云南罗平中三叠世翼鳕属一新种及早期辐鳍鱼类系统发育关系

doi:10.19615/j.cnki.2096-9899.210518

摘要/Abstract

摘要：

辐鳍鱼亚纲是现存脊椎动物中最大的类群,包括腕鳍鱼次亚纲、辐鳍鱼次亚纲(包括软骨硬鳞类和新鳍鱼类)和亲缘关系密切的化石类群。已灭绝的翼鳕属(Pteronisculus)是隶属于辐鳍鱼亚纲的一个干群,包括产于马达加斯加、欧洲和北美下三叠统的11个种和中国中三叠统的一个种。根据滇东罗平中三叠世(安尼期)海相地层中发现的5块保存完好的化石,命名翼鳕属一新种,张氏翼鳕(Pteronisculus changae sp. nov.)。这是翼鳕属在中三叠世的第二个确切种,最大体长达295 mm, 代表了罗平生物群中已知体型最大的辐鳍鱼亚纲干群物种。新种具有翼鳕属的独特衍征,泪骨具有牙齿,但它又有明显区别于本属其他种的自近裔特征,如间颞骨中部有一个内突起,21根上神经骨,83列侧线鳞。分支分析结果为早期辐鳍鱼类系统发育关系提供了新的见解,认为翼鳕属是Cyranorhis的姐妹群。根据体型和口缘牙齿等特征推测张氏翼鳕是一个快速游动的捕食者,以浮游无脊椎动物和体型较小的鱼类或鱼类幼体为食。作为翼鳕属最年轻的成员之一,张氏翼鳕的发现进一步表明翼鳕的多样性比我们过去认识的要高,古特提斯洋东缘可能是该属在中三叠世早期的避难所。

关键词: 云南罗平, 中三叠世, 翼鳕属, 辐鳍鱼类, 系统发育

Abstract:

Actinopterygii, the largest group of extant vertebrates, includes Cladistia, Actinopteri (Chondrostei plus Neopterygii) and closely related fossil taxa. The extinct genus Pteronisculus belongs to a stem lineage of actinopterygian fishes represented by 11 species from the Early Triassic of Madagascar, Europe and North America, and a single species from the early Middle Triassic of China. Here, we report the discovery of a new species of this genus, Pteronisculus changae, on the basis of five well-preserved specimens from the Middle Triassic (Anisian) marine deposits exposed in Luoping, eastern Yunnan, China. The discovery documents the second convincing species of Pteronisculus in the Middle Triassic and the largest stem actinopterygian fish in the Luoping Biota, having a maximum total length of up to 295 mm. The new species possesses a toothed lacrimal, which is characteristic ofPteronisculus, but it is easily distinguished from other species of the genus by some autapomorphies, e.g., a medial process at the middle portion of the intertemporal, 21 supraneurals, and 83 lateral line scales. The results of our cladistic analysis provide new insights into the relationships of early actinopterygians and recover Pteronisculus as a sister taxon of the Carboniferous rhadinichthyid Cyranorhis at the actinopterygian stem. Based on the body form, teeth and other features, it can be deduced that Pteronisculus changae is likely a relatively fast-swimming predator, feeding on planktonic invertebrates and smaller or younger fishes known to occur in the same biota. As one of the youngest species of the genus, the new species provides additional evidence to suggest that the diversity of Pteronisculus is higher than previously thought and that the eastern Paleotethys Ocean likely constituted a refuge for species of this genus during the early Middle Triassic.

Key words: Luoping, Yunnan, Middle Triassic, Pteronisculus, Actinopterygii, phylogeny

中图分类号:

Q915.862

任艺, 徐光辉. 云南罗平中三叠世翼鳕属一新种及早期辐鳍鱼类系统发育关系. 古脊椎动物学报, 2021, 59(3): 169-199.

REN Yi, XU Guang-Hui. A new species of Pteronisculus from the Middle Triassic (Anisian) of Luoping, Yunnan, China, and phylogenetic relationships of early actinopterygian fishes. Vertebrata Palasiatica, 2021, 59(3): 169-199.

