珠江三角洲平原螃蟹遗迹特征及其对古环境的指示意义<sup>*</sup>

doi:10.7605/gdlxb.2025.00.027

摘要/Abstract

摘要： 螃蟹是生态环境工程师,可以灵敏反映和改造其所在的沉积和生态环境。珠江三角洲平原地区沉积环境复杂,螃蟹种类多样且数量丰富,但区域内不同种类螃蟹的分布和组成特征及其对古环境的指示意义尚缺乏研究。本文在珠江三角洲平原选取92个采样点进行水样、土样采集,个别采样点进行生物潜穴的取心,同时观察和拍照记录各个采样点螃蟹遗迹的分布与特征,利用盐度、粒度分析、CT扫描和三维重构等方法对沉积环境和螃蟹遗迹进行研究。结果表明: (1)珠江三角洲平原主要生活的螃蟹有方蟹总科相手蟹属无齿东方相手蟹、中华东方相手蟹和沙蟹总科沙蟹属丽彩拟瘦招潮等,它们形成的遗迹包括足辙迹、觅食迹、排泄迹、潜穴等,潜穴形状大多为I形、J形,还有少量Y形。(2)螃蟹遗迹主要分布在受潮汐影响的潮间带位置,其中河口区域螃蟹遗迹的丰度最高,分流河道和滨海湿地相对较少。(3)类比发现,螃蟹遗迹的形态特征和沉积环境与遗迹化石Psilonichnus类似,主要分布在潮间带的坚固地面环境中,与螃蟹遗迹相关的遗迹相为Glossifungites相、Psilonichnus相和Skolithos相。该研究不仅补充了珠江三角洲平原的现代沉积学资料,而且还揭示出螃蟹遗迹对古环境的指示意义,为地质学和生态学研究提供了重要线索和依据。

关键词: 现代生物遗迹, 遗迹学, 螃蟹, 遗迹化石, 珠江三角洲

Abstract:

Crabs serve as ecological engineers capable of sensitively reflecting and modifying sedimentary and ecological environments in which they reside. The sedimentary environment of the Pearl River Delta Plain is highly complex,which is characterized by a diverse and abundant crab population. This paper focuses on investigating the distribution and compositional characteristics of different crab species within the region,as well as their significance in indicating paleoenvironments. A total of 92 sampling points in the Pearl River Delta Plain were established to collect water samples,soil samples,and biogenic submerged burrow cores. Observations and photographic documentation were conducted to record the distribution and characteristics of crab traces at each sampling point. Subsequently,sedimentary environments and crab traces within the study area were analyzed using methods such as salinity measurement,granulometric analysis,CT scanning,and three-dimensional reconstruction. Key findings include: (1)The predominant crab species identified were Orisarma dehaani and Orisarma sinense from the family Grapsidae,Grapsoidea,and Paraleptuca splendida from the family Ocypodidae,Ocypodoidea. Crab traces included tracks,fodinichnia,waste trails,and burrows,with most burrows exhibiting I-and J-shaped configurations and a few Y-shaped structures. (2)Crab traces were predominantly distributed in intertidal zones influenced by tides,with the highest abundance observed in estuarine areas,and relatively fewer traces in diversion channels and coastal wetlands. (3)Through analogy,it was determined that the morphological characteristics and depositional environments of crab traces resemble those of the trace fossil Psilonichnus,which is primarily found in shallow marine firmground environments. Additionally,trace phases associated with crab traces include Glossifungites,Psilonichnus,and Skolithos. This study contributes modern sedimentological data for the Pearl River Delta Plain,reveals the paleoenvironmental significance of crab traces,and provides critical evidence for geological and ecological research.

Key words: modern biological traces, ichnology, crabs, trace fossil, Pearl River Delta

中图分类号:

Q958.1

王媛媛, 张亚斌, 勾松林. 珠江三角洲平原螃蟹遗迹特征及其对古环境的指示意义^*[J]. 古地理学报, 2025, 27(3): 667-683.

