Tremarctinae: Versiyalar orasidagi farq

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„Tremarctinae“ sahifasi tarjima qilib yaratildi
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8-Oktyabr 2022, 05:18 dagi koʻrinishi

Tremarctinae

Temporal range: late Miocene–present
A spectacled bear in Tennōji Zoo, Osaka.
Arctodus simus ning skleti
Ilmiy tasniflash e
Olam: Hayvonlar
Tip: Xordalilar
Sinf: Sutemizuvchilar
Oila Ursidae
Kemja tur: Tremarctinae

Merriam & Stock, 1925
Urug'

†Plionarctos†Arctodus†Arctotherium'Tremarctos

Tremarctinae yoki kalta yuzli ayiqlar - Ursidae turkumining kichik oilasi a'zosi boʻlib, uning bitta tirik vakili, Janubiy Amerikaning koʻzoynakli ayigʻi (Tremarctos ornatus) va toʻrt avlodga mansub bir nechta yoʻqolib ketgan turlar: Florida koʻzoynakli ayigʻi (Tremarctos floridanus), Shimoliy Amerika giganti - kalta yuzli ayiqlar Arctodus ( A. pristinus va A. simus ), Janubiy Amerika giganti - kalta yuzli ayiq Arctotherium (jumladan, A. angustidens, A. vetustum, A. bonariense, A. wingei va A. tarijense) hamda Plionarktos (P. edensis va P. harroldorum), qolgan uchta avlodning ajdodi hisoblanadi[1]. Ular orasidagi qisqa yuzli gigant ayiqlari (Arctodus simus va Arctotherium angustidens) Amerikadagi eng yirik yirtqich hayvonlar bo'lgan bo'lishi mumkin. Guruh Shimoliy Amerikaning sharqiy qismida yashagan va keyinchalik Buyuk Amerika almashinuvining bir qismi sifatida Janubiy Amerika hududiga ko'chib o'tgan deya taxmin qilinadi. Ko'pchilik qisqa yuzli ayiqlar pleystotsen oxirida yo'q bo'lib ketishgan.

Evolyutsiya

Arctotherium bonariense

Tremarktinalar Shimoliy Amerikaning O'rta Gemfiliya davrida (eng qadimgi so'nggi Miosen, ~10M) umumiy ajdodi Plionarktosdan kelib chiqqan; Plionarktosning oxirgi vakili Blankanning boshida (erta Pliotsen, ~3,3 milya) yashagnligi qayd etilgan. Miosen - Pliotsen chegarasi (~5 million) atrofida tremarktinlar boshqa ursidlar qatori xilma-xillikda portlovchi nurlanishni boshdan kechirdi, chunki usha davrda C4 o'simliklari ( o'tlar ) va ochiq yashash joylari katta maydonni qamrab olgan, dunyoda harorat sezilarli darajada pasaydi va mavsumiylik kuchaydi shuningdek, fauna aylanmasi butun Yevrosiyo faunasining 60-70 foizini va Shimoliy Amerika o'simliklarining 70-80 foizini yo'q bo'lib ketishiga sabab bo'ldi[1]. Shunga mos ravishda, 2008-yilgi tadqiqot shuni ko'rsatdiki, Arctotherium, Arctodus va Tremarctos uchun o'rtacha farqlanish sanalari 4,8 million yilni, Arctotherium va Tremarktos o'rtasidagi farq esa 4,1 million yilni tashkil etadi[2]. Keyingi tadqiqot Arctodus va Tremarktos o'rtasidagi ajralish vaqtini taxminan 5,5 million yil ichida sodir bo'lganligini hisoblab chiqdi[3]. Har uchala avlod birinchi boʻlib Shimoliy Amerikaning Blankan (Pliotsen/Pleystotsen chegarasi) davridan qayd etilgan[4].

