云南拟步甲区系与地理分布特征（鞘翅目：拟步甲总科）
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摘要
本研究在多年野外考察和对河北大学馆藏云南拟步甲标本的鉴定和整理的基础上，结合有关文献资料，对云南省拟步甲的物种多样性及其区系组成、地理分布和区系起源等进行了讨论，取得如下初步成果：
一、种类多样性
经对云南15000余件拟步甲标本进行分类鉴定和资料整理，共记录6亚科29族101属438种（亚种），分别描述了各级分类阶元特征，编制了分亚科、族、属和种（亚种）的检索表，列出了属、种文献引证、检视标本信息及分布地，按最新分类体系编排了云南拟步甲系统分布名录，包括2新种（已发表）、1中国新纪录亚属、2中国新纪录种、云南新纪录6属5种、并提出1新名。
2新种为：徐氏壑轴甲Hexarhopalus (Leprocaulus) xui Ren et Xu,2011和具齿光轴甲Augolesthus（Augolesthus）dentatus Xu et Ren,2012。
1中国新纪录亚属为：光轴甲指名亚属Augolesthus (Augolesthus) Motschulsky,1872。
2中国新纪录种为：纹翅壑轴甲Hexarhopalus (Hexarhopalus) sculptilis Kaszab,1960和哑黑涅轴甲Necrobioides gibbicollis (Fleutiaux,1887)。
云南6新纪录属为：沟轴甲属Cleomis Fairmaire,1893、闽轴甲属Foochounus Pic,1921、涅轴甲属Necrobioides Fairmaire,1882、棒轴甲属Rhopalobates Fairmaire,1897、越轴甲属Tonkinius Fairmaire,1903及斐轴甲属Phaedis Pascoe,1866。
云南5新纪录种为：宽边沟轴甲Cleomis marginicollis (Gebien,1914)、多毛棒轴甲Rhopalobatesvillardi Fairmaire,1896、刻纹越轴甲指名亚种Tonkinius sculptilis sculptilis Fairmaire,1903、隆线异土甲Heterotarsus carinula Marseul,1876、瘤翅异土甲Heterotarsus pustulifer Fairmaire,1889。
为Leprocaulinus Kaszab,1982(nec Leprocaulinus Uvarov,1940:176)提出新名：Pseudoleprocaulusnom. nov.
二、区系成分及分布类型
1、云南拟步甲以东洋成分占主要地位（298种），占总数的68.0%，其次是东洋+古北成分（64种），占总数的14.6%，古北成分58种，占总数的13.2%，仓储害虫10种，占总数的2.2%，其它不同的跨界组合共8种，占总数的2.0%。
2、将29族的地理分布归纳为6种分布类型，101属的地理分布归纳为8种分布类型及7种变型，其中，典型的干旱区成分少，如漠甲亚科种类仅为4种，族、属均以热带成分占有绝对优势，表明云南拟步甲区系具明显的热带亲缘。
三、生物地理学
将云南拟步甲和邻近地区的区系相似性进行了比较研究，结果显示：
1、在世界动物地理区划中，除东洋界及古北界外，云南拟步甲与非洲界特别是热带非洲及北非具有较多的共布族、属，表明它们可能有较密切的亲缘关系。
2、在东洋界内，云南拟步甲可能与印度亚界有较近的渊源关系，共有种最多（60种）>印马亚界（32种）>斯里兰卡（10种）。
3、在中印亚界内，云南拟步甲可能与四川有较近的渊源关系，与其共有种最多：四川80种>越南70种>广西53种>缅甸52种>老挝41种>贵州36种；相似性系数（QS）亦最大：四川（0.23）>广西（0.2）>贵州（0.15）。
四、空间分布特点
1、栅格分析结果表明：云南的拟步甲由南向北，由西向东种类均呈减少的态势，但在不同纬度带上有较大波动，分布显著地受云南独特的地貌分异所影响。
2、古北界（尤其是青藏高原）拟步甲可沿高山向南延伸，部分类群可向南延伸至中南半岛；东洋界的种类，则可沿峡谷向北延伸，其分布也可达海拔3375m的高山地带，两界拟步甲呈垂直相混分布。
五、区域分布特点
1、云南拟步甲特有成分丰富，共有3特有属和171特有种，特有种数占总数的39.0%，但略低于横断山地区和青藏高原地区，这3个地区特有成分占总数的比例为：青藏高原58.9%﹥横断山区55.9%﹥云南39.0%。
2、采用栅格聚类方法确定了云南拟步甲的4个特有分布区，即滇南山地小区、南部横断山小区、中部横断山小区、滇东高原小区。
六、区系起源与适应特性
综合考量相关类群的生物学特性、形态与栖境间的相互关系，并结合地质历史，对云南拟步甲的区系起源、重要适应特性等问题进行讨论和推断。初步推测现代云南拟步甲区系是以第3纪古热带区系为基础演化而来，但深受青藏高原隆升及第四纪冰川的强烈影响，其区系性质发生了深刻改变，带有明显的过渡性。
This dissertation deals with taxonomic and faunistic studies on the Tenebrionidae (Coleoptera) fromYunnan Province, also refers to the origin, diversity, and distribution pattern. The main results of this workare listed as follows:
I. Based on the data from the original manuscript and the results of identifying tenebrionid specimensdeposited in MHBU, in total,6subfamilies,29tribes,101genera and438species (subspecies) arerecorded from Yunnan. The diagnoses of each subfamily, tribe and genus, as well as the keys tosubfamilies, tribes, genera and species are provided. The citations and information of the examinedmaterials are provided for each genus and species.
