Ants of the Hengduan Mountains: a new altitudinal survey and updated checklist for Yunnan Province highlight an understudied insect biodiversity hotspot.

China's Hengduan Mountain region has been considered one of the most diverse regions in the northern hemisphere. Its stunning topography with many deep valleys and impassable mountain barriers has promoted an astonishing diversification in many groups of organisms including plants, birds, mammals, and amphibians. However, the insect biodiversity in this region is still poorly known. Here, the first checklist of ant species from the Southern Hengduan Mountain region is presented, generated by sampling ant diversity using a wide array of collection methods, including Winkler leaf litter extraction, vegetation beating, and hand collection. 130 species/morphospecies from nine subfamilies and 49 genera were identified. Among them, 17 species from 13 genera represent new records for Yunnan province, and eight species are newly recorded for China. Moreover, we believe 41 novel morphospecies (31% of the total collected taxa) will prove to be new to science. These results highlight the rich ant fauna of this region and strongly support its status as a biodiversity hotspot. The current ant species checklist for the whole of Yunnan Province was updated by recording 550 named species from 99 genera. Taken together, our results suggest that the Yunnan ant fauna still remains under-sampled, and future sampling will likely yield many more species, among them many undescribed ones. Cong Liu, Georg Fischer, Francisco Hita Garcia, Seiki Yamane, Qing Liu, Yan Qiong Peng, Evan P. Economo, Benoit Guénard, Naomi E. Pierce.

Introduction

The Hengduan Mountain region, located in the southeastern part of the Qinghai-Tibet Plateau, is one of the 35 recognized biodiversity hotspots in the world (Myers et al. 2000). The unique landscape, geomorphology, microhabitat differentiation and geographic isolation created by tectonic uplift during the last eight million years has promoted an astonishing diversification in many groups of organisms, making this region one of the most diverse temperate regions in the northern hemisphere (Boufford 2014; Price et al 2014; Xing and Ree 2017). For example, it harbors nearly 40 percent of China’s vascular plant diversity (ca. 12,000 species), including more than 3,000 endemic species (Boufford 2014). However, aside from the well-documented plants and some vertebrates, the diversity of other groups, especially invertebrates in this region remains largely unknown. Insect taxonomic groups in particular have received limited attention, and our understanding of their diversity in the Hengduan Mountains is extremely fragmented.

Ants are an ecologically dominant component of many ecosystems in terms of their abundance, richness, and ecosystem function (Hölldobler and Wilson 1990). Globally, about 15,600 ant species and subspecies have been described (Bolton 2020), making them the most diverse group of social insects and one of the most diverse families of insects. Despite the fact that ant diversity is mainly concentrated within tropical regions (Dunn et al. 2009; Guénard et al. 2012; Economo et al. 2018), the ant fauna of many other regions is still poorly known, especially in Asia (Guénard et al. 2010). Compiling and curating comprehensive and accurate ant species checklists for these regions is essential not only for insights into ant taxonomy and systematics, but also for long-term monitoring and conservation of these ecosystems (Guénard et al. 2017). The goal of this study is to provide a better understanding of the poorly known ant biodiversity in China’s Hengduan Mountains. The ultra-variable topography of this region, ideal for creating numerous vicariance events, combined with its wide range of climatic zones has contributed to the exceptional richness of endemic species inhabiting this area. Nevertheless, the rough topography has also made access and exploration rather challenging in the past. Against the background of extraordinary levels of plant diversity harbored by the Hengduan Mountains, it remains unclear whether or not ants and other insects display similar patterns of high diversity and endemism in this region.

To address this gap, we here present the results of an ant biodiversity survey conducted in the Gaoligong Shan mountains (part of the Hengduan Mountains), Yunnan Province, southwest China undertaken in 2019. Our goal is to present a complete species checklist of ants from the Gaoligong Mountains, including new records, as well as to update the current ant species checklist for the whole of Yunnan Province.

The Gaoligong Shan mountains (lat. 24°560'–28°220'N, long. 98°080'–98°500'E) comprise the western-most part of the Hengduan Mountain Range, and are among the most biodiversity-rich areas in Yunnan (Li et al. 2008; Dumbecher et al. 2011; Lo and Bi 2019). The ant fauna in the Gaoligong Shan mountains remains poorly understood, despite several studies focusing on ant diversity patterns that have recorded 62 ant species from 31 genera (Xu 2001a, b), but lack a comprehensive list of species collected.

Yunnan province is the richest province of China in terms of ant diversity (Guénard and Dunn 2012). The latest ant checklist of Yunnan was compiled almost 10 years ago and consisted of 462 ant species. Since then, new ant inventories have been conducted (e.g. Liu et al. 2015a), as well as new species descriptions (e.g., Guénard et al. 2013; Xu et al. 2014a, b; Liu et al. 2015b; Staab et al. 2018), and the identification of previously dubious records have sensibly modified our understanding of Yunnan’s ant diversity and species composition. Therefore, in this study, we also provide an update to the ant species checklist of Yunnan province and discuss future trends.

Materials and methods

Ant specimens were collected from natural forests along an elevational gradient on both the eastern and western slopes of the Gaoligong Mountains in July 2019. We sampled leaf litter ants from 16 sites at roughly 150 m elevational intervals from 600 m to 3000 m, following the standardized sampling protocol developed in Liu et al. 2016. At each site, we established a 400 m2 quadrat (20 m × 20 m) and collected leaf litter samples at the four corners of the quadrat (1 m2). We also collected leaf litter within the quadrat to cover a variety of microhabitats. Finally, ants on the ground, lower vegetation, and tree branches were collected both by hand and using a beating sheet. Leaf litter samples were extracted using mini Winkler extractors for 72 hours using the shuffling method described in Guénard and Lucky (2011).

Ant specimens were first placed in 99% ethanol and later sorted into morphospecies and point mounted. Each mounted specimen was assigned a unique Museum of Comparative Zoology, Harvard University () specimen code and collection labels. Extended depth of field specimen images were taken with a Leica DFC400 digital camera mounted on a Leica M205C stereomicroscope through the Leica Application Suite V4 software in the Ant Room at the MCZ. Specimens were identified to species / morphospecies using available keys, the digital resources on Antwiki (http://www.antwiki.org) and AntWeb (http://www.antweb.org), as well as reference museum material. All mounted and alcohol-preserved ant specimens are currently deposited in the Ant Room of the MCZ.

