Arxeopteriks - Archaeopteryx

Ushbu maqola qadimgi hayvon haqida. Qadimgi o'simlik uchun qarang Archaeopteris.
Boshqa maqsadlar uchun qarang Arxeopteriks (ajralish).

Arxeopteriks
Vaqtinchalik diapazon: Kech yura, 150.8–148.5 Ma
To'liq Arxeopteriks qoldiqlari, shu jumladan qanot va dumidagi patlarning chuqurliklari
Berlin Arxeopteriks namuna (A. siemensii).
Ilmiy tasnif e
Qirollik:Animalia
Filum:Chordata
Klade:Dinozavrlar
Klade:Saurischia
Klade:Theropoda
Klade:Avialae
Oila:†Archaeopterygidae
Tur:†Arxeopteriks
Meyer, 1861
(saqlangan ism )
Tur turlari
†Arxeopteriks litografikasi
Meyer, 1861 yil (saqlangan ism)
Yo'naltirilgan turlar
  • †A. siemensii
    Dames, 1897 yil
  • †A. albersdoerferi
    Kundrat va boshq. 2018 yil
Sinonimlar[1]

Arxeopteriks (/ˌ.rkiːˈɒptarɪks/ ba'zan "nemis nomi bilan ataladigan" eski qanot "), Urvogel ("asl qush" yoki "birinchi qush"), a tur ning qush o'xshash dinozavrlar. Ism qadimgi yunoncha χrχabos (arxaoslar), "qadimiy" degan ma'noni anglatadi va υξrυξ (ptéryx), "tuk" yoki "qanot" ma'nosini anglatadi. 19-asr oxiri va 21-asr boshlari orasida Arxeopteriks tomonidan odatda qabul qilingan paleoontologlar va eng qadimgi qush sifatida mashhur ma'lumotnomalar (guruh a'zosi) Avialae ).[2] Keyinchalik potentsial avialanslar aniqlandi, shu jumladan Anchiornis, Xiaotingia va Aurornis.[3]

Arxeopteriks da yashagan Kech yura taxminan 150 million yil oldin, hozirgi janubiy Germaniya hududida va Portugaliya, Evropa sayoz iliq tropik dengizdagi orollar arxipelagi bo'lgan davrda, unga ancha yaqin bo'lgan ekvator hozirgiga qaraganda. A o'lchamiga o'xshash Evroosiyo magpini, ehtimol a kattaligiga erishgan eng katta shaxslar bilan qarg'a,[4] ning eng katta turlari Arxeopteriks uzunligi taxminan 0,5 metrgacha o'sishi mumkin. Kichkina o'lchamlari, keng qanotlari va uchish yoki sirpanish qobiliyatiga qaramay, Arxeopteriks boshqa kichiklar bilan ko'proq o'xshashliklari bor edi Mezozoy dinozavrlar zamonaviy qushlarga qaraganda. Xususan, ular quyidagi xususiyatlarni dromaeosauridlar va troodontidlar: o'tkir jag'lar tish, uchta barmoq bilan tirnoqlari, uzun suyakli dum, giperekstensiv ikkinchi barmoqlar ("tirnoqni o'ldirish"), patlar (ular ham iliq qonlik ) va turli xil xususiyatlari skelet.[5][6]

Ushbu xususiyatlar Arxeopteriks a uchun aniq nomzod o'tish davri fotoalbomlari qush bo'lmagan dinozavrlar va qushlar o'rtasida.[7][8] Shunday qilib, Arxeopteriks nafaqat o'rganishda, balki muhim rol o'ynaydi qushlarning kelib chiqishi, ammo dinozavrlarni o'rganishda. Bittadan nomlangan patlar 1861 yilda,[9] kimligi qarama-qarshi bo'lgan.[10][11] O'sha yili, birinchi to'liq namunasi Arxeopteriks e'lon qilindi. Yillar davomida yana o'nta qoldiq Arxeopteriks yuzaga chiqdi. Ushbu qoldiqlarning xilma-xilligiga qaramay, ko'pchilik mutaxassislar topilgan qoldiqlarni bitta turga tegishli deb hisoblashadi, ammo bu hali ham muhokama qilinmoqda.

Arxeopteriks uzoq vaqtdan beri qushlarning evolyutsion daraxtining boshlanishi deb hisoblangan. Uning qush bo'lishni qanday aniqlashga yordam beradigan fazilatlari bor, masalan, uning uzun, qudratli oldingi oyoq-qo'llari. Biroq, so'nggi yillarda bir nechta mayda tukli dinozavrlarning kashf etilishi paleontologlar uchun sirni yaratib, qaysi hayvonlar zamonaviy qushlarning ajdodlari va qaysi biri ularning qarindoshlari ekanligi haqida savollar tug'dirdi.[12]

Qoldiq xususiyatlarini tahlil qilish shuni ko'rsatadiki Arxeopteriks umuman qush emas. Qushlar orasidagi chegarani bosib o'tgan so'nggi qazilma topilma parranda bo'lmagan dinozavrlar paleontologlarni ikkalasi o'rtasidagi evolyutsion aloqani hisobga olgan jonzotni qayta baholashda etakchilik qilmoqda.[12]

Ushbu o'n bitta qoldiqning aksariyat qismida patlarning taassurotlari mavjud. Ushbu tuklar rivojlangan shaklda bo'lgani uchun (uchish patlari ), bu qoldiqlar patlarning evolyutsiyasi So'nggi Yura davridan oldin boshlanganligining dalilidir.[13] The turdagi namunalar ning Arxeopteriks atigi ikki yildan keyin topilgan Charlz Darvin nashr etilgan Turlarning kelib chiqishi to'g'risida. Arxeopteriks Darvin nazariyalarini tasdiqlaganday tuyuldi va shu vaqtdan boshlab qushlarning kelib chiqishi, o'tish davri toshqotganliklari munozarasi va tasdiqlanganligining asosiy daliliga aylandi. evolyutsiya.

Kashfiyot tarixi

Asosiy maqola: Arxeopteriks namunalari
Vaqt xati Arxeopteriks 2007 yilgacha kashfiyotlar

O'tgan yillar davomida, o'n ikki tanadagi fotoalbom namunalari Arxeopteriks topildi. Qoldiqlarning barchasi ohaktosh yaqin asrlar davomida qazib olinadigan konlar Solnhofen, Germaniya.[14][15]

Yagona tuk

Dastlabki kashfiyot, bitta tuk, 1860 yoki 1861 yillarda topilgan va 1861 yilda tasvirlangan Hermann fon Meyer.[16] Ayni paytda u joylashgan Berlin tabiiy tarix muzeyi. Garchi bu boshlang'ich edi holotip, u tana qoldiqlari bilan bir xil hayvondan bo'lmasligi mumkinligiga ishora bor edi.[9] 2019 yilda lazer yordamida ko'rish kvilingning tuzilishini aniqladi (pat tasvirlanganidan keyin bir muncha vaqt o'tgach ko'rinmas edi) va tuklar boshqa morfologiyaga mos kelmadi. Arxeopteriks tuklar ma'lum bo'lib, u boshqa dinozavrdan kelib chiqqan degan xulosaga keladi.[10] Ushbu xulosa 2020 yilda dargumon deb taxmin qilingan; tuk morfologiya asosida aniqlangan, chunki u yuqori asosiy birlamchi bo'lgan yashirin tuklar.[11]