During the study, two species, H.(Leprocaulus) xui Ren et Xu,2011, Augolesthus（Augolesthus）dentatus Xu et Ren,2012have been described from Yunnan; subgenus Augolesthus (A.) Motschulsky,1872, Hexarhopalus (H.) sculptilis Kaszab,1960and Necrobioides gibbicollis (Fleutiaux,1887) wererecorded from China for the first time.6genera（Cleomis Fairmaire,1893、Foochounus Pic,1921、Necrobioides Fairmaire,1882、Rhopalobates Fairmaire,1897、Tonkinius Fairmaire,1903、Phaedis Pascoe,1866）and5species（Cleomis marginicollis (Gebien,1914)、Rhopalobates villardi Fairmaire,1896、Tonkinius sculptilis sculptilis Fairmaire,1903、Heterotarsus carinula Marseul,1876、Heterotarsuspustulifer Fairmaire,1889）are newly recorded from Yunnan.
Pseudoleprocaulus nom. nov., new replacement name for the genus Leprocaulinus Kaszab,1982[nec Leprocaulinus Uvarov,1940(Phasmatodea)], is proposed.
II. Within this region,438tenebrionid species are recorded, of which298species are Orientalcomponents (68.0%of the total species),58species are the Palaearctic ones (13.2%). These speciesbelongs to Alleculinae (2tribes,10genera), Diaperinae (1tribe,14genera), Lagriinae (6tribes,21genera),Pimeliinae (3tribes,4genera), Stenochiinae (2tribes,23genera) and Tenebrioninae (11tribes,29genera).
A comprehensive analysis was made of the distribution of29already recorded tribes and101generaof Yunnan. As a result, distribution patterns of29tribes can be classified into6types, and these of101genera can be classified into8types and7subtypes. Most of the Yunnan tenebrionid tribes and genera areof tropical elements, typical arid elements are rare and only4species of Pimeliinae are in this region.
III. The tenebrionid fauna of Yunnan is the closest to that of Ethiopian realm in world zoo-geographical realm and closest to that of Indian subrealm in Oriental realm. In Indo-Chinese subrealm,it is the closest to Shichuan Province, descending of quotient of similarity (QS): Yunnan-Sichuan (0.23),Yunnan-Guangxi (0.20), Yunnan-Guizhou (0.15). And it has80species in common with Sichuan Province,70with Vietnam,53with Guangxi Zhuang Autonomous Region,52with Myanmar,41with Laos, and36with Guizhou Province.
IV. By using species statistics and GIS mapping, it shows that: as a whole, from south to north, andalso west to east, the species richness decreases with the increase of latitude and longitude, but it alsoshows greater volatility in different latitudes. The distribution is notably restricted by natural geographicallocation and special landscape.
Typical Palaearctic elements distribute in the northern parts of this region, but some groups mayspread southward to Indo-China Peninsula, but usually confined to the high altitude mountains; sometypical Oriental elements may advance to the north through deep gorges, even to3375m altitude zone; ThePalaearctic and Oriental elements overlapped with each other. It is remarkable that there is a gradualtransition and strong introgression of the Oriental and Palaearctic elements in a moutain, which is of greatinterest to the studies on world biogeography.
V. Within this region,438tenebrionid species are recorded, of which171species are endimic ones(possessing39%of the total species). Grid-Cluster analysis is applied to investigate patterns of endemismfor tenebrionid beetles. As a result, there are4areas of endemism: i.e. Southern Subregion of theHengduan Mountains, Middle Subregion of the Hengduan Mountains, Eastern Plateau Subregion ofYunnan, Southern Mountainous Subregion of Yunnan.
VI. Referring to the biogeographical history, the origin and evolution of the tenebrionid beetles of theYunnan is analyzed and discussed. The preliminary results show that the dakling beetle fauna of Yunnanare the descendant of the tropical Tertiary tenebrionid fauna, but it has undergone profound change, sincestrongly influenced by the uplift of the Qinghai-Tibet Plateau and the Quaternary glacial, and its faunisticcomponents obviously with transitional characteristics. At last, the adaptations to arboreal life of darklingbeetles are discussed.