Distribution maps of species were generated from records included within the Global Ant Biodiversity Informatics () database and available at https://antmaps.org (Janicki et al, 2016; Guénard et al. 2017). These maps are based on records reported at the country level, or at the first administrative division for the larger countries (China, India, Japan). For larger islands that form their own natural biogeographic units like Borneo, Sumatra, New Guinea, the distribution maps used the island boundary instead of political boundaries (see also Guénard et al. 2012).

Results

Ants of the Hengduan Mountain region

More than 3000 specimens were collected during this survey, and 130 species and morphospecies in 49 genera and nine subfamilies were identified. After identification of 88 valid species from the 130 total collected species, a total of 17 new species records are presented for Yunnan province and eight represent new records for China (see Table 1). The newly recorded species belong to 13 genera from four subfamilies. Moreover, the 41 morphospecies that could not be identified are likely to represent new species.

Table 1.

List of ant species (Formicidae) in the Gaoligong Shan mountains, Yunnan with their respective illustrations. * New to Yunnan province; **New to China.

Species	Figure
Dorylinae
Aenictus artipus Wilson, 1964	Fig. 1
** Aenictus brevinodus Jaitrong & Yamane, 2011	Fig. 2
Aenictus hodgsoni Forel, 1901	Fig. 3
Aenictus paradentatus Jaitrong, Yamane & Tasen, 2012	Fig. 4
* Aenictus watanasiti Jaitrong & Yamane, 2013	Fig. 5
Cerapachys sulcinodis Emery, 1889	Fig. 6
Cerapachys sp. clm01	Fig. 7
Chrysapace costatus (Bharti & Wachkoo, 2013)	Fig. 8
Dorylus orientalis Westwood, 1835	Figs 9, 10
Ooceraea biroi (Forel, 1907)	Fig. 11
Amblyoponinae
Stigmatomma octodentatum (Xu, 2006)	Fig. 12
Dolichoderinae
Dolichoderus feae Emery, 1889	Fig. 13
Dolichoderus squamanodus Xu, 2001	Fig. 14
Dolichoderus taprobanae (Smith, 1858)	Fig. 15
Ochetellus glaber (Mayr, 1862)	Fig. 16
Tapinoma melanocephalum (Fabricius, 1793)	Fig. 17
Ectatomminae
Gnamptogenys quadrutinodules Chen, Lattke & Zhou, 2017	Fig. 18
Formicinae
Anoplolepis gracilipes (Smith, 1857)	Fig. 19
** Camponotus bellus leucodiscus Wheeler, 1919	Fig. 20
** Camponotus keihitoi Forel, 1913	Fig. 21
Camponotus lasiselene Wang & Wu, 1994	Figs 22, 23
Camponotus mitis (Smith, 1858)	Fig. 24
Camponotus nicobarensis Mayr, 1865	Fig. 25
Camponotus sp. clm01	Fig. 26
Camponotus sp. clm02	Fig. 27
Camponotus sp. clm03	Fig. 28
Camponotus sp. clm04	Fig. 29
Camponotus sp. clm05	Fig. 30
Formica cunicularia Latreille, 1798	Fig. 31
Formica japonica Motschoulsky, 1866	Fig. 32
* Lasius obscuratus Stitz, 1930	Fig. 33
* Lasius himalayanus Bingham, 1903	Fig. 34
Nylanderia bourbonica (Forel, 1886)	Fig. 35
Nylanderia sp. clm01	Fig. 36
Nylanderia sp. clm02	Fig. 37
Oecophylla smaragdina (Fabricius, 1775)	Fig. 38
Paraparatrechina sakurae (Ito, 1914)	Fig. 39
Paraparatrechina sp. clm01	Fig. 40
Paraparatrechina sp. clm02	Fig. 41
Polyrhachis armata (Le Guillou, 1842)	Fig. 42
Polyrhachis bihamata (Drury, 1773)	Fig. 43
Polyrhachis dives Smith, 1857	Fig. 44
Polyrhachis furcata Smith, 1858	Fig. 45
Polyrhachis halidayi Emery, 1889	Fig. 46
Polyrhachis illaudata Walker, 1859	Fig. 47
Polyrhachis laevigata Smith, 1857	Fig. 48
Polyrhachis tibialis Smith, 1858	Fig. 49
* Prenolepis angularis Zhou, 2001	Fig. 50
* Prenolepis fustinoda Williams & LaPolla, 2016	Fig. 51
Prenolepis sp. clm01	Fig. 52
Prenolepis sp. clm02	Fig. 53
Pseudolasius emeryi Forel, 1915	Fig. 54
Pseudolasius silvestrii Wheeler, 1927	Fig. 55
Myrmicinae
Aphaenogaster feae Emery, 1889	Fig. 56
Aphaenogaster sp. clm01	Fig. 57
Aphaenogaster sp. clm02	Fig. 58
Aphaenogaster sp. clm03	Fig. 59
Aphaenogaster sp. clm04	Fig. 60
Aphaenogaster sp. clm05	Fig. 61
* Cardiocondyla itsukii Seifert, Okita & Heinze, 2017	Fig. 62
Cardiocondyla sp. clm01	Fig. 63
Carebara acutispina (Xu, 2003)	Fig. 64
Carebara affinis (Jerdon, 1851)	Fig. 65
Carebara altinoda (Xu, 2003)	Fig. 66
Carebara bihornata (Xu, 2003)	Fig. 67
Carebara sp. clm01	Fig. 68
* Cataulacus marginatus Bolton, 1974	Fig. 69
Crematogaster quadriruga Forel, 1911	Fig. 70
Crematogaster sp. clm01	Fig. 71
Crematogaster sp. clm02	Fig. 72
** Dilobocondyla eguchii Bharti & Kumar, 2013	Fig. 73
Gaoligongidris planodorsa Xu, 2012	Fig. 74
Gauromyrmex sp. clm01	Fig. 75
Lordomyrma sp. clm01	Fig. 76
Monomorium pharaonis (Linnaeus, 1758)	Fig. 77
Monomorium sp. clm01	Fig. 78
Myrmica draco Radchenko, Zhou & Elmes, 2001	Fig. 79
Myrmica pleiorhytida Radchenko & Elmes, 2009	Fig. 80
Myrmica sp. clm01	Fig. 81
Myrmecina sp. clm01	Fig. 82
Myrmecina sp. clm02	Fig. 83
Myrmecina sp. clm03	Fig. 84
Pheidole allani Bingham, 1903	Figs 85, 86
Pheidole fervens Smith, 1858	Fig. 87
Pheidole fervida Smith, 1874	Fig. 88, 89
Pheidole gatesi (Wheeler, 1927)	Fig. 90
Pheidole indica Mayr, 1879	Fig. 91
Pheidole magna Eguchi, 2006	Figs 92, 93
* Pheidole nodifera (Smith 1858)	Fig. 94
Pheidole zoceana Santschi, 1925	Figs 95, 96
Pristomyrmex brevispinosus Emery, 1887	Fig. 97
Pristomyrmex hamatus Xu & Zhang, 2002	Fig. 98
Stenamma wumengense Liu & Xu, 2011	Fig. 99
Strumigenys assamensis De Andrade, 1994	Fig. 100
Strumigenys strygax Bolton, 2000	Fig. 101
** Strumigenys taphra (Bolton, 2000)	Fig. 102
Strumigenys sp. clm01	Fig. 103
Strumigenys sp. clm02	Fig. 104
Strumigenys sp. clm03	Fig. 105
* Temnothorax striatus Zhou, Huang, Yu & Liu, 2010	Fig. 106
Temnothorax sp. clm01	Fig. 107
Temnothorax sp. clm03	Fig. 108
Tetramorium tonganum Mayr, 1870	Fig. 109
Tetramorium sp. clm01	Fig. 110
Tetramorium sp. clm02	Fig. 111
Tetramorium sp. clm03	Fig. 112
Tetramorium sp. clm04	Fig. 113
Vollenhovia pyrrhoria Wu & Xiao, 1989	Fig. 114
Vollenhovia sp. clm03	Fig. 115
Ponerinae
Brachyponera luteipes (Mayr, 1862)	Fig. 116
Ectomomyrmex lobocarenus (Xu, 1995)	Fig. 117
** Ectomomyrmex obtusus Emery, 1900	Fig. 118
Hypoponera sp. clm01	Fig. 119
Hypoponera sp. clm02	Fig. 120
Hypoponera sp. clm03	Fig. 121
Leptogenys birmana Forel, 1900	Fig. 122
Leptogenys kitteli (Mayr, 1870)	Fig. 123
Odontomachus circulus Wang, 1993	Fig. 124
* Odontomachus fulgidus Wang, 1993	Fig. 125
Platythyrea parallela (Smith, 1859)	Fig. 126
Ponera bawana Xu, 2001	Fig. 127
Ponera xantha Xu, 2001	Fig. 128
Proceratinae
Discothyrea banna Xu, Burwell & Nakamura, 2014	Fig. 129
Discothyrea diana Xu, Burwell & Nakamura, 2014	Fig. 130
Proceratium longigaster Karavaiev, 1935	Fig. 131
Proceratium longmenense Xu, 2006	Fig. 132
Proceratium zhaoi Xu, 2000	Fig. 133
Pseudomyrmecinae
Tetraponera allaborans (Walker, 1859)	Fig. 134
Tetraponera attenuata Smith, 1877	Fig. 135
Tetraponera protensa Xu & Chai, 2004	Fig. 136