Deb nomlanuvchi birinchi skelet London namunasi (BMNH 37001),[17] yaqinida 1861 yilda topilgan Langenaltxaym, Germaniya va, ehtimol, tibbiy xizmat evaziga mahalliy shifokor Karl Häberleinga berilgan. Keyin u uni 700 funtga sotdi (2020 yilda taxminan 83000 funt.) [18]) uchun Tabiiy tarix muzeyi Londonda, u erda qoladi.[14] Bosh va bo'yinning katta qismini yo'qotib, 1863 yilda tasvirlangan Richard Ouen kabi Arxeopteriks makrurasi, bu tuklar bilan bir xil turga tegishli emasligiga imkon beradi. Uning keyingi to'rtinchi nashrida Turlarning kelib chiqishi to'g'risida,[19] Charlz Darvin ba'zi bir mualliflar "qushlarning butun sinfi eosen davrida to'satdan paydo bo'lganligini" ta'kidlagan edilar; ammo endi biz professor Ouenning so'zlariga ko'ra qush yuqori yashil maydon cho'kkan paytda yashaganligini bilamiz; va yana ko'p narsalar yaqinda, o'sha g'alati qush, Arxeopteriks, uzun kaltakesakka o'xshash quyruq bilan, har bir bo'g'inida juft tuklar va qanotlari bilan ikkita erkin tirnoq bilan jihozlangan, Solnhofen oolitik slanetsidan topilgan. Yaqinda o'tkazilgan har qanday kashfiyot dunyoning sobiq aholisi haqida biz hali ham bilmaganligimizni bundan ham kuchliroq ko'rsatmaydi. "[20]

The Yunoncha "pteryx" atamasi (υξrυξ) birinchi navbatda "qanot" degan ma'noni anglatadi, lekin shunchaki "tuklar" ni ham belgilashi mumkin. Meyer buni tavsifida taklif qildi. Avvaliga u zamonaviy qushlarga o'xshab ko'rinadigan bitta tukni eslatib o'tdi remex (qanotli tuklar), lekin u bu haqda eskizini eshitgan va ko'rsatgan London u "deb nomlagan namunaSkelett eines mit ähnlichen Federn bedeckten Darajalar"(" o'xshash tuklar bilan qoplangan hayvon skeletlari "). Nemis tilida bu noaniqlik atama bilan hal qilinadi Shvinge bu uchish uchun ishlatiladigan qanot degani emas. Urshvin ning maqbul tarjimasi edi Arxeopteriks XIX asr oxirlarida nemis olimlari orasida. Ingliz tilida "antik pinion" taxminiy taxminni taklif qiladi.

O'shandan beri o'n ikkita nusxa qayta tiklandi:

The Berlin namunasi (HMN 1880/81) 1874 yoki 1875 yillarda Blumenberg yaqinida topilgan Eichstätt, Germaniya, fermer Yakob Nimeyer tomonidan. U bu qimmatbaho buyumni sotdi fotoalbom 1876 ​​yilda sigir sotib olish uchun pulni, xo'jayin Yoxann Dörrga berdi, u uni yana K. Häberleinning o'g'li Ernst Otto Häberleinga sotdi. 1877 yildan 1881 yilgacha sotuvga qo'yilgan, potentsial xaridorlar bilan O. C. Marsh ning Yel universiteti Peabody muzeyi, oxir-oqibat, uni 20 mingga sotib olishdi Goldmark hozirda namoyish etiladigan Berlin tabiiy tarix muzeyi tomonidan. Bitim moliyalashtirildi Ernst Verner fon Simens, asoschisi mashhur kompaniya uning nomi bilan atalgan.[14] 1884 yilda tasvirlangan Wilhelm Dames, bu eng to'liq namunadir va to'liq bosh bilan birinchi. 1897 yilda u Dames tomonidan yangi tur deb nomlangan, A. siemensii; ko'pincha sinonimi deb qaralsa ham A. litografik, 21-asrning bir qancha tadqiqotlari natijasida u Berlin, Myunxen va Termopolis namunalarini o'z ichiga olgan alohida tur ekanligi to'g'risida xulosa qilishdi.[21][22]

Maxberg namunasini quyish

Daraxt tanasidan iborat Maxberg namunasi (S5) 1956 yilda Langenaltxaym yaqinida topilgan; bu professorning e'tiboriga havola etildi Florian Xeller 1958 yilda va u tomonidan 1959 yilda tasvirlangan. Namunaning boshi va dumi yo'q, garchi skeletning qolgan qismi asosan buzilmagan. Garchi u bir vaqtlar namoyish etilgan bo'lsa ham Maxberg muzeyi Solnhofenda u hozirda yo'qolgan. Bu tegishli edi Eduard Opitsch, uni muzeyga 1974 yilgacha ijaraga bergan. 1991 yilda vafot etganidan so'ng, namunaning yo'qligi va o'g'irlangan yoki sotilgan bo'lishi mumkinligi aniqlandi.

The Haarlem namunasi (TM 6428/29, shuningdek Teylers namunasi) yaqinida 1855 yilda topilgan Riedenburg, Germaniya va a Pterodaktil qassoblar 1857 yilda Meyer tomonidan. 1970 yilda qayta tasniflangan Jon Ostrom va hozirda joylashgan Teylers muzeyi yilda Haarlem, Nederlandiya. Bu topilgan birinchi namunadir, ammo o'sha paytda noto'g'ri tasniflangan. Bundan tashqari, bu asosan oyoq-qo'l suyaklari, ajratilgan bachadon bo'yni umurtqalari va qovurg'alardan tashkil topgan eng kam namunalardan biridir. 2017 yilda u alohida tur sifatida nomlandi Ostromiya bilan chambarchas bog'liq deb hisoblanadi Anchiornis Xitoydan.[23]

Bir vaqtlar alohida tur deb hisoblangan Eichstätt namunasi, Jurapteryx

The Eichstätt namunasi (JM 2257) 1951 yilda Germaniyaning Workingzell shahri yaqinida topilgan va uni tasvirlab bergan Piter Uellnhofer 1974 yilda. Hozirda joylashgan Yura muzeyi yilda Eichstätt, Germaniya, u tanilgan eng kichik namunadir va boshi bo'yicha ikkinchi o'rinda turadi. Ehtimol, bu alohida tur (Jurapteryx recurva) yoki turlar (A. recurva).

The Solnhofen namunasi (raqamlanmagan namunalar) yaqin 70-yillarda topilgan Eichstätt, Germaniya va 1988 yilda Wellnhofer tomonidan tasvirlangan. Hozirda joylashgan Byurgermeister-Myuller-muzeyi Solnhofenda u dastlab quyidagicha tasniflangan Kompsognatus havaskor kollektsioner tomonidan, o'sha mer Fridrix Myuller tomonidan muzey nomi berilgan. Bu ma'lum bo'lgan eng katta namunadir va alohida tur va turlarga mansub bo'lishi mumkin, Wellnhoferia grandis. Unda faqat bo'yin, dum, umurtqa pog'onasi va bosh qismlari yo'q.

