[Ornithology • 2020] Unforeseen Diversity of Quails (Galliformes: Phasianidae: Coturnix) in Oceanic Islands provided by the Fossil Record of Macaronesia

[A] Coturnix lignorum, [B] C. alabrevis & [C] C. centensis  

Rando, Alcover, Pieper, Olson, Hernández & López-Jurado. 2020. 

DOI: 10.1093/zoolinnean/zlz107 

 twitter.com/ZoolJLinnSoc 

 Artwork by Pau Oliver. 

Abstract

The original bird fauna of most oceanic islands has been affected by recent extinction processes associated with human arrival and its subsequent impacts. In the volcanic Macaronesian archipelagos (Azores, Madeira, Selvagens, Canary Islands and Cape Verde), in the North Atlantic, the Late Quaternary fossil record indicates that there was formerly a higher avian diversity, including a high number of now extinct endemic species. This assemblage of extinct birds includes endemic insular quails (Galliformes: Phasianidae). In this study, we describe three newly discovered extinct species of quails, two of which inhabited the archipelago of Madeira (Coturnix lignorum sp. nov. from Madeira Island and Coturnix alabrevis sp. nov. from Porto Santo Island) and one from Cape Verde (Coturnix centensis sp. nov.). The fossil record also indicates the presence of additional species of extinct endemic quails on other Macaronesian islands. These birds plus the extinct Canary Island quail (Coturnix gomerae) indicate a high former endemic diversity of this genus in Macaronesia, a feature unique among oceanic archipelagos. Anatomical traits show that the new taxa were flightless ground dwellers, making them vulnerable to human interference, with their extinction being linked to human arrival and subsequent habitat alterations and the introduction of invasive species.

Keywords: anatomy, extinction, fossil birds, island biogeography, morphometrics, Quaternary

SYSTEMATIC PALAEONTOLOGY 

Galliformes Temminck, 1820 

Phasianidae Horsfield, 1821 

Coturnix Garsault, 1764

Figure 3. Premaxilla, coracoid, scapula, sternum and synsacrum of Coturnix lignorum from Madeira (A1 –A5), Coturnix alabrevis from Porto Santo (B1 , B2), Coturnix coturnix (C1 –C5), Coturnix gomerae from La Gomera (D2 ), Coturnix sp. A from Bugio (E2 ) and Coturnix sp. C from Graciosa (F2 ).
 A1 –C1 , premaxilla dorsal view (top) and left lateral view (bottom). A2 –F2 , coracoid, dorsal view. A3 , C3 , scapula, left lateral view. A4 , C4 , sternum ventral view (top) and left lateral view (bottom) (the high of apex carinae is shown). A5 , C5 , synsacrum ventral view. Scapulas and coracoids are from the left side, except for E2 . A1 , MMF 47329; B1 , MMF 47413; C1 , IMEDEA 106905; F1 , MCMa 2354.018; A2 , MMF 47325; B2 , MMF 47411; C2 , IMEDEA 106904; D2 , DZUL 1830; E2 , MMF 47416; F2 , MCMa FC-1368; A3 , MMF 47330; C3 , IMEDEA 106907; A4 , MMF 47328; C4 , IMEDEA 106905; A5 , MMF 47327; C5 , IMEDEA 106906.

Figure 1. Map of the Macaronesian Islands. Silhouettes indicate specimen records discussed in this paper: (1) Coturnix lignorum; (2) Coturnix alabrevis; (3) Coturnix centensis; (4) C. gomerae; and (5–7) Coturnix sp.

Figure 7. Artistic reconstruction of: A, Coturnix lignorum; B, Coturnix alabrevis; C, Coturnix centensis; and D, the silhouette of the common quail Coturnix coturnix for comparison, based on bone measurements, when available (wing and leg bones of the three species, sternum of C. lignorum, and premaxillae of C. lignorum and Coturnix alabrevis). Colours are speculative. All drawings are depicted at the same scale. Artwork by Pau Oliver.

