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Topic:Evolutionary Biology

Evolutionary biology and horses focus on the study of the evolutionary processes that have shaped the development, adaptation, and diversification of the Equus genus. This field examines the genetic, morphological, and ecological changes that have occurred over millions of years, leading to the modern horse. Researchers explore the transition from small, multi-toed ancestors to the large, single-toed horses of today, analyzing fossil records, genetic data, and environmental factors that influenced these changes. This page compiles peer-reviewed research studies and scholarly articles that investigate the evolutionary history, genetic adaptations, and ecological interactions of horses, providing insights into their development and survival strategies throughout history.
Climate warming and humans played different roles in triggering Late Quaternary extinctions in east and west Eurasia.
Proceedings. Biological sciences    March 24, 2017   Volume 284, Issue 1851 20162438 doi: 10.1098/rspb.2016.2438
Wan X, Zhang Z.Climate change and humans are proposed as the two key drivers of total extinction of many large mammals in the Late Pleistocene and Early Holocene, but disentangling their relative roles remains challenging owing to a lack of quantitative evaluation of human impact and climate-driven distribution changes on the extinctions of these large mammals in a continuous temporal-spatial dimension. Here, our analyses showed that temperature change had significant effects on mammoth (genus ), rhinoceros (Rhinocerotidae), horse (Equidae) and deer (Cervidae). Rapid global warming was the predominant factor...
Genome-Wide Identification and Evolutionary Analysis of Sarcocystis neurona Protein Kinases.
Pathogens (Basel, Switzerland)    March 21, 2017   Volume 6, Issue 1 12 doi: 10.3390/pathogens6010012
Murungi EK, Kariithi HM.The apicomplexan parasite Sarcocystis neurona causes equine protozoal myeloencephalitis (EPM), a degenerative neurological disease of horses. Due to its host range expansion, S. neurona is an emerging threat that requires close monitoring. In apicomplexans, protein kinases (PKs) have been implicated in a myriad of critical functions, such as host cell invasion, cell cycle progression and host immune response evasion. Here, we used various bioinformatics methods to define the kinome of S. neurona and phylogenetic relatedness of its PKs to other apicomplexans. We identified 97 putative PKs clust...
Repetitive mammalian dwarfing during ancient greenhouse warming events.
Science advances    March 15, 2017   Volume 3, Issue 3 e1601430 doi: 10.1126/sciadv.1601430
D'Ambrosia AR, Clyde WC, Fricke HC, Gingerich PD, Abels HA.Abrupt perturbations of the global carbon cycle during the early Eocene are associated with rapid global warming events, which are analogous in many ways to present greenhouse warming. Mammal dwarfing has been observed, along with other changes in community structure, during the largest of these ancient global warming events, known as the Paleocene-Eocene Thermal Maximum [PETM; ~56 million years ago (Ma)]. We show that mammalian dwarfing accompanied the subsequent, smaller-magnitude warming event known as Eocene Thermal Maximum 2 [ETM2 (~53 Ma)]. Statistically significant decrease in body size...
The legacy of Columbus in American horse populations assessed by microsatellite markers.
Journal of animal breeding and genetics = Zeitschrift fur Tierzuchtung und Zuchtungsbiologie    February 14, 2017   Volume 134, Issue 4 340-350 doi: 10.1111/jbg.12255
Cortés O, Dunner S, Gama LT, Martínez AM, Delgado JV, Ginja C, Jiménez LM, Jordana J, Luis C, Oom MM, Sponenberg DP, Zaragoza P, Vega-Pla JL.Criollo horse populations descend from horses brought from the Iberian Peninsula over the period of colonization (15th to 17th century). They are spread throughout the Americas and have potentially undergone genetic hybridization with other breeds in the recent past. In this study, 25 autosomal microsatellites were genotyped in 50 horse breeds representing Criollo populations from 12 American countries (27 breeds), breeds from the Iberian Peninsula (19), one breed each from France and Morocco and two cosmopolitan horse breeds (Thoroughbred and Arabian). The genetic relationships among breeds i...
Decoupled ecomorphological evolution and diversification in Neogene-Quaternary horses.
