PROC. ENTOMOL. SOC. WASH. 112(2), 2010, pp. 214–228

NOTES AND NEW RECORDS OF (INSECTA: SIPHONAPTERA) FROM AND MAMMALS COLLECTED IN SOUTHERN

MICHAEL W. HASTRITER AND SARAH E. BUSH

(MWH) Monte L. Bean Life Science Museum, Brigham Young University, 290 MLBM, P.O. Box 20200, Provo, Utah 84602-0200, U.S.A. (e-mail: [email protected]); (SEB) Natural History Museum and Biodiversity Research Center, The University of Kansas, Lawrence, Kansas 66045, U.S.A.

Abstract.—During 2004–2007, 1,222 birds comprising 184 and 540 small mammals comprising 61 species were examined for ectoparasites as part of a broader biodiversity survey and inventory program conducted in Guangxi and Guizhou Provinces in southern China. A collection of 87 fleas was obtained. They included eight species representing seven genera and four families: gallinulae gallinulae (Dale), Macrostylophora jiangkouensis Li and Huang (); incurva Jordan, Palaeopsylla remota Jordan (Ctenophthalmidae); Nycteridopsylla iae Beaucournu and Kock, Thaumapsylla breviceps Rothschld (); and Aviostivalius klossi Jordan and Rothschild, and Lentistivalius insolli Traub (Pygiopsyllidae). Illustrations are provided to supplement those in the original description of M. jiangkouensis. The host preference for L. insolli for birds is established with certainty and its derivation and distribution are discussed. The potential impact of predatory ants on flea populations at low elevations in tropical latitudes is considered. Key Words: Ceratophyllidae, Ctenophthalmidae, ectoparasites, Ischnopsyllidae, Pygiopsyllidae, Aviostivalius, Macrostylophora, Lentistivalius, Nyc- teridopsylla, Palaeopsylla, Thaumopsylla DOI: 10.4289/0013-8797-112.2.214

The tropical region of southern China five nature reserves in Guangxi and is part of the Indo-Burma biodiversity Guizhou Provinces. Ectoparasites were hotspot, one of the richest and most sampled from terrestrial vertebrates as threatened centers of biodiversity on part of the survey. Here we report on the earth (Myers et al. 2000). In the past, fleas (Siphonaptera) that were collected the political landscape of China made from small mammals and birds. the description and documentation of biodiversity in this region difficult; MATERIALS AND METHODS however, in 2004–2007 we conducted Fleas were collected from the fol- biotic surveys of terrestrial vertebrates in lowing sites (Fig. 1). * Edited by Jerry Cook; accepted by Michael W. Site 1. Jing Xi County Nature Gates Preserve, located in Guangxi Province, VOLUME 112, NUMBER 2 215

