華南寒武紀(jì)鰓曳動物形態(tài)分類與譜系演化研究
發(fā)布時間：2018-07-13 19:10
【摘要】：鰓曳動物是原口動物亞界蛻皮類群中的一個小門,現(xiàn)生類型迄今只描述了7屬19種。然而,軟軀體鰓曳動物卻是寒武紀(jì)海洋中底內(nèi)生物群落的優(yōu)勢類群,具有非常高的形態(tài)差異度和個體豐度。華南寒武紀(jì)臺地相和斜坡相一系列特異埋藏化石群保存有大量鰓曳動物化石,是研究鰓曳動物早期起源與輻射、蛻皮動物起源與演化以及“寒武紀(jì)大爆發(fā)”等基礎(chǔ)前沿課題的重要化石材料。但是,目前學(xué)術(shù)界對早期鰓曳動物的系統(tǒng)分類、輻射演化、古生態(tài)學(xué)特征和系統(tǒng)演化關(guān)系等問題仍有較大爭議。在系統(tǒng)總結(jié)前人研究成果基礎(chǔ)上,本文對湖北三峽地區(qū)的石牌生物群(臺地相,寒武系第4階)、陜西西鄉(xiāng)三郎鋪地區(qū)的西鄉(xiāng)生物群(臺地相,寒武系第4階)、貴州劍河革東地區(qū)的劍河生物群(斜坡相,寒武系第4階晚期)和凱里生物群(斜坡相,寒武系第5階早期)等化石群中的鰓曳動物化石材料進(jìn)行研究,力求對上述學(xué)術(shù)界長期爭議的科學(xué)問題有所解答。本文系統(tǒng)描述了四個生物群中的古蠕蟲類化石5屬8種,包括3個新屬新種、2個新種和2個未定種。其中,Sanxiascolex papillogyrus gen. et sp. nov., Wronascolex yichangensis sp. nov.和W. spinosus (Ivanstsov and Wrona,2004)描述于石牌生物群；Shaanxiscolex xixiangensis gen. et sp. nov.為西鄉(xiāng)生物群的特有類型：Guizhouscolex balangensis gen. et sp. nov.和Yunnanscolex sp.是劍河生物群的兩個新類型；W. jianhensis sp. nov.和W.sp.則是凱里生物群的化石分子。此外,Gen. et sp. uncertain A和Gen. et sp. uncertain B是兩個分別描述于劍河生物群和凱里生物群的未正式命名新類型。這些化石屬種都具有常見于早古生代地層的Hadimopanella類骨板,但不同類型之間骨板系(scleritomes)特征均顯示出一定差異性,這與地史時期不同產(chǎn)地和層位古蠕蟲類的骨板特征多不相同的情形相一致,說明古蠕蟲類的表皮骨板構(gòu)造可能與海底沉積物的性質(zhì)有關(guān)。論文系統(tǒng)討論了Wronascolex屬的分類原則、古地理分布和地史延限,結(jié)果表明該屬是寒武紀(jì)分異度最高(含8種和3未定種)、分布范圍最廣(華南、西伯利亞、澳大利亞、西班牙和北美)、延限時間最長(寒武系第4階至鼓山階)的古蠕蟲類實體化石。但是,Wronascolex的屬征仍不明確,而且Hadimopanella孤立骨板的形態(tài)分類研究爭議較大,大大削弱了該化石屬及其骨板的地層學(xué)意義。另外,石牌生物群中W. spinosus的骨板具有獨特的排列方式,表明古蠕蟲類的骨板可能具有多種增長模式。本文還研究了云南澄江動物群中的古蠕蟲類Mafangscolex yunnanensis (Luo et Zhang),1986,首次識別出該種軀干腹側(cè)有圓錐狀感覺構(gòu)造和可能的腿肢,為古蠕蟲類形態(tài)學(xué)、古生態(tài)學(xué)和系統(tǒng)演化研究提供了新信息。化石證據(jù)顯示,古蠕蟲類具有較強(qiáng)的掘穴能力,部分類型可能兼有潛穴和底表爬行生活方式,并偶爾食含腐質(zhì)沉積物。盡管古蠕蟲類部分孤立骨板類型(如Hadimopanella)的最早化石記錄可追溯到寒武系第二階晚期,但處于第三階的澄江動物群代表鰓曳動物的首次輻射演化事件。寒武紀(jì)時期不同化石群中鰓曳動物的組合面貌差異很大,多數(shù)類型為各個生物群的特有分子,只有少數(shù)屬種具有較長的地史延限,說明早期鰓曳動物在不同生活環(huán)境下有較好的適應(yīng)能力并發(fā)生快速分異演化。寒武紀(jì)早期揚子臺地斜坡相至臺地相地層都有大量鰓曳動物化石產(chǎn)出,表明華南滇東淺海地區(qū)未必是這一時期鰓曳動物的輻射中心。凱里生物群產(chǎn)有重要鰓曳動物類型Ottoia和Sicyophorus。研究結(jié)果顯示,凱里生物群的奧托蟲體長50-80 mm,外伸的翻吻有25列縱列吻刺,后部短柱形；軀干寬度較均勻,具密集環(huán)紋,達(dá)13條/5mm；咽部和腸道之間有一個可能的肌胃；腸道壁見肌纖維；軀干后部有1對長的后伸縮肌。這些特征明顯不同于布爾吉斯頁巖生物群(Burgess Shale)中的O. prolifica Walcott,1911,命名為貴州奧托蟲O. guizhouensis Yang, Zhao et Zhang,2015。原地埋藏的O. guizhouensis保存有蟲體正在鉆泥掘穴的證據(jù),而且腸道有泥質(zhì)充填物,表明Ottoia是一種能在底表沉積物中多維度自由穿梭、偶爾食泥質(zhì)生活的底內(nèi)動物,并非此前推測那樣穴居于U型管穴中。通過類比澄江動物群中的特殊鰓曳動物類型Sicyophorus rara Luo et al.,1999和Palaeopriapulites parvus Hou et al.,1999,識別出凱里生物群中葫蘆蟲的軀干大約有50條縱列隔板,頸部有2-3條環(huán)紋,末端有一條較短的尾突構(gòu)造,因而建立貴州葫蘆蟲(新種)S. guizhouensis sp. nov.。形態(tài)學(xué)和解剖學(xué)證據(jù)表明,Sicyophorus既有相似于鰓曳動物的形態(tài)構(gòu)造(翻吻和尾突),也有類似于鎧甲動物的軀干特征(具縱列隔板),而卷曲盤繞的腸道區(qū)別于其他環(huán)神經(jīng)類動物,可能是鰓曳動物向鎧甲動物演化過程中幼態(tài)持續(xù)發(fā)育而來的一個特化類型。本文運用TNT軟件對O. guizhouensis和S. guizhouensis的系統(tǒng)演化位置進(jìn)行分析。譜系分支樹顯示,Ottoia屬于Scalidophora干群類型,而Sicyophorus位于Scalidophora冠群底部靠近鎧甲動物的位置,支持由鰓曳動物幼態(tài)持續(xù)發(fā)育而來的觀點。此外,古蠕蟲類不同屬種的系統(tǒng)演化位置差別很大。根據(jù)形態(tài)學(xué)、解剖學(xué)證據(jù)和譜系分析,論文討論了兩側(cè)對稱動物祖先類型Urbilateria-葉足動物Xenusia- Scalidophora干群類型Fieldiai和Ancalagonia—古蠕蟲類Palaeoscolecida—Louisellia—鰓曳動物Priapulida—葫蘆蟲Sicyophourus—鎧甲動物L(fēng)oricifera整個演化線系中各個節(jié)點動物代表的身體構(gòu)型特征,認(rèn)為部分古蠕蟲類(如M.yunnanensis)可能具有相似于蛻皮動物祖先類型的形態(tài)特征。