Coturnix lignorum Rando, Alcover, Pieper, Olson, Hernández & López-Jurado sp. nov.

Etymology: The specific name lignorum is derived from from the Latin genitive plural of lignum, wood, a direct translation of the Portuguese madeira.

Coturnix alabrevis Rando, Alcover, Pieper, Olson, Hernández & López-Jurado sp. nov.

Etymology: From Latin ala, wing, and brevis, short, in reference to the small size of the forelimb (wing) bones.

Coturnix centensis Rando, Alcover, Pieper, Olson, Hernández & López-Jurado sp. nov.

Etymology: In the Portuguese-based creole language spoken in Cape Verde, the island of São Vicente is called ‘Son Cent'. We based our epithet on this, adding the Latin locative suffix -ensis.

Juan C. Rando, Josep A. Alcover, Harald Pieper, Storrs L. Olson, C. Nayra Hernández and L. Felipe López-Jurado. 2020. Unforeseen Diversity of Quails (Galliformes: Phasianidae: Coturnix) in Oceanic Islands provided by the Fossil Record of Macaronesia. Zoological Journal of the Linnean Society. 188(4); 1296–1317. DOI: 10.1093/zoolinnean/zlz107

  twitter.com/ZoolJLinnSoc/status/1247507061753155585

     

[Ornithology • 2017] On Temminck’s Tailless Ceylon Junglefowl, Gallus ecaudatus, and How Darwin denied their Existence

 Lithograph of Gallus ecaudatus, based on specimen RMNH. AVES.224888, by Jean-Gabriel Prêtre prepared c.1806 for an illustrated work in three volumes that Temminck intended to publish on pigeons and Galliformes.

in van Grouw, Dekkers & Rookmaaker, 2017.

Bull. B.O.C. 137(4) 

  Summary

Ceylon Junglefowl was described in 1807 by the Dutch ornithologist Coenraad Jacob Temminck. The specimens he examined were tailless (‘rumpless’) and therefore he named them Gallus ecaudatus. In 1831 the French naturalist René Primevère Lesson described a Ceylon Junglefowl with a tail as Gallus lafayetii (= lafayetii), apparently unaware of Temminck’s ecaudatus. Subsequently, ecaudatus and lafayetii were realised to be the same species, of which G. stanleyi and G. lineatus are junior synonyms. However, Charles Darwin tried to disprove the existence of wild tailless junglefowl on Ceylon in favour of his theory on the origin of the domestic chicken. 

‘The tailless cock inhabits the immense forests of the island of Ceylon’ (Temminck 1813: 268). 

 ‘… but this statement [tailless fowls are wild in Ceylon] … is uterly false’ (Darwin 1868: 259).

Figure 2. Lithograph of Gallus ecaudatus, based on specimen RMNH. AVES.224888, by Jean-Gabriel Prêtre prepared c.1806 for an illustrated work in three volumes that Temminck intended to publish on pigeons and Galliformes. Only the volume on pigeons was published, in 1808, and the two volumes on Galliformes never appeared due to a confict between Temminck and the French illustrator of the frst volume, Pauline Knip (Dickinson et al. 2010). Instead, Temminck later published Histoire naturelle générale des pigeons et des gallinacés in three volumes (1813–15) without any colour illustrations. The reference ‘Gall. v. 1. pl. Enl.’ in Temminck’s published catalogue (1807) refers to the frst of the two unpublished volumes on Galliformes, which would have been vol. 2 of the complete work (Naturalis Biodiversity Center, Leiden)

Hein van Grouw, Wim Dekkers and Kees Rookmaaker. 2017. On Temminck’s Tailless Ceylon Junglefowl, and How Darwin denied their Existence. Bull. B.O.C. 137(4); 261-271

 boc-online.org/bulletins/downloads/BBOC1374-02vanGrouw.pdf

[PaleoOrnithology • 2018] Panraogallus hezhengensis • Vocal Specialization through Tracheal Elongation in An Extinct Miocene Pheasant from China