Fig. 1. Map illustrating biodiversity study areas: Jing Xi County Nature Preserve, Shiwandashan National Nature Preserve, Kuan Kuoshui Nature Preserve, Dashahe Nature Preserve, and Shuipu (Maolan National Nature Preserve). elevation 900–1400 m, 2380701200N, forests of many large trees 20–30 m in 10585703600E, September–October height with some reaching ;50 m. 2004. The fieldwork was based along Site 2. Shiwandashan National Na- a rapid stream in secondary forest at ture Preserve, located in Guangxi Prov- 950 m. Old banana cultivation was ince, elevation 300–900 m, 2181304800N, evident with less disturbed ridge top 10785204800E, March–May 2005. Field- 216 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON work was based along a winding paved are divided among the collections of road, through steep mountains of the both authors. Nomenclature for birds preserve at 500 m. Secondary forests and mammals follow those of Inskipp et had maximum tree heights of ;25 m. al. (2001) and Wilson and Reeder Site 3. Kuan Kuoshui Nature Pre- (2005), respectively. All associated serve, located in Guizhou Province, hosts were collected and prepared as elevation 1450–1750 m, 2881304800N, museum specimens. Mammal speci- 10780903600E, March–May 2006. Field- mens are deposited at the Royal Ontario work was based in deciduous and Museum, Toronto, Canada, except for a evergreen forests with some trees subsample of specimens that remained achieving ;25 m. Valley floor and with local authorities in China. The surrounding hills were cultivated. specimens are deposited at the Kansas Streams drained into a small lake on Museum of Natural History, Lawrence, the valley floor. KS (identified herein by KU#), except Site 4. Dashahe Nature Preserve, for a subsample of specimens that located in Guizhou Province, elevation remained with local authorities in China 1350–1650 m, 2981001200N, (identified herein by BWB#, RLB#, or 10783401200E, March–May 2006. Field- SEB#). work was based in heavily disturbed natural vegetation at 1350 m. Maximum RESULTS AND DISCUSSION height of trees was ;15 m, secondary A total of 540 mammals representing growth surrounded the valley, much of 61 species (Table 1) and 1,222 birds which was agricultural plots of tobacco, representing 174 species were exam- etc. Secondary vegetation surrounded ined (Table 2). Eight species of fleas the valley and some native vegetation were recovered: two from bats, four lined a small river running through the from ground dwelling mammals, and steep karst landscape. two from birds. Only three of the 17 bat Site 5. Shuipu village, Libo County, genera represented in our study har- near Maolan National Nature Preserve, bored fleas. With the exception of located in Guizhou Province, elevation Myotis Kaup and Pipistrellus Kaup, 635–850 m, 2582900500N, 10785205400E, other genera examined are not known March–May 2007. Fieldwork was to harbor fleas in China, nor have any of based in and around Shuipu village in the Myotis and Pipistrellus species in gently sloping cultivated valley ap- our study been documented previously proximately 500 m wide and several as hosts of fleas. kilometers long. The valley was sur- Fleas were recovered from only five rounded by steep, rugged, karst forma- of 16 ground dwelling mammal species. tions. Perennial streams flowing into Of those five infested species, only 8 of valley were diverted to agricultural 85 specimens harbored fleas, and then fields. only in low numbers. These low Methods for the collection and pres- numbers were especially true at the ervation of ectoparasites were detailed three lowest elevation sites (Jing Xi, in Hastriter and Bush (2006) and Shiwandashan and Shuipu), where fleas Hastriter and Whiting (2003). Proce- were only recovered from three (4.5%) dures for preparing illustrations are of 45 positive host species (6.7%). Fleas outlined in Hastriter and Bush (2006). were more common on ground-dwell- DNA voucher specimens are deposited ing mammals at the two highest eleva- in the Brigham Young University flea tion sites (Kuan Kuoshui and Dashahe), collection and the remaining specimens where fleas were collected from 5 of 40 VOLUME 112, NUMBER 2 217

Table 1. Mammals examined for fleas from five field sites in southern China: Jing Xi (J.X.), Shiwandashan (Shiw.), Kuan Kuoshui (K.K.), Dashahe (Dash.), and Shuipu (Shui.). Numbers in parentheses indicate individual hosts that harbored fleas.

Order/Family Scientific Name J.X. Shiw. K.K. Dash. Shui. Total Fleas

Chiroptera Hipposideridae Aselliscus stoliczkanus 000033 Hipposideros armiger 204039 H. larvatus 0 1 0 0 29 30 H. pomona 300025 Megadermatidae Megaderma lyra 002046 Pteropodidae Cynopterus sphinx 20 7 0 0 0 27 Rousettus leschenaulti 16 (4) 0 0 0 0 16 *1 Sphaerias blanfordi 19 0 0 0 0 19 Rhinolophidae Rhinolophus affinis 26 19 0 0 0 45 R. lepidus 000022 R. luctus 010012 R. macrotis 100067 R. paradoxalophus 1 0 1 0 10 12 R. pearsonii 44911735 R. pusillus 1401814 R. rouxii 311921641 R. yunnanensis 5000611 Rhinolophus n. sp.001001 Vespertilionidae Eptesicus serotinus 000101 Harpiocephalus harpia 1 3 0 0 2 (1) 6 *2 Ia io 0 0 2 0 1 (1) 3 *2 Kerivoula hardwickii 103004 Miniopterus magnater 111003 Murina aenea 100001 M.aurata 610029 M. cyclotis 41 7 0 0 0 48 M. leucogaster 11 2 0 0 0 13 M. tubinaris 900009 Murina sp.000011 Myotis altarium 000011 M. daubentonii 000033 M. montivagus 060006 M. muricola 320207 M. mystacinus 000088 M. ricketti 003003 M. siligorensis 102104 Pipistrellus cadornae 100001 P. ceylonicus 000101 P. coromandra 010001 P. paterculus 120003 P. pulveratus 010001 Pipistrellus sp.100001 Scotomanes ornatus 040004 Vespertilio sinensis 000303 Erinaceomorpha Erinaceidae Neotetracus sinensis 000011 Rodentia Cricetidae Eothenomys melanogaster 002002 218 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued.