一、种类多样性
经对云南15000余件拟步甲标本进行分类鉴定和资料整理，共记录6亚科29族101属438种（亚种），分别描述了各级分类阶元特征，编制了分亚科、族、属和种（亚种）的检索表，列出了属、种文献引证、检视标本信息及分布地，按最新分类体系编排了云南拟步甲系统分布名录，包括2新种（已发表）、1中国新纪录亚属、2中国新纪录种、云南新纪录6属5种、并提出1新名。
2新种为：徐氏壑轴甲Hexarhopalus (Leprocaulus) xui Ren et Xu,2011和具齿光轴甲Augolesthus（Augolesthus）dentatus Xu et Ren,2012。
1中国新纪录亚属为：光轴甲指名亚属Augolesthus (Augolesthus) Motschulsky,1872。
2中国新纪录种为：纹翅壑轴甲Hexarhopalus (Hexarhopalus) sculptilis Kaszab,1960和哑黑涅轴甲Necrobioides gibbicollis (Fleutiaux,1887)。
云南6新纪录属为：沟轴甲属Cleomis Fairmaire,1893、闽轴甲属Foochounus Pic,1921、涅轴甲属Necrobioides Fairmaire,1882、棒轴甲属Rhopalobates Fairmaire,1897、越轴甲属Tonkinius Fairmaire,1903及斐轴甲属Phaedis Pascoe,1866。
云南5新纪录种为：宽边沟轴甲Cleomis marginicollis (Gebien,1914)、多毛棒轴甲Rhopalobatesvillardi Fairmaire,1896、刻纹越轴甲指名亚种Tonkinius sculptilis sculptilis Fairmaire,1903、隆线异土甲Heterotarsus carinula Marseul,1876、瘤翅异土甲Heterotarsus pustulifer Fairmaire,1889。
为Leprocaulinus Kaszab,1982(nec Leprocaulinus Uvarov,1940:176)提出新名：Pseudoleprocaulusnom. nov.
二、区系成分及分布类型
1、云南拟步甲以东洋成分占主要地位（298种），占总数的68.0%，其次是东洋+古北成分（64种），占总数的14.6%，古北成分58种，占总数的13.2%，仓储害虫10种，占总数的2.2%，其它不同的跨界组合共8种，占总数的2.0%。
2、将29族的地理分布归纳为6种分布类型，101属的地理分布归纳为8种分布类型及7种变型，其中，典型的干旱区成分少，如漠甲亚科种类仅为4种，族、属均以热带成分占有绝对优势，表明云南拟步甲区系具明显的热带亲缘。
三、生物地理学
将云南拟步甲和邻近地区的区系相似性进行了比较研究，结果显示：
1、在世界动物地理区划中，除东洋界及古北界外，云南拟步甲与非洲界特别是热带非洲及北非具有较多的共布族、属，表明它们可能有较密切的亲缘关系。
2、在东洋界内，云南拟步甲可能与印度亚界有较近的渊源关系，共有种最多（60种）>印马亚界（32种）>斯里兰卡（10种）。
3、在中印亚界内，云南拟步甲可能与四川有较近的渊源关系，与其共有种最多：四川80种>越南70种>广西53种>缅甸52种>老挝41种>贵州36种；相似性系数（QS）亦最大：四川（0.23）>广西（0.2）>贵州（0.15）。
四、空间分布特点
1、栅格分析结果表明：云南的拟步甲由南向北，由西向东种类均呈减少的态势，但在不同纬度带上有较大波动，分布显著地受云南独特的地貌分异所影响。
2、古北界（尤其是青藏高原）拟步甲可沿高山向南延伸，部分类群可向南延伸至中南半岛；东洋界的种类，则可沿峡谷向北延伸，其分布也可达海拔3375m的高山地带，两界拟步甲呈垂直相混分布。
五、区域分布特点
1、云南拟步甲特有成分丰富，共有3特有属和171特有种，特有种数占总数的39.0%，但略低于横断山地区和青藏高原地区，这3个地区特有成分占总数的比例为：青藏高原58.9%﹥横断山区55.9%﹥云南39.0%。
2、采用栅格聚类方法确定了云南拟步甲的4个特有分布区，即滇南山地小区、南部横断山小区、中部横断山小区、滇东高原小区。
六、区系起源与适应特性
综合考量相关类群的生物学特性、形态与栖境间的相互关系，并结合地质历史，对云南拟步甲的区系起源、重要适应特性等问题进行讨论和推断。初步推测现代云南拟步甲区系是以第3纪古热带区系为基础演化而来，但深受青藏高原隆升及第四纪冰川的强烈影响，其区系性质发生了深刻改变，带有明显的过渡性。
This dissertation deals with taxonomic and faunistic studies on the Tenebrionidae (Coleoptera) fromYunnan Province, also refers to the origin, diversity, and distribution pattern. The main results of this workare listed as follows:
I. Based on the data from the original manuscript and the results of identifying tenebrionid specimensdeposited in MHBU, in total,6subfamilies,29tribes,101genera and438species (subspecies) arerecorded from Yunnan. The diagnoses of each subfamily, tribe and genus, as well as the keys tosubfamilies, tribes, genera and species are provided. The citations and information of the examinedmaterials are provided for each genus and species.