Within the recent collection, the most speciose ant genus is with eleven species (8.5% of the total species collected in the survey), followed by (ten species, 7.7%), and (seven species, 5.4%). Other diverse genera include (6 species, 4.6%), (six species, 4.6%), (six species, 4.6%), (five species, 3.8%), and (five species, 3.8%). More details are presented in Table 2.

Table 2.

Number of ant species of per genus collected in this survey as well the total number of each species per genus in Yunnan province.

Genus	Gaoligongshan Mt.	Yunnan	Genus	Gaoligongshan Mt.	Yunnan
Camponotus	10	30	Leptogenys	2	17
Pheidole	8	42	Monomorium	2	6
Polyrhachis	8	32	Odontomachus	2	6
Aphaenogaster	6	10	Ponera	2	14
Strumigenys	6	24	Pristomyrmex	2	4
Tetramorium	5	29	Pseudolasius	2	6
Aenictus	5	19	Vollenhovia	2	3
Carebara	5	19	Anoplolepis	1	1
Prenolepis	4	7	Brachyponera	1	3
Crematogaster	3	25	Cataulacus	1	4
Dolichoderus	3	9	Chrysapace	1	1
Hypoponera	3	7	Dilobocondyla	1	3
Lasius	2	6	Dorylus	1	3
Myrmica	3	12	Gaoligongidris	1	1
Myrmecina	3	7	Gauromyrmex	1	1
Nylanderia	3	10	Gnamptogenys	1	7
Paraparatrechina	3	2	Lordomyrma	1	1
Proceratium	3	4	Ochetellus	1	1
Temnothorax	3	7	Oecophylla	1	1
Tetraponera	3	12	Ooceraea	1	1
Cardiocondyla	2	4	Platythyrea	1	2
Cerapachys	2	1	Stenamma	1	4
Discothyrea	2	3	Stigmatoma	1	11
Ectomomyrmex	2	8	Tapinoma	1	4
Formica	2	7	Total	130	550

Here, we present the list of ant species that were collected in the Gaoligong Shan mountains (Table 1), as well as images for each species (Figs 1–136).

Figure 1.

worker (MCZ-ENT00763651) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

Figure 136.

worker (MCZ-ENT00763526) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

Updated ant checklist in Yunnan

The ant species list of Yunnan Province was generated using records from GABI available at https://antmaps.org (Janicki et al. 2016; Guénard et al. 2017). In total, the Yunnan ant fauna is composed of 99 genera and 550 named species and subspecies. Among them, the ant genera , , and are only known from unidentified morphospecies. Through our collection and the records from GABI, we have added 125 species and subspecies to the list of ants of Yunnan since the last ant checklist (Guénard et al. 2012). We also excluded 26 species records from the previous list and explained our rationale in each case (Table 3).

Table 3.