Order/Family Scientific Name J.X. Shiw. K.K. Dash. Shui. Total Fleas

Muridiae Chiropodomys gliroides 10 0 0 0 1 Leopoldamys edwardsi 00 0 0 1 1 Niviventer confucianus 2 12 (2)> 17 8 4 43 *3 N. fulvescens 9 11 3 0 4 (1)> 27 *3 N. montivagus 01 0 0 0 1 Rattus nitidus 01 0 0 1 2 R. tanezumi 00 0 0 1 1 Platacanthomyidae Typhlomys cinereus 00 0 0 1 1 Sciuridae Callosciurus swinhoei 10 0 0 0 1 Tamiops swinhoei 0 2 1 (1) 0 0 3 *4 Soricomorpha Soricidiae Anourosorex squamiceps 0 0 4 (2) 0 0 4 *5 Crocidura attenuata 13 0 0 3 7 C. fuliginosa 0 0 7 (2) 0 1 8 *6 C. horsfieldi 32 0 0 0 5 Crocidura sp.110002 Total species 32 26 17 9 29 61 Total individuals 197 110 71 30 132 540

1 Thaumapsylla breviceps. 2 Nycteridopsylla iae. 3 Aviostivalius klossi. 4 Macrostylophora jiangkouensis. 5 Palaeopsylla remota. 6 Palaeopsylla incurva. positive host species (12.5%). Similar Panama (Olson 1994), and Venezuela effects of elevation were also docu- (Janzen et al. 1976). Second, during mented in a survey of the Hengduan field-work in Papua New Guinea, the Mountains in southwestern China. senior author noted that ants were Gong et al. (2005) found that flea commonly collected from traps and diversity was low at 1000 m, but both nests of ground dwelling mammals species richness and abundance in- below 1,000 m elevation, where fleas creased with elevation up to 3800 m, were virtually absent. Third, in Bexar where flea diversity peaked. Minimal County, Texas, nests of Cotton Rats numbers of fleas at low elevations may (Sigmodon hispidus Say and Ord) are be explained by the presence of pred- typically infested by mites and fleas; atory ants. In areas where fleas and however, if fire ants (Solenopsis sp.) are predatory ants coexist, ants may reduce present, the nests are devoid of mites flea populations by feeding on their and all stages of fleas (pers. obs. senior susceptible immature stages. Several author, 1998). Although other factors facts support this hypothesis. First, may contribute to a dearth of fleas at tropical ant species diversity and abun- lower elevations, the potential impact of dance is known to decrease with predatory ants on flea populations increased elevation in tropical areas, should be more thoroughly investigat- as has been shown in Costa Rica ed. (Janzen 1973), India (Sabu et al. Two flea species from birds were 2008), Malaysia (Bru¨hl et al. 1998), found among 19 bird species that were VOLUME 112, NUMBER 2 219

Table 2. Birds examined for fleas from five field sites in southern China: Jing Xi (J.X.), Shiwandashan (Shiw.), Kuan Kuoshui (K.K.), Dashahe (Dash.), and Shuipu (Shui.). Numbers in parentheses indicate individual hosts that harbored fleas.

Order/Family Avian Species J.X. Shiw. K.K. Dash. Shui. Total Fleas

Apodiformes Apodidae Apus pacificus 00 04 04 Caprimulgiformes Caprimulgidae Caprimulgus indicus 12 00 14 Charadriiformes Scolopacidae Scolopax rusticola 10 00 01 Ciconiiformes Ardeidae Butorides striatus 10 00 01 Egretta garzetta 00 10 01 Gorsachius melanolophus 01 00 01 Columbiformes Columbidae Chalcophaps indica 32 00 05 Streptopelia chinensis 00 02 02 S. orientalis 00 10 01 Treron sieboldii 10 00 01 Coraciiformes Alcedinidae Alcedo hercules 10 00 01 A. atthis 00 00 44 Ceyx erithacus 01 00 01 Cuculiformes Cuculidae Clamator coromandus 01 00 01 Cuculus micropterus 00 10 01 Hierococcyx fugax 01 00 01 Phaenicophaeus tristis 01 00 01 Falconiformes Accipitridae Accipiter virgatus 10 00 01 Gruiformes Rallidae Amaurornis phoenicurus 00 00 11 Passeriformes Aegithalidae Aegithalos concinnus 0 0 0 3 12 15 Aegithinidae Chloropsis hardwickii 10 00 23 Campephagidae Coracina macei 20 00 02 C. melaschistos 30 10 04 Hemipus picatus 11 00 02 Pericrocotus divaricatus 02 00 02 P. ethologous 00 02 02 P. flammeus 14 00 27 P. solaris 00 00 22 Cinclidae Cinclus pallasii 00 05 05 Corvidae Dendrocitta formosae 10 00 01 Garrulus glandarius 00 10 01 Urocissa erythrorhyncha 00 22 04 Dicaeidae Dicaeum concolor 05 00 05 Dicruridae Dicrurus hottentottus 81 00 09 D. leucophaeus 51 00 06 Emberizidae Emberiza elegans 0 0 11 5 0 16 E. godlewskii 00 01 01 E. pusilla 00 71 715 E. spodocephala 0 0 4 2 (1) 7 13 *1 220 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Continued.

