae. Megapaloelodus is primarily known from Miocene America, from South Dakota and Oregon in the north to Argentina in the south, but the species Megapaloelodus goliath was found in Europe. Additionally, one unnamed species was discovered in Miocene sediments from Namibia. Due to a lack of skull material, little can be said about the ecology of Megapaloelodus. Species of this genus are typically larger than those of Palaelodus and appear to have inhabited similar brackish lake environments. Additionally, they may have been capable of "locking" their legs in a standing position.

Phoeniconotius is an extinct genus of flamingo that lived in Australia from the late Oligocene to the early Miocene. Unlike modern flamingos and the contemporary Phoenicopterus novaehollandiae, it was likely less well adapted for swimming and deep water wading. Phoeniconotius was a robust flamingo with bones more massive than those of the modern greater flamingo. Only a single species is recognized, Phoeniconotius eyrensis.

<span class="mw-page-title-main">Mirandornithes</span> Taxon of birds

Mirandornithes is a clade that consists of flamingos and grebes. Many scholars use the term Phoenicopterimorphae for the superorder containing flamingoes and grebes.

<i>Palaeortyx</i> Extinct genus of birds

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<i>Juncitarsus</i> Extinct genus of birds

Juncitarsus is an extinct genus of wading birds from the Eocene of the United States and Germany. Though previously considered a flamingo, it is likely a stem-flamingo, possibly a relative of the group which contains both flamingos and grebes (Mirandornithes).

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Pelargopappus is an extinct genus of raptor related to the secretarybird that lived in early Miocene France. Only one species, the type species P. magnus is officially recognized. A second species, P. schlosseri from the mid-and late Oligocene, was split off into the genus Amphisagittarius.

Harrisonavis is an extinct genus of flamingo that lived during the Late Oligocene and Early Miocene periods in what is now France. Despite being one of the oldest known members of the flamingo family, it already shows a skull remarkably similar to that of the modern greater flamingo. Although generally similar, it subtly differs in the curvature of the bill and the size of the ventral keel of the maxilla, both signs that Harrisonavis was not yet as adapted towards filter feeding as modern species are. Harrisonavis inhabited brackish lakes alongside the more basal Palaelodidae. It contains the single species Harrisonavis croizeti, first described in 1852.

<i>Miobaptus</i> Extinct water bird

Miobaptus is a fossil genus of grebe that is known from several specimens collected from Czechia and Lake Baikal dating from the Early Miocene to Middle Miocene. Considered to be one of the most primitive genera of grebes, the anatomy of Miobaptus suggests it was less adapted for the aquatic mode of life than modern grebes, but had better flight maneuverability.

Adelalopus is an extinct genus of palaelodid bird from the lowermost Oligocene of Belgium. It is the oldest member of its family and the largest known palaelodid from Europe, slightly larger than Megapaloelodus goliath. It contains a single species, Adelalopus hoogbutseliensis.

Laricola is a genus of extinct gull-like birds that lived during the late Oligocene and early Miocene in what is now Europe.

References

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  24. ↑ Mayr, Gerald (2006). "The contribution of fossils to the reconstruction of the higher-level phylogeny of birds" (PDF). Species, Phylogeny and Evolution. 1: 59–64. ISSN   1864-211X . Retrieved 12 August 2009.
  25. 1 2 Mayr, Gerald (2004). "Morphological evidence for sister group relationship between flamingos (Aves: Phoenicopteridae) and grebes (Podicipedidae)" (PDF). Zoological Journal of the Linnean Society . 140 (2): 157–169. doi: 10.1111/j.1096-3642.2003.00094.x . ISSN   0024-4082 . Retrieved November 3, 2009.
  26. ↑ Cheneval, J. (1989). "Fossil bird study, and paleoecological and paleoenvironmental consequences: Example from the Saint-Gérand-le-Puy deposits (lower miocene, Allier, France)". Palaeogeography, Palaeoclimatology, Palaeoecology. 73 (3–4): 295–309. Bibcode:1989PPP....73..295C. doi:10.1016/0031-0182(89)90010-2.
  27. ↑ Mikhailov, K.E. (1997). Avian Eggshells: An Atlas of Scanning Electron Micrographs. Vol. 3. British Ornithologists' Club Occasional Publications. ISBN   0-9522886-2-1.
  28. ↑ Rasmussen, D. T.; Simons, E.L.; Hertel, F.; Judd, A. (2001). "Hindlimb of a giant terrestrial bird from the upper Eocene, Fayum, Egypt". Palaeontology. 44 (2): 325–337. Bibcode:2001Palgy..44..325R. doi:10.1111/1475-4983.00182. S2CID   130033734.
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