Nedyopodini 族只包含一屬，就是Nedyopus 屬。這個屬是由Attems 在1914 年建立。因為發表者對種描述的不詳盡，或者沒有圖形，甚至是錯誤鑑定，該屬確實包含了幾種有效種一直很混亂。利用解剖光學顯微鏡及電子顯微鏡觀察比對模式標本、新採集標本及文獻發現，原紀錄有二十二個種/亞種/變種的Nedyopus 屬，其中一個亞種、一個變種提升為種的階層、有七個種/雅種為同種異名、而N. dawydoffiae (Attems, 1953)須從該屬剃除，該屬確定含有十四種。此外發表五個新種，故目前Nedyopus 屬共有十九種。文中亦修訂了Nedyopus 屬也就是Nedyopodini 族的鑑別特徵。本研究中建置該屬十九個物種的檢索表以及世界分布圖。從該屬的世界分布圖可知分佈於東方到東南亞的Nedyopus 屬主要是暖溫帶的物種，也分佈於熱帶，從圖中顯示出Nedyopus 屬的物種多樣性在台灣很高，據此推測台灣應是該屬的熱點(hotspot)。利用型態特徵使用paup 進行最大簡約法(maximum parsimony)分析、貝葉氏導出式分析(Bayesian inference analysis)則以Mr.Bayes 3.2.2 分析Nedyopus 屬的親緣關係，結果支持Nedyopus 屬是單系群 (在最大簡約法中，bootstrap 值是100%，且有兩個完全沒有同塑性質的近裔共性之特徵支持)，而N. pictus (Brölemann, 1916)是Nedyopus 屬中較原始的物種。許多後衍而出的Nedyopus 物種分布於日本、台灣，顯示出這兩地的生態環境有利於Nedyopus 屬的種化。根據最大簡約法分析的結果，利用馬陸生殖肢作親緣關係之研究仍較使用身體特徵更好(生殖肢每一特徵有較高RC值)。 研究顯示Nedyopus 屬生殖肢末端的B瓣在演化中最早發展出來，顯示出該特徵在交配中應占重要功能。

Tribe Nedyopodini Jeekel, 1968 only contained a genus Nedyopus which was established byAttems (1914) contained how many valid species is a puzzle all along because the poor descriptions, lacking illustrations, or misidentifications. Making use of stereomicroscope and scanning electron microscope to comparing type specimens and new materials, and checking the old literatures, there were twenty-two species/subspecies/variety turn out fourteen species are valid, among them, a subspecies and a variety are promoted as species status, seven species/subspecies are new synonyms, one species N. dawydoffiae (Attems, 1953) is deleted. Besides, there are five new species being added. Totally, there are nineteen species in genus Nedyopus so far. The diagnosis of genus Nedyopus/tribe Nedyopodini is amended. Provide a key and distribution map in the world which shows genus Nedyopus are warm temperate (mainly)–tropical East and Southeast Asian creatures, and Taiwan
should be its main hotspot by owing a high species diversities in genus Nedyopus. Making use of paup running maximum parsimony analysis and Mr.Bayes 3.2.2 running Bayesian inference analysis to develop the phyletic relationships of genus Nedyopus based on morphological characters, and the results supported genus Nedyopus is monophylic with 100% bootstrap value and two nonhomoplastic synapomorphies in maximum parsimony analysis, and N. pictus (Brölemann, 1916) is basal lineage. Quite a few deepest derived species placed in Japan and Taiwan which shows these two localities are advantages for genus Nedyopus speciation. According the cladistics analysis, gonopodal traits are still better characters than somatic traits in study phylogeny because higher RC values. Solenophore lobe B of genus Nedyopus developed first in the evolutionary history showed its importance in copulation.