Order/Family Avian Species J.X. Shiw. K.K. Dash. Shui. Total Fleas

E. tristrami 000 0 11 Latoucheornis siemsseni 000 0 22 Melophus lathami 000 0 44 Estrilididae Lonchura striata 0 0 0 0 14 14 Fringillidae Carduelis sinica 001 0 01 Carpodacus erythrinus 000 0 66 Hirundinidae Hirundo daurica 000 0 55 Hirundo rustica 000 0 11 Laniidae Lanius schach 000 0 11 Monarchidae Hypothymis azurea 010 0 01 Rhipidura albicollis 600 0 06 Terpsiphone paradisi 1100 0 011 Motacillidae Anthus hodgsoni 1 0 0 2 10 13 Motacilla alba 200 0 24 M. cinerea 100 4 05 Muscicapidae Copsychus saularis 001 0 01 Culicicapa ceylonensis 9 1 4 0 16 30 Cyanoptila cyanomelana 410 0 05 Cyornis banyumas 12 7 0 0 4 23 C. hainanus 2 5 3 (1) 0 1 11 *1 C. rubeculoides 100 0 01 Enicurus leschenaulti 1 0 3 (1) 0 2 6 *1 E. schistaceus 450 0 413 Eumyias thalassina 301 1 05 Ficedula hyperythra 110 0 02 F. mugimaki 000 0 11 F. parva 110 0 24 F. tricolor 000 0 22 F. zanthopygia 020 0 02 Luscinia cyane 300 0 03 L. sibilans 000 0 11 Muscicapa ferruginea 300 0 03 M. muttui 430 4 112 M. sibirica 100 0 01 Myiomela leucura 2 2 3 (2) 0 0 7 *1 Myophonus caeruleus 1 0 1 (1) 0 4 6 *1 davidi 0 0 7 (1) 1 10 18 *1 N. grandis 700 0 07 N. macgrigoriae 18 2 0 0 0 20 Phoenicurus auroreus 002 1 03 Rhinomyias brunneatus 210 0 03 Rhyacornis fuliginosus 0 2 0 11 (1) 0 13 *1 Saxicola ferrea 100 1 13 S. torquata 000 0 11 Zoothera citrina 470 0 516 Z. dauma 000 0 33 Z. sibirica 200 0 02 Zoothera sp.100001 Nectariniidae Aethopyga christinae 070 0 411 A. gouldiae 001 1 02 Arachnothera magna 110 0 02 VOLUME 112, NUMBER 2 221

Table 2. Continued.

Order/Family Avian Species J.X. Shiw. K.K. Dash. Shui. Total Fleas

Oriolidae Oriolus chinensis 010 0 01 Panuridae Paradoxornis alphonsianus 004 0 15 P. gularis 131 0 05 P. verreauxi 007 0 07 P. webbianus 0 0 0 2 (2) 5 7 *1 Paridae Melanochlora sultanea 100 0 01 Parus major 002 0 46 P. monticolus 0 0 16 (1) 2 0 18 *1 P. spilonotus 000 0 11 P. venustulus 000 0 77 Parus sp.100001 Sylviparus modestus 002 0 02 Passeridae Passer rutilans 0 0 1 1 12 14 Pycnonotidae flaveolus 600 0 06 Alophoixus pallidus 380 0 011 Hemixos castanonotus 2 9 0 0 11 22 leucocephalus 020 0 13 H. mcclellandii 10 4 3 2 4 23 Pycnonotus jocosus 150 0 06 P. sinensis 020 0 35 P. xanthorrhous 0 0 1 2 12 15 Spizixos semitorques 0 0 0 6 16 22 Sittidae Sitta frontalis 100 0 01 Sturnidae Acridotheres cristatellus 000 1 01 Cettia fortipes 103 0 15 Orthotomus cucullatus 110 0 02 O. sutorius 010 0 12 Phylloscopus spp. 18 12 6 10 13 59 Prinia atrogularis 020 0 24 P. inornata 000 0 11 Seicercus spp. 16 1 13 11 5 46 Tesia cyaniventer 500 0 05 Urosphena squameiceps 500 0 05 Timaliidae Alcippe chrysotis 008 1 09 A. dubia 000 0 11 A. morrisonia 18 5 20 4 (1) 36 83 *1 Babax lanceolatus 0 0 0 5 (1) 0 5 *2 Garrulax canorus 010 0 01 G. chinensis 210 0 03 G. cineraceus 0 0 5 (1) 0 4 9 *1 G. maesi 240 0 06 G. milnei 2 0 11 (2) 0 0 13 *1 G. ocellatus 0 0 0 1 (1) 0 1 *1 G. pectoralis 002 0 02 G. poecilorhynchus 0 0 3 (2) 0 0 3 *1 G. sannio 003 0 14 Leiothrix argentarus 030 0 03 L. lutea 4 0 22 (4) 21 (1) 2 49 *1 Liocichla phoenicea 100 0 01 Minla cyanouroptera 006 1 07 M. ignotincta 001 2 03 222 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Continued.