Science (New York, N.Y.)    February 12, 2017   Volume 355, Issue 6325 627-630 doi: 10.1126/science.aag1772
Cantalapiedra JL, Prado JL, Hernández Fernández M, Alberdi MT.Evolutionary theory has long proposed a connection between trait evolution and diversification rates. In this work, we used phylogenetic methods to evaluate the relationship of lineage-specific speciation rates and the mode of evolution of body size and tooth morphology in the Neogene and Quaternary radiation of horses (7 living and 131 extinct species). We show that diversification pulses are a recurrent feature of equid evolution but that these pulses are not correlated with rapid bursts in phenotypic evolution. Instead, rapid cladogenesis seems repeatedly associated with extrinsic factors t...
Complete mitochondrial genome of an extinct Equus (Sussemionus) ovodovi specimen from Denisova cave (Altai, Russia).
Mitochondrial DNA. Part B, Resources    February 6, 2017   Volume 2, Issue 1 79-81 doi: 10.1080/23802359.2017.1285209
Druzhkova AS, Makunin AI, Vorobieva NV, Vasiliev SK, Ovodov ND, Shunkov MV, Trifonov VA, Graphodatsky AS. is an extinct subgenus of first characterized and delineated in 2010. The almost complete mitochondrial genome is available only for a single specimen of - a 40,000 years old from Proskuryakova cave (Khakassia, Russia). Our studies of ancient horses from Denisova cave (Altai, Russia) revealed mitochondrial DNA of this species in a 32,000 years old sample. Using alignments to multiple mitochondrial genomes of non-caballine equids, we recovered 100% complete mitochondrial genome of for the first time. Phylogenetic analysis demonstrates close relationship between this individual and the one ...
[The importance of conservation of rare domestic animal breeds using the example of Arc Warder].
Tierarztliche Praxis. Ausgabe G, Grosstiere/Nutztiere    January 10, 2017   Volume 45, Issue 1 33-41 doi: 10.15653/TPG-160777
Frölich K, Jandowsky A.The domestic animals descend from various wild ancestors. Thus, for example, of the wolf, the dog (15  000 BC), of the Bezoar goat the goat (10  000 BC), of the Asian mouflon the sheep (10  000 BC), of the wild boar the pig (8000 BC) and of the aurochs the cattle (7000 BC). Domestication has dramatically changed our culture and led to a great diversity of animal breeds. This is a unique cultural and historical treasure, which we have to preserve for future generations. The zoological park Arc Warder is Europe's largest center for rare and endangered domestic animal breeds. Arc Ward...
Differential Infection Patterns and Recent Evolutionary Origins of Equine Hepaciviruses in Donkeys.
Journal of virology    December 16, 2016   Volume 91, Issue 1 e01711-16 doi: 10.1128/JVI.01711-16
Walter S, Rasche A, Moreira-Soto A, Pfaender S, Bletsa M, Corman VM, Aguilar-Setien A, García-Lacy F, Hans A, Todt D, Schuler G....The hepatitis C virus (HCV) is a major human pathogen. Genetically related viruses in animals suggest a zoonotic origin of HCV. The closest relative of HCV is found in horses (termed equine hepacivirus [EqHV]). However, low EqHV genetic diversity implies relatively recent acquisition of EqHV by horses, making a derivation of HCV from EqHV unlikely. To unravel the EqHV evolutionary history within equid sister species, we analyzed 829 donkeys and 53 mules sampled in nine European, Asian, African, and American countries by molecular and serologic tools for EqHV infection. Antibodies were found in...
Diversity of mitochondrial DNA in three Arabian horse strains.
Journal of applied genetics    December 14, 2016   Volume 58, Issue 2 273-276 doi: 10.1007/s13353-016-0384-z
Almarzook S, Reissmann M, Brockmann GA.Arabian horse registries classify Arabian horses based on their dam lineages into five main strains. To test the maternal origin of Syrian Arabian horses, 192 horses representing the three major strains Saglawi, Kahlawi, and Hamdani were sequenced for 353 bp of their mitochondrial displacement loop (D-loop) region. Sequencing revealed 28 haplotypes comprising 38 sequence variations. The haplotype diversity values were 0.95, 0.91, and 0.90 in Kahlawi, Hamdani, and Saglawi strains, respectively. The pair-wise population differentiation estimates (Fst) between strains were low, ranging between 0...
Spotted phenotypes in horses lost attractiveness in the Middle Ages.