The Myunxen namunasi (BSP 1999 I 50, ilgari sifatida tanilgan Solenhofer-Aktien-Verein namunasi) 1992 yil 3 avgustda Langenaltxaym yaqinida topilgan va 1993 yilda Wellnhofer tomonidan tasvirlangan. Ayni paytda u joylashgan Myunxendagi Paläontologisches muzeyi 1999 yilda 1,9 mln.ga sotilgan Myunxenda Deutschmark. Dastlab suyak deb ishonilgan narsa ko'krak suyagi ning bir qismi bo'lib chiqdi korakoid,[24] lekin a xaftaga oid sternum mavjud bo'lishi mumkin. Yuzining faqat old qismi yo'qolgan. U alohida turlar uchun asos sifatida ishlatilgan, A. bavarica,[25] ammo yaqinda o'tkazilgan tadqiqotlar shuni ko'rsatmoqdaki A. siemensii.[22]

Kutish namunasi, ning holotipi A. albersdoerferi

Sakkizinchi, bo'lakcha namunasi 1990 yilda yoshroqida topilgan Mörnsheim shakllanishi da Kutilmoqda, Sueviya. Shuning uchun, u sifatida tanilgan Namunani kutishva 1996 yildan beri Naturkundemuseumda qisqacha namoyish qilingan aktyorlar guruhidan tanilgan Bamberg. Asl nusxasini paleontolog Raimund Albertsdörfer 2009 yilda sotib olgan.[26] U yana oltita asl qoldiq bilan birinchi marta namoyish etildi Arxeopteriks 2009 yil oktyabr oyida Myunxen mineral ko'rgazmasida.[27] Keyinchalik "Kutish namunasi" nomi berildi Arxeopteriks albersdoerferi Kundrat va boshq. (2018).[28][29]

Burgermeister-Myuller ("tovuq qanoti") Namuna

2000 yilda yana bir bo'lak qoldiq topilgan. U shaxsiy mulkda va 2004 yildan beri Solnhofendagi Byurgermeister-Myuller muzeyiga ijaraga berilgan, shuning uchun uni " Byurgermeister-Myuller namunasi; institutning o'zi rasmiy ravishda uni "Ottman va Stayl, Solnhofen oilalarining namunasi" deb ataydi. Parcha bitta qanotining qoldiqlarini ifodalaydi Arxeopteriks, ushbu qoldiqning mashhur nomi "tovuq qanoti" dir.

Shveytsariyadagi shaxsiy to'plamda uzoq vaqt Termopolis namunasi (WDC CSG 100) Bavariyada topilgan va 2005 yilda Mayr, Pohl va Piters tomonidan tasvirlangan. Xayriya qilingan Vayominning dinozavr markazi yilda Termopolis, Vayoming, u eng yaxshi saqlanib qolgan bosh va oyoqlarga ega; bo'yinning va pastki jag'ning ko'p qismi saqlanib qolmagan. "Termopolis" namunasi 2005 yil 2-dekabrda tasvirlangan Ilm-fan jurnal maqolasi sifatida "Yaxshi saqlanib qolgan Arxeopteriks "Theropod xususiyatlari bilan namuna"; bu shuni ko'rsatadiki Arxeopteriks qushlarning universal xususiyati - orqaga burilgan barmog'i yo'q edi - uning shoxlarga o'tirish qobiliyatini cheklaydi va quruqlikda yoki magistralda ko'tarilish turmush tarzini anglatadi.[30] Bu dalil sifatida talqin qilingan teropod ajdodlar. 1988 yilda, Gregori S. Pol giperekstensiv ikkinchi barmoqning dalillarini topdik,[31] ammo bu boshqa olimlar tomonidan Termopolis namunasi tavsiflanmaguncha tekshirilmagan va qabul qilinmagan. "Hozirgacha bu xususiyat faqat turning yaqin qarindoshlari, deinonixosaurlarga tegishli deb o'ylar edi."[15] Termopolis namunasi tayinlangan Arxeopteriks siemensii 2007 yilda.[22] Namuna eng to'liq va eng yaxshi saqlanib qolgan deb hisoblanadi Arxeopteriks hali qolmoqda.[22]

O'n birinchi namuna

O'n birinchi namunaning kashf etilishi 2011 yilda e'lon qilingan va u 2014 yilda tasvirlangan. Bu to'liqroq namunalardan biri, ammo bosh suyagining katta qismi va bitta old qismi yo'qolgan. U xususiy mulk bo'lib, unga hali nom berilmagan.[32][33]Paleontologlar Myudxenning Lyudvig Maksimilian universiteti shilimshiqning ilgari noma'lum xususiyatlarini, masalan, yuqori va pastki oyoqlarning patlari va metatarsusni va saqlanib qolgan yagona quyruq uchini ochib bergan namunani o'rganib chiqdi.[34][35]

O'n ikkinchi namunani havaskor kollektsioner 2010 yilda Shamxaupten karerida topgan, ammo topilma faqat 2014 yil fevralida e'lon qilingan.[36] Bu 2018 yilda ilmiy tavsiflangan. Bu bosh suyagi bilan to'la va asosan bo'g'inli skeletni aks ettiradi. Bu saqlanib qolgan tuklar etishmaydigan yagona namunadir. Bu Bo'yalgan shakllanish va boshqa namunalardan biroz kattaroqdir.[37]

Haqiqiylik

1985 yildan boshlab havaskorlik guruhi, shu jumladan astronom Fred Xoyl va fizik Li Spetner Berlin va London namunalaridagi patlar deb da'vo qilgan bir qator hujjatlarni nashr etdi Arxeopteriks qalbaki edi.[38][39][40][41] Ularning da'volari rad etildi Alan J. Charig va boshqalar Londondagi tabiiy tarix muzeyi.[42] Ularning qalbakilashtirilganligini isbotlovchi dalillarning aksariyati jarayonlar bilan tanish emaslikka asoslangan edi litifikatsiya; Masalan, ular tuklar bilan bog'liq bo'lgan to'qimalar farqiga asoslanib, tuklar taassurotlari yupqa qatlamga qo'llanilishini taklif qilishdi. tsement,[39] patlar o'zlari to'qima farqini keltirib chiqarganini bilmasdan.[42] Shuningdek, ular toshqotganlarni noto'g'ri talqin qilib, dumni bitta katta tuklar singari to'qilgan deb da'vo qilishdi.[39] qachon ko'rinadigan bo'lsa, bunday emas.[42] Bundan tashqari, ular boshqa namunalar deb da'vo qilishdi Arxeopteriks o'sha paytda ma'lum bo'lgan patlar yo'q edi,[38][39] bu noto'g'ri; Maxberg va Eichstätt namunalarida aniq tuklar bor.[42]

Ular, shuningdek, plitalar shunchalik mayin bo'linishiga yoki toshqotgan toshlar bo'lgan plitaning yarmi yaxshi saqlanib qolishiga ishonmasliklarini bildirishdi, lekin dastgoh.[38][40] Bu Solnhofen qoldiqlarining umumiy xususiyatlari, chunki o'lik hayvonlar qotib qolgan yuzalarga tushib, kelajakdagi plitalarning bo'linishi uchun tabiiy tekislikni hosil qiladi va qoldiqlarning asosiy qismini bir tomonda, ikkinchisida esa ozgina qoldiradi.[42]

Va nihoyat, ular qalbakilashtirishga taklif qilgan motivlar kuchli emas va qarama-qarshi; bittasi shu Richard Ouen qo'llab-quvvatlovchi dalillarni soxtalashtirmoqchi edi Charlz Darvin nazariyasi evolyutsiya, bu Ouenning Darvin va uning nazariyasiga nisbatan qarashlari bilan bog'liq emas. Ikkinchisi, Ouen Darvin uchun tuzoq qo'ymoqchi edi, chunki ikkinchisi toshqotganlarni qo'llab-quvvatlaydi, shuning uchun Ouen uni soxtalashtirish bilan obro'sizlantirishi mumkin; bu ehtimoldan yiroq, chunki Ouen London namunasi bo'yicha batafsil maqola yozgan, shuning uchun bunday harakat albatta o'z samarasini beradi.[43]