Order/Family Avian Species J.X. Shiw. K.K. Dash. Shui. Total Fleas

Napothera brevicaudata 320 0 0 5 albiventre 320 0 0 5 P. tickelli 100 0 0 1 Pomatorhinus ruficollis 7 6 9 (2) 6 (1) 14 42 *1 Pteruthius flaviscapis 500 0 0 5 Spelaeornis chocolatinus 400 0 0 4 Stachyris nigriceps 11 0 0 0 0 11 S. ruficeps 1 10 9 (2) 1 (1) 17 38 *1 S. striolata 17 2 0 0 0 19 Stachyris sp.10000010 Yuhina castaniceps 060 0 410 Y. diademata 001 0 0 1 Y. nigrimenta 302 1 1 7 Y. zantholeuca 430 0 2 9 Turdidae Turdus dissimilis 000 0 2 2 Turdus sp.100001 Zosteropidae Zosterops japonicus 0 0 2 0 12 14 Capitonidae Megalaima asiatica 020 0 0 2 Megalaima franklinii 030 0 0 3 Megalaima virens 100 0 0 1 Piciformes Picidae Blythipicus pyrrhotis 100 0 0 1 Dendrocopos canicapillus 001 0 1 2 D. hyperythrus 100 0 0 1 Picumnus innominatus 001 1 5 7 Picus canus 0 0 2 (1) 0 0 2 *1 Sasia ochracea 550 0 515 Podicipediformes Podicipedidae Tachybaptus ruficollis 001 0 0 1 Stigiformes Strigidae Glaucidium brodiei 100 0 1 2 G. cuculoides 100 0 2 3 Ninox scutulata 000 0 1 1 Otus letti 100 0 0 1 O. spilocephalus 300 0 0 3 O. sunia 000 0 2 2 Trogoniformes Trogonidae Harpactes erythrocephalus 200 0 0 2 Total species 87 59 38 30 73 174 Total individuals 317 188 224 134 359 1222

1 Lentistivalius insolli. 2 Dasypsyllus gallinulae gallinulae.

infested. Of particular interest among Ceratophyllidae the two bird fleas reported herein was Dasypsyllus gallinulae gallinulae (Dale) Lentistivalius insolli (Traub 1950), a pygiopsyllid bird flea, which was wide- Material examined.—Dashahe: ex ly distributed on passerine birds. Babax lanceolatus (Verreaux) (BWB- VOLUME 112, NUMBER 2 223