Index
Chapter 1. Introduction 1
References……………………………………………………………………………………………… 19
The key to 16 orders of Diplopoda………………………………………………………………… 25
Appendix 1. A classification of the Class Diplopoda (after Shelly 2002) and the
biogeography of illipedes………………………………………………… 31

Chapter 2 Review the East and Southeast Asian genus Nedyopus Attems, 1914 and
tribe Nedyopodini Jeekel, 1968
2.1 Abstract…………………………………………………………………………………………… 32
2.2 Introduction……………………………………………………………………………………… 33
The diagnosis of genus Nedyopus Attems, 1914, tribe Nedyopodini Jeekel, 1968… 36
2.3 Material and Methods………………………………………………………………………… 39
2.4 Results
2.41 Species needs description again-
(1) Nedyopu cingulatus (Attems, 1898)……………………………………………. 43
Remarks………………………………………………………………………… 50
(2a) Nedyopus tambamus (Attems, 1901)…………………………………………. 55
Remarks……………………………………………………………………….. 60
(2b) Nedyopus tambanus mangaesinus (Attems, 1909)……………………………. 61
Remarks………………………………………………………………………. 62
(3) Nedyopus ikaonus (Attems, 1909), new status……………………………… 63
Remarks………………………………………………………………………… 67
(4) Nedyopus unicolor (Carl, 1902), new status……………………………… 69
Remarks………………………………………………………………………… 75
(5) Nedyopus boninensis Verhoeff, 1940………………………………………… 79
Remarks……………………………………………………………………… 84
2.42 Species were alreadys (re)described in detail- 85
(6) Nedyoups beroni (Golovatch, 1995)…………………………………………… 86
(7) Nedyoups picturatus (Golovatch, 1995) ………………………………………87
(8) Nedyopus hsientienensis (Wang, 1957)…………………………………………88
(9) Nedyopus pectinatus (Wang, 1957)………………………………………………90
(10) Nedyopus caudatus Chen, Golovath and Chang, 2006……………………… 92
(11) Nedyopus latus Chen, Golovath and Chang, 2006……………………………93
(12) Nedyopus wui Chen, Golovath and Chang, 2006………………………………95
(13) Nedyopus pictus (Brölemann, 1916)……………………………………………97
(14) Nedyopus mahunkai (Korsós and Golovatch, 1989)………………………… 98
2.43 Species deleted from genus Nedyopus-
“Helicorthomorpha” dawydoffiae (Attems, 1953)………………………………99
2.5 Discussion
Species in genus Nedyopus Attems, 1914……………………………………………103
The amend diagnosis of the tribe Nedyopodini and genus Nedyopus………… 105
Biogeography of genus Nedyopus Attems, 1914…………………………………… 106
The possible dispersal direction of Nedyopus unicolor (Carl, 1902),
new status……… 109
Acknowledgments………………………………………………………………………………………… 111
References ……………………………………………………………………………………………… 114
Appendix 2.1 A letter about the holotype Nedyopus hiburizimensis Miyosi, 1955
was replied from 藤江孝利……………………………………………………………… 190
Appendix 2.2 The Map of Humid continental climate………………………………………… 191
Appendix 2.3 The Map of Humid subtropical climate ……………………………………….. 192
Appendix 2.4 The Map of Tropical climate………………………………………………….. 193

Chapter 3 Describe five new species of genus Nedyopus from Japan, Taiwan, and Hong Kong, with special reference to
Taiwan ecological protection
3.1 Abstract……………………………………………………………………………………………… 194
3.2 Introduction……………………………………………………………………………….. 194
3.3 Material………………...…………………………………………………………………. 196
3.4 Results
(15) Nedyopus nov. sp. 1……….……………………………………………………….. 196
(16) Nedyopus nov. sp. 2……….……………………………………………………….. 200
(17) Nedyopus nov. sp. 3………..………………………………………………………. 206
(18) Nedyopus nov. sp. 4 ……….………………………………………………………. 210
(19) Nedyopus nov. sp. 5 …………….…………………………………………………. 215
3.5 Discussion
Species of genus Nedyopus Attems, 1914………………….…………………………….. 220
Key to nineteen Nedyopus species ………………...……………………………………. 220
Ecology of the genus Nedyopus Attems, 1914..…………...………………………………. 225
The localities in Taiwan needed to be protected…………...…………………………… 228
Acknowledgments…………………………………………………………………………………………… 230
References………………………………………………………………………………………………… 231
Appendix 3.1 Taiwan millipedes in Wang’s work (1955~1964), Korsós (2004),
and New species added/corrected…………………………………………257
References…………………………………………………………………….263
Appendix 3.2 Fauna of Taiwan millipedes 2016…………………………………………………… 268
Species recovered…………………………………………………………… 272
Species which became junior synonyms……………………………………272
Deleted species …………………………………………………………… 272
Chapter 4. Phylogeny of genus Nedyopus Attems, 1914 based on morphological analysis