Charig va boshq. ikkala tosh va fotoalbom taassurotlari bo'ylab o'tadigan plitalardagi sochlar chizig'idagi yoriqlar va kashf etilishidan oldin yuzaga kelgan minerallar o'sishi va patlarning asl ekanligidan dalolat sifatida.[42] Spetner va boshq. keyin yoriqlar ularning postulyatsiyalangan tsement qatlami orqali tabiiy ravishda tarqalishini ko'rsatishga urindi,[44] ammo yoriqlar eskirganligi va to'ldirilganligi sababli hisobga olishni e'tiborsiz qoldirgan kaltsit va shu tariqa targ'ib qila olmadilar.[43] Shuningdek, ular London namunasida tsement borligini ko'rsatishga harakat qilishdi Rentgen spektroskopiyasi, va tosh bo'lmagan narsani topdi;[44] u ham tsement emas edi va, ehtimol, namunadan qoliplar yasalganda orqada qolgan silikon kauchukning bo'lagi.[43] Ularning takliflari paleontologlar tomonidan jiddiy qabul qilinmadi, chunki ularning dalillari asosan geologiyani noto'g'ri tushunishga asoslangan edi va ular shu vaqtdan beri soni ko'paygan boshqa tuklar namunalarini hech qachon muhokama qilmadilar. Charig va boshq. rang o'zgarishi haqida xabar bergan: ohaktoshning ikki qatlami orasidagi qorong'u tasma - ular cho'kindi hosilasi deyishadi.[42] Ohaktosh atrofining rangini olishi tabiiy va aksariyat ohaktoshlar ma'lum darajada ranglanadi (agar rang bantlanmagan bo'lsa), shuning uchun qorong'ulik bunday iflosliklarga bog'liq edi.[45] Shuningdek, ular tosh plitalarida havo pufakchalarining to'liq yo'qligi namunaning haqiqiy ekanligiga yana bir dalil ekanligini ta'kidlashadi.[42]

Tavsif

Arxeoperteriksning o'lchamlari uzunligi taxminan 25 dan 50 sm gacha va qanotlarining uzunligi 25 dan 60 sm gacha
A bilan taqqoslanadigan namunalar inson miqyosda

Ko'pgina namunalar Arxeopteriks Germaniyaning janubiy Bavariyasidagi Solnhofen ohaktoshidan kelib chiqqan, bu lagerstätte, noyob va ajoyib geologik shakllanishdir, bu Yura davrining dastlabki titoniyalik bosqichida yotqizilgan ajoyib detallari bilan tanilgan,[46] taxminan 150,8–148,5 million yil oldin.[47]

Arxeopteriks taxminan a ga teng edi qarg'a,[4] tanasining uzunligiga nisbatan keng dumaloq uchlari va uzun dumlari bilan. Tana uzunligi 500 millimetrga (20 dyuym) etishi mumkin, taxminiy massasi 0,8 dan 1 kilogrammgacha (1,8 dan 2,2 funtgacha).[4] Arxeopteriks tuklar, boshqa xususiyatlaridan kamroq hujjatlashtirilgan bo'lsa-da, tuzilishi jihatidan hozirgi qushlarning patlariga juda o'xshash edi.[46] Ko'p sonli parrandalar mavjudligiga qaramay,[48] Arxeopteriks ko'plab parranda bo'lmagan tropod dinozavri xususiyatlari. Zamonaviy qushlardan farqli o'laroq, Arxeopteriks kichik tishlari bor edi,[46] shuningdek, uzun suyak dumi, xususiyatlari Arxeopteriks vaqtning boshqa dinozavrlari bilan bo'lishgan.[49]

Unda qushlarga ham, parranda bo'lmagan dinozavrlarga ham xos xususiyatlar mavjud, Arxeopteriks ko'pincha ular orasidagi bog'lovchi deb hisoblangan.[46] 1970-yillarda, Jon Ostrom, quyidagi Tomas Genri Xaksli 1868 yilda olib borilgan qo'rg'oshinlar, qushlarning dinozavrlar ekologik rivojlanganligini va Arxeopteriks ushbu dalil uchun juda muhim dalil edi; u qushlarning bir nechta xususiyatlariga ega edi, masalan tilak suyagi, uchish tuklari, qanotlar va qisman teskari o'girilgan birinchi barmog'i bilan birga dinozavr va terropod xususiyatlari. Masalan, ning uzoq ko'tarilish jarayoni bor oyoq suyagi, tishlararo plitalar, an obturator jarayoni iskiyum va dumidagi uzun chevronlar. Xususan, Ostrom buni aniqladi Arxeopteriks Theropod oilasiga juda o'xshash edi Dromaeosauridae.[50][51][52][53][54][55][56][57][58]Arxeopteriksning har bir oyog'ida "tirnoq" bilan tugaydigan uchta alohida raqam bor edi. Bunday xususiyatlarga ega bo'lgan ozgina qushlar. O'rdak, oqqush va Jacanas kabi ba'zi qushlar (Jakana sp.) va hoatzin (Opisthocomus hoazin) ularni patlarning ostiga berkitsin.[59]

Tuklar

"Frond-tail" ni taqqoslaydigan anatomik illyustratsiya Arxeopteriks zamonaviy qushning "fan-dumi" bilan

Namunalari Arxeopteriks eng yaxshi rivojlanganligi bilan ajralib turardi uchish patlari. Ular sezilarli darajada assimetrik edilar va zamonaviy qushlardagi parvozlar patlarining tuzilishini ko'rsatib berdilar, qanotlari a tomonidan barqarorlik berildi tikan -barbule -barbitsel joylashuvi.[60] Quyruq patlari kamroq assimetrik bo'lib, yana zamonaviy qushlarning holatiga mos keladi va shuningdek, qattiq qanotlarga ega edi. The bosh barmog'i hanuzgacha alohida harakatlanuvchi tutamga ega emas edi qattiq tuklar.

Tanasining tuklari Arxeopteriks kamroq hujjatlashtirilgan va faqat yaxshi saqlanib qolgan holda tegishli ravishda o'rganilgan Berlin namunasi. Shunday qilib, bir nechta tur ishtirok etgandek tuyuladi, chunki Berlin namunasining patlari bo'yicha tadqiqotlar qolgan turlar uchun to'g'ri kelishi shart emas. Arxeopteriks. Berlin namunasida oyoqlarda yaxshi rivojlangan patlarning "shimlari" mavjud; bu patlarning ba'zilari asosiy kontur patlarni tuzilishiga o'xshaydi, ammo biroz parchalanib ketgan (ular etishmayapti) barbitsellar kabi ratitlar ).[61] Qisman ular qat'iy va shuning uchun parvozni qo'llab-quvvatlashga qodir.[62]

Yamoq tukli patlarni uning orqa tomoni bo'ylab yugurib yurganligi aniqlangan, bu zamonaviy qushlarning tana patlarining kontur patlariga juda o'xshash, garchi parvoz bilan bog'liq tuklar kabi qattiq bo'lmasa ham. Bundan tashqari, Berlin namunasidagi tuklar izlari bir xil "proto-" bilan cheklangan.pastga "dinozavrda topilganga o'xshamaydi Sinozauropteriks: chirigan va paxmoq, va hattoki hayotdagi patlardan ko'ra ko'proq mo'ynaga o'xshaydi (garchi ularning mikroskopik tuzilishida emas). Bular tananing qolgan qismida sodir bo'ladi - garchi ba'zi tuklar toshib ketmagan bo'lsa, boshqalari esa tayyorgarlik paytida yo'q bo'lib, namunalarda yalang'och yamalar qoldirgan va pastki bo'ynida.[61]