Noaniq Arctotherium birinchi marta El Salvadorning so'nggi Pliotsen davridan boshlab uchrashi aniqlangan[2]. Erta pleystosenda qisqa yuzli ayiqlar Shimoliy va Janubiy Amerikada katta populyatsiyasini qayd etishgan[5]. O'rta kattalikdagi Arctodus pristinus Shimoliy Amerika qit'asida keng miqyosda yashagan, Tremarctos floridanus Fors ko'rfazi sohiliga xos jonivordir. Janubiy Amerikadagi Arctotheriumning uchrash vaqtiga doir qaydlar Argentinada taxminan 1 million yil davomida ulkan Arctotherium angustidens bo'lganligini hamda ehtimol, u quruqlikdagi eng katta yirtqich sutemizuvchilardan biri ekanligi taxmin qilinadi. Arctotheriumning oldingi 1,5 million yildagi evolyutsion tarixi va ularning Janubiy Amerikadagi tarixi fanga noma'lum. O'rta pleystosenda Arctodus ham<i id="mwXQ">, Arctotherium</i> ham yangi shakllarga yo'l berdi; Arctodus pristinus o'z o'rnini Alyaskadan Meksikagacha bo'lgan butun qit'a oralig'ida yashovchi ulkan Arctodus simusiga berdi[6]. Arctotherium angustidens, boshqa tomondan, bir qator kichikroq, o'rta o'lchamli tur - Arctotherium vetustumlar bilan almashildi birozdan keyin Arctotherium bonariense va Arctotherium tarijense turlari rivojlandi[5]. Kichikroq Arctotherium wingei faqat kech Pleystotsen yozuvlaridan ma'lum bo'lsa-da, Arctotherium oila daraxtida turning ko'proq arxaik mavqei O'rta Pleystotsenda kelib chiqqanligini ko'rsatadi. Arctotherium wingei Arctotheriumning yagona ma'lum turi bo'lib, asosan janubiy konusning shimolida joylashgan[7] va Markaziy Amerika hududlarida uchragan[8].

Pleystotsen terminalida Arctodus simus, Tremarctos floridanus, Arctotherium tarijense va Arctotherium wingei birgalikda Alyaskadan Patagoniyaning janubiy qismigacha bo'lgan hududlarda yashagan. Bu shakllarning barchasi erta golosen oxirida yo'q bo'lib ketgan. Taxminan o'sha paytda, ko'zoynakli ayiq deb nomlanuvchi Tremarctos ornatus Janubiy Amerikaning fotoalbom yozuvlarida uchraydi[9]. Olimlarning ta'kidlashicha, ko'zoynakli ayiqlar Arctotherium wingei yo'q bo'lib ketganidan keyin Markaziy va Janubiy Amerikaga ko'chib o'tgan, Andean yozuvlari Arctotherium sp.ni ko'zoynakli ayiq sifatida tasvirlamagan[10][11].

Anatomiya

Tremarctinae zamonaviy ayiqlarga nisbatan nomutanosib ravishda qisqa tumshuqqa ega bo'lib, bu holat ularga "qisqa yuzli" nomini berilishiga sabab bo'laadi. Bu zohiriy kaltalik ursinae ayiqlari bilan solishtirganda tremarktin ayiqlarining chuqur tumshug'i va kalta burun suyaklari tufayli yuzaga kelgan illyuziya ekanligini anglash mumkin; Tremarctinae ko'pchilik tirik ayiqlarga qaraganda chuqurroq, ammo qisqaroq yuz tuzilishiga ega bo'lgan[12]. Tremarktin va ursin ayiqlari o'rtasidagi osteoologik farqlarga, m1 molaridagi trigonid va talonid o'rtasidagi qo'shimcha yonboshdagi do'nglik, mandibulada premasseterik chuqurcha va ko'pincha tremarktin ayiqlarining son suyagidagi entepikondilar teshik kabilar kiradi[13]. Bundan tashqari, tremarktin ayiqlarning bosh suyagi chuqurroq va braxitsefal bo'lib, ularning zigomatik yoylari va glenoid chuqurchalari yaxshi rivojlangan va ursinlarga nisbatan kattaroq molarlarga ega. Bundan tashqari, tremarktin ayiqlarining orbitalari ham kattaroq, yumaloqroq va laterallashgan[12].