Ant species records that have been excluded from Yunnan when compared to the previous list. The explanation “Needs verification” usually signifies that the species has never been recorded before in this region and/or is easily mistaken for another species and likely to have been misidentified. “Dubious” means that the record occurrence is highly unlikely given the known species distribution. Notes provide additional references regarding records and/or further information.

Excluded species records	Explanations	Notes
Camponotus aethiops	Needs verification	A Palearctic species with distribution in Asia needs confirmation
Camponotus spenceri	Dubious	An Australian species misreported previously
Cardiocondyla nuda	Dubious	Could be C. kagutsuchi, see Seifert 2003
Discothyrea clavicornis	Dubious	A misidentification of D. diana
Discothyrea kamiteta	Dubious	A misidentification of D. banna
Formica fusca	Needs verification	A Palearctic species with distribution in Asia needs confirmation
Hypoponera exoecata	Needs verification	Species with distribution limited to East Asia
Lasius alienus	Dubious	See Seifert 2020
Lasius emarginatus	Dubious	A West Palearctic species with distribution in Asia doubtful
Lasius fuliginosus	Dubious	See Espadaler et al. 2001
Lasius niger	Dubious	See Seifert 1992
Lasius productus	Needs verification	Species with distribution limited to Japan and the Korean Peninsula
Lasius spathepus	Needs verification	Species with distribution limited to Japan, the Korean Peninsula and Eastern Russia
Leptogenys yerburyi	Dubious	See Xu and He 2015
Myrmica inezae	Needs verification	See Chen et al. 2016.
Odontoponera transversa	Dubious	See Yamane 2009
Proceratium deelemani	Dubious	Record represented a new species subsequently described in Staab et al. 2018.
Proceratium japonicum	Dubious	A misidentification of P. longigaster
Temnothorax melleus	Needs verification	A central Asian species which presence in Yunnan requires confirmation
Tetramorium inglebyi	Dubious	An Indian species that is restricted to the Southwest.
Tetramorium globulinode	Dubious	An Afrotropical species incorrectly reported in Asia
Tetramorium khnum	Dubious	An endemic species in the Philippines
Tetramorium melleum	Dubious	A misidentification of T. wroughtonii
Tetraponera aitkenii	Dubious	Phil Ward (Personal communication, 18 August 2015)
Tetraponera nigra	Dubious	Phil Ward (Personal communication, 18 August 2015)
Vollenhovia emeryi	Dubious	See Wetterer et al. 2015

In Yunnan, the most diverse ant genus is with 42 named species, followed by (33 species), (30 species), and (29 species). Other diverse genera include (25 species), and (25 species). Although 15 ant genera contain more than ten named species in Yunnan, the majority of ant genera occurring in Yunnan seem to be not particularly diverse. For example, 35 genera are represented by only one species in Yunnan (Table 4).

Table 4.

Number of ant species (both native and exotic species) in Yunnan Province. * Ant genus only known from morphospecies records.

Genus	Native	Exotic	Genus	Native	Exotic
Pheidole	42	0	Solenopsis	2	1
Polyrhachis	32	0	Acanthomyrmex	2	0
Camponotus	30	0	Acropyga	2	0
Tetramorium	28	1	Echinopla	2	0
Crematogaster	25	0	Meranoplus	2	0
Strumigenys	24	1	Myrmoteras	2	0
Aenictus	19	0	Paraparatrechina	2	0
Carebara	19	0	Perissomyrmex	2	0
Leptogenys	17	0	Platythyrea	2	0
Ponera	14	0	Pseudoneoponera	2	0
Tetraponera	12	0	Rhopalomastix	2	0
Myrmica	12	0	Trichomyrmex	0	2
Stigmatomma	11	0	Vollenhovia	2	0
Technomyrmex	11	0	Anoplolepis	1	0
Aphaenogaster	10	0	Buniapone	1	0
Nylanderia	9	1	Centromyrmex	1	0
Dolichoderus	9	0	Cerapachys	1	0
Ectomomyrmex	8	0	Chrysapace	1	0
Lepisiota	8	0	Diacamma	1	0
Colobopsis	7	0	Emeryopone	1	0
Hypoponera	5	2	Erromyrma	1	0
Prenolepis	7	0	Euponera	1	0
Temnothorax	7	0	Gaoligongidris	1	0
Formica	7	0	Gauromyrmex	1	0
Gnamptogenys	7	0	Gesomyrmex	1	0
Myrmecina	7	0	Harpegnathos	1	0
Anochetus	6	0	Iridomyrmex	1	0
Lasius	6	0	Lasiomyrma*	1	0
Odontomachus	6	0	Liometopum	1	0
Pseudolasius	6	0	Lioponera	1	0
Cryptopone	5	0	Lordomyrma*	1	0
Monomorium	5	0	Mesoponera	1	0
Proceratium	4	0	Messor	1	0
Cataulacus	4	0	Myrmicaria	1	0
Plagiolepis	3	1	Mystrium	1	0
Pristomyrmex	4	0	Ochetellus	1	0
Protanilla	4	0	Odontoponera	1	0
Stenamma	4	0	Oecophylla	1	0
Tapinoma	4	0	Ooceraea	1	0
Brachyponera	3	0	Parasyscia	1	0
Cardiocondyla	2	1	Paratrechina	0	1
Chronoxenus	3	0	Philidris	1	0
Dilobocondyla	3	0	Prionopelta*	1	0
Discothyrea	3	0	Probolomyrmex	1	0
Dorylus	3	0	Rotastruma	1	0
Kartidris	3	0	Simopone	1	0
Leptanilla	3	0	Syscia	1	0
Lophomyrmex	3	0	Vombisidris	1	0
Myopias	3	0	Yunodorylus	1	0
Recurvidris	3	0

worker (MCZ-ENT00763491, new to China) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763191) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763384) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00764608, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759751) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01worker (MCZ-ENT00763371) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00763341) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

minor worker (MCZ-ENT00760027) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

major worker (MCZ-ENT00760028) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759984) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759880) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763272) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00762839) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763246) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763401) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760062) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759741) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760060) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760068, new to China) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763692, new to China) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

minor worker (MCZ-ENT00763190) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

major worker (MCZ-ENT00763247) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763213) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763198) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00762843) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00759861) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm03 worker (MCZ-ENT00762821) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm04 worker (MCZ-ENT00762978) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm05 worker (MCZ-ENT00763312) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00759967) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760066) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760025, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763360, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760019) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00759776) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm02 worker (MCZ-ENT00759968) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763551) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759953) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00763500) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm02 worker (MCZ-ENT00763427) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763282) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763176). A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760042). A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763549) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763195) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760071) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763568) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763284). A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763328, new to Yunnan). A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763200, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00763220) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00763467) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00762951) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00762838) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763554) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map..