Scientific reports    December 7, 2016   Volume 6 38548 doi: 10.1038/srep38548
Wutke S, Benecke N, Sandoval-Castellanos E, Döhle HJ, Friederich S, Gonzalez J, Hallsson JH, Hofreiter M, Lõugas L, Magnell O, Morales-Muniz A....Horses have been valued for their diversity of coat colour since prehistoric times; this is especially the case since their domestication in the Caspian steppe in ~3,500 BC. Although we can assume that human preferences were not constant, we have only anecdotal information about how domestic horses were influenced by humans. Our results from genotype analyses show a significant increase in spotted coats in early domestic horses (Copper Age to Iron Age). In contrast, medieval horses carried significantly fewer alleles for these phenotypes, whereas solid phenotypes (i.e., chestnut) became domina...
Runs of homozygosity: current knowledge and applications in livestock.
Animal genetics    December 1, 2016   Volume 48, Issue 3 255-271 doi: 10.1111/age.12526
Peripolli E, Munari DP, Silva MVGB, Lima ALF, Irgang R, Baldi F.This review presents a broader approach to the implementation and study of runs of homozygosity (ROH) in animal populations, focusing on identifying and characterizing ROH and their practical implications. ROH are continuous homozygous segments that are common in individuals and populations. The ability of these homozygous segments to give insight into a population's genetic events makes them a useful tool that can provide information about the demographic evolution of a population over time. Furthermore, ROH provide useful information about the genetic relatedness among individuals, helping t...
Polymorphism at expressed DQ and DR loci in five common equine MHC haplotypes.
Immunogenetics    November 26, 2016   Volume 69, Issue 3 145-156 doi: 10.1007/s00251-016-0964-4
Miller D, Tallmadge RL, Binns M, Zhu B, Mohamoud YA, Ahmed A, Brooks SA, Antczak DF.The polymorphism of major histocompatibility complex (MHC) class II DQ and DR genes in five common equine leukocyte antigen (ELA) haplotypes was determined through sequencing of mRNA transcripts isolated from lymphocytes of eight ELA homozygous horses. Ten expressed MHC class II genes were detected in horses of the ELA-A3 haplotype carried by the donor horses of the equine bacterial artificial chromosome (BAC) library and the reference genome sequence: four DR genes and six DQ genes. The other four ELA haplotypes contained at least eight expressed polymorphic MHC class II loci. Next generation...
Density-dependent resource selection by a terrestrial herbivore in response to sea-to-land nutrient transfer by seals.
Ecology    November 20, 2016   Volume 97, Issue 8 1929-1937 doi: 10.1002/ecy.1451
McLoughlin PD, Lysak K, Debeffe L, Perry T, Hobson KA.Sea-to-land nutrient transfers can connect marine food webs to those on land, creating a dependence on marine webs by opportunistic species. We show how nitrogen, imported by gray seals, Halichoerus grypus, and traced through stable isotope (δ N) measurements in marram grass, Ammophila breviligulata, significantly alters foraging behavior of a free-roaming megaherbivore (feral horses, Equus ferus caballus) on Sable Island, Canada. Values of δ N correlated with protein content of marram and strongly related to pupping-seal densities, and positively influenced selective foraging by horses. The...
A pilot study on interaction between donkey tetherin and EIAV stains with different virulent and replication characteristics.
Microbial pathogenesis    November 2, 2016   Volume 106 65-68 doi: 10.1016/j.micpath.2016.10.019
Yao Q, Ma J, Wang X, Guo M, Li Y, Wang X.Tetherin (BST-2) is an important host restriction factor that can inhibit the release of a diverse array of enveloped viruses from infected cells. Conversely, to facilitate their release and spread, many viruses have evolved various strategies to overcome the antiviral effect of tetherin in a species-specific manner. During the development of an attenuated equine infectious anemia virus (EIAV) vaccine in our laboratory, we found that serial passage of a field-isolated virulent EIAV strains in horse and donkey as well as the cultivated donkey cells, produces several typical EIAV strains, includ...
The Evolutionary Origin and Genetic Makeup of Domestic Horses.