Bo'yinning yuqori qismida va boshda patlarni ko'rsatadigan narsa yo'q. Ehtimol, bu yalang'och bo'lishi mumkin edi, ammo bu hali ham saqlanib qolishi mumkin. Bu eng ko'p ko'rinadi Arxeopteriks namunalar ichiga joylashtirildi anoksik dengizda bir muncha vaqt orqalarida siljiganidan keyin cho'kindi - bosh, bo'yin va dum odatda pastga egilgan, bu esa namunalar singdirilgandan so'ng chiriy boshlaganligini, tendonlar va mushaklarning bo'shashishi bilan xarakterli shakli (o'lim pozasi ) fotoalbom namunalarga erishildi.[63] Bu terining allaqachon yumshatilganligini va bo'shashganligini anglatar edi, bu ba'zi namunalarda parvoz patlarining cho'kindiga singib ketish joyidan ajrala boshlagani bilan mustahkamlanadi. Shunday qilib, tegishli namunalar dafn qilinishidan oldin dengiz tubi bo'ylab sayoz suvda bir muncha vaqt harakat qilgan, faraz qilinganki, bosh va yuqori bo'yin patlari yulqalanib, mahkam o'rnashgan quyruq patlari qolgan.[21]

Rangi

Rassomning restavratsiyasi Karni tadqiqotining bir talqinini aks ettiradi[64]

2011 yilda aspirant Rayan Karni va uning hamkasblari birinchi rangli tadqiqotni an Arxeopteriks namuna.[64] Foydalanish skanerlash elektron mikroskopi texnologiya va energetik dispersiv rentgen tahlil qilish, jamoa tuzilishini aniqlashga muvaffaq bo'ldi melanosomalar Natijada olingan o'lchovlar 87 zamonaviy qush turlari bilan taqqoslandi va asl rang 95% qora bo'lish ehtimoli bilan hisoblab chiqildi. Tuklar qora rangda, distal uchida og'irroq pigmentatsiya borligi aniqlandi. O'rganilgan pat, ehtimol, dorsal edi yashirin, bu qanotlarda asosiy patlarni qisman qoplagan bo'lar edi. Tadqiqot bu degani emas Arxeopteriks butunlay qora edi, lekin uning qopqog'ini o'z ichiga olgan qora rangga ega ekanligini taxmin qilmoqda. Karni, bu bizning zamonaviy parvoz xususiyatlari haqida bilgan narsalarga mos kelishini ta'kidladi, chunki qora melanosomalar parvoz uchun patlarni mustahkamlovchi tuzilish xususiyatlariga ega.[65] 2013 yilda chop etilgan bir tadqiqotda Analitik atom spektrometriyasi jurnali, ning yangi tahlillari Arxeopteriks 'patlaridan ma'lum bo'lishicha, hayvonning ochiq va quyuq rangli tuklari bo'lgan, distal uchlari va tashqi qanotlarida og'irroq pigmentatsiya bo'lgan.[66] Rang taqsimotining ushbu tahlili, avvalo, sulfat qoldiqlari bilan taqsimlanishiga asoslangan edi. Oldingi muallif Arxeopteriks rang tadqiqotida misning to'liq Archeopteryx namunasidagi eumelanin ko'rsatkichi sifatida talqin qilinishiga qarshi bahs yuritildi, ammo sulfat tarqalishi haqida hech narsa aytilmagan. Carney tomonidan chop etilgan press-reliz va konferentsiya avtoulovi 2013 yilgi tadqiqotga qarshi chiqmoqda, ammo yangi dalillar nashr etilmagan.[67][68]

Tasnifi

Termopolis namunasi

Bugungi kunda ushbu jinsning qoldiqlari Arxeopteriks odatda bir yoki ikki turga beriladi, A. litografik va A. siemensii, ammo ularning taksonomik tarixi murakkab. Bir nechta namunalar uchun o'nta nom nashr etilgan. Bugungi kunda talqin qilinganidek, ism A. litografik faqat tomonidan tasvirlangan bitta tuklarga ishora qiladi Meyer. 1954 yilda Gavin de Pivo London namunasi bu degan xulosaga keldi holotip. 1960 yilda Svinton shunga ko'ra bu nomni taklif qildi Arxeopteriks litografikasi muqobil nomlarni rasmiylashtirgan rasmiy nasllar ro'yxatiga kiritilgan Griphosaurus va Griphornis yaroqsiz.[69] The ICZN, de Beerning nuqtai nazarini bevosita qabul qilgan holda, birinchi skelet namunalari uchun dastlab taklif qilingan muqobil nomlarning ko'pligini bostirgan,[70] Bu asosan Meyer va uning raqibi o'rtasidagi kelishmovchiliklar natijasida kelib chiqqan Yoxann Andreas Vagner (kimning Griphosaurus problematicus - "muammoli topishmoq -lizard "- bu Meyerda vitriolik kinoya edi Arxeopteriks).[71] Bundan tashqari, 1977 yilda Komissiya Haarlem namunasining birinchi tur nomi, qassoblar, Meyer tomonidan tasvirlangan a pterosaur uning asl mohiyati amalga oshmasdan oldin, unga ustunlik berilmasligi kerak edi litografiya olimlar ularni bir xil turni vakili deb hisoblagan holatlarda.[7][72]

Birinchi namunasi bo'lgan patlar ekanligi ta'kidlangan Arxeopteriks tasvirlangan, parvoz bilan bog'liq bo'lgan patlarga yaxshi mos kelmaydi Arxeopteriks. Bu albatta parvoz tuklari zamonaviy turdagi, ammo uning kattaligi va nisbati uning boshqa kichikroq turlariga tegishli bo'lishi mumkinligini ko'rsatadi tukli teropod, ulardan qaysi biri hozirgacha faqat shu tuklar ma'lum.[9] Tuklar belgilab qo'yilganidek turdagi namunalar, ism Arxeopteriks bundan keyin skeletlarga qo'llanilmasligi kerak va shu bilan ahamiyatli bo'ladi nomenklatura chalkashlik. 2007 yilda ikkita olim olimlar ICZN-ga London namunasini yangi holotip namunasi sifatida belgilash orqali aniq qilib tayyorlashni iltimos qilib, neotip.[73] Ushbu taklif ICZN tomonidan to'rt yillik bahs-munozaralardan so'ng qo'llab-quvvatlandi va London namunasi 2011 yil 3 oktyabrda neotip sifatida belgilandi.[74]

O'n ikkinchi namuna

Quyida a kladogramma tomonidan 2013 yilda nashr etilgan Godefroit va boshq.[3]

Avialae
 

Aurornis

 
 
 

Anchiornis

 
 
 

Arxeopteriks

 
 
 

Xiaotingia

 
 
 
 

Shenzhouraptor

 
 

Rahonavis

 
 
 
 

Balaur

 
 
 

Avebrevikauda (zamonaviy qushlarni o'z ichiga oladi)

 
 
 
 
 
 
 
 