sp. clm01 worker (MCZ-ENT00762870) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00763366) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm03 worker (MCZ-ENT00763603) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm04 worker (MCZ-ENT00764622) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm05 worker (MCZ-ENT00762809) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00762820, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01worker (MCZ-ENT00763607) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00759841) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759773) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759928) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759796) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00759855) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00760045, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759778) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00762837) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00762875) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00763656, new to China) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759792) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00763514) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760064) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01worker (MCZ-ENT00759771) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00759985) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759935) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00763256) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00759803). A mesosoma in profile view B mesosoma in dorsal view C head in front view.

minor worker (MCZ-ENT00759865) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

major worker (MCZ-ENT00759866) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00764619) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

minor worker (MCZ-ENT00759918) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

major worker (MCZ-ENT00760026) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763577) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00762822) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

minor worker (MCZ-ENT00759762) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

major worker (MCZ-ENT00759980) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759837, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

minor worker (MCZ-ENT00760015) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

major worker (MCZ-ENT00760016) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763505) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763502) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00762907) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759885) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763507) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759758, new to China) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00763511) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00759897) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm03 worker (MCZ-ENT00759991) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00759763, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00759977) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm03 worker (MCZ-ENT00763303) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00764651) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00759754) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00763454) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm03 worker (MCZ-ENT00760040) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm04 worker (MCZ-ENT00759856) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00759854) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm03 worker (MCZ-ENT00764617) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00759752) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759748) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759859, new to China) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

sp. clm01 worker (MCZ-ENT00759780) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm02 worker (MCZ-ENT00759849) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

sp. clm03 worker (MCZ-ENT00759808) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

worker (MCZ-ENT00763178) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763321). A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00762856). A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00760009, new to Yunnan) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763657) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759807) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759845) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759809) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759806) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759931) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763325) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00759857) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763523) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

worker (MCZ-ENT00763165) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

Yunnan ant list

AMBLYOPONINAE

: 1 species

Emery, 1989

: 1 species (undescribed)

sp.

: 11 species

* Karavaiev, 1935

* (Xu, 2012)

(Xu, 2001)

* (Xu, 2012)

* (Xu, 2012)

* (Xu, 2000)

(Xu, 2006)

(Forel, 1900)

* (Guénard, 2013)

(Wheeler, 1928)

(Xu, 2001)

DOLICHODERINAE

: 3 species

(Forel, 1895)

(Forel, 1895)

(Forel, 1895)

: 9 species

Emery, 1889

Emery, 1889

Xu, 1995

* Santschi, 1920

Forel, 1886

Xu, 2001

Xu, 2001

(Smith, 1858)

(Smith, 1860)

: 1 species

(Roger, 1863)

: 1 species

Wheeler, 1921

: 1 species

(Mayr, 1862)

: 1 species

(Emery, 1895)

: 4 species

Wheeler, 1927

Wheeler, 1895

(Fabricius, 1793)

Forel, 1904

: 11 species

(Smith, 1861)

Zhou, 2001

Emery, 1893

Forel, 1895

Forel, 1902

Forel, 1912

* Forel, 1905

Wheeler, 1928

(Smith, 1860)

* Mann, 1921

* Bolton, 2007

DORYLINAE

: 19 species

* Wilson, 1964

Forel, 1900

* Jaitrong & Yamane, 2011

(Mayr, 1866)

Forel, 1911

Emery, 1889

Forel, 1901

Bingham, 1903

Forel, 1901

(Smith, 1857)

* Jaitrong & Yamane, 2013

* Jaitrong & Yamane, 2012

Wheeler & Chapman, 1930

Forel, 1901

Forel, 1901

Terayama & Kubota, 1993

* Jaitrong & Yamane, 2013

Forel, 1901

* Liu, 2015

: 1 species

Emery, 1889

: 1 species

* (Bharti & Wachkoo, 2013)

: 3 species

(Smith, 1857)

Westwood, 1835

Wheeler, 1913

: 1 species

(Mayr, 1879)

: 1 species

(Forel, 1907)

: 1 species

(Forel, 1895)

: 1 species

* Chen, 2015

: 1 species

Roger, 1861

: 1 species

Xu, 2000

ECTATOMMINAE

: 6 species

(Emery, 1889)

Zhou, 2001

* (Roger, 1860)

* Chen, 2017

Lattke, 2004

* Wu & Xiao, 1987

* Lattke, 2004

FORMICINAE

: 2 species

Terayama, 1985

Terayama & Hashimoto, 1996

: 1 species

(Smith, 1857)

: 28 species

Bingham, 1903

Wu & Wang, 1989

Zhou, 2001

Forel, 1892

* Wheeler, 1919

* Forel, 1894

Wu & Wang, 1989

(Fabricius, 1787)

Forel, 1894

Wang & Wu, 1994

* Forel, 1902

Forel, 1892

Forel, 1886

* Xiao & Wang, 1989

Emery, 1889

* Forel, 1892

* Forel, 1912

Mayr, 1866

Xiao & Wang, 1989

Wang & Wu, 1994

Wang & Wu, 1994

(Smith, 1858)

Mayr, 1865

Emery, 1889

Wu & Wang, 1989

Wu & Wang, 1989

* Forel, 1892

Forel, 1901

(Smith, 1858)

Santschi, 1925

(Smith, 1874)

: 7 species

(Smith, 1857)

* (Emery, 1925)

(Forel, 1893)

(Emery, 1889)

(Wu & Wang, 1994)

(Forel, 1893)

(Smithi, 1860)

: 2 species

* Bharti & Gul, 2012

* Smith, 1857

: 5 species

Latreille, 1798

* Ruzsky, 1904

Santschi, 1925

* Bondroit, 1917

Motschoulsky, 1866

Latreille, 1798

Wheeler, 1913

: 1 species

* Wheeler, 1929

: 6 species

Collingwood, 1982

(Fabricius, 1782)