WANG Yuanyuan, ZHANG Yabin, GOU Songlin. Characteristics of modern crab traces in the Pearl River Delta Plain and their significance as indicators of palaeoenvironments[J]. Journal of Palaeogeography（Chinese Edition）, 2025, 27(3): 667-683.

http://www.gdlxb.cn/CN/Y2025/V27/I3/667

图/表 15

图1 珠江三角洲平原位置图(卫星底图来自于Google Earth Pro)

Fig.1 Location map of the Pearl River Delta Plain(the satellite basemap from Google Earth Pro)

图2 珠江三角洲平原采样点分布

Fig.2 Distribution map of sampling sites in the Pearl River Delta Plain

图3 珠江三角洲平原沉积构造特征 A—平行层理; B—浪成波痕; C—干涉波痕; D—纵向波痕

Fig.3 Sedimentary structure characteristics of the Pearl River Delta Plain

图4 无齿东方相手蟹营造遗迹 A—无齿东方相手蟹; B—无齿东方相手蟹性别图,上位雌蟹,下为雄蟹; C—近圆形居住潜穴,潜穴口周围有挖出的泥浆; D—螃蟹排泄迹,为细条状物; E—J形潜穴; F—Y形潜穴口; G—I形潜穴; H—与图F相对应的Y形潜穴; I—CT扫描潜穴,近乎垂直的I形

Fig.4 Construction traces by Orisarma dehaani

图5 中华东方相手蟹营造遗迹 A—中华东方相手蟹; B—中华东方相手蟹潜穴遗迹; C—有树枝遮挡的潜穴; D—在潜穴口观望的中华东方相手蟹

Fig.5 Construction traces by Orisarma sinense

图6 字纹弓蟹营造遗迹 A—觅食迹,在潜穴口围成一圈的近圆形泥球颗粒; B—围墙形潜穴口,离地层表面1~3 cm;C—I形潜穴; D—J形潜穴

Fig.6 Construction traces by Varuna litterata

图7 大陆拟相手蟹营造遗迹 A—I形潜穴,里边居住着大陆拟相手蟹; B—分布密集的潜穴及螃蟹进食过的泥球颗粒

Fig.7 Construction traces by Parasesarma continentale

图8 长足长方蟹营造遗迹 A—斜向潜穴; B—伴有水坑的潜穴; C—点坑状的无规则足辙迹; D—成排足辙迹

Fig.8 Construction traces by Metaplax longipe

图9 丽彩拟瘦招潮营造遗迹 A—觅食迹; B—烟囱状潜穴,离地层表面1~4 cm;C—I形潜穴; D—潜穴周围密集的点坑状足辙迹

Fig.9 Construction traces by Paraleptuca splendida

图10 珠江三角洲平原盐度分布图

Fig.10 Salinity distribution in the Pearl River Delta Plain

图11 珠江三角洲平原粒度分布图

Fig.11 Grain size distribution in the Pearl River Delta Plain

图12 珠江三角洲平原螃蟹遗迹分布三维模式

Fig.12 Three-dimensional model of crab construction traces distribution in the Pearl River Delta Plain

图13 Psilonichnus 的不同形状 A-D—Psilonichnus quietis的4种不同形状的隆起,A为半圆形隆起,B为半三角形隆起,C为鞋形隆起,D为腿形隆起,根据 Myint(2001,图5,第 7页)重新绘制; E—Psilonichnus tubiformis 的Y形潜穴,根据Neto de Carvalho(2016,图3-c,第5页)重新绘制; F—Psilonichnus upsilon的 J形潜穴,根据Frey(1984,图9,第344页)重新绘制; G—Psilonichnus 显示向下变细,根据Humphreys和Balson(1988,图3,第170页)重新绘制; H—Psilonichnus 的分枝,根据Doyle等(2021, 图2-b,第4页)重新绘制。均未按比例绘制

Fig.13 Various shapes of Psilonichnus

表2 珠江三角洲平原螃蟹遗迹与类似遗迹化石的形态特征和参数对比

Table2 Comparison of morphological features and parameters between crab traces in the Pearl River Delta Plain and similar trace fossils