Paleobiologiya

Tremarktinlar chuqurliklarda yashagan - Arctotherium wingei hamda Tremarctos ornatus kabi ko'proq o'rmon hududida yashaydigan kichik va asosan o'txo'r ayiqlardan ulkan go'shtxo'r va moslashuvchan ayiq turlari Arctotherium angustidens va Arctodus simusgacha bo'lgan barcha ayiq turlarini qamrab oladi. Ikki gigant tur yuzaki jihatlariga kora o'xshash bo'lib ko'rinsa-da, aslida ikkala tur ham asosiy, sezilarli farqlarga ega edi. Arctodus simus Shimoliy Amerika hududida 800 000 yil davomida yashagan, Arctotherium angustidens ochiq tekisliklarda joylashgan Janubiy konus maydonlari bilan cheklangan hududlarda uchragan. Bundan tashqari, Arctodus simus o'z ovqatlanish ratsionini Alyaskadagi kvazi yirtqich hayvonlar ovqatlanishidan klassik omnivor holatiga o'zgartirgan bo'lsa, izotop tadqiqotlariga ko'ra, Arctotherium angustidens namunalari ularning haqiiqy yirtqich va go'shtxo'r bo'lganliklarini ko'rsatadi[14][15]. Bundan tashqari, kuchli Arctotherium angustidensdan farqli o'laroq, Arctodusning ancha nozik tana shakli tadqiqotchilarni hayratda qoldirgan[16][17]. Biroq, Janubiy Amerikadagi <i>Chapalmalaniya</i> prokionidlari hamda Shimoliy Amerikadagi Borofag va Agrioterium kabi yirtqich hayvonlarning pliotsen davrida yo'q bo'lib ketishi Arktod va Arktoteriy singari turlarda gigantizmga umumiy turtki bo'lganligi taxmin qilinadi[4].

Arctodus va Tremarktos turlari o'txo'r ayiqlarga xos xususiyatlarga ega. Katta yonoq tishlari, chuqur ostki jag' va katta yuqorigi jag' mushaklariga ko'ra shunday xulosaga kelish mumkin. O'txo'r yirtqich hayvonlarda mikroblar ta'sirida o'simlik moddalarini parchalash uchun samarali ovqat hazm qilish tizimi yo'qligi sababli, ular o'simlik moddalarini uzoq chaynash yoki panjalarida maydalash orqali parchalashlari kerak bo'lgan hamda shu bilan jag'ning yuqori mexanik ustunligini [aydo qilish uchun ayrim xususiyatlarga ega bo'lishi kerak bo'lgan. Agarda jag'ning bunday tuzilishi ovqatlanish bilan bog'liq bo'lmasa, u holda bu uning ajdodlaridan o'tgan bo'lishi mumkin degan taxminlarga sabab bo'ladi[18].

Sistematika

Tremarctinae within Ursidae
 

daggerHemicyoninae

 
 
 

daggerUrsavinae

 
 
 

daggerAgriotheriinae

 
 
 

Ailuropodinae

 
 
 

Ursinae

 
 
Tremarctinae (short‑faced bears)
 

daggerPlionarctos

 
 
 

daggerArctodus

 
 
 

Tremarctos

 
 

daggerArctotherium

 
 
 
 
 
 
 
 
 
Tremarctinae within Ursidae
 

daggerHemicyoninae

 
 
 

daggerUrsavinae

 
 
 

daggerAgriotheriinae

 
 
 

Ailuropodinae

 
 
 

Ursinae

 
 
Tremarctinae (short‑faced bears)
 

daggerPlionarctos

 
 
 

daggerArctodus

 
 
 

Tremarctos

 
 

daggerArctotherium

 
 
 
 
 
 
 
 
 

An'anaga ko'ra, tremarktinlarning filogenetik ichki munosabatlarini tahlil qilish davomida, Arctodus va Arctotheriumning qisqa yuzli ayiq to'dasiga nisbatan basal guruhlar sifatida shakllanishi Plionarktos va Tremarktos davrida kuzatilganligi aniqlangan[11][17]. Arctotheriumga tegishli ayiqlarning yaqinliklarini o'rganish bilan bog'liq tadqiqotlar shuni ko'rsatadiki, ular Arktoddan ko'ra ko'zoynakli ayiq bilan chambarchas bog'liq bo'lib, bu ikki nasldagi katta hajmdagi konvergent evolyutsiyani paydo bo'lgan[19].

Tremarktinlar vaqti-vaqti bilan eski ilmiy adabiyotlarda arktodontlar yoki arktoterlar[20] singari nomlar bilan uchraydi, ammo bu nomlar bugungi kunda qo'llanilmaydi.