Genetics    October 13, 2016   Volume 204, Issue 2 423-434 doi: 10.1534/genetics.116.194860
Librado P, Fages A, Gaunitz C, Leonardi M, Wagner S, Khan N, Hanghøj K, Alquraishi SA, Alfarhan AH, Al-Rasheid KA, Der Sarkissian C, Schubert M....The horse was domesticated only 5.5 KYA, thousands of years after dogs, cattle, pigs, sheep, and goats. The horse nonetheless represents the domestic animal that most impacted human history; providing us with rapid transportation, which has considerably changed the speed and magnitude of the circulation of goods and people, as well as their cultures and diseases. By revolutionizing warfare and agriculture, horses also deeply influenced the politico-economic trajectory of human societies. Reciprocally, human activities have circled back on the recent evolution of the horse, by creating hundreds...
Genetic diversity and population structure of Kazakh horses (Equus caballus) inferred from mtDNA sequences.
Genetics and molecular research : GMR    October 5, 2016   Volume 15, Issue 4 doi: 10.4238/gmr.15048618
Gemingguli M, Iskhan KR, Li Y, Qi A, Wunirifu W, Ding LY, Wumaierjiang A.The Kazakh horse is an important old horse breed in Xinjiang. They have contributed greatly to the breeding and improvement of other local horse breeds, yet their genetic diversity and population structure are not well understood. In the present study, we evaluated the genetic diversity of Kazakh horses and their relationship with other horse breeds using the mtDNA D-loop region, Cyt b gene, and a DNA fragment (nps 7974-9963, containing COX3, tRNA-Gly, ND3, and tRNA-Arg). A total of 130 Kazakh horses from 8 populations in China and Kazakhstan were analyzed. A total of 88 haplotypes (haplotype ...
Identification, characterization and bioactivity of tumor necrosis factor (TNF)-related apoptosis-inducing ligand from Equus caballus.
Developmental and comparative immunology    August 31, 2016   Volume 67 340-349 doi: 10.1016/j.dci.2016.08.018
Ma L, Sang M, Zhang J, Zhang S.Tumor necrosis factor (TNF)-related apoptosis-inducing ligand (TRAIL) is a member of the TNF superfamily and plays multifunctional roles in the immune system. In the present study, a homolog of TRAIL from the Mongolian horse (named ecTRAIL) was identified and characterized. The 870-bp open reading frame encodes a polypeptide of 289 amino acid residues with a predicted molecular weight of 33.47 kDa and pI of 8.47. The genomic structure of ecTRAIL shares a five-exon/six-intron arrangement similar to its orthologs. Multiple alignments show that ecTRAIL is a type II transmembrane protein with a t...
[Horse, cow and reindeer were converted into arctic domestic animals].
Duodecim; laaketieteellinen aikakauskirja    August 16, 2016   Volume 132, Issue 13-14 1231-1236 
Kantanen J.Domestic animal production in the arctic region is often thought to be based exclusively on reindeer herding. There are, however, regions in Northern Europe and Siberia having a long tradition in rearing breeds of cattle and horse adapted to the northers conditions also. The development of these arctic animal breeds has been largely founded on old tradition rather than on the programs of breeding organizations. As a result of the selection carried out by nature and man, the domestic animals of arctic regions express characteristics that are metabolic, structural, associated with reproductive p...
The origin of ambling horses.
Current biology : CB    August 10, 2016   Volume 26, Issue 15 R697-R699 doi: 10.1016/j.cub.2016.07.001
Wutke S, Andersson L, Benecke N, Sandoval-Castellanos E, Gonzalez J, Hallsson JH, Lõugas L, Magnell O, Morales-Muniz A, Orlando L, Pálsdóttir AH....Horseback riding is the most fundamental use of domestic horses and has had a huge influence on the development of human societies for millennia. Over time, riding techniques and the style of riding improved. Therefore, horses with the ability to perform comfortable gaits (e.g. ambling or pacing), so-called 'gaited' horses, have been highly valued by humans, especially for long distance travel. Recently, the causative mutation for gaitedness in horses has been linked to a substitution causing a premature stop codon in the DMRT3 gene (DMRT3_Ser301STOP) [1]. In mice, Dmrt3 is expressed in spinal...
ANIMAL BEHAVIOR AND WELL-BEING SYMPOSIUM: Interaction between coping style/personality, stress, and welfare: Relevance for domestic farm animals.