遗迹和遗迹化石		珠江三角洲平原螃蟹遗迹	Psilonichnus tubiformis	Psilonichnus upsilon	Psilonichnus lutimuratus	Psilonichnus upsilon
基本描述和重要特征		简单的I形、J形和Y形潜穴; 横截面为圆形,少有分枝; 潜穴无衬里,内部光滑; 密度较大	上部为垂直的Y形或U形,向下延伸为直管或略微弯曲的管状; 圆柱形潜穴,外侧有短的分枝	略微倾斜,几乎不分枝; J形或Y形潜穴; 无衬里的潜穴	横截面呈圆形; 基底渐细且稍弯曲; 光滑的内外壁; 少有侧枝或没有侧枝; 密集、重叠、不规则	略倾斜; 主要为I形,极少的Y形; 无衬里的圆柱形潜穴; 整个结构在一个垂直平面内
测量参数	长度/cm	5~15	最大超200	多为120	80	10~208
	直径/cm	0.4~3.5	3~5	2.5~4.5	0.5~3	1.2~4
沉积环境特征		沙和粉沙; 近海岸线潮间带、分流河道	粉砂质黏土岩、粉砂岩; 潮间带、低能环境	中到细粒的钙钛矿(碳酸盐岩); 前滨上部至后滨,海滩上部至沙丘下部,潮间带	砂岩和泥岩; 开阔的、以风暴和波浪为主的河口点条状沉积物	中砂岩; 前滨到后滨; 很可能是后滨
资料来源		本文	Fürsich,1981; Frey et al., 1984; Gingras et al., 2000	Frey et al., 1984; Gingras et al., 2000	Nesbit and Campbell,2002	Olszewska-Nejbert et al., 2020