Taksonomiya

Tremarktin ayiqlarining quyidagi taksonomiyasi Mitchell va boshqa olimlar tomonidan o'rganilgan (2016-yil)[19]:

  • Tremarctinae kenja oilasi (Merriam & Stock, 1925)
    • † Plionartos (Frik, 1926)
      • † Plionactos harroldorum (Tedfored va Martin, 2001)
      • † Plionactos edensis (Frik, 1926)
    • † Arctodus (Leidy, 1854)
      • † Arctodus simus (Cope, 1879)
      • † Arctodus pristinus (Leidy, 1854)
    • † Arctotherium (Burmeister, 1879)
      • † Arctotherium angustidens (Gervais & Ameghino, 1880)
      • † Arctotherium vetustum (Ameghino, 1885)
      • † Arctotherium wingei (Ameghino, 1902)
      • † Arctotherium bonariense (Gervais, 1852)
      • † Arctotherium tarijense (Ameghino, 1902)
    • Tremarctos (Gervais, 1855)

Manbalar

  1. ↑ 1,0 1,1 Krause, J.; Unger, T.; Noçon, A.; Malaspinas, A.; Kolokotronis, S.; Stiller, M.; Soibelzon, L.; Spriggs, H. et al. (2008-07-28). "Mitochondrial genomes reveal an explosive radiation of extinct and extant bears near the Miocene-Pliocene boundary". BMC Evolutionary Biology 8: 220. doi:10.1186/1471-2148-8-220. PMID 18662376. PMC 2518930. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=2518930. 
  2. ↑ 2,0 2,1 Soibelzon, Leopoldo H.; Romero, M.R. Aguilar (2008-10-14). "A Blancan (Pliocene) short-faced bear from El Salvador and its implications for Tremarctines in South America" (en). Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 250 (1): 1–8. doi:10.1127/0077-7749/2008/0250-0001. ISSN 0077-7749. http://www.schweizerbart.de/papers/njgpa/detail/250/59574/A_Blancan_Pliocene_short_faced_bear_from_El_Salvad?af=crossref. 
  3. ↑ Pedersen, Mikkel Winther; De Sanctis, Bianca; Saremi, Nedda F.; Sikora, Martin; Puckett, Emily E.; Gu, Zhenquan; Moon, Katherine L.; Kapp, Joshua D. et al. (June 2021). "Environmental genomics of Late Pleistocene black bears and giant short-faced bears". Current Biology 31 (12): 2728–2736.e8. doi:10.1016/j.cub.2021.04.027. ISSN 0960-9822. https://doi.org/10.1016/j.cub.2021.04.027. 
  4. ↑ 4,0 4,1 Fowler, Nicholas L.; Spady, Thomas J.; Wang, Guiming; Leopold, Bruce D.; Belant, Jerrold L. (2021). "Denning, metabolic suppression, and the realisation of ecological opportunities in Ursidae" (en). Mammal Review 51 (4): 465–481. doi:10.1111/mam.12246. ISSN 1365-2907. https://onlinelibrary.wiley.com/doi/abs/10.1111/mam.12246. 
  5. ↑ 5,0 5,1 Soibelzon, L.H.; Tonni, E.P.; Bond, M. (2005). "The fossil record of South American short-faced bears (Ursidae, Tremarctinae)". Journal of South American Earth Sciences 20 (1–2): 105–113. doi:10.1016/j.jsames.2005.07.005. https://www.researchgate.net/publication/223390695. Qaraldi: 2019-02-21. Tremarctinae]]
  6. ↑ „FIGURE 1—Map showing locations of reported Arctodus simus localities...“ (en). ResearchGate. Qaraldi: 2022-yil 12-fevral.
  7. ↑ „Figura 5. Distribución inferida de algunos mamíferos grandes y...“ (en). ResearchGate. Qaraldi: 2022-yil 12-fevral.
  8. ↑ Schubert, Blaine W.; Chatters, James C.; Arroyo-Cabrales, Joaquin; Samuels, Joshua X.; Soibelzon, Leopoldo H.; Prevosti, Francisco J.; Widga, Christopher; Nava, Alberto et al. (May 2019). "Yucatán carnivorans shed light on the Great American Biotic Interchange". Biology Letters 15 (5): 20190148. doi:10.1098/rsbl.2019.0148. ISSN 1744-9561. PMID 31039726. PMC 6548739. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=6548739. 
  9. ↑ Access Suspended. doi:10.2192/08gr017r1.1. https://bioone.org/access-suspended. Qaraldi: 2022-02-12. Tremarctinae]]
  10. ↑ García-López, Daniel; Ortiz, Pablo; Madozzo Jaén, M.; Moyano, M. (2008-12-12). "First Record of Arctotherium (Ursidae, Tremarctinae) in Northwestern Argentina and its Paleobiogeographic Significance". Journal of Vertebrate Paleontology 28 (4): 1232–1237. doi:10.1671/0272-4634-28.4.1232. https://www.researchgate.net/publication/249023497. 