Journal of animal science    June 11, 2016   Volume 94, Issue 6 2284-2296 doi: 10.2527/jas.2015-0125
Koolhaas JM, Van Reenen CG.This paper will argue that understanding animal welfare and the individual vulnerability to stress-related disease requires a fundamental understanding of functional individual variation as it occurs in nature as well as the underlying neurobiology and neuroendocrinology. Ecological studies in feral populations of mice, fish, and birds start to recognize the functional significance of phenotypes that individually differ in their behavioral and neuroendocrine response to environmental challenge. Recent studies indicate that the individual variation within a species may buffer the species for st...
Convergent evolution of pregnancy-specific glycoproteins in human and horse.
Reproduction (Cambridge, England)    June 8, 2016   Volume 152, Issue 3 171-184 doi: 10.1530/REP-16-0236
Aleksic D, Blaschke L, Mißbach S, Hänske J, Weiß W, Handler J, Zimmermann W, Cabrera-Sharp V, Read JE, de Mestre AM, O'Riordan R, Moore T....Pregnancy-specific glycoproteins (PSGs) are members of the carcinoembryonic antigen cell adhesion molecule (CEACAM) family that are secreted by trophoblast cells. PSGs may modulate immune, angiogenic and platelet responses during pregnancy. Until now, PSGs are only found in species that have a highly invasive (hemochorial) placentation including humans, mice and rats. Surprisingly, analyzing the CEACAM gene family of the horse, which has a non-invasive epitheliochorial placenta, with the exception of the transient endometrial cups, we identified equine CEACAM family members that seem to be rel...
Equine rhinitis B viruses in horse fecal samples from the Middle East.
Virology journal    June 7, 2016   Volume 13 94 doi: 10.1186/s12985-016-0547-x
Woo PC, Lau SK, Choi GK, Huang Y, Wernery R, Joseph S, Wong EY, Elizabeth SK, Patteril NA, Li T, Wernery U, Yuen KY.Among all known picornaviruses, only two species, equine rhinitis A virus and equine rhinitis B virus (ERBV) are known to infect horses, causing respiratory infections. No reports have described the detection of ERBV in fecal samples of horses and no complete genome sequences of ERBV3 are available. Methods: We performed a molecular epidemiology study to detect ERBVs in horses from Dubai and Hong Kong. Complete genome sequencing of the ERBVs as well as viral loads and genome, phylogenetic and evolutionary analysis were performed on the positive samples. Results: ERBV was detected in four (13.8...
Leaping eels electrify threats, supporting Humboldt’s account of a battle with horses.
Proceedings of the National Academy of Sciences of the United States of America    June 6, 2016   Volume 113, Issue 25 6979-6984 doi: 10.1073/pnas.1604009113
Catania KC.In March 1800, Alexander von Humboldt observed the extraordinary spectacle of native fisherman collecting electric eels (Electrophorus electricus) by "fishing with horses" [von Humboldt A (1807) Ann Phys 25:34-43]. The strategy was to herd horses into a pool containing electric eels, provoking the eels to attack by pressing themselves against the horses while discharging. Once the eels were exhausted, they could be safely collected. This legendary tale of South American adventures helped propel Humboldt to fame and has been recounted and illustrated in many publications, but subsequent investi...
Mitochondrial diversity in mountain horse population from the South-Eastern Europe.
Mitochondrial DNA. Part A, DNA mapping, sequencing, and analysis    June 1, 2016   Volume 28, Issue 6 787-792 doi: 10.1080/24701394.2016.1186667
Hristov P, Yordanov G, Ivanova A, Mitkov I, Sirakova D, Mehandzyiski I, Radoslavov G.It is believed that population structure of mountain horse breeds is preserved from any genetic introgression, because of their geographical isolation and specific semi-wild life style of husbandry. Till date there are no molecular data for the Balkan horses. In this study we try to give information about some autochthonous mountain horse breeds from Bulgaria. A total of 121 horses from three different mountain massifs are presented: Stara Planina (the Balkan mountain), the Rhodopes and Rila-Pirin massif were genotyped according to mitochondrial D-loop region. The results showed huge diversity...
Polymorphism and preadaptation of horses (Equus) of the mountains surrounding Lake Baikal in Pleistocene.
Doklady biological sciences : proceedings of the Academy of Sciences of the USSR, Biological sciences sections    May 20, 2016   Volume 467, Issue 1 59-62 doi: 10.1134/S0012496616020010
Kalmykov NP.The first data on polymorphism and preadaptation of the Pleistocene horses in the mountains surrounding Lake Baikal are presented. It has been shown that disregard of intraspecific polymorphism leads to unreasonable ignoring of their diversity, phylogenetically false constructions, and incorrect interpretation of natural environment.