[Abstract]:Branchial tracer is a small door in the subboundary molt group of the original animal, and the present type has only described 7 genera and 19 species so far. However, the soft somatic branchial tracer is the dominant group in the Cambrian middle subsea biological community, with very high morphological difference and individual abundance. A series of special burial of the Cambrian platform and slope facies in Southern China. The stone group has a large number of branchial fossils. It is an important fossil material for the study of the early origin and radiation of the branchial tracers, the origin and evolution of molt, and the "Cambrian outbreak". However, the systematic classification, radiological evolution, paleo ecological characteristics and systematic evolution of the early branchial tracers in the academic circles are at present. On the basis of a systematic summary of the achievements of previous studies, this paper is based on the stone brand biota in the Three Gorges area (terracostage, Cambrian fourth), Xixiang biota (terrace, Cambrian, fourth) in the Sanlun area, Xixiang, Xixiang, and the Jianhe biota (slope facies, late Cambrian in the late fourth order) and Kay in Jianhe, Guizhou. The study of the branchial fossil materials in the fossils of the fossils of the biota (slope facies and the Early Cambrian fifth order) has been studied in order to solve the scientific problem of the long term controversy in the academic circles. This paper systematically describes the 5 genera and 8 species of the paleo worms of the four biota, including 3 new species, 2 new species and 2 undetermined species. Among them, Sanxiasc Olex papillogyrus Gen. et sp. Nov., Wronascolex yichangensis sp. Nov. and W. spinosus, described in the Xixiang biota of biota. The two new types, W. jianhensis sp. Nov. and W.sp. are the fossil molecules of the Kaili biota. In addition, Gen. et sp. uncertain A and Gen. et are two unofficially named new types described respectively in the biota and the Kaili biota. These species are all common in the early Paleozoic strata. Bone plate, but the characteristics of different types of bone plate system (scleritomes) show a certain difference, which is consistent with the different characteristics of the bone plate characteristics of different habitats and layers of ancient worms in the period of geological history, indicating that the structure of the epidermal bone plate of the ancient worms may be related to the nature of the seabed sediments. This paper systematically discusses the Wronascolex genus. The classification principle, palaeogeographic distribution and earth history extension limit indicate that the genus is the highest (8 species and 3 undetermined species) in Cambrian (including 8 species and 3 undetermined species), and the most widely distributed (Southern China, Siberia, Australia, Spain and North America) and the oldest (Cambrian, fourth to GUSHAN) paleogeic solid fossils. However, the genera of Wronascolex are still not clear. Moreover, the study of the morphological classification of the Hadimopanella isolated bone plates has greatly weakened the Stratigraphic Significance of the fossil genus and its bone plate. In addition, the bone plate of the W. spinosus in the stone brand biota has a unique arrangement, indicating that the bone plates of the ancient worms may have a variety of growth patterns. This paper also studied the Chengjiang fauna of Yunnan. The ancient worms Mafangscolex yunnanensis (Luo et Zhang), 1986, identified the conical structure and possible leg limbs on the ventral side of the trunk for the first time. It provides new information for the study of the morphological, paleecology and systematic evolution of the ancient worms. Fossil evidence shows that the ancient worms have strong point digging ability and some types may have both potential and potential. Although the earliest fossil records of partial solitary bone plate types (such as Hadimopanella) can be traced back to the late Cambrian second stage of the Cambrian, the third order Chengjiang fauna represented the first radiative evolution of the branchial tract. The combination features of the tractable