  11. ↑ 11,0 11,1 Soibelzon, Leopoldo H.; Rincón, Ascanio D. (2007-07-27). "The fossil record of the short-faced bears (Ursidae, Tremarctinae) from Venezuela. Systematic, biogeographic, and paleoecological implications" (en). Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen 244 (3): 287–298. doi:10.1127/0077-7749/2007/0244-0287. ISSN 0077-7749. http://www.schweizerbart.de/papers/njgpa/detail/244/59121/The_fossil_record_of_the_short_faced_bears_Ursidae?af=crossref. 
  12. ↑ 12,0 12,1 Figueirido, Borja; Soibelzon, Leopoldo H. (2010). "Inferring palaeoecology in extinct tremarctine bears (Carnivora, Ursidae) using geometric morphometrics" (en). Lethaia 43 (2): 209–222. doi:10.1111/j.1502-3931.2009.00184.x. ISSN 1502-3931. https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1502-3931.2009.00184.x. 
  13. ↑ Steffen, Martina L.; Fulton, Tara L. (February 2018). "On the association of giant short-faced bear ( Arctodus simus ) and brown bear ( Ursus arctos ) in late Pleistocene North America" (en). Geobios 51 (1): 61–74. doi:10.1016/j.geobios.2017.12.001. https://linkinghub.elsevier.com/retrieve/pii/S0016699517300384. 
  14. ↑ Soibelzon, Leopoldo H.; Grinspan, Gustavo A.; Bocherens, Hervé; Acosta, Walter G.; Jones, Washington; Blanco, Ernesto R.; Prevosti, Francisco (November 2014). "South American giant short-faced bear (Arctotherium angustidens) diet: evidence from pathology, morphology, stable isotopes, and biomechanics" (en). Journal of Paleontology 88 (6): 1240–1250. doi:10.1666/13-143. ISSN 0022-3360. https://www.cambridge.org/core/journals/journal-of-paleontology/article/abs/south-american-giant-shortfaced-bear-arctotherium-angustidens-diet-evidence-from-pathology-morphology-stable-isotopes-and-biomechanics/2616CEF2B348B1453C68EF26CF2858C9. 
  15. ↑ Chichkoyan, Karina; Figueirido, Borja; Belinchón, Margarita; Lanata, José; Moigne, Anne-Marie; Martínez-Navarro, Bienvenido (2017-05-09). "Direct evidence of megamammal-carnivore interaction decoded from bone marks in historical fossil collections from the Pampean region". PeerJ 5: e3117. doi:10.7717/peerj.3117. PMID 28503369. PMC 5426367. https://www.researchgate.net/publication/316944512. 
  16. ↑ Lynch, Eric (2012-08-15). "Cursorial Adaptations in the Forelimb of the Giant Short-Faced Bear, Arctodus simus, Revealed by Traditional and 3D Landmark Morphometrics". Electronic Theses and Dissertations. https://dc.etsu.edu/etd/1477. 
  17. ↑ 17,0 17,1 Soibelzon, Leopoldo H.; Schubert, Blaine W. (2011). "The largest known bear, Arctotherium angustidens, from the early Pleistocene pampean region of Argentina: with a discussion of size and diet trends in bears". Journal of Paleontology 85 (1): 69–75. doi:10.1666/10-037.1. 
  18. ↑ Donohue, Shelly L.; DeSantis, Larisa R. G.; Schubert, Blaine W.; Ungar, Peter S. (2013-10-30). "Was the Giant Short-Faced Bear a Hyper-Scavenger? A New Approach to the Dietary Study of Ursids Using Dental Microwear Textures". PLOS ONE 8 (10): e77531. doi:10.1371/journal.pone.0077531. ISSN 1932-6203. PMID 24204860. PMC 3813673. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=3813673. 
  19. ↑ 19,0 19,1 Mitchell, Kieren J.; Bray, Sarah C.; Bover, Pere; Soibelzon, Leopoldo; Schubert, Blaine W.; Prevosti, Francisco; Prieto, Alfredo; Martin, Fabiana et al. (2016-04-30). "Ancient mitochondrial DNA reveals convergent evolution of giant short-faced bears (Tremarctinae) in North and South America". Biology Letters 12 (4): 20160062. doi:10.1098/rsbl.2016.0062. PMID 27095265. PMC 4881349. //www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=4881349. 
  20. ↑ Packard. „Mammalian Tracks in the Late Pliocene or Early Pleistocene Beds of Lake County Oregon“. Oregon Geology. Qaraldi: 2017-yil 11-iyun. 
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