参考文献 59

[1]	陈顺洋, 陈光程, 陈彬, 叶勇, 马志远. 2014. 红树林湿地相手蟹科动物摄食生态研究进展. 生态学报, 34(19): 5349-5359.
	[Chen S Y, Chen G C, Chen B, Ye Y, Ma Z Y. 2014. Feeding ecology of sesarmid crabs in mangroves. Acta Ecologica Sinica, 34(19): 5349-5359]
[2]	陈晓旋, 陈淑云, 曾从盛, 王维奇. 2018. 螃蟹对闽江河口湿地土壤碳氮磷含量及其生态化学计量学特征影响. 环境科学学报, 38(3): 1179-1188.
	[Chen X X, Chen S Y, Zeng C S, Wang W Q. 2018. Effects of crabs on soil carbon,nitrogen,phosphorus concentration and ecological stoichiometry in Minjiang River estuarine wetlands. Acta Scientiae Circumstantiae, 38(3): 1179-1188]
[3]	陈耀泰. 1992. 珠江口现代沉积速率与沉积环境. 中山大学学报(自然科学版), 31(2): 100-107.
	[Chen Y T. 1992. Modern sedimentary velocity and sedimetary environment in the Pearl River Mouth. Acta Scientiarum Natuaralium Uniniversitatis Sunyatseni, 31(2): 100-107]
[4]	程明豪. 1984. 西江河口(磨刀门)沉积环境分析. 地理研究, 3(1): 51-65. doi: 10.11821/yj1984010009
	[Cheng M H. 1984. A study of the sedimentary environment in Modaomen estuary,the main river-mouth of Xijiang(west river). Geographical Research, 3(1): 51-65]
[5]	崔伟中. 2004. 珠江河口水环境时空变异对河口生态系统的影响. 水科学进展, 15(4): 472-478.
	[Cui W Z. 2004. Impact of temporal and spatial variation of water environment on ecosystem of the Pearl River estuary. Advances in Water Science, 15(4): 472-478]
[6]	戴仕宝, 杨世伦, 蔡爱民. 2007. 51年来珠江流域输沙量的变化. 地理学报, 62(5): 545-554.
	[Dai S B, Yang S L, Cai A M. 2007. Variation of sediment discharge of the Pearl River Basin from 1955 to 2005. Acta Geographica Sinica, 62(5): 545-554]
[7]	龚一鸣, 胡斌, 卢宗盛, 齐永安, 张国成. 2009. 中国遗迹化石研究80年. 古生物学报, 48(3): 322-337.
	[Gong Y M, Hu B, Lu Z S, Qi Y A, Zhang G C. 2009. Study of trace fossils in the past eighty years in China. Acta Palaeontologica Sinica, 48(3): 322-337]
[8]	胡斌, 王冠忠, 齐永安. 1997. 痕迹学理论与应用. 江苏徐州: 中国矿业大学出版社.
	[Hu B, Wang G Z, Qi Y A. 1997. Theory and Application of Trace Science. Jiangsu Xuzhou: China University of Mining and Technology Press]
[9]	胡斌, 王媛媛, 张璐, 单婷婷. 2012. 黄河中下游焦作区段现代边滩沉积中的生物遗迹. 古地理学报, 14(5): 628-638. doi: 10.7605/gdlxb.2012.05.008
	[Hu B, Wang Y Y, Zhang L, Shan T T. 2012. Biogenic traces in modern point bar deposits of the Middle-Lower Reaches of Yellow River in Jiaozuo area,Henan Province. Journal of Palaeogeography(Chinese Edition), 14(5): 628-638]
[10]	胡斌, 齐永安, 宋慧波, 牛永斌, 张立军, 郑伟, 王长征. 2021. 中国遗迹学研究十年进展. 古地理学报, 23(2): 284-320. doi: 10.7605/gdlxb.2021.02.021
	[Hu B, Qi Y A, Song H B, Niu Y B, Zhang L J, Zheng W, Wang C Z. 2021. Research progress of Chinese ichnology in recent ten years. Journal of Palaeogeography(Chinese Edition), 23(2): 284-320]
[11]	梁伟诺, 何克宏, 胡亮. 2023. 珠江口淇澳岛潮间带蟹类(十足目: 短尾下目)物种多样性与分布. 海洋学报, 45(11): 76-87.
	[Liang W N, He K H, Hu L. 2023. Intertidal crabs(Decapoda: Brachyura)species diversity and distribution in Qi'ao Island,Zhujiang River Estuary. Haiyang Xuebao, 45(11): 76-87]
[12]	龙云作, 霍春兰, 司桂贤, 杨聿容, 朱卫勤, 石瑞祥, 牛作民, 马道修, 徐明广, 周青伟, 张光威, 蓝先洪. 1985. 对珠江三角洲沉积特征和沉积模式的一些认识. 海洋地质与第四纪地质, 5(4): 49-57.
	[Long Y Z, Huo C L, Si G X, Yang Y R, Zhu W Q, Shi R X, Niu Z M, Ma D X, Xu M G, Zhou Q W, Zhang G W, Lan X H. 1985. On sedimentary characteristics and model of Zhujiang River Delta. Marine Geology & Quaternary Geology, 5(4): 49-57]
[13]	龙云作, 霍春兰, 杨胜雄. 1989. 珠江三角洲现代沉积环境及沉积特征. 海洋地质与第四纪地质, 9(4): 15-27.
	[Long Z Y, Huo C L, Yang S X. 1989. Modern sedimentary environment and characteristics of the Zhujiang River Delta. Marine Geology & Quaternary Geology, 9(4): 15-27]
[14]	罗秉征. 1992. 河口及近海的生态特点与渔业资源. 长江流域资源与环境, 1(1): 24-30.