During the study, two species, H.(Leprocaulus) xui Ren et Xu,2011, Augolesthus（Augolesthus）dentatus Xu et Ren,2012have been described from Yunnan; subgenus Augolesthus (A.) Motschulsky,1872, Hexarhopalus (H.) sculptilis Kaszab,1960and Necrobioides gibbicollis (Fleutiaux,1887) wererecorded from China for the first time.6genera（Cleomis Fairmaire,1893、Foochounus Pic,1921、Necrobioides Fairmaire,1882、Rhopalobates Fairmaire,1897、Tonkinius Fairmaire,1903、Phaedis Pascoe,1866）and5species（Cleomis marginicollis (Gebien,1914)、Rhopalobates villardi Fairmaire,1896、Tonkinius sculptilis sculptilis Fairmaire,1903、Heterotarsus carinula Marseul,1876、Heterotarsuspustulifer Fairmaire,1889）are newly recorded from Yunnan.
Pseudoleprocaulus nom. nov., new replacement name for the genus Leprocaulinus Kaszab,1982[nec Leprocaulinus Uvarov,1940(Phasmatodea)], is proposed.
II. Within this region,438tenebrionid species are recorded, of which298species are Orientalcomponents (68.0%of the total species),58species are the Palaearctic ones (13.2%). These speciesbelongs to Alleculinae (2tribes,10genera), Diaperinae (1tribe,14genera), Lagriinae (6tribes,21genera),Pimeliinae (3tribes,4genera), Stenochiinae (2tribes,23genera) and Tenebrioninae (11tribes,29genera).
A comprehensive analysis was made of the distribution of29already recorded tribes and101generaof Yunnan. As a result, distribution patterns of29tribes can be classified into6types, and these of101genera can be classified into8types and7subtypes. Most of the Yunnan tenebrionid tribes and genera areof tropical elements, typical arid elements are rare and only4species of Pimeliinae are in this region.
III. The tenebrionid fauna of Yunnan is the closest to that of Ethiopian realm in world zoo-geographical realm and closest to that of Indian subrealm in Oriental realm. In Indo-Chinese subrealm,it is the closest to Shichuan Province, descending of quotient of similarity (QS): Yunnan-Sichuan (0.23),Yunnan-Guangxi (0.20), Yunnan-Guizhou (0.15). And it has80species in common with Sichuan Province,70with Vietnam,53with Guangxi Zhuang Autonomous Region,52with Myanmar,41with Laos, and36with Guizhou Province.
IV. By using species statistics and GIS mapping, it shows that: as a whole, from south to north, andalso west to east, the species richness decreases with the increase of latitude and longitude, but it alsoshows greater volatility in different latitudes. The distribution is notably restricted by natural geographicallocation and special landscape.
Typical Palaearctic elements distribute in the northern parts of this region, but some groups mayspread southward to Indo-China Peninsula, but usually confined to the high altitude mountains; sometypical Oriental elements may advance to the north through deep gorges, even to3375m altitude zone; ThePalaearctic and Oriental elements overlapped with each other. It is remarkable that there is a gradualtransition and strong introgression of the Oriental and Palaearctic elements in a moutain, which is of greatinterest to the studies on world biogeography.
V. Within this region,438tenebrionid species are recorded, of which171species are endimic ones(possessing39%of the total species). Grid-Cluster analysis is applied to investigate patterns of endemismfor tenebrionid beetles. As a result, there are4areas of endemism: i.e. Southern Subregion of theHengduan Mountains, Middle Subregion of the Hengduan Mountains, Eastern Plateau Subregion ofYunnan, Southern Mountainous Subregion of Yunnan.
VI. Referring to the biogeographical history, the origin and evolution of the tenebrionid beetles of theYunnan is analyzed and discussed. The preliminary results show that the dakling beetle fauna of Yunnanare the descendant of the tropical Tertiary tenebrionid fauna, but it has undergone profound change, sincestrongly influenced by the uplift of the Qinghai-Tibet Plateau and the Quaternary glacial, and its faunisticcomponents obviously with transitional characteristics. At last, the adaptations to arboreal life of darklingbeetles are discussed.
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