Panraogallus hezhengensis 

Li, Clarke, Eliason, Stidham, Deng & Zhou, 2018

DOI: 10.1038/ncomms12739 

Abstract

Modifications to the upper vocal tract involving hyper-elongated tracheae have evolved many times within crown birds, and their evolution has been linked to a ‘size exaggeration’ hypothesis in acoustic signaling and communication, whereby smaller-sized birds can produce louder sounds. A fossil skeleton of a new extinct species of wildfowl (Galliformes: Phasianidae) from the late Miocene of China, preserves an elongated, coiled trachea that represents the oldest fossil record of this vocal modification in birds and the first documentation of its evolution within pheasants. The phylogenetic position of this species within Phasianidae has not been fully resolved, but appears to document a separate independent origination of this vocal modification within Galliformes. The fossil preserves a coiled section of the trachea and other remains supporting a tracheal length longer than the bird’s body. This extinct species likely produced vocalizations with a lower fundamental frequency and reduced harmonics compared to similarly-sized pheasants. The independent evolution of this vocal feature in galliforms living in both open and closed habitats does not appear to be correlated with other factors of biology or its open savanna-like habitat. Features present in the fossil that are typically associated with sexual dimorphism suggest that sexual selection may have resulted in the evolution of both the morphology and vocalization mechanism in this extinct species.

Systematic paleontology

AVES Linnaeus, 1758

GALLIFORMES Linnaeus, 1758

PHASIANIDAE Vigors, 1825

Panraogallus hezhengensis gen et sp. nov.

Etymology: The genus name is the pinyin of the Chinese characters meaning ‘coiling’ and the Latin for ‘chicken,’ referring to the preserved elongate trachea in this species. The specific epithet, ‘hezhengensis’ refers to Hezheng area in the Linxia Basin of Gansu Province where abundant fossils, including the holotype, have been collected.

Specimen number: HMV 1876 (HMV, Hezheng Paleozoological Museum, Vertebrate Collection, Gansu Province, China).

Locality and age: Baihua area near Zhuangkeji Township, Guanghe County, Gansu Province, China exposes the Liushu Formation that is late Miocene (7.25–11.1 Ma).

Fig.1 Type specimen of Panraogallus hezhengensis (HMV 1876). It has coiled, super-elongated windpipe and lived in the late Miocene of Gansu Province in northwest China (Photo by Z. LI and reconstruction by X. GUO at IVPP)

Zhiheng Li, Julia A. Clarke, Chad M. Eliason, Thomas A. Stidham, Tao Deng and Zhonghe Zhou. 2018. Vocal Specialization through Tracheal Elongation in An Extinct Miocene Pheasant from China. Scientific Reports. 8, Article number: 8099. DOI: 10.1038/ncomms12739

Song From The Distant Past, A New Fossil Pheasant From China Preserves A Super-Elongated Windpipe english.ivpp.cas.cn/rh/rp/201805/t20180525_192866.html

Song from the distant past, a new fossil pheasant from China preserves a super-elongated windpipe phys.org/news/2018-06-song-distant-fossil-pheasant-china.html via @physorg_com

Song from the distant past, a new fossil pheasant from China preserves a...  bit.ly/2xqnJn4 via @EurekAlert

El esqueleto fosilizado de un faisán con una tráquea súper larga  nationalgeographic.com.es/ciencia/actualidad/esqueleto-fosilizado-faisan-traquea-super-larga_12769 via @natgeoesp

[Ornithology • 2017] The Role of Niche Divergence and Geographic Arrangement in the Speciation of Eared Pheasants (Crossoptilon, Hodgson 1938)

 Wang, Liu, Liu, Chang, Wang & Zhang, 2017  DOI: 10.1016/j.ympev.2017.05.003 

Highlights

• The phylogenetic relationship of Eared Pheasants was resolved based on 45 loci.