	[Luo B Z. 1992. Ecological characteristics and fishery resources of the Yangtze River Estuary and adjacent sea. Resources and Environment in the Yangtze Valley, 1(1): 24-30]
[15]	潘保柱, 王兆印, 何雪宝. 2011. 西江大型底栖动物群落特征研究. 水生生物学报, 35(5): 851-856.
	[Pan B Z, Wang Z Y, He X B. 2011. Studies on assemblage characteristics of macrozoobenthos in the west river. Acta Hydrobiologica Sinica, 35(5): 851-856]
[16]	宋慧波, 于会新, 王海邻, 范代读, 胡斌, 王芳. 2014. 杭州湾庵东浅滩现代沉积物中的生物遗迹. 古地理学报, 16(5): 703-714. doi: 10.7605/gdlxb.2014.05.56
	[Song H B, Yu H X, Wang H L, Fan D D, Hu B, Wang F. 2014. Biogenic traces in modern shoal deposits of Andong area,Hangzhou Bay. Journal of Palaeogeography(Chinese Edition), 16(5): 703-714]
[17]	苏亚芳. 1992. 谈珠江三角洲的形成与发展. 遥感信息,(3): 29-30.
	[Su Y F. 1992. Talking about the formation and development of the Pearl River Delta. Remote Sensing Information,(3): 29-30]
[18]	王翠. 2022. 黄河三角洲潮坪环境中现代生物遗迹的组成和分布特征. 河南理工大学硕士学位论文.
	[Wang C. 2022. The composition and distribution characteristics of biogenic sedimentary structures in tidal flat of the Yellow River Delta. Masteral dissertation of Henan Polytechnic University]
[19]	王翠, 王媛媛, 胡斌. 2023. 黄河三角洲潮坪环境现代生物遗迹与物化条件的响应关系. 沉积学报, 41(3): 748-762.
	[Wang C, Wang Y Y, Hu B. 2023. The response relationship between biogenic structures and physicochemical stresses of the Yellow River deltaic tidal flat. Acta Sedimentologica Sinica, 41(3): 748-762]
[20]	王海邻, 张彬, 宋慧波, 王长征, 胡斌, 张立军. 2022. 滦河三角洲潮坪环境现代生物遗迹组合及其分布特征. 古地理学报, 24(6): 1179-1192. doi: 10.7605/gdlxb.2022.06.074
	[Wang H L, Zhang B, Song H B, Wang C Z, Hu B, Zhang L J. 2022. Assemblages of lebensspuren and distribution characteristics in tidal flat of Luanhe delta. Journal of Palaeogeography(Chinese Edition), 24(6): 1179-1192]
[21]	王建华, 曹玲珑, 王晓静, 杨小强, 阳杰, 苏志华. 2009. 珠江三角洲万顷沙地区晚第四纪沉积相与古环境演变. 海洋地质与第四纪地质, 29(6): 35-41.
	[Wang J H, Cao L L, Wang X J, Yang X Q, Yang J, Su Z H. 2009. Evolution of sedimentary facies and paleoenvironment during the late Quaternary in Wanqingsha area of the Pearl River delta. Marine Geology & Quaternary Geology, 29(6): 35-41]
[22]	王金庆. 2008. 长江口盐沼优势蟹类的生境选择与生态系统工程师效应. 复旦大学博士学位论文.
	[Wang J Q. 2008. Habitat selection and ecosystem engineering effects of dominant burrowing crabs in salt marshes in the Yangtze River Estuary,China. Doctoral dissertation of Fudan University]
[23]	王珊珊. 2008. 珠江三角洲和近岸河口海域现代沉积环境及晚更新世以来的环境演变. 中国海洋大学博士学位论文.
	[Wang S S. 2008. Modern sedimentary environment and environmental evolvement since Late Pleistocene for the Pearl River Delta and intracoastal estuary and sea area. Doctoral dissertation of Ocean University of China]
[24]	王媛媛, 王学芹, 胡斌. 2019. 黄河三角洲潮坪环境中现代生物遗迹组成与分布特征. 沉积学报, 37(6): 1244-1257.
	[Wang Y Y, Wang X Q, Hu B. 2019. The composition and distribution characteristics of biogenic sedimentary structures in tidal flat of Yellow River Delta. Acta Sedimentologica Sinica, 37(6): 1244-1257]
[25]	王媛媛, 王翠, 王学芹, 胡斌, 王长征. 2020. X射线计算机断层扫描技术在生物遗迹识别方面的应用. 现代地质, 34(6): 1221-1229.
	[Wang Y Y, Wang C, Wang X Q, Hi B, Wang C Z. 2020. Application of X-ray computed tomography in identification of biogenic structures. Geoscience, 34(6): 1221-1229]
[26]	王媛媛, 勾松林, 张国成. 2024. 现代珠江三角洲前缘生物遗迹组成与分布特征. 沉积学报, 42(5): 1512-1529.
	[Wang Y Y, Gou S L, Zhang G C. 2024. The composition and distribution characteristics of biological traces in the Pearl River Delta front. Acta Sedimentologica Sinica, 42(5): 1512-1529]
[27]	韦惺, 吴超羽. 2018. 珠江三角洲沉积体与河网干流河道的形成发育. 海洋学报, 40(7): 66-78.