4.1 Abstract…………………………………………………………………………………………… 273
4.2 Introduction…………………………………………………………………………………… 274
4.3 Material and Methods
4.31 Taxon sampling…………………………………………………………………………… 278
4.32 Character assessment…………………………………………………………………… 278
4.33 Phylogenetic analysis………………………………………………………………… 279
4.34 Analytical methods……………………………………………………………………… 281
4.35 Morphological characters discussion………………………………………………… 282
4.4 Results
The phylogenies reconstructed by MP
(1) On the basis of 52 somatic and gonopodal characters…………………………… 309
(1.1) In Equal Weighting analysis (EW)…………………………………………… 309
(1.2) In both analyses (EW and SW)…………………………………………………. 310
(1.2a) In EW analysis ……………………………………………………… 311
(1.2b) In SW analysis ……………………………………………………… 312
(2) Besed on 18 gonopodal characters…………………………………………………… 313
(2.1) In Equal Weighting analysis (EW) ………………………………………… 313
(2.2) Clades A, B, C are recovered as well with a strong bootstrap
support in both analyses………………………………………………… 313
The phylogenies reconstructed by BI………………………………………………………… 314
(1) Anaylsis by 52 somatic and gonopodal characters………………………………….314
(2) Anaylsis only by 18 gonopodal characters……………………………………………315
4.5 Discussion………………………………………………………………………………………………315
The classification of “H.” dawydoffiae (Attems, 1953) is unfixed…………………316
Genus Nedyopus Attems, 1914 is a monophyly which is documented
and N. pictus (Brölemann, 1916) is the basal element of genus
Nedyopus by phylogenetic study ………………………………………………316
The clade (N. caudatus, N. wui)………………………………………………………………318
The possible phylogenetic relationships……………………………………………………319
The possible phylogenetic relationships of Nedyopus Attems, 1914……...........323
The speciation of genus Nedyopus Attems, 1914………………………………329
The possible source of Taiwan Nedyopus species………………………………………… 330
Morphology discussed…………………………………………………………………………… 331
Prospective in the future……………………………………………………………………… 334
Acknowledgments…………………………………………………………………………………………… 336
References……………………………………………………………………………………………………337
Appendix 4.1 Map of Taiwan connected to Ryukyu Islands (Japan), mainland
China, and Philippines by shallow seabed (After Wallace 1880)… 399
Appendix 4.2 Maps of tropical Southeast Asia and Austral-Asia iIlustrating depth
contours of 10, 50 and 120 m below present level…………………… 400
Appendix 4.3 Character change lists at every branch (18 gonopodal characters)………… 403
Appendix 4.4 Apomorphy lists at every branch (18 gonopodal characters)………………… 406
Appendix 4.5 Eighteen gonopodal character diagnostics………………………………………… 405
Appendix 4.6 Character change lists at each branch (52 morphological characters)…… 406
Appendix 4.7 Apomorphy lists at each branch (52 morphological characters)……………… 423
Appendix 4.8 Fifty-two somatic and gonopodal character diagnostics……………………… 427