Turlar

Har xil namunalarning skelet tiklanishi

Barcha namunalar bir xil turga mansub, A. litografik.[75] Namunalar orasida farqlar mavjud va ba'zi tadqiqotchilar buni namunalarning turli yoshiga qarab deb hisoblasa, ba'zilari haqiqiy turlarning xilma-xilligi bilan bog'liq bo'lishi mumkin. Xususan, Myunxen, Eichstätt, Solnhofen va Termopolis namunalari London, Berlin va Haarlem namunalaridan kichikroq yoki kattaroqligi, barmoqlarning turli nisbatiga ega bo'lishi, oldinga yo'naltirilgan tishlar bilan o'ralgan ingichka tumshuqlarga ega bo'lishi va mavjud bo'lishi mumkinligi bilan farq qiladi. ko'krak suyagi. Ushbu farqlar tufayli aksariyat individual namunalarga u yoki bu nuqtada o'z turlarining nomi berilgan. Berlin namunasi sifatida belgilangan Archaeornis siemensii, Eichstätt namunasi sifatida Jurapteryx recurva, Myunxen namunasi Arxeopteriks bavaricava Solnhofen namunasi Wellnhoferia grandis.[21]

2007 yilda barcha yaxshi saqlangan namunalarni, shu jumladan, o'sha paytda yangi kashf etilgan Termopolis namunalarini ko'rib chiqishda ikkita alohida turdagi Arxeopteriks qo'llab-quvvatlanishi mumkin: A. litografik (kamida London va Solnhofen namunalaridan iborat) va A. siemensii (kamida Berlin, Myunxen va Termopolis namunalaridan iborat). Ikkala tur birinchi navbatda oyoq tirnoqlarida katta fleksor tubercles bilan ajralib turadi A. litografik (tirnoqlari A. siemensii namunalar nisbatan sodda va to'g'ri). A. litografik shuningdek, ba'zi tishlarda tojning qisqargan qismi va dag'al metatars bor edi. Taxminan qo'shimcha turlar, Wellnhoferia grandis (Solnhofen namunasi asosida) bilan ajralib turmaydiganga o'xshaydi A. litografik uning kattaligi bundan mustasno.[22]

Sinonimlar

Solnhofen namunasi, ba'zilar tomonidan bu turga mansub deb hisoblanadi Wellnhoferia

Agar ikkita ism berilgan bo'lsa, birinchisi "tur" ning asl kelib chiqishini bildiradi, ikkinchisi ushbu ism birikmasi asosidagi muallif. Har doimgidek zoologik nomenklatura, Qavslar ichiga muallifning ismini qo'yish bu degani takson dastlab boshqa turda tasvirlangan.

  • Arxeopteriks litografikasi Meyer, 1861 yil [saqlangan ism]
    • Archaeopterix lithographica Anon., 1861 yil [lapsus]
    • Griphosaurus problematicus Vagner, 1862 yil [ICZN 607 fikri bo'yicha rad etilgan ism 1961]
    • Griphornis longicaudatus Ouen video Vudvord, 1862 yil [ICZN 607 fikri bo'yicha rad etilgan ism 1961]
    • Arxeopteriks makrurasi Ouen, 1862 yil [ICZN 607 fikri bo'yicha rad etilgan ism 1961]
    • Arxeopteriks oweni Petronievika, 1917 yil [ICZN 607 fikri bo'yicha rad etilgan ism 1961]
    • Arxeopteriks qaytalanishi Howgate, 1984 yil
    • Jurapteryx recurva (Howgate, 1984) Howgate, 1985 yil
    • Wellnhoferia grandis Elżanovskiy, 2001 yil
  • Arxeopteriks siemensii Dames, 1897 yil
    • Archaeornis siemensii (Dames, 1897) Petronievika, 1917 yil[22]
    • Arxeopteriks bavarica Wellnhofer, 1993 yil

"Arxeopteriks" vitsensensisi (Anon.) fide Lambrecht, 1933) a nomen nudum chunki ta'riflanmagan pterosaurga o'xshagan narsa.

Filogenetik holat

Deinonychus va Archeopteryx oldingi suyaklaridagi tasvirlar, ikkalasi ham ikkita barmog'i va bir-biriga qarama-qarshi tirnoqlari bilan juda o'xshash tuzilishga ega, ammo Arxeopteriks suyaklari ingichka bo'lsa ham
Oldingi taqqoslash Arxeopteriks (o'ngda) bilan Deinonychus (chapda)

Zamonaviy paleontologiya ko'pincha tasniflangan Arxeopteriks eng ibtidoiy qush sifatida. Bu zamonaviy qushlarning haqiqiy ajdodi deb o'ylanmaydi, aksincha, bu ajdodning yaqin qarindoshi.[76] Shunga qaramay, Arxeopteriks ko'pincha haqiqiy ajdod qushining namunasi sifatida ishlatilgan. Bir nechta mualliflar buni qildilar.[77] Lou (1935)[78] va Thulborn (1984)[79] yo'qmi degan savol tug'dirdi Arxeopteriks chindan ham birinchi qush edi. Ular buni taklif qilishdi Arxeopteriks boshqa dinozavr guruhlariga qaraganda qushlar bilan chambarchas bog'liq bo'lmagan dinozavr edi. Kurzanov (1987) buni taklif qildi Avimimus ga qaraganda barcha qushlarning ajdodlari bo'lish ehtimoli ko'proq edi Arxeopteriks.[80] Barsbold (1983)[81] va Zweers va Van den Berge (1997)[82] ko'pchilik ta'kidladi maniraptoran nasablar nihoyatda qushga o'xshashdir va ular turli xil qushlar guruhlari turli dinozavr ajdodlaridan kelib chiqqan bo'lishi mumkin deb taxmin qilishgan.

Yaqindan bog'liq bo'lgan kashfiyot Xiaotingia 2011 yilda yangi filogenetik tahlillarga olib keldi Arxeopteriks a deinonixozaur avialan o'rniga, shuning uchun bu atamada eng ko'p ishlatiladigan "qush" emas.[2] Ushbu farazni sinab ko'rish uchun ko'p o'tmay batafsilroq tahlil e'lon qilindi va bir xil natijaga erishilmadi; topildi Arxeopteriks Avialae bazasida an'anaviy holatida, ammo Xiaotingia bazal dromaeosaurid yoki troodontid sifatida tiklandi. Keyingi tadqiqot mualliflari noaniqliklar hali ham mavjudligini va ishonch bilan aytish mumkin emas yoki yo'qligini ta'kidladilar. Arxeopteriks tegishli turlarning yangi va yaxshiroq namunalarini taqiqlovchi Avialae a'zosi yoki emas.[83]

Senter tomonidan o'tkazilgan filogenetik tadqiqotlar, va boshq. (2012) va Turner, Makovicky va Norell (2012) Arxeopteriks dromaeosauridlar va troodontidlarga qaraganda tirik qushlar bilan yaqinroq aloqada bo'lish.[84][85] Boshqa tomondan, Godefroit, va boshq. (2013) tiklandi Arxeopteriks dromaeozauridlar va troodontidlar bilan chambarchas bog'liq bo'lib, ularning tavsifiga kiritilgan Eozinopteryx brevipenna. Mualliflar matritsaning o'zgartirilgan versiyasidan foydalanib, tadqiqotni tasvirlab berishgan Xiaotingia, qo'shib Jinfengopteryx elegans va Eozinopteryx brevipenna unga, shuningdek, tuklarning rivojlanishi bilan bog'liq to'rtta qo'shimcha belgini qo'shish. Ning tavsifidan tahlildan farqli o'laroq Xiaotingia, Godefroit tomonidan o'tkazilgan tahlil, va boshq. topmadim Arxeopteriks ayniqsa yaqindan bog'liq bo'lishi kerak Anchiornis va Xiaotingia, ularning o'rniga bazal troodontidlar sifatida tiklangan.[86]