	[Wei X, Wu C Y. 2018. The formation and development of the deposition bodies and main channels in the Zhujiang River Delta. Haiyang Xuebao, 40(7): 66-78]
[28]	吴超羽, 韦惺. 2021. 从溺谷湾到三角洲: 现代珠江三角洲形成演变研究辨析. 海洋学报, 43(1): 1-26.
	[Wu C Y, Wei X. 2021. From drowned valley to delta: discrimination and analysis on issues of the formation and evolution of the Zhujiang River Delta. Haiyang Xuebao, 43(1): 1-26]
[29]	吴洁, 刘春莲, 张素青, 杨婷婷, 殷鉴. 2014. 珠江三角洲东部晚第四纪微体动物群的古环境意义. 中山大学学报(自然科学版), 53(6): 45-52.
	[Wu J, Liu C L, Zhang S Q, Yang T T, Yin J. 2014. Palaeoenvironmental significance of Late Quaternary microfaunas in the eastern Pearl River Delta. Acta Scientiarum Naturalium Universitatis Sunyatseni, 53(6): 45-52]
[30]	吴月琴, 刘春莲, 杨小强, 黄毅, 殷鉴, 张恺. 2019. 珠江三角洲中部全新世以来的微体动物群记录与古环境重建. 海洋地质与第四纪地质, 39(2): 31-43.
	[Wu Y Q, Liu C L, Yang X Q, Hang Y, Yin J, Zhang K. 2019. Holocene microfaunal records in the central Pearl River Delta and implications for palaeoenvironmental changes. Marine Geology & Quaternary Geology, 39(2): 31-43]
[31]	徐永健, 申屠基康, 丁张妮. 2015. 砂粒粒径与砂层厚度对单体筐养养殖系统中三疣梭子蟹(Portunus trituberculatus)幼蟹摄食行为与生长特性的影响研究. 海洋与湖沼, 46(1): 233-239.
	[Xu Y J, Shentu J K, Ding Z N. 2015. Grain size and thickness of sand bottom affect the growth of swimming crab Portunus trituberculatus in single-crab basket-culture system. Oceanologia et Limnologia Sinica, 46(1): 233-239]
[32]	杨洪, 易朝路, 邢阳平, 谢平. 2004. 武汉东湖沉积物孔隙度与含水量. 湖泊科学, 16(1): 68-72.
	[Yang H, Yi C L, Xing Y P, Xie P. 2004. Porosity and water content of sediments in Lake Donghu,Wuhan. Journal of Lake Sciences, 16(1): 68-72]
[33]	殷鉴, 刘春莲, 吴洁, 黄毅, 吴月琴. 2016. 珠江三角洲中部晚更新世以来的有孔虫记录与古环境演化. 古地理学报, 18(4): 677-690. doi: 10.7605/gdlxb.2016.04.050
	[Yin J, Liu C L, Wu J, Huang Y, Wu Y Q. 2016. Foraminiferal records and palaeoenvironmental changes since the Late Pleistocene in central Pearl River Delta. Journal of Palaeogeography(Chinese Edition), 18(4): 677-690]
[34]	袁菲, 何用, 许劼婧. 2022. 近期珠江三角洲地形演变特征及趋势. 泥沙研究, 47(1): 59-64.
	[Yuan F, He Y, Xu J J. 2022. Recent topographical evolution characteristics and trend of the Pearl River Delta. Journal of Sediment Research, 47(1): 59-64]
[35]	张光威. 1988. 珠江口现代沉积环境的沉积物特征. 海洋地质动态, 4(2): 4-6.
	[Zhang G W. 1988. Sediment characteristics of modern sedimentary environments in the Pearl River Estuary. Marine Geology Frontiers, 4(2): 4-6]
[36]	张光威, 马道修, 徐明广, 周青伟, 蓝先洪. 1988. 珠江口现代沉积物构造特征及其沉积环境. 海洋地质与第四纪地质, 8(3): 71-83.
	[Zhang G W, Ma D X, Xu M G, Zhou Q W, Lan X H. 1988. Sedimentary environments and structures of modern sediments in the Mouth of Zhujiang River. Marine Geology & Quaternary Geology, 8(3): 71-83]
[37]	张绍轩, 汤永杰, 郑翠美, 陈震, 郑卓. 2020. 珠江三角洲全新世海-陆沉积模式转换及其年代. 海洋地质与第四纪地质, 40(5): 107-117.
	[Zhang S X, Tang Y J, Zheng C M, Chen Z, Zheng Z. 2020. Holocene sedimentary environment transform and onset time of Pearl River Delta progradation. Marine Geology & Quaternary Geology, 40(5): 107-117]
[38]	张蔚, 严以新, 郑金海, 吴宏旭. 2010. 珠江三角洲年际潮差长期变化趋势. 水科学进展, 21(1): 77-83.
	[Zhang W, Yan Y X, Zheng J H, Wu H X. 2010. Interannual tidal range trend in Pearl River Delta. Advances in Water Science, 21(1): 77-83]
[39]	赵焕庭. 1982. 珠江三角洲的形成和发展. 海洋学报(中文版), 4(5): 595-607.
	[Zhao H T. 1982. The formation and development of the Pearl River Delta. Acta Oceanologica Sinica, 4(5): 595-607]
[40]	赵焕庭. 1983. 珠江三角洲的水文特征. 热带海洋, 2(2): 108-117.
	[Zhao H T. 1983. Hydrological characteristics of the Zhujiang(Pearl River)Delta. Tropic Oceanology, 2(2): 108-117]
[41]	赵焕庭. 1984. 珠江河口演变的基本过程. 热带海洋, 3(4): 1-10.