• Asymmetric historical gene flow occurred between both parapatric and allopatric sister species.

• Allopatric sister species exhibit significantly divergent ecological niches whereas parapatric sister species show niche conservatism.

• Ecological divergence may have been the main factor that promoted ecological niche divergence.

Abstract

One of the most contentious theories in current ecology is the ecological niche conservatism, which is defined as conservatism among closely related species; however, the ecological niche can also be shifted, as documented in several cases. Genetic drift and ecological divergent selection may cause ecological niche divergence. The current study aims to test whether the ecological niche is conserved or divergent and to determine the main factor that drives ecological niche divergence or conservation. We analyzed the phylogenetic relationship, ecological niche model (ENM) and demographic history of Eared Pheasants in the genus Crossoptilon (Galliformes: Phasianidae) to test niche conservatism with respect to different geographically distributed patterns. The phylogenetic relationship was reconstructed using ∗BEAST with mitochondrial cytochrome b (cyt b) and 44 unlinked autosomal exonic loci, and ENMs were reconstructed in MAXENT using an average of 41 occurrence sites in each species and 22 bioclimatic variables. A background similarity test was used to detect whether the ecological niche is conserved. Demographic history was estimated using the isolation with migration (IM) model. We found that there was asymmetric gene flow between the allopatric sister species Crossoptilon mantchuricum and C. auritum and the parapatric sister species C. harmani and C. crossoptilon. We found that ecological niches were divergent, not conserved, between C. mantchuricum and C. auritum, which began to diverge at approximately 0.3 million years ago. However, the ecological niches were conserved between C. crossoptilon and C. harmani, which gradually diverged approximately half a million years ago. Ecological niches can be either conserved or divergent, and ecological divergent selection for local adaptation is probably an important factor that promotes and maintains niche divergence in the face of gene flow. This study provides a better understanding of the role that divergent selection has in the initial speciation process. The platform combined demographic processes and ecological niches to offer new insights into the mechanism of biogeography patterns.

Keywords: Crossoptilon; Eared-pheasant; Divergent selection; Ecological niche modeling; Genetic drift; Gene flow

Fig. 1. Map of the study area indicating the occurrence points used in for the background similarity test of Ecological Niche Models (ENMs) and the location of DNA samples used in demographic analyses. (The occurrence points (circles) were from bird-watching records (http://birdtalker.net), the Global Biodiversity Information Facility (http://www.gbif.org) and our unpublished survey data. Occurrence points that were far from each other (at least 10 km) and were randomly chosen in ArcGIS software were used for the background similarity test. The study area was the minimal convex polygon of those occurrence points with an additional 200 km. Triangles represent the locations of DNA samples. The area surrounded by the black dashed line was the study area used for the background similarity test. 

Pengcheng Wang, Yang Liu, Yinong Liu, Yajing Chang, Nan Wang and Zhengwang Zhang. 2017. The Role of Niche Divergence and Geographic Arrangement in the Speciation of Eared Pheasants (Crossoptilon, Hodgson 1938). Molecular Phylogenetics and Evolution. In Press. DOI: 10.1016/j.ympev.2017.05.003

[PaleoOrnithology • 2016] Osteology Supports a Stem-Galliform Affinity for the Giant Extinct Flightless Bird Sylviornis neocaledoniae (Sylviornithidae, Galloanseres)

Fig 15. A skeletal reconstruction for Sylviornis neocaledoniae in a resting pose.
 The missing skeletal parts are estimated (shaded bones), with the pelvis based on proportions of Leipoa ocellata. The bill rhamphotheca is not reconstructed so as to not obscure the underlying skeletal morphology.