List of Tables
Table
3.1 Information for type specimens of five new Nedyopus species…………………………………………234
3.2 Taiwan endemic species placed at only one locality…………………………………………………… 237
4.1 The differences of characters of genus Nedyopus species and outgroups………………………… 343
4.2 The states and codes of all traits used (based on adult males) in phylogenetic analysis… 376
4.3 Data matrix for the phylogenetic analysis………………………………………………………………… 380

List of Figures
Chapter 1
Figures
1–8 Orders Polyxenida, Glomerida, Sphaerotheriida, Polydesmida,
Glomeridesmida, Chordeumatida, Polyzoniida, and Chordeumatida………………………………… 27
9–15 Orders Siphonocryptida, Siphonophorida, Siphoniulida, and Platydesmida…………………………… 28
16–22 Orders Stemmiulida, Callipodida,and Spirobolida…………………………………………………………… 29
23–29 Show gnathochilarium, Orders Spirostreptida and Julida………………………………………………… 30

Chapter 2
Figures
2.1–25, 257–266, 381, 388–390 Nedyopus cingulatus (Attems, 1898)
and Orthomorpha cingulata Attems, 1898, syntype……………………………………………………122–124, 161,176, 177
2.26–38, 382, 383 Strongylosoma patrioticus Attems, 1898, syntypes………………………………………… 125, 126, 176
2.39–50, 384, 385 Nedyopus koreanus Verhoeff, 1936, monotype………………………………………………… 127, 128, 176
2.51–55, 386, 387 Nedyopus koreanus niponianus Verhoeff, 1940, syntypes………………………………… 129, 176
2.56–68, 267–281 Nedyopus tambanus (Attems, 1901)…………………………………………………………… 130, 131,162,163
2.69, 393, 394 Strongylosoma tambanum Attems, 1901, holotype………………………………………………… 131, 178
2.70–75, 395 Strongylosoma tambanus mangaesinus Attems, 1909, cotype……………………………………… 132, 178
2.76–85, 396, 397 Strongylosoma tambanum ikaonum Attems, 1909, syntype…………………………………… 133, 134, 178
2.86–105, 282–300, 398, 399 Nedyopus unicolor (Carl, 1902), new status non-type…………………… 135, 136, 164,165, 179
2.106–111, 400, 401 Nedyopus venustus Shinohara, 1963, holotype…...…………………………………… 137, 179
2.402–406 Nedyopus hiburizimensis Miyosi, 1955…………………………………………………………………… 179
2.109–111 Nedyopus hiburizimensis aquilus Haga, 1968…………………………………………………………… 137
2.112–124, 391, 392 Nedyopus boninensis Verhoeff, 1940, monotype…………………………………………… 138, 139, 177
2.125–140, 407–411 Vaulogerodesmus beroni Golovatch, 1995, holotype……………………………………… 140, 141, 180
2.141–155, 412–414 Vaulogerodesmus picturatus Golovatch, 1995, holotype………………………………… 142, 143, 180
2.156–165, 301–318, 415–420 Nedyopus hsientienensis (Wang, 1957)……………………………………… 144, 166, 1167, 181
2.166–185, 319–331, 421–428 Nedyopus pectinatus (Wang, 1957) ……………………………………………… 145–148, 168, 169, 182
2.186–196, 332–339, 429–446 Nedyopus caudatus Chen, Golovath and Chang, 2006………………………… 149, 150, 170, 183
2.197–208, 340–348, 447–455 Nedyopus latus Chen, Golovath and Chang, 2006…………………………… 151, 152, 171, 184
2.209–220, 349–356, 456–465 Nedyopus wui Chen, Golovath and Chang, 2006……………………………… 153, 154, 172, 185
2.221–231, 357–368, 466, 467 Nedyopus pictus (Brölemann, 1916)…………………………………………… 155, 156, 173, 174, 186
2.232–242, 369–380, 468–471 Nedyopus mahunkai (Korsós and Golovatch, 1989)………………………… 157, 158, 174, 175, 186
2.243–256 Nedyopus dawydoffiae (Attems, 1953)….……………………………………………………………… 159, 160
2.472&473 Helicorthomorpha dawydoffiae Attems, 1953…………………………………………………………… 187
2.474–480 Vaulogerodesnus dawydoffiae (Attems, 1953)………………………………………………………… 187