* Bingham, 1903

Forel, 1912

* Stitz, 1930

Seifert, 2020

: 8 species

Xu, 1994

(Mayr, 1862)

(Forel, 1892)

* (Forel, 1892)

Xu, 1994

(Forel, 1894)

(Forel, 1902)

(Wu & Wang, 1995)

: 2 species

Forel, 1893

Xu, 1998

: 10 species

(Forel, 1902)

(Forel, 1886)

* (Forel, 1894)

* (Wang, 1997)

(Smith, 1874)

(Forel, 1894)

(Forel, 1899)

(Forel, 1894)

(Nylander, 1846) (Exotic)

(Forel, 1894)

: 1 species

(Fabricius, 1775)

: 2 species

* (Ito, 1914)

(Forel, 1913)

: 1 species

(Latreille, 1802) (Exotic)

: 4 species

Emery, 1894 (Exotic)

Santschi, 1920

Forel, 1894

* Forel, 1894

: 32 species

(Le Guillou, 1842)

Xu, 1998

Smith, 1858

(Drury, 1773)

Xu, 2002

Donisthorpe, 1938

Xu, 2002

Zhou & Huang, 2002

Xu, 1998

Xu, 2002

Smith, 1857

* (Walker, 1859)

Smith, 1858

Emery, 1901

Emery, 1889

* Emery, 1861

Emery, 1893

Walker, 1859

Wang & Wu, 1991

Smith, 1857

Emery, 1887

Xu, 2002

Wang & Wu, 1991

Roger, 1863

Mayr, 1879

Roger, 1863

(Latreille, 1802)

Xu, 2002

Smith, 1858

Bingham, 1903

Roger, 1863

Smith, 1858

: 7 species

Zhou, 2001

* Williams & LaPolla, 2016

* Williams & LaPolla, 2016

Emery, 1893

Forel, 1902

* Williams & LaPolla, 2016

* Chen & Zhou, 2018

: 6 species

Xu, 1997

Wu & Wang, 1992

Forel, 1911

(Smith, 1860)

Forel, 1894

Wheeler, 1927

LEPTANILLINAE

: 3 species

Tang, Li & Chen, 1992

Xu & Zhang, 2002

Xu, 2002

: 4 species

Xu, 2002

Xu, 2002

* Xu, 2002

Xu, 2012

MYRMICINAE

: 2 species

Zhou & Zheng, 1997

Emery, 1893

: 9 species

Emery, 1887

(Smith, 1921)

(Smith, 1874)

Emery, 1889

* Wheeler, 1921

Forel, 1911

Wheeler, 1930

(Forel, 1902)

(Forel, 1902)

(Forel, 1902)

: 3 species

* Seifert, Okita & Heinze, 2017 (Exotic)

Wheeler, 1929

(Forel, 1890)

: 18 species

(Xu, 2003)

(Jerdon, 1951)

(Xu, 2003)

(Forel, 1902)

(Forel, 1902)

(Xu, 2003)

(Xu, 2003)

* (Jerdon, 1851)

* Wu & Wang, 1995

Westwood, 1840

* (Zhou & Zheng, 1997)

(Xu, 2003)

(Wheeler, 1928)

(Xu, 2003)

(Xu, 2003)

(Forel, 2003)

(Wheeler, 1928)

(Zhou & Zheng, 1997)

(Ettershank, 1966)

: 4 species

(Latreille,1802)

* Bolton, 1974

Emery, 1893

Smith, 1853

: 25 species

Smith, 1857

* Mayr, 1879

Forel, 1904

Mayr, 1897

* Mayr, 1879

Forel, 1902

Mayr, 1879

Forel, 1902

Emery, 1888

Forel, 1902

* Smith, 1857

Wu & Wang, 1995

Forel, 1901

Ito, 1914

Forel, 1900

Forel, 1902

* Forel, 1911

Mayr, 1879

Mayr, 1879

Mayr, 1879

Forel, 1902

Emery, 1896

Forel, 1902

Forel, 1902

Santschi, 1925

: 3 species

* Bharti & Kumar, 2013

Santschi, 1910

* Bharti & Kumar, 2013

: 1 species

(Mayr, 1872)

: 1 species

Xu, 2012

: 1 species

(Karavaiev, 1935)

: 3 species

Xu & Zheng, 1995

Bolton, 1991

Xu, 1999

: 1 species (undescribed)

sp.

: 3 species

Emery, 1893

Emery, 1893

(Jerdon, 1851)

: 1 species (undescribed)

sp.

: 2 speices

(Guérin-Méneville, 1844)

Emery, 1889

: 1 species

* (Smith, 1874)

: 5 species

Santschi, 1925

(Jerdon, 1851)

* Wu & Wang, 1995

Mayr, 1879

(Linnaeus, 1758)

: 5 species

* Okido, Ogata & Hosoishsi, 2020

* Okido, Ogata & Hosoishsi, 2020

Zhou, Huang & Ma, 2008

Zhou, 2001

* Wheeler, 1921

Emery, 1889

Terayama, 1995

: 11 species

Radchenko, Zhou & Elmes, 2008

Radchenko, Zhou & Elmes, 2008

* Kupyanskaya, 1990

* Radchenko & Elmes, 2008

Emery, 1889

Radchenko & Elmes, 2009

Radchenko & Rigato, 2008

Emery, 1889

Wheeler, 1928

Radchenko, Zhou & Elmes, 2008

Mayr, 2001

Radchenko & Elmes, 2008

: 1 species

Saunders, 1842

: 2 species

Zhou & Huang, 2006

Xu & Wang, 2004

: 42 species

Bingham, 1903

Forel, 1902

Emery, 1902

Forel, 1902

Eguchi, 2008

(Mayr, 1866)

Smith, 1858

Smith, 1874

* Eguchi, 2006

(Wheeler, 1927)

* Wheeler, 1928

Mayr, 1879

* Wheeler, 1928

Forel, 1885

* Eguchi, 2006

* Eguchi, 2006

Forel, 1902

Emery, 1901

* Smith, 1858

Smith, 1874

* Eguchi, 2008

* Mayr, 1865

Santschi, 1925

* Smith, 1860

* Santschi, 1920

* Forel, 1913

Forel, 1902

* Eguchi, 2008

Forel, 1902

* Özdikmen, 2010

(Wu & Wang, 1992)