975) (1? DNA voucher F-272), 29 IV the original description and to further 06, S.E. Bush. validate the identification of this taxon. Remarks.—This flea occurs com- monly in Europe on many species of Ctenophthalmidae birds. This bird flea virtually parasitizes Palaeopsylla incurva Jordan any bird (or mammal) for which it comes into contact. The species is less Material examined.—Kuan Kuoshui: documented in Eastern Asia but has ex Crocidura dracula ¼ Crocidura been reported in China, Japan, the fuliginosa (Blyth) (Southeast Asian Philippine Islands and the Himalayas Shrew), F-47633, F-47642, 25, 27 IV (Sikkim) (Haddow et al. 1983). 06, S.E. Bush and D.H. Clayton (1? DNA voucher F-271, BYU; 1?). Macrostylophora jiangkouensis Li Remarks.—Palaeopsylla incurva has and Huang been collected infrequently and the (Figs. 2A–E) preferred host is uncertain. Guo et al. (2000) found it statistically in very low Material examined.—Kuan Kuoshui: numbers on Crocidura attenuata Milne- ex Tamiops swinhoei (Milne-Edwards) Edwards in neighboring Yunnan Prov- (Swinhoe’s Striped Squirrel), F-47640, ince. If the host preference of this flea is 26 IV 06, S.E. Bush and D.H. Clayton to be determined, future collections (7?,3/). might focus on the nests of various species of Crocidura in the known Remarks.—The ranges from range of P. incurva (northern Myanmar Pakistan through China, Indonesia, and and the southern Chinese Provinces of the Philippine Islands occurring primarily Yunnan and Guangxi). on squirrels, especially members of the Genus Callosciurus Gray. Li and Traub (1998) erected the subgenus Songshu- Palaeopsylla remota Jordan psylla, and assigned M. jiangkouensis to the euteles group (southern China) of that Materials examined.—Kuan Kuosh- subgenus, opposed to the hebeiensis ui: ex Anourosorex squamipes Milne- group (northern China). Specialists con- Edwards (Chinese Mole Shrew) F- tinue to describe new species and subspe- 47579, F-47587, 16, 18 IV 06, S.E. cies without developing and publishing Bush and D.H. Clayton (4?,2/). identification keys. The genus has 44 Remarks.—It is not surprising to find species including 12 subspecies and is in P. remota on A. squamipes. Hopkins need of revision. Within the genus Macro- and Rothschild (1966) reported this flea stylophora, the anatomy of the aedeagus from northeastern India and Myanmar provides important characters on which to from A. squamipes and Guo et al. distinguish species. Combined with both (2000) found A. squamipes to be the the complexity and the difficulty of dominant host of P. remota in their viewing the aedeagus (often hidden study in Yunnan Province, China. beneath other structures), it is often omitted from discussion or illustration. Ischnopsyllidae (Ischnopsyllinae) Such is the case with M. jiangkouensis. Nycteridopsylla iae Beaucournu and Illustrations (Figs. 2A–E) of the male Kock genitaliaof M.jiangkouensisare therefore provided to supplement those included in Material examined.—Shuipu: ex Har- 224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Macrostylophora jiangkouensis, male. A. Aedeagus. B. Apex of aedeagus. C. Tergite eight and clasper (basimere and telomere). D. Sternites seven and eight, ventral view. E. Proximal and distal arms of ninth sternite, ventral view. Scale A, C–E ¼ 200 l,B¼ 100 l. piocephalus harpia (Temminck) (Hairy- Remarks.—This flea has been col- winged Bat, F-47763), 2 IV 07, S.E. lected exclusively on I. io with the Bush (1/); same data except ex Ia io exception of this sole record on H. Thomas (F-47871), 15 IV 07 (1?,2/). harpia (see Hastriter et al. 2009). VOLUME 112, NUMBER 2 225