Agnolin va Novas (2013) topildi Arxeopteriks va (ehtimol sinonim) Wellnhoferia eng oddiy avialans bo'lish (Avialae mualliflar tomonidan shu jumladan, deb belgilanadi Arxeopteriks litografikasi va Passer, ularning eng so'nggi umumiy ajdodi va uning barcha avlodlari), bilan Mikroraptoriya, Unenlagiinae va o'z ichiga olgan qoplama Anchiornis va Xiaotingia Avialae bilan ketma-ket yaqin guruhlar bo'lish.[87] Godefroit tomonidan o'tkazilgan yana bir filogenetik tadqiqot, va boshq., tavsifidagi tahlildan ko'ra ko'proq inklyuziv matritsadan foydalanish Eozinopteryx brevipenna, shuningdek topildi Arxeopteriks Avialae-ning a'zosi bo'lish (mualliflar tomonidan eng ko'p qamrab olingan qoplama sifatida belgilangan Passer domesticus, lekin emas Dromaeosaurus albertensis yoki Troodon formosi ). Arxeopteriks hosil qilganligi aniqlandi sinf bilan Avialae bazasida Xiaotingia, Anchiornisva Aurornis. Ga solishtirganda Arxeopteriks, Xiaotingia mavjud qushlar bilan chambarchas bog'liqligi aniqlandi, ikkalasi ham Anchiornis va Aurornis uzoqroq bo'lganligi aniqlandi.[3]

Paleobiologiya

Parvoz

1880 yilda Berlin namunasining fotosurati, unda tayyorlanish paytida olib tashlangan oyoq patlari ko'rsatilgan

Zamonaviy qushlarning qanotlarida bo'lgani kabi, uchish patlari Arxeopteriks biroz assimetrik va quyruq patlari ancha keng edi. Bu shuni anglatadiki, qanotlari va dumlari liftni ishlab chiqarish uchun ishlatilgan, ammo bu aniq emas Arxeopteriks parvozni yoki shunchaki planerni uchirishga qodir edi. Suyakning etishmasligi ko'krak suyagi buni taklif qiladi Arxeopteriks juda kuchli uchuvchi emas edi, lekin parvoz mushaklari bumerang shaklidagi qalin tilim suyagiga, plastinkaga o'xshash bo'lishi mumkin edi korakoidlar yoki, ehtimol, a xaftaga oid ko'krak suyagi. Glenoid (elka) qo'shimchasining yon tomon yo'nalishi skapula, korakoid va humerus - zamonaviy qushlarda uchraydigan dorsal burchakli tartib o'rniga - buni ko'rsatishi mumkin Arxeopteriks qanotlarini orqa tomondan ko'tarolmadi, zamonaviy qaltirash parvozida ko'tarilish uchun talab. Tomonidan o'tkazilgan tadqiqotga ko'ra Filipp Senter 2006 yilda, Arxeopteriks chindan ham zamonaviy qushlar singari flapping parvozidan foydalana olmadi, lekin u pastga tushirish uchun faqatgina flap yordamida sirpanish texnikasini qo'llagan bo'lishi mumkin.[88] Biroq, yaqinda o'tkazilgan tadqiqotlar ushbu masalani parranda bo'lmagan uchadigan theropodlar uchun parvoz strokining boshqa konfiguratsiyasini taklif qilish orqali hal qildi.[89]

Arxeopteriks qanotlari nisbatan katta bo'lgan, bu esa to'xtash tezligining past bo'lishiga va pasayishiga olib keladi burilish radiusi. Qanotlarning qisqa va yumaloq shakli tortishni kuchaytirar edi, shuningdek, daraxtlar va cho'tka kabi tartibsiz muhitda uchish qobiliyatini yaxshilashi mumkin edi (shunga o'xshash qanot shakllari daraxtlar va cho'tkalar orqali uchadigan qushlarda ko'rinadi, masalan. qarg'alar va qirg'ovullar ). Kabi dromaeosauridlarda kuzatilgan oyoqlarga o'xshash "orqa qanotlar", assimetrik uchish patlari borligi. Mikroraptor, shuningdek, havo harakatlanishiga qo'shilgan bo'lar edi Arxeopteriks. Longrich tomonidan 2006 yilda orqa qanotlarni birinchi batafsil o'rganish natijasida tuzilmalar umumiy sonning 12 foizigacha hosil bo'lishi kerak plyonka. Bu to'xtash tezligini 6% gacha va burilish radiusini 12% gacha qisqartirishi mumkin edi.[62]

Ning patlari Arxeopteriks assimetrik edi. Bu uning varaqasi ekanligining isboti sifatida talqin qilingan, chunki uchmaydigan qushlar nosimmetrik patlarga ega. Tomson va Spikmen kabi ba'zi olimlar bunga shubha bilan qarashgan. Ular 70 dan ortiq tirik qushlarning oilalarini o'rganishdi va ba'zi uchib ketmaydigan turlarining patlarida assimetriya diapazoni borligini va Arxeopteriks ushbu qatorga kiring.[90] Asimmetriya darajasi Arxeopteriks parvoz qilmaydigan qushlarga qaraganda sekinroq uchuvchilar uchun ko'proq xosdir.[91]

Myunxen namunasi

2010 yilda Robert L. Nudds va Garet J. Dayk jurnalda Ilm-fan tahlil qilgan maqolani nashr etdi rachises of the primary feathers of Konfutsiyornis va Arxeopteriks. The analysis suggested that the rachises on these two genera were thinner and weaker than those of modern birds relative to body mass. The authors determined that Arxeopteriks va Konfutsiyornis, were unable to use flapping flight.[92] This study was criticized by Filipp J. Kurri and Luis Chiappe. Chiappe suggested that it is difficult to measure the rachises of fossilized feathers, and Currie speculated that Arxeopteriks va Konfutsiyornis must have been able to fly to some degree, as their fossils are preserved in what is believed to have been marine or lake sediments, suggesting that they must have been able to fly over deep water.[93] Gregori Pol also disagreed with the study, arguing in a 2010 response that Nudds and Dyke had overestimated the masses of these early birds, and that more accurate mass estimates allowed powered flight even with relatively narrow rachises. Nudds and Dyke had assumed a mass of 250 g (8.8 oz) for the Munich specimen Arxeopteriks, a young juvenile, based on published mass estimates of larger specimens. Paul argued that a more reasonable body mass estimate for the Munich specimen is about 140 g (4.9 oz). Paul also criticized the measurements of the rachises themselves, noting that the feathers in the Munich specimen are poorly preserved. Nudds and Dyke reported a diameter of 0.75 mm (0.03 in) for the longest primary feather, which Paul could not confirm using photographs. Paul measured some of the inner primary feathers, finding rachises 1.25–1.4 mm (0.049–0.055 in) across.[94] Despite these criticisms, Nudds and Dyke stood by their original conclusions. They claimed that Paul's statement, that an adult Arxeopteriks would have been a better flyer than the juvenile Munich specimen, was dubious. This, they reasoned, would require an even thicker rachis, evidence for which has not yet been presented.[95] Another possibility is that they had not achieved true flight, but instead used their wings as aids for extra lift while running over water after the fashion of the basilisk lizard, which could explain their presence in lake and marine deposits (see Evolution of bird flight ).[96][97]