* Forel, 1902

Forel, 1902

Roger, 1863

* Forel, 1913

* Forel, 1905

* Eguchi, 2008

* Eguchi, 2008

* Eguchi, 2006

Forel, 1902

Forel, 1902

* Santschi, 1925

: 4 species

Emery, 1887

Xu & Zhang, 2002

(Smith, 1860)

Emery, 1895

: 3 species

* (Wheeler, 1954)

Xu & Zheng, 1995

(Forel, 1890)

: 2 species

* Forel, 1900

Xu, 1999

: 1 species

* Bolton, 1991

: 3 species

Wheeler, 1928

Buren, 1972 (Exotic)

Wheeler, 1923

: 4 species

Liu & Xiu, 2011

DuBois, 1998

* DuBois, 1998

Liu & Xiu, 2011

: 24 species

Xu & Zhou, 2004

* De Andrade, 1994

(Xu, 2000)

* Emery, 1887

* (Bolton, 2000)

Bolton, 2000

Emery, 1895

* (Terayama, 1996)

* Wheeler, 1929

Cameron, 1886

* (Roger, 1862)

* Emery, 1869 (Exotic)

* (Brown, 2000)

(Brown, 1949)

Lin & Wu, 1996

* De Andrade, 1994

(Xu & Zhou, 2004)

* Bolton, 2000

* Bolton, 2000

* (Forel, 1912)

Bolton, 2000

* Bolton, 2000

* (Bolton, 2000)

* Bolton, 2000

(Xu & Zhou, 2004)

: 7 species

Zhou, 2010

(Smith, 1874)

(Huang, 2004)

(Ruzsky, 1895)

Zhou, 2010

Zhou, 2010

(Wheeler, 1929)

: 29 species

Bolton, 1977

(Nylander, 1846)

Xu & Zheng, 1994

Bolton, 1977

Wang & Wu, 1988

Bolton, 1977

Xu & Zheng, 1994

* Bolton, 1977

* Emery, 1893

Wheeler, 1927

(Smith, 1861)

(Bolton, 1976)

Forel, 1905

Mayr, 1870

Bolton, 1977

Wheeler, 1928

Bingham, 1903

Viehmeyer, 1916

Mayr, 1870

* (Emery, 1893)

* Yamane & Jaitrong, 2011

Wang & Xiao, 1988

(Smith, 1851) (Exotic)

Mayr, 1879

* Mayr, 1870

(Forel, 1890)

* (Forel, 1902)

Forel, 1902

Xu & Zheng, 1994

: 2 species

(Jerdon, 1851) (Exotic)

(Forel, 1902) (Exotic)

: 2 species

* Wang, 2005

Wu & Xiao, 1989

: 1 species

* Xu & Yu, 2012

PONERINAE

: 6 species

Mayr, 1870

* Mayr, 1897

Radchenko, 1993

* Emery, 1893

Forel, 1900

Forel, 1912

: 2 species

Xu, 1994

* (Emery, 1895)

(Mayr, 1862)

: 1 species

(Emery, 1887)

: 1 species

(Emery, 1889)

: 5 species

Wu & Wang, 1995

Xu, 1998

* (Wheeler, 1906)

(Forel, 1930)

Emery, 1893

: 1 species

* (Le Guillou, 1842)

: 8 species

(André, 1892)

(Smith, 1858)

Mayr, 1867

(Forel, 1886)

(Xu, 1995)

* Emery, 1900

(Forel, 1912)

(Xu, 1995)

: 1 species

Xu, 1998

: 1 species

(Wheeler, 1928)

: 1 species

(Smith, 1858)

: 7 species

(Mayr, 1897)

(Roger, 1860)

* (Forel, 1893) (Exotic)

(Santschi, 1937)

(Roger, 1859) (Exotic)

Onoyama, 1989

(Smith, 1860)

: 17 species

Forel, 1900

Forel, 1900

(Mayr, 1870)

Emery, 1895

* Karavaiev, 1935

(Smith, 1857)

* (Mayr, 1870)

* Forel, 1905

Xu, 2000

Emery, 1895

Xu, 2000

Xu, 2000

* (André, 1887)

* (Jerdon, 1851)

* Zhou, 2012

* Xu, 2015

Xu, 2000

: 1 species

(Emery, 1893)

: 3 species

Xu, 1998

* Xu, 2014

* Xu, 2012

: 6 species

Wang, 1993

* Wang, 1993

Wang, 1993

Emery, 1892

Smith, 1857

Wang, 1993

: 1 species

* (Smith, 1858)

: 2 species

Forel, 1911

* (Smith, 1859)

: 14 species

Terayama, 1986

Xu, 2001

Xu, 2001

* Terayama, 1986

Xu, 2001

Xu, 2001

Xu, 2001

Xu, 2001

* Zhou, 2001

Xu, 2001

Xu, 2001

Wheeler, 1928

Wheeler,1928

Xu, 2001

: 2 species

(Smith, 1858)

(Jerdon, 1851)

PROCERATIINAE

: 3 species

* Xu, 2014

* Xu, 2014

Forel, 1912

: 1 species

Xu & Zeng, 2000

: 4 species

Karavaiev, 1935

Xu, 2006

* Staab, 2018

Xu, 2000

PSEUDOMYRMECINAE

: 12 species

(Walker, 1859)

Xu & Chai, 2004

Smith, 1877

(Forel, 1902)

Xu & Chai, 2004

Xu & Chai, 2004

Xu & Chai, 2004

Wu & Wang, 1990

(Smith, 1860)

Ward, 2001

Xu & Chai, 2004

(Jerdon, 1851)

Discussion

Ants in the Hengduan Mountain region

Field inventories and data synthesis efforts are essential for our understanding of ant diversity in ‘hotspots’ that harbor most of Earth’s biodiversity. Our study represents new survey data from an understudied region. We produce the first ant species checklist from China’s Hengduan Mountains (130 species).