DOI: 10.1371/journal.pone.0150871 

Abstract

The giant flightless bird Sylviornis neocaledoniae (Aves: Sylviornithidae) existed on La Grande Terre and Ile des Pins, New Caledonia, until the late Holocene when it went extinct shortly after human arrival on these islands. The species was generally considered to be a megapode (Megapodiidae) until the family Sylviornithidae was erected for it in 2005 to reflect multiple cranial autapomorphies. However, despite thousands of bones having been reported for this unique and enigmatic taxon, the postcranial anatomy has remained largely unknown. We rectify this deficiency and describe the postcranial skeleton of S. neocaledoniae based on ~600 fossils and use data from this and its cranial anatomy to make a comprehensive assessment of its phylogenetic affinities. Sylviornis neocaledoniae is found to be a stem galliform, distant from megapodiids, and the sister taxon to the extinct flightless Megavitiornis altirostris from Fiji, which we transfer to the family Sylviornithidae. These two species form the sister group to extant crown-group galliforms. Several other fossil galloanseres also included in the phylogenetic analysis reveal novel hypotheses of their relationships as follows: Dromornis planei (Dromornithidae) is recovered as a stem galliform rather than a stem anseriform; Presbyornis pervetus (Presbyornithidae) is the sister group to Anseranatidae, not to Anatidae; Vegavis iaai is a crown anseriform but remains unresolved relative to Presbyornis pervetus, Anseranatidae and Anatidae. Sylviornis neocaledoniae was reconstructed herein to be 0.8 m tall in a resting stance and weigh 27–34 kg. The postcranial anatomy of S. neocaledoniae shows no indication of the specialised adaptation to digging seen in megapodiids, with for example, its ungual morphology differing little from that of chicken Gallus gallus. These observations and its phylogenetic placement as stem galliforms makes it improbable that this species employed ectothermic incubation or was a mound-builder. Sylviornis neocaledoniae can therefore be excluded as the constructor of tumuli in New Caledonia.

Summary

Our new anatomical information and phylogenetic analysis finds that the large flightless bird Sylviornis neocaledoniae, originally described as a ratite, but then long regarded as a megapode, to be a stem galliform that forms a clade with Megavitiornis altirostris, forming Sylviornithidae. Neither species are megapodes, but the former perception that they were has led to the suggestion that S. neocaledoniae constructed the large mounds or tumuli in New Caledonia. Ectothermic incubation uniquely characterises Megapodiidae[21], with all included taxa depositing their eggs either in mounds, where heat from composting vegetation warms the eggs, or in holes dug in thermally heated soil or sand where sun can heat it such as in beach dunes. Our finding that it is a stem galliform (and thus not a megapode) makes it most unlikely to have been a mound-builder, as this scenario would require both mound-building and ectothermic incubation to have evolved twice. Furthermore, Sylviornis neocaledoniae shows no specific adaptation for digging to facilitate mound-building, unlike all extant megapodiids, making it even more unlikely that it exhibited ectothermic incubation.

Trevor H. Worthy, Miyess Mitri, Warren D. Handley, Michael S. Y. Lee, Atholl Anderson and Christophe Sand. 2016. Osteology Supports a Stem-Galliform Affinity for the Giant Extinct Flightless Bird Sylviornis neocaledoniae (Sylviornithidae, Galloanseres). PLoS ONE. DOI: 10.1371/journal.pone.0150871

http://biostamps.narod.ru/worldlist/fiji/fiji_2006_ext/fiji_2006_ext.htm

[PaleoOrnithology • 2016] Is the “Genyornis” Egg of A Mihirung or Another Extinct Bird from the Australian Dreamtime?

Fig. 1. A, Comparison of SAM P.42421, the ‘Spooner Egg’, left, attributed to Genyornis newtoni, and an emu egg (Dromaius novaehollandiae) SAM B.9899. B, Comparison of femora of Genyornis newtoni SAM P13864 and a 22 cm long Dromaius novaehollandiae femur (FUR 058). C, The Spooner Egg as partly excavated revealing its intact nature; photograph by Gifford Miller, INSTAAR, Colorado. D, The Spooner Egg in situ as found by NS on 23rd July 2000, photograph by Gifford Miller using a reflex camera, and is the best image taken of the egg prior to excavation.