Transposable Element Targeting by piRNAs in Laurasiatherians with Distinct Transposable Element Histories.
Genome biology and evolution    May 9, 2016   Volume 8, Issue 5 1327-1337 doi: 10.1093/gbe/evw078
Vandewege MW, Platt RN, Ray DA, Hoffmann FG.PIWI proteins and PIWI-interacting RNAs (piRNAs) are part of a cellular pathway that has evolved to protect genomes against the proliferation of transposable elements (TEs). PIWIs and piRNAs assemble into complexes that are involved in epigenetic and post-transcriptional repression of TEs. Most of our understanding of the mechanisms of piRNA-mediated TE silencing comes from fruit fly and mouse models. However, even in these well-studied animals it is unclear how piRNA responses relate to variable TE expression and whether the strength of the piRNA response affects TE content over time. Here, w...
The Brain of the Domestic Bos taurus: Weight, Encephalization and Cerebellar Quotients, and Comparison with Other Domestic and Wild Cetartiodactyla.
PloS one    April 29, 2016   Volume 11, Issue 4 e0154580 doi: 10.1371/journal.pone.0154580
Ballarin C, Povinelli M, Granato A, Panin M, Corain L, Peruffo A, Cozzi B.The domestic bovine Bos taurus is raised worldwide for meat and milk production, or even for field work. However the functional anatomy of its central nervous system has received limited attention and most of the reported data in textbooks and reviews are derived from single specimens or relatively old literature. Here we report information on the brain of Bos taurus obtained by sampling 158 individuals, 150 of which at local abattoirs and 8 in the dissecting room, these latter subsequently formalin-fixed. Using body weight and fresh brain weight we calculated the Encephalization Quotient (EQ)...
New insights on equid locomotor evolution from the lumbar region of fossil horses.
Proceedings. Biological sciences    April 29, 2016   Volume 283, Issue 1829 doi: 10.1098/rspb.2015.2947
Jones KE.The specialization of equid limbs for cursoriality is a classic case of adaptive evolution, but the role of the axial skeleton in this famous transition is not well understood. Extant horses are extremely fast and efficient runners, which use a stiff-backed gallop with reduced bending of the lumbar region relative to other mammals. This study tests the hypothesis that stiff-backed running in horses evolved in response to evolutionary increases in body size by examining lumbar joint shape from a broad sample of fossil equids in a phylogenetic context. Lumbar joint shape scaling suggests that st...
Revelation of Influencing Factors in Overall Codon Usage Bias of Equine Influenza Viruses.
PloS one    April 27, 2016   Volume 11, Issue 4 e0154376 doi: 10.1371/journal.pone.0154376
Kumar N, Bera BC, Greenbaum BD, Bhatia S, Sood R, Selvaraj P, Anand T, Tripathi BN, Virmani N.Equine influenza viruses (EIVs) of H3N8 subtype are culprits of severe acute respiratory infections in horses, and are still responsible for significant outbreaks worldwide. Adaptability of influenza viruses to a particular host is significantly influenced by their codon usage preference, due to an absolute dependence on the host cellular machinery for their replication. In the present study, we analyzed genome-wide codon usage patterns in 92 EIV strains, including both H3N8 and H7N7 subtypes by computing several codon usage indices and applying multivariate statistical methods. Relative synon...
Comparative genomics of hormonal signaling in the chorioallantoic membrane of oviparous and viviparous amniotes.
General and comparative endocrinology    April 19, 2016   Volume 244 19-29 doi: 10.1016/j.ygcen.2016.04.017
Griffith OW, Brandley MC, Whittington CM, Belov K, Thompson MB.In oviparous amniotes (reptiles, birds, and mammals) the chorioallantoic membrane (CAM) lines the inside of the egg and acts as the living point of contact between the embryo and the outside world. In livebearing (viviparous) amniotes, communication during embryonic development occurs across placental tissues, which form between the uterine tissue of the mother and the CAM of the embryo. In both oviparous and viviparous taxa, the CAM is at the interface of the embryo and the external environment and can transfer signals from there to the embryo proper. To understand the evolution of placental ...
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