Thaumapsyllinae 97548, 25 IV 06 (2?,6/); ex Garrulax Thaumapsylla breviceps Rothschild cineraceus (Godwin-Austen) (Mous- tached Laughing Thrush), KU-97590, Material examined.—Jing Xi: ex 22 IV 06 (1?,1/); ex Garrulax milnei, Rousettus leschenaulti (Desmarest) (Le- (David) (Red-Tailed Laughing Thrush), schenault’s Rousette) F-47412, 6 X 04, KU-97564, 18 IV 06, S.E. Bush and S.E. Bush (3/, DNA voucher, BYU; D.H. Clayton (1?,1/); ex G. milnei, 1?,1/); same data except ex R. KU-97559, 23 IV 06 (1?); ex Garrulax leschenaulti, F-47437, F-47442, F- poecilerhynchus (Rusty Laughing 47443, 8 X 04 (1?,2/). Thrush), KU-97553, 20 IV 06 (1/); Remarks.—A common parasite of ex G. poecilerhynchus, KU-97522, 23 fruit bats of the genus Rousettus Gray IV 2006 (2?); ex Leiothrix lutea (Pteropidae), it is restricted to tropical (Scopoli) (Red-Billed Leiothrix), SEB- and semitropical habitats of South 510, SEB-5112, 19 IV 06 (2/); ex L. Africa, Southeast Asia and Wallacea. lutea, BWB-922, 21 IV 06 (1?); ex L Beaucournu and Kock (1994) provided lutea, RLB-867, 22 IV 06 (1/); ex excellent pictorial details of the distri- Myiomela leucura (Hodgson) (White- bution of this species. This flea record Tailed Robin), KU-97551, 21 IV 06 represents the northern fringe of the (2/); ex M. leucura, KU-97550, 27 IV species and the authors are not aware of 2006 (1?,1/); ex Myophonus caer- any records of T. breviceps heretofore uleus (Scopoli) (Blue-Whistling reported from China. Thrush), KU-97521, 23 IV 06 (1?); ex Niltava davidi (Fujian Niltava), SEB-518, 24 IV 06 (1?); ex Parus Pygiopsyllidae monticolus Vigors (Green-Backed Tit), Aviostivalius klossi (Jordan and KU-97597, 25 IV 06 (1?); ex Picus Rothschild) canus J.F. Gmelin (Gray-Headed Wood- pecker), KU-97408, 21 IV 06 (2/); ex Material examined.—Shiwandashan: Pomatorhinus ruficollis (Streak-Breast- ex Niviventer confucianus (Milne-Ed- ed Scimitar Babbler), KU-97577, 20 IV wards) (Chinese White-bellied Rat), F- 06 (2?); ex P. ruficollis, KU-97578, 23 ? 47469, 18 IV 05, S.E. Bush (1 ,DNA IV 06 (1/); ex Stachyris ruficeps Blyth voucher F-255, BYU; 1?); same data (Rufous-Capped Babbler), KU-97454, except ex N. confucianus, (F-47548), 30 20 IV 06 (1?); ex S. ruficeps, KU- IV 05 (1/). Shuipu: ex Niviventer 97602, 24 IV 06 (1/); ex M. leucura, fulvescens (Gray) (F-47774), 20 II 07, 27 IV 2006 (1?,1/). Dashahe: ex S.E. Bush (1?DNA voucher F-280, Alcippe morrisonia Swinhoe (Gray- BYU). Cheeked ), BWB-985, 30 IV Remarks.—A common lowland spe- 06 (2/); ex Emberiza spodocephala cies wide spread throughout Southeast (Black-Faced Bunting), BWB-994, 1 V Asia on murid . 06 (1/); ex Garrulax ocellatus (Vigors) (Spotted Laughing Thrush), KU-97504, Lentistivalius insolli (Traub) 5 V 06 (1?); ex L. lutea, SEB-534, 1 V 06 (1/); ex Paradoxornis webbianus Material examined.—Kuan Kuoshui: (Gould) (Vinous-Throated Parrot Bill), ex Cyornis hainanus (Ogilvie-Grant) KU-97482, 1 V 06 (1/); ex P. webbia- (Hainan Blue-Fly Catcher) KU-97520, nus, KU-97483, 2 V 2006 (1?); ex 24 IV 06 (1/); ex Enicurus lesche- Rhyacornis fuliginosus (Vigors) naultii (White-Crowned Fork-Tail) KU- (Plumbeous Water-Redstart), KU- 226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