Chapter 3
Figures
3.1–3.11, 3.61–3.68 Nedyopus nov. sp. 1……….……………………………………………………………… 238, 246
3.12–3.27, 3.69–3.76 Nedyopus nov. sp. 2……….……………………………………………………………… 239, 240, 247
3.28–3.40, 3.77–3.84 Nedyopus nov. sp. 3 ……….……………………………………………………………. 241, 242, 248
3.41–3.49, 3.85–3.103 Nedyopus nov. sp. 4 ……….…………………………………………………………… 243, 249, 250
3.50–3.60, 3.104–3.109 Nedyopus nov. sp. 5…………….…………………………………………………… 244, 245, 251
3.110 The elevations of genus Nedyopus species in the world………………………………………………. 252

Chapter 4
Figures
4.1–4.8, Tectoporus fugilil Golovach and Stoev, 2009…………………………………………………………… 381, 382
4.9–4.16, Tectoporus bispinosusGolovach and Stoev, 2010…………………………………………………… 383, 384
4.17–4.27, 4.52–4.55 Helicorthomorpha holstii (Pocock, 1895)……………………………………………. 385, 386, 391
4.28–4.43, 4.56, 4.57 Aponedyopus montanus Verhoeff, 1939…………………………………………………… 387, 389, 391
4.44–4.51, 4.58–4.63 Orthomorpha coarctata (De Saussure, 1860)…………………………………………… 390, 392

Figure 4.60 The most-parsimony cladogram derived under the equal weighting
procedure based on 52 morphological characters…………………………………………… 393
Figure 4.61 The most-parsimony cladogram derived under the successive
weighting procedure based on 52 morphological characters……………………………… 394
Figure 4.62 The most-parsimony cladogram derived under the equal weighting
procedure based on 18 gonopodal characters only……………………………………………395
Figure 4.63 Phylogenetic relationships of Nedyopus species using Maximum
Parsimony (MP) analysis under the successive weighting procedure
based on 18 gonopodal characters only…………………………………………………………396
Figure 4.64 Bayesian inference topology of 52 somatic and gonopodal characters.
Numbers below branches are Bayesian posterior probability (BPP)………………………397
Figure 4.65 Bayesian inference topology of 18 gonopodal characters only.
Numbers below branches are posterior probability values…………………………………398

List of Map
Map 2.1. The distributions of genus Nedyopusspecies in the world………………………………………………188
Map 2.2 The distributions of Nedyopus species in Taiwan………………………………………………………… 189
Map 3.1 The distributions of new species of genus Nedyopusin the world (2016)…………………………… 253
Map 3.2 The distributions of genus Nedyopusspecies in the world up to present (2016)……………………254
Map 3.3 The distribution of seven Nedyopus species in Taiwan……………………………………………………255
Map 3.4 Thirty-three Taiwan endemic species placed at only one locality:
locality 1 to locality 23………………………………………………………………………………256

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Humid continental climate (溫帶大陸性濕潤氣候):
Available at: https://en.wikipedia.org/wiki/Humid_continental_climate
Humid subtropical climate (副熱帶濕潤氣候):
Available at:https://en.wikipedia.org/wiki/Humid_subtropical_climate
Japan's Tropical Timber Imports-An Overview, Japan Tropical Forest Action Network (JATAN). Available at: http://www.jatan.org/eng/index-e.html
Köppen climate classification (柯本氣候分類法):
Available at: https://en.wikipedia.org/wiki/Köppen_climate_classification
The project of MILLI-PEET. Available at: http://www.fieldmuseum.org/research_
collections/zoology/zoo_sites/millipeet/milli_key.html
Tropical monsoon climate (熱帶季風氣候):
Available at: https://zh.wikipedia.org/wiki/热带