A majority of the ant species were only collected below 1500 m, consistent with the strong effect of elevation on ant diversity observed elsewhere (Suppl. material 1, Fig. S1). This also suggests that future sampling in low elevation areas may increase species detection. For example, the number of species recovered in this survey is relatively low compared to the overall richness of this genus. This could be because we have relatively few collection events at low elevations where many of these species are known to occur. Indeed, all six species were collected below 1000 m from only three independent Winkler sampling sites.

Many of the new records in our collection such as , , , , , , and represent the northern-most records of their known distributional ranges. Species records such as , , , , , and show a disjunction from the rest of their known distributions. It is unclear whether those records represent true biogeographic disjunctions, or sampling / taxonomic artifacts. Another potential reason could be that they were collected in the past, but have not been reported due to the lack of taxonomic infrastructure and species check lists from this region (Guénard et al. 2017). Additional inventories of ant diversity and taxonomic treatments are needed to answer these questions.

Despite the comparatively small area of China’s Hengduan Mountains that we explored for this inventory of myrmecofauna, we were able to collect 130 species, which accounts for more than 24 % of the total number of ant species (N = 550) for Yunnan province. Among them, more than 10% of the ant species that were collected in this survey represent new records for Yunnan province. Moreover, there are still more than 41 morphospecies (32% of the total collected) that we believe are undescribed and new to science. To date, three species (Figs 82–84) and one species (Fig. 75) are undergoing taxonomic revision, and species descriptions are being prepared.

Figure 82.

sp. clm01 worker (MCZ-ENT00759959) A mesosoma in profile view B mesosoma in dorsal view C head in front view.

Figure 84.

sp. clm03 worker (MCZ-ENT00763515). A mesosoma in profile view B mesosoma in dorsal view C head in front view.

Figure 75.

sp. clm01 worker (MCZ-ENT00764656) A mesosoma in profile view B mesosoma in dorsal view C head in front view D global distribution map.

Our sampling of the full ant diversity of the Hengduan mountain region is still relatively limited. For example, we only had one sampling site per elevation, which is insufficient to cover the complex topology of the Hengduan Mountains. We also only used leaf litter extraction and hand collection, which is unlikely to recover complete ant assemblages. The incorporation of additional sampling techniques into our methodology, such as pitfall trapping, soil baiting, twig sampling, light trapping and canopy fogging, will cover more strata and lifestyles, and thus significantly increase our rate of species discovery. Overall, our results highlight how little was previously known about the ant fauna in this region and emphasize the need for further collecting in order to better understand the hidden ant biodiversity in China’s Hengduan Mountains, and Yunnan Province overall.

Ants in Yunnan

If the total species richness of ants in Yunnan, with 550 species, is still an underestimate of the full species numbers, the exceptional diversity of genera encountered in this region needs to be highlighted. With 99 genera, Yunnan generic diversity is only matched globally by a few regions in South East Asia, and Queensland, Australia. A major difference with other Asian regions lies in the composition of the genera retrieved and their origin. For instance, genera found in Borneo, Sumatra, Java, Vietnam, Thailand, Peninsular Malaysia are almost exclusively derived from tropical, Oriental origins. In contrast, the geographic location and topography of Yunnan province has promoted an intermixing of taxa from several biogeographic regions: the Oriental realm from the south, the Palearctic realm from the northwest, and the Sino-Japanese realm from the northeast. As a result, the composition of the Yunnan ant fauna includes both tropical, subtropical, and temperate elements. Such intermixed communities are evident even at a small scale. For instance, during previous fieldwork conducted in the Gaoligongshan Mountains in 2015 by two of the authors (BG and CL), for which specimens were unfortunately lost, the coexistence of tropical (, ), subtropical () and temperate genera (, ) was observed on a hillside at an elevation of about 1900 m on an ~ 250 m² patch of grassland. Interestingly, while this area exhibited a transition where fauna from distinctly different origins coexisted along a thin band of altitude, at lower elevations, tropical genera were dominant and at higher elevations, temperate genera became dominant. Overall, this generated an unexpectedly diverse faunal composition, with such mixed communities contemplated by Wheeler (1915) to explain the generic composition and diversity of fossil ants observed in Baltic amber. Possibly, the ant composition of genera now retrieved within Yunnan might represent the remains of a once more widespread assemblage found within Asia and Europe during the Miocene (Guénard et al. 2015). This highlights the specific nature of the Yunnan ant fauna and its importance in studying ant biogeography within Asia. It also serves as an excellent example of the formation and stability of ant community assemblages over time. Other neighboring regions such as Myanmar, Bhutan, Nepal or northeast India are likely to exhibit similar features, but to this point, the myrmecological exploration of these regions has been largely fragmentary (Guénard et al. 2010, 2012).

The diversity of Yunnan ants is also remarkable for particular ant genera for which their global peak of diversity is encountered in the region. While it is important to note that the global diversity of specific genera as well as their overall taxonomic descriptions remain incomplete, seven genera present their highest currently known global diversity in Yunnan (: 5 species, : 8 species, : 3 species, : 2 species, : 14 species, : 7 species, : 11 species), while six others are remarkable by the level of global diversity there, among the highest observed globally (: 19 species, : 19 species, : 3 species, : 7 species, : 4 species, : 3 species).

In conclusion, the important topographic variation, with mountain ranges aligned along a north-south axis combined with the presence of multiple climatic zones, including tropical rainforest in the lowland areas of the southern part of Yunnan create a diversity of microhabitats for supporting a diverse ant fauna. Moreover, the geographic position of Yunnan at the confluence of three biogeographic realms may promote ant diversity in the region. The collection of these 16 new ant records for Yunnan together with our previous discovery of 40 new ant records for Yunnan (Liu et al. 2015a) suggest that the true ant diversity in Yunnan is significantly higher. Moreover, the species diversity of some ant genera in nearby regions (based on data from GABI) also suggests that some ant genera sampled will ultimately be much more diverse in Yunnan (Guénard et al. 2017). For example, Hong Kong has recorded a similar diversity of species while having an area nearly 350 times smaller and a much less diverse topography compare to Yunnan (Tang et al. 2019), perhaps because leaf litter extraction has not been widely used for sampling ants in Yunnan. Thus, our survey to date indicates that further intensive sampling focused on different ecological strata (arboreal, leaf litter, subterranean) and combining various methods of extraction in both tropical and mountain habitats should yield many additional records and new species discovery in this region.