97479, 1 V 06 (1/); ex P. ruficollis, (Timaliidae). The flea infested 13 KU-97513, 5 V 06 (2/); and ex S. (12.7%) of the individual timalid hosts ruficeps, SEB-567, 5 V 06 (1?). collected at Kuan Kuoshui and 5 Remarks.—Traub (1950) described (11.6%) of the individuals collected at L. insolli from a large series collected Dashahe. Of the 13 genera of Timalii- from three bird nests in the Malaysian dae sampled, five (38.5%) harbored this Cameron Highlands. Mardon (1981) flea. Lentistivalius insolli was found on reported a single male from a human only one non-passerine species: the at Fraser’s Hill (a resort town near the Grey-Headed Woodpecker (Picus Cameron Highlands, Pahang State, Ma- canus). This bird occupies and nests in laysia). A single specimen (representing tree cavities that may be frequented by the same specimen reported in each passerine birds. It is undoubtedly an report) was subsequently reported from accidental host and probably plays little Da Lat, South by Suntsov and or no role in the propagation of L. Suntsova (1999), Beaucournu and insolli. In the future, examining the Sountsov (1999), and Adler et al. nests of passerine birds may be far more (2001). The latter two identified the productive in collecting this flea than host as the Chestnut White-bellied Rat attempting to collect them from the (Niviventer fulvescens). Traub’s data body of their hosts, where the fleas from the type series indicated that L. occur in low prevalence and abundance. insolli was a bird flea, but a bird species Additional sampling of birds and nests was not associated with the sizable in Southeast Asia is necessary to series taken from the three ‘‘small bird elucidate flea populations and distribu- nests’’ (Traub 1950, 1972). tion of L. insolli on passerine bird Our collections definitively associate species. Because of the low infestation this flea with birds. The current collec- rate, many passerine bird species that tion of 18 males and 28 females of have not been found to harbor L insolli Lentistivalius insolli were collected may be found in the future to harbor from 30 individual birds represented this flea. Elevation may be a limiting by 18 species and 6 avian families factor in the distribution of this species (Passeriformes: Emberizidae, Muscica- since all collections of L. insolli oc- pidae, Panuridae, Paridae, Timaliidae, curred at elevations above 1350 m, and Piciformes: Picidae). This flea was (none occurred on passerine birds at found almost exclusively on lower elevation study sites). at the two sites (Kuan Kuoshui and There are five other species within Dashahe) where it was collected. At the genus Lentistivalius Traub. Two are these sites, the flea occurred at a low parasites of ground dwelling soricids [L. prevalence and low intensity. Among occidentayunnanus Li, Xie and Gong, the passerines at these sites, L. insolli China and L. ferinus (Rothschild), occurred on 29 (8.5%)ofthe343 India, Nepal, and Sri Lanka], two of passerines collected (Table 2). Rarely non-arboreal murine rodents [L. alienus was there more than one flea for any Smit, Africa and L. aestivalis (Jameson infested host specimen. Ninety-seven and Sakiguti), Japan], and one of percent of infested hosts had only one scansorial tree shrews of the genus or two fleas; however, one bird, a Tupaia Raffles (L. vomerus Traub, White-Crowned Fork-Tail (Enicurus Borneo and Sarawak). Lentistivalius leschenaulti) was infested with eight vomerus Traub is remarkably similar fleas. Lentistivalius insolli was fre- to L. insolli. Their close morphological quently found on Old World Babblers affinity would suggest that L. insolli VOLUME 112, NUMBER 2 227 might have been derived from L. les Ischnopsyllidae de la Re´gion Orientale, vomerus and spread through Malaysia II. Stations ine´dites et description d’une to southern China on highly mobile espe`ce nouvelle du genre Lagaropsylla Jordan and Rothschild 1921 (Insecta: Si- volant avian hosts, while L. vomerus phonaptera). Senckenbergiana biologica remained isolated in Borneo and con- 73(1–2): 67–75. tiguous Sarawak. Lentistivalius insolli Beaucournu, J. C. and V. V. Sountsov. 1999. has the broadest distribution of all Puces nouvelles ou peu connues du Vietnam. species of Lentistivalius and likely Ctenophthalmidae et Pygiopsyllidae (Si- occurs on passerine birds in all coun- phonaptera). Bulletin de la Socie´te´ entomo- logique de France 104(1): 77–83. tries lying between Malaysia and south- Bru¨hl, C. A., G. Gusalam, and K. E. Linsenmair. ern China (Cambodia, Thailand, Re- 1998. Stratification of ants in a rain forest of public of , Myanmar, and Borneo. Journal of Tropical Ecology 14: Vietnam). Its lack of preference for 285–297. any one avian host species, an almost Gong, Z., H. Wu, X. Duan, X. Feng, Y. Zhang, exclusive preference for passerine birds, and Q. Liu. 2005. Species richness and vertical distribution pattern of flea fauna in the distinction of the only avian parasite Hengduan Mountains of western Yunnan, in the genus, and the lack of morpho- China. Biodiversity Science 13: 279–289. logical differentiation of characters in Guo, X. G., Z. D. Gong, T. J. Qian, X. G. Feng, disjunct populations from China and X. D. Duan, W. Li, and X. K. Zhang. 2000. those from Malaysia might suggest a fauna investigation in some foci of rather recent switch from mammals to human plague in Yunnan, China. Acta birds. Zootaxonomica Sinica 25(3): 291–297. Haddow, J., R. Traub, and M. Rothschild. 1983. Distribution of ceratophyllid fleas and notes ACKNOWLEDGMENTS on their hosts, pp. 42–163. In R. Traub, M. Rothschild, and J. F. Haddow, eds. The We thank Brett W. Benz, Roger L. Rothschild Collection of Fleas. The Cerato- Boyd, Guojun Chen, Dale H. Clayton, phyllidae: Key to the Genera and Host Tristan J. Davis, Judith Eger, Burton Relationships. University Press, Cambridge. Lim, A. Townsend Peterson, Arpad Hastriter, M. W. and S. E. Bush. 2006. Maabella Nyari and Mark B. Robbins for various gen. nov. (Streblidae: Ascodipterinae) from forms of assistance, including the Guangxi Province, China and Vietnam with notes on preservation of Ascodipterinae. collection of important specimens. For Zootaxa 1176: 27–40. the continuous support of M. F. Whit- Hastriter, M. W., S. E. Bush, K. Dittmar, S. S. ing, Curator of and the staff of Hla Bu, and M. F. Whiting. 2009. Nycter- the Monte L. Bean Life Science Muse- idopsylla quadrispina Lu and Wu, a junior um for providing space, equipment, and synonym of Nycteridopsylla iae Beaucournu supplies needed to continue work on and Kock (Siphonaptera: Ischnopsyllidae) as fleas, we are indebted. This work was determined by morphological and DNA analyses. 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