animals are very different. Most of them are endemic molecules of the various biota. Only a few species have a long history delay limit. It shows that the early branchial tracer had better adaptability and rapid differentiation in different living environments. The tracer fossils are produced, indicating that the Southern China East Yunnan shallow sea area is not necessarily the radiant center of the branchial tracts in this period. The Kaili biota produces an important branchial type Ottoia and Sicyophorus., which shows that the Otto body of the Kaili biota has a long 50-80 mm length, and the extended kiss has 25 columns of spines, a short column in the rear, and the trunk width of the trunk. Uniform, dense ring lines, up to 13 /5mm; there is a possible muscle stomach between the pharynx and the intestine; the intestinal wall appears muscle fiber; the trunk of the trunk has 1 pairs of long hind extention muscles. These characteristics are distinctly different from the O. prolifica Walcott in the Burgess Shale biota (Shale), 1911, named the O. guizhouensis Yang, Zhao of the Guizhou Otto worm, Zhao. Et Zhang, 2015. buried O. guizhouensis preserved the evidence that the body is drilling mud, and there is a muddy filling in the intestines, indicating that Ottoia is a kind of inward animal that can freely shuttle in the bottom surface sediments and occasionally live in mud. It was not previously assumed that the cavern was in the U cavern. By analogy with the Chengjiang fauna. The special branchial tracer types, Sicyophorus rara Luo et al., 1999 and Palaeopriapulites parvus Hou et al., 1999, identified the trunk of the cucurbits in the Kaili biota with about 50 column separators, 2-3 rings in the neck and a short tail process at the end, thus establishing the Guizhou cucurbits (new species) S. guizhouensis. Morphological and anatomical evidence shows that Sicyophorus has both morphological structure (kissing and caudate) similar to the branchial tracers, and also the trunk characteristics of the armour (with a column partition), while the curly coiled intestines are different from other annulus, and may be the continuous development of the branchlets during the evolution of the armour to the armour. This article uses TNT software to analyze the system evolution position of O. guizhouensis and S. guizhouensis. The branch tree of the pedigree shows that Ottoia belongs to the Scalidophora group type, and Sicyophorus is located at the bottom of the Scalidophora canopy near the position of the armour, supporting the continuous development of the branchial tractable immature state. In addition, the system evolution position of different species of ancient worms is very different. According to the morphological, anatomical evidence and genealogical analysis, the paper discusses the Xenusia- Scalidophora trunk type Fieldiai of the Urbilateria- leaf feet of bilaterally symmetrical animals and Ancalagonia - the Palaeoscolecida - Louisellia - branchial Priap Ulida - the body configuration characteristics of the representative of each node in the entire evolution line of the Sicyophourus - armour Loricifera of cucurbit cucurbit. It is believed that some of the ancient worms (such as M.yunnanensis) may have similar morphological characteristics to the ancestral types of the molting animals.
【學(xué)位授予單位】：西北大學(xué)
【學(xué)位級別】：博士
【學(xué)位授予年份】：2016
【分類號】：Q915
本文編號：2120457