Replica of the London Specimen

In 2004, scientists analysing a detailed KTni tekshirish ning braincase London Arxeopteriks concluded that its brain was significantly larger than that of most dinosaurs, indicating that it possessed the brain size necessary for flying. The overall brain anatomy was reconstructed using the scan. The reconstruction showed that the regions associated with vision took up nearly one-third of the brain. Other well-developed areas involved hearing and muscle coordination.[98] The skull scan also revealed the structure of its inner ear. The structure more closely resembles that of modern birds than the inner ear of non-avian reptiles. These characteristics taken together suggest that Arxeopteriks had the keen sense of hearing, balance, spatial perception, and coordination needed to fly.[99] Arxeopteriks had a cerebrum-to-brain-volume ratio 78% of the way to modern birds from the condition of non-coelurosaurian kabi dinozavrlar Carcharodontosaurus yoki Allosaurus, which had a crocodile-like anatomy of the brain and inner ear.[100] Newer research shows that while the Arxeopteriks brain was more complex than that of more primitive theropods, it had a more generalized brain volume among Maniraptora dinosaurs, even smaller than that of other non-avian dinosaurs in several instances, which indicates the neurological development required for flight was already a common trait in the maniraptoran clade.[101]

Recent studies of flight feather barb geometry reveal that modern birds possess a larger barb angle in the trailing vane of the feather, whereas Arxeopteriks lacks this large barb angle, indicating potentially weak flight abilities.[102]

Qayta qurilgan skelet, Dreksel universiteti Tabiiy fanlar akademiyasi

Arxeopteriks continues to play an important part in scientific debates about the origin and evolution of birds. Some scientists see it as a semi-arboreal climbing animal, following the idea that birds evolved from tree-dwelling gliders (the "trees down" hypothesis for the evolution of flight proposed by O. C. Marsh ). Other scientists see Arxeopteriks as running quickly along the ground, supporting the idea that birds evolved flight by running (the "ground up" hypothesis proposed by Samuel Wendell Williston ). Still others suggest that Arxeopteriks might have been at home both in the trees and on the ground, like modern crows, and this latter view is what currently is considered best-supported by morphological characters. Altogether, it appears that the species was not particularly specialized for running on the ground or for perching. A scenario outlined by Elżanowski in 2002 suggested that Arxeopteriks used its wings mainly to escape yirtqichlar by glides punctuated with shallow downstrokes to reach successively higher perches, and alternatively, to cover longer distances (mainly) by gliding down from cliffs or treetops.[21]

In March 2018, scientists reported that Arxeopteriks was likely capable of parvoz, but in a manner distinct and substantially different from that of zamonaviy qushlar.[103][104] This study on Arxeopteriks 's bone histology suggests that it was closest to true flying birds, and in particular to pheasants and other burst flyers.

O'sish

Growth trends compared with other dinosaurs and birds

A gistologik study by Erickson, Norell, Zhongue, and others in 2009 estimated that Arxeopteriks grew relatively slowly compared to modern birds, presumably because the outermost portions of Arxeopteriks bones appear poorly vascularized;[4] in living vertebrates, poorly vascularized bone is correlated with slow growth rate. They also assume that all known skeletons of Arxeopteriks come from juvenile specimens. Because the bones of Arxeopteriks could not be histologically sectioned in a formal skeletochronological (growth ring) analysis, Erickson and colleagues used bone vascularity (porosity) to estimate bone growth rate. They assumed that poorly vascularized bone grows at similar rates in all birds and in Arxeopteriks. The poorly vascularized bone of Arxeopteriks might have grown as slowly as that in a mallard (2.5 micrometres per day) or as fast as that in an ostrich (4.2 micrometres per day). Using this range of bone growth rates, they calculated how long it would take to "grow" each specimen of Arxeopteriks to the observed size; it may have taken at least 970 days (there were 375 days in a Late Jurassic year) to reach an adult size of 0.8–1 kg (1.8–2.2 lb). The study also found that the avialans Jeholornis va Sapeornis grew relatively slowly, as did the dromaeosaurid Mahakala. The avialans Konfutsiyornis va Ixtyornis grew relatively quickly, following a growth trend similar to that of modern birds.[105] One of the few modern birds that exhibit slow growth is the flightless kivi, and the authors speculated that Arxeopteriks and the kiwi had similar bazal metabolizm darajasi.[4]

Kundalik faoliyat turlari

O'rtasidagi taqqoslashlar scleral rings ning Arxeopteriks and modern birds and reptiles indicate that it may have been kunduzgi, similar to most modern birds.[106]

Paleoekologiya

Qayta tiklash Arxeopteriks chasing a juvenile Kompsognatus

The richness and diversity of the Solnhofen ohaktoshlari in which all specimens of Arxeopteriks have been found have shed light on an ancient Jurassic Bavaria strikingly different from the present day. The latitude was similar to Florida, though the climate was likely to have been drier, as evidenced by fossils of plants with adaptations for arid conditions and a lack of terrestrial sediments characteristic of rivers. Evidence of plants, although scarce, include tsikllar and conifers while animals found include a large number of insects, small lizards, pterozavrlar va Kompsognatus.[14]

The excellent preservation of Arxeopteriks fossils and other terrestrial fossils found at Solnhofen indicates that they did not travel far before becoming preserved.[107] The Arxeopteriks specimens found were therefore likely to have lived on the low islands surrounding the Solnhofen lagoon rather than to have been corpses that drifted in from farther away. Arxeopteriks skeletons are considerably less numerous in the deposits of Solnhofen than those of pterosaurs, of which seven genera have been found.[108] The pterosaurs included species such as Ramforinxus ga tegishli Ramphorhynchidae, the group which dominated the joy hozirda egallab olgan dengiz qushlari, and which became extinct at the end of the Jurassic. The pterosaurs, which also included Pterodaktil, were common enough that it is unlikely that the specimens found are sarson-sargardonlar from the larger islands 50 km (31 mi) to the north.[109]

The islands that surrounded the Solnhofen lagoon were low lying, yarim quruq, and sub-tropik uzoq bilan quruq mavsum and little rain.[110] The closest modern analogue for the Solnhofen conditions is said to be Orca Basin shimolda Meksika ko'rfazi, although it is much deeper than the Solnhofen lagoons.[108] The flora of these islands was adapted to these dry conditions and consisted mostly of low (3 m (10 ft)) shrubs.[109] Contrary to reconstructions of Arxeopteriks climbing large trees, these seem to have been mostly absent from the islands; few trunks have been found in the sediments and fossilized tree polen also is absent.

Turmush tarzi Arxeopteriks is difficult to reconstruct and there are several theories regarding it. Some researchers suggest that it was primarily adapted to life on the ground,[111] while other researchers suggest that it was principally arboreal on the basis of the curvature of the claws[112] which has since been questioned.[113] The absence of trees does not preclude Arxeopteriks from an arboreal lifestyle, as several species of bird live exclusively in low shrubs. Various aspects of the morphology of Arxeopteriks point to either an arboreal or ground existence, including the length of its legs and the elongation in its feet; some authorities consider it likely to have been a generalist capable of feeding in both shrubs and open ground, as well as along the shores of the lagoon.[109] It most likely hunted small prey, seizing it with its jaws if it was small enough, or with its claws if it was larger.

Shuningdek qarang

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Qo'shimcha o'qish

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