图/表 17

Fig. 1 Fossil localities of Pteronisculus A. Map showing the global distribution of fossil localities of Pteronisculus: ① Yunnan, China, ② Spitsbergen (Svalbard, Norway), ③ East Greenland, ④ Nevada, USA, ⑤ Madagascar; B. Geographic locality yielding Pteronisculus changae sp. nov. in Luoping, Yunnan, China; C. Map of eastern Yunnan showing the fossil locality (World map: GS(2016)No. 1611; China map: GS(2016)No. 1549)

Fig. 2 Measurements adopted to describe the new species

Fig. 3 Specimens and reconstruction of Pteronisculus changae sp. nov. A. IVPP V 18994, holotype; B. IVPP V 20493; C. reconstruction

Fig. 4 Two specimens of Pteronisculus changae sp. nov. A. IVPP V 24340; B. IVPP V 25615

Table 1 Measurements of Pteronisculus changae sp. nov. from Luoping, Yunnan (mm)

Specimen	ORB	POL	DD	HD	HL	HA	CD	CL	SL	TL
V 18994	9.3	30.2	46.5	56.7	49.1	61.6	21.7	61.2	155.5	221.0
V 24340	10.6	26.7	41.8	49.8	40.1	57.6	19.4	-	138.2	-
V 20493	7.0	23.4	34.0	38.8	38.2	43.5	15.8	44.3	118.7	163.3
V 25615	13.2	31.7	51.5	57.7	52.5	60.1	23.4	65.5	156.0	221.5
V 25618	17.4	34.1	58.4	76.8	64.1	83.1	30.0	80.3	215.0	295.3

Fig. 5 Photograph (A) and line-drawing (B) of skull and pectoral girdle of Pteronisculus changae sp. nov., IVPP V 18994 (holotype)

Fig. 6 Photograph (A) and line-drawing (B) of skull and pectoral girdle of Pteronisculus changae sp. nov., IVPP V 25615

Fig. 7 Photograph (A) and line-drawing (B) of skull and pectoral girdle of Pteronisculus changae sp. nov., IVPP V 24340

Fig. 8 Anterior portion of skull and cheek region of Pteronisculus changae sp. nov. A. anterior potion of skull, IVPP V 20493, dusted with ammonium chloride; B. line-drawing of A; C. maxilla in medial view, IVPP V 18994; D. line-drawing of C

Fig. 9 Paired fins of Pteronisculus changae sp. nov. A. left pectoral fin, IVPP V 18994; B. right pelvic fin, V 25615; C. right pelvic plate, V 24340; D. line-drawing of C

Fig. 10 Fins and scales of Pteronisculus changae sp. nov. A. caudal fin, IVPP V 25615; B. close up of epichordal lobe of caudal fin in A; C. anal fin, V 25615; D. dorsal fin, V 18994; E. pit organs in pre-dorsal region, V 18994

Fig. 11 Scales of Pteronisculus changae sp. nov., IVPP V 25618, dusted with ammonium chloride A. scutes anterior to the anal fin; B. scutes anterior to the ventral lobe of the caudal fin; C. peg-socket articulation between scales; D. lateral line scales; E. scales with white arrows indicating sensory pores; F. scales with black arrows indicating pit organs

Fig. 12 Photograph (A) and line-drawing (B) of supraneural bones and pterygiophores supporting the dorsal fin of Pteronisculus changae sp. nov., IVPP V 25615

Fig. 13 Reconstructions of skulls and pectoral girdles of selected species of Pteronisculus in lateral view A. Pteronisculus changae sp. nov.; B. P. nielseni (from Xu et al., 2014b); C. P. cicatrosus (from Lehman, 1952). D. P. magnus (from Nielsen, 1942); E. P. stensioi (from Nielsen, 1942)

Fig. 14 Reconstructions of skulls of selected species of Pteronisculus in dorsal view A. Pteronisculus changae sp. nov.; B. P. stensioi (from Nielsen, 1942); C. P. magnus (from Nielsen, 1942); D. P. cicatrosus (from Lehman, 1952)

Fig. 15 Comparisons of preopercles (A) and maxillae (B) in different species ofPteronisculus Not to scale

Fig. 16 Strict consensus of 12 most parsimonious trees Tree length = 996 steps, consistency index = 0.3002 and retention index = 0.6715 Bremer indexes larger than 1 are indicated with numbers

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