	[Zhao H T. 1984. The general evolution process of Zhujiang(Pearl River)Mouth. Tropic Oceanology, 3(4): 1-10]
[42]	赵焕庭. 1989. 珠江河口的水文和泥沙特征. 热带地理, 9(3): 201-212.
	[Zhao H T. 1989. Hydrological and sedimentary characteristics of the Pearl River estuary. Tropical Geography, 9(3): 201-212]
[43]	Doyle E, Orr P, Murray J. 2021. The earliest occurrence of the ichnogenus Psilonichnus: a new record from the Mississippian of the west of Ireland. Ichnos, 28(3): 208-216.
[44]	Frey R W, Curran H A, Pemberton S G. 1984. Tracemaking activities of crabs and their environmental significance: the ichnogenus Psilonichnus. Journal of Paleontology, 58(2): 333-350.
[45]	Fürsich F T. 1981. Invertebrate trace fossils from the Upper Jurassic of Portugal. Comunicações do Serviço Geológico de Portugal, 67(2): 153-168.
[46]	Gingras M K, Hubbard S M, Pemberton S G, Saunders T. 2000. The significance of Pleistocene psilonichnus at Willapa Bay. Washington. Palaios, 15: 142-151.
[47]	Humphreys B, Balson P S. 1988. Psilonichnus(Fürsich)in late Pliocene subtidal marine sands of eastern England. Journal of Paleontology, 62(2): 168-172.
[48]	Kristensen E. 2008. Mangrove crabs as ecosystem engineers: with emphasis on sediment processes. Journal of Sea Research, 59: 30-43.
[49]	Li G, Yan W, Zhong L F. 2016. Element geochemistry of offshore sediments in the northwestern South China Sea and the dispersal of Pearl River sediments. Progress in Oceanography, 141: 17-29.
[50]	Li Y, Wang W, Ma X Z, Zhang Y J. 2013. Effect of rice-crab culture system on zoobenthos diversity in paddy field. Chinese Journal of Eco-Agriculture, 21(7): 838-843.
[51]	Myint M. 2001. Psilonichnus quietis isp. nov. from the Eocene Iwaki formation,Shiramizu group,Joban coal field,Japan. Ichnos, 8(1): 1-14.
[52]	Nesbitt E A, Campbell K A. 2002. A new Psilonichnus ichnospecies attributed to mud-shrimp Upogebia in estuarine settings. Journal of Paleontology, 76(5): 892-901.
[53]	Neto de Carvalho C. 2016. Psilonichnus Fürsich,1981 in its type-locality(Praia do Salgado,western Portugal). Comunicações Geológicas, 103: 13-21.
[54]	Olszewska-Nejbert D, Kotowski J, Nejbert K. 2020. Psilonichnus upsilon Frey,Curran and Pemberton,1984 burrows and their environmental significance in transgressive Albian(Lower Cretaceous)sands of Glanów-Stroniczki,Cracow Upland,southern Poland. Palaeogeography,Palaeoclimatology,Palaeoecology, 538: 109388.
[55]	Pervesler P, Uchman A. 2009. A new Y-shaped trace fossil attributed to Upogebiid Crustaceans from Early Pleistocene of Italy. Acta Palaeontologica Polonica, 54(1): 135-142.
[56]	Pemberton S G, Jones B. 1988. Ichnology of the Pleistocene ironshore formation,Grand Cayman island,British west Indies. Journal of Paleontolog, 62(4): 495-505.
[57]	Rodríguez-Tovar F J, Seike K, Allen Curran H. 2014. Characteristics,distribution patterns,and implications for ichnology of modern burrows of Uca(Leptuca)speciosa,San Salvador Island,Bahamas. Journal of Crustacean Biology, 34(5): 565-572.
[58]	Tiwari R P, Rajkonwar C, Patel S J. 2013. Funalichnus bhubani isp. nov. from Bhuban Formation,Surma Group(Lower-Middle Miocene)of Aizawl,Mizoram,India. PLoS One, 8(10): e77839.
[59]	Wang Y Y, Wang X Q, Hu B, Luo M. 2019. Tomographic reconstructions of crab burrows from deltaic tidal flat: contribution to palaeoecology of decapod trace fossils in coastal settings. Palaeoworld, 28(4): 514-524.

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