[Abstract]:Branchial tracer is a small door in the subboundary molt group of the original animal, and the present type has only described 7 genera and 19 species so far. However, the soft somatic branchial tracer is the dominant group in the Cambrian middle subsea biological community, with very high morphological difference and individual abundance. A series of special burial of the Cambrian platform and slope facies in Southern China. The stone group has a large number of branchial fossils. It is an important fossil material for the study of the early origin and radiation of the branchial tracers, the origin and evolution of molt, and the "Cambrian outbreak". However, the systematic classification, radiological evolution, paleo ecological characteristics and systematic evolution of the early branchial tracers in the academic circles are at present. On the basis of a systematic summary of the achievements of previous studies, this paper is based on the stone brand biota in the Three Gorges area (terracostage, Cambrian fourth), Xixiang biota (terrace, Cambrian, fourth) in the Sanlun area, Xixiang, Xixiang, and the Jianhe biota (slope facies, late Cambrian in the late fourth order) and Kay in Jianhe, Guizhou. The study of the branchial fossil materials in the fossils of the fossils of the biota (slope facies and the Early Cambrian fifth order) has been studied in order to solve the scientific problem of the long term controversy in the academic circles. This paper systematically describes the 5 genera and 8 species of the paleo worms of the four biota, including 3 new species, 2 new species and 2 undetermined species. Among them, Sanxiasc Olex papillogyrus Gen. et sp. Nov., Wronascolex yichangensis sp. Nov. and W. spinosus, described in the Xixiang biota of biota. The two new types, W. jianhensis sp. Nov. and W.sp. are the fossil molecules of the Kaili biota. In addition, Gen. et sp. uncertain A and Gen. et are two unofficially named new types described respectively in the biota and the Kaili biota. These species are all common in the early Paleozoic strata. Bone plate, but the characteristics of different types of bone plate system (scleritomes) show a certain difference, which is consistent with the different characteristics of the bone plate characteristics of different habitats and layers of ancient worms in the period of geological history, indicating that the structure of the epidermal bone plate of the ancient worms may be related to the nature of the seabed sediments. This paper systematically discusses the Wronascolex genus. The classification principle, palaeogeographic distribution and earth history extension limit indicate that the genus is the highest (8 species and 3 undetermined species) in Cambrian (including 8 species and 3 undetermined species), and the most widely distributed (Southern China, Siberia, Australia, Spain and North America) and the oldest (Cambrian, fourth to GUSHAN) paleogeic solid fossils. However, the genera of Wronascolex are still not clear. Moreover, the study of the morphological classification of the Hadimopanella isolated bone plates has greatly weakened the Stratigraphic Significance of the fossil genus and its bone plate. In addition, the bone plate of the W. spinosus in the stone brand biota has a unique arrangement, indicating that the bone plates of the ancient worms may have a variety of growth patterns. This paper also studied the Chengjiang fauna of Yunnan. The ancient worms Mafangscolex yunnanensis (Luo et Zhang), 1986, identified the conical structure and possible leg limbs on the ventral side of the trunk for the first time. It provides new information for the study of the morphological, paleecology and systematic evolution of the ancient worms. Fossil evidence shows that the ancient worms have strong point digging ability and some types may have both potential and potential. Although the earliest fossil records of partial solitary bone plate types (such as Hadimopanella) can be traced back to the late Cambrian second stage of the Cambrian, the third order Chengjiang fauna represented the first radiative evolution of the branchial tract. The combination features of the tractable animals are very different. Most of them are endemic molecules of the various biota. Only a few species have a long history delay limit. It shows that the early branchial tracer had better adaptability and rapid differentiation in different living environments. The tracer fossils are produced, indicating that the Southern China East Yunnan shallow sea area is not necessarily the radiant center of the branchial tracts in this period. The Kaili biota produces an important branchial type Ottoia and Sicyophorus., which shows that the Otto body of the Kaili biota has a long 50-80 mm length, and the extended kiss has 25 columns of spines, a short column in the rear, and the trunk width of the trunk. Uniform, dense ring lines, up to 13 /5mm; there is a possible muscle stomach between the pharynx and the intestine; the intestinal wall appears muscle fiber; the trunk of the trunk has 1 pairs of long hind extention muscles. These characteristics are distinctly different from the O. prolifica Walcott in the Burgess Shale biota (Shale), 1911, named the O. guizhouensis Yang, Zhao of the Guizhou Otto worm, Zhao. Et Zhang, 2015. buried O. guizhouensis preserved the evidence that the body is drilling mud, and there is a muddy filling in the intestines, indicating that Ottoia is a kind of inward animal that can freely shuttle in the bottom surface sediments and occasionally live in mud. It was not previously assumed that the cavern was in the U cavern. By analogy with the Chengjiang fauna. The special branchial tracer types, Sicyophorus rara Luo et al., 1999 and Palaeopriapulites parvus Hou et al., 1999, identified the trunk of the cucurbits in the Kaili biota with about 50 column separators, 2-3 rings in the neck and a short tail process at the end, thus establishing the Guizhou cucurbits (new species) S. guizhouensis. Morphological and anatomical evidence shows that Sicyophorus has both morphological structure (kissing and caudate) similar to the branchial tracers, and also the trunk characteristics of the armour (with a column partition), while the curly coiled intestines are different from other annulus, and may be the continuous development of the branchlets during the evolution of the armour to the armour. This article uses TNT software to analyze the system evolution position of O. guizhouensis and S. guizhouensis. The branch tree of the pedigree shows that Ottoia belongs to the Scalidophora group type, and Sicyophorus is located at the bottom of the Scalidophora canopy near the position of the armour, supporting the continuous development of the branchial tractable immature state. In addition, the system evolution position of different species of ancient worms is very different. According to the morphological, anatomical evidence and genealogical analysis, the paper discusses the Xenusia- Scalidophora trunk type Fieldiai of the Urbilateria- leaf feet of bilaterally symmetrical animals and Ancalagonia - the Palaeoscolecida - Louisellia - branchial Priap Ulida - the body configuration characteristics of the representative of each node in the entire evolution line of the Sicyophourus - armour Loricifera of cucurbit cucurbit. It is believed that some of the ancient worms (such as M.yunnanensis) may have similar morphological characteristics to the ancestral types of the molting animals.
【學(xué)位授予單位】：西北大學(xué)
【學(xué)位級別】：博士
【學(xué)位授予年份】：2016
【分類號】：Q915
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