Scale bars in A and B = 10 cm. DOI:  10.1016/j.quascirev.2015.12.011  

Highlights

• Eggshell previously identified as from Genyornis newtoni is reassessed.

• Egg size and microstructure is not conducive with an identity as an dromornithid.

• We suggest that this eggshell is from one of the extinct megapodes in Progura.

• Previous assessments of the timing of Genyornis extinction relate to Progura species.

Abstract

The iconic Australian Genyornis newtoni (Dromornithidae, Aves) is the sole Pleistocene member of an avian clade now hypothesized to be alternatively in Anseriformes or the sister group of crown Galloanseres. A distinctive type of fossil eggshell commonly found in eroding sand dunes, has been referred to Genyornis newtoni since the 1980s. The 126 by 97 mm Spooner Egg, dated at 54.7 ± 3.1 ka by optical dating of its enclosing sediments, is a complete specimen of this eggshell type that was reconstructed from fragments of a broken egg. We show that the size of the eggs from which this ‘Genyornis’ eggshell derives, either as predicted from measurements of fragments, or as indicated by the Spooner Egg, is unexpectedly small given the size of G. newtoni, which has an estimated mass of 275 kg, or about seven times the mass of the emu that has a similar sized egg. We compared the microstructure of the putative Genyornis eggshell to that of other dromornithids and a range of galloanseriform taxa using several microcharacterisation techniques. The ‘Genyornis’ eggshell displays a mosaic of oological characters that do not unambiguously support referral to any known modern bird. Its shell structure, coupled with chemical compounds in the accessory layer, makes it unlikely to have been laid by a dromornithid, whereas several characters support a megapode origin. A potential candidate for the bird that laid the putative ‘Genyornis’ eggs in the Pleistocene fossil avifaunal record has been ignored: Progura, a genus of extinct giant megapodes, whose species were widespread in Australia. Regression of egg size of megapodes and body mass shows that the Spooner Egg approximates the expected size for eggs laid by species of Progura. We advance the suggestion that the fossil eggshell hitherto referred to Genyornis newtoni, is more likely to have been laid by species of the giant extinct Progura. As megapodes, the species of Progura were obligate ectothermic incubators, which we suggest laid their eggs into a hole dug in sand like the modern megapode Macrocephalon maleo, thus explaining the abundant ‘Genyornis’ eggshell in sand dunes. Referral of this eggshell to Progura means that the fossil record of Genyornis newtoni is limited to bones and the timing of the extinction of this last dromornithid is unknown. In addition, structural similarities of eggshell in megapodes, the putative Genyornis eggshell and dromornithids, raise the possibility that these taxa are phylogenetically more closely related to each other than any is to anseriforms. Specifically, this means that dromornithids might be a sister group to galliforms rather than to or within anseriforms.

Keywords: Eggs; Eggshells; Paleoenvironments; Genyornis newtoni; Dromornithids; Megapodes; Progura; Micro-CT; EBSD; Quaternary; Australia

Conclusions

We have described in detail the structure of putative Genyornis eggshell and raised several obstacles to the hypothesis first advanced by Williams (1981) and accepted thereafter ( Miller et al., 1999 and Miller et al., 2005), that this ootype was laid by the giant dromornithid G. newtoni. Rather, we think it more likely that it was laid by one of the several species of giant megapodes in the genus Progura that were widespread in the Pleistocene in Australia.

Gerald Grellet-Tinner, Nigel A. Spooner and Trevor H. Worthy. 2016. Is the “Genyornis” Egg of A Mihirung or Another Extinct Bird from the Australian Dreamtime? QUATERNARY SCIENCE REVIEWS. 133:147-164   DOI:  10.1016/j.quascirev.2015.12.011

ResearchGate.net/publication/289500812_Is_the_Genyornis_egg_of_a_mihirung_or_another_extinct_bird_from_the_Australian_dreamtime