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Journal of Medical Entomology, XX(X), 2020, 1–22 doi: 10.1093/jme/tjaa206 Morphology, Systematics, Evolution Research

Eight New Species of Sucking Lice (Psocodea: Phthiraptera) From Endemic Murine Rodents in Australia and an Updated Identification Key Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020

Wei Wang,1, Lance A Durden,2 Haylee Weaver,3 and Renfu Shao1,4

1GeneCology Research Centre, School of Science and Engineering, University of the Sunshine Coast, Sippy Downs, Queensland AADate 4556, Australia, 2Department of Biology, Georgia Southern University, Statesboro, GA 30458, 3Australian Biological Resources Study, Department of Agriculture, Water and the Environment, GPO Box 787, Canberra ACT 2601, Australia, and 4Corresponding author, e-mail: [email protected] AAMonth Subject Editor: Richard Wilkerson

AAYear Received 12 July 2020; Editorial decision 27 August 2020

Abstract Based on a comprehensive study of museum specimens, eight new species of sucking lice of the genus Hoplopleura Enderlein, 1904 (Psocodea: Phthiraptera: Hoplopleuridae), are described from six genera of Australian Old Endemic rodents: Conilurus Ogilby, 1838 (Rodentia: Muridae), LeggadinaThomas, 1910 (Rodentia: Muridae), Leporillus Thomas, 1906 (Rodentia: Muridae), Mesembriomys Palmer, 1906 (Rodentia: Muridae), Pogonomys Milne-Edwards, 1877 (Rodentia: Muridae), and Xeromys Thomas, 1889 (Rodentia: Muridae). The description of these new species increases the number of sucking louse species from endemic Australian rodents from 13 to 21 and extends the records of sucking lice to all of the 14 genera of endemic rodents in Australia. Our results show that sucking lice are much more diverse among rodents in Australia than previously known. Furthermore, the Australian Hoplopleura species are host specific—eachHoplopleura species, including the eight new species described in the present study, parasitizes only a single host species, except Hoplopleura irritans Kuhn and Ludwig, 1967 (Psocodea: Phthiraptera: Hoplopleuridae) and Hoplopleura melomydis Weaver, 2017 (Psocodea: Phthiraptera: Hoplopleuridae), each of which is found on two host species. An updated dichotomous key for identifying Australian Hoplopleura species is included.

Key words: Phthiraptera, Anoplura, Hoplopleura identification key, endemic murine rodents, new species

Sucking lice (Psocodea: Phthiraptera: Anoplura) are obligate wingless and Carleton 2005, Van Dyck and Strahan 2008). All of these species ectoparasites of eutherian mammals (Kim and Ludwig 1978) with are in the Murinae, one of the five subfamilies of Muridae (Musser piercing, sucking mouthparts and feed entirely on the blood of their and Carleton 2005, Breed and Ford 2007, Van Dyck and Strahan hosts. To date, more than 540 species of sucking lice have been de- 2008). Murine rodents colonized the Australian continent in two scribed from 840 species of eutherian mammals worldwide (Durden major migration waves referred to as Old and New Endemics. The and Musser 1994a). Sucking lice have high host specificity com- Old Endemics entered Australia 5–8 million years ago (MYA) and pared with most other ectoparasites: more than half of the recorded diversified into 56 species in 13 genera (including 10 extinct spe- sucking louse species are found only on a single host species (Durden cies). Fifty of the Old Endemic species are found only in Australia and Musser 1994a, b). Hoplopleura Enderlein, 1904 (Psocodea: but Conilurus penicillatus (Gould, 1842) (Rodentia: Muridae), Phthiraptera: Hoplopleuridae) is the largest genus of Anoplura with Hydromys chrysogaster Geoffroy, 1804 (Rodentia: Muridae), more than 140 species (Durden and Musser 1994a, Light et al. 2010) Melomys burtoni (Ramsay, 1887) (Rodentia: Muridae), Pogonomys and has been recorded from rodents (Rodentia) worldwide and from mollipilosus Peters and Doria, 1881 (Rodentia: Muridae), Uromys pikas (Lagomorpha: Ochotonidae) in the Palearctic region (Durden caudimaculatus (Krefft, 1867) (Rodentia: Muridae), and Xeromys and Musser 1994a,b). Biogeographically isolated by oceans, the myoides Thomas, 1889 (Rodentia: Muridae) are also found in New Australian continent has a distinct endemic fauna of rodents (Tate Guinea (H. chrysogaster and subspecies of U. caudimaculatus occur 1951, Watts and Aslin 1981). Sixty-three rodent species (including on some Indonesian islands as well). The New Endemics arrived in extinct species) have been recognized as endemic to Australia (Musser Australia 1–2 MYA and diversified into seven species, all in the genus

© The Author(s) 2020. Published by Oxford University Press on behalf of Entomological Society of America. 1 All rights reserved. For permissions, please e-mail: [email protected]. Version of Record, first published online November 12, 2020 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN. 2 Journal of Medical Entomology, 2020, Vol. XX, No. XX

Rattus Fischer de Waldheim, 1803 (Rodentia: Muridae) (Musser and under a dissecting microscope (Nikon SMZ800N); the lice col- Carleton 2005, Breed and Ford 2007, Van Dyck and Strahan 2008). lected were kept in 2-ml screw-cap vials (one vial per host specimen) Five species of New Endemics are found only in Australia; the other filled with ethanol (80%) and stored in −20°C freezer until further two species, Rattus leucopus (Gray, 1867) (Rodentia: Muridae) and examination. Rattus sordidus (Gould, 1858) (Rodentia: Muridae) are also found on New Guinea and on some Indonesian islands. Morphological Examination Prior to the present study, only 13 louse species had been re- Louse specimens were mounted on microscope slides in Canada corded from 15 endemic rodent species in Australia (Neumann balsam as described in detail in Wang et al. (2020). Intact louse spe- 1909; Johnson 1960; Kuhn and Ludwig 1966; Kim 1972; Palma cimens with minimal gut contents were selected for slide mounting. and Barker 1996; Weaver and Barton 2008; Weaver 2017; Wang The specimens were cleared and softened in KOH (20%) for 24–48 h Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 et al. 2018, 2020). All of these species are in the genus Hoplopleura at room temperature, sequentially immersed in distilled water for 30 except for the introduced spiny rat louse, Polyplax spinulosa mins, and then in acetic acid (10%) for 30 mins to neutralize re- (Burmeister, 1839) (Psocodea: Phthiraptera: Polyplacidae), which sidual KOH. The specimens were then stained in acid fuchsin (1%) has been found on six endemic rodent species and two introduced for 6–12 h, and dehydrated in ethanol of different concentrations rodent species in Australia (Palma and Barker 1996; Wang et al. (40, 70, and 100%), each for 30 min. The specimens were clari- 2018, 2020). Twelve Hoplopleura species have been described from fied with pure clove oil for 24–48 h, and finally mounted on slides 13 endemic rodent species in Australia. Each Hoplopleura spe- using Canada balsam. Specimens mounted on slides were examined cies is found on a single host species except for H. irritans, Kuhn and measured with a photomicroscope (Nikon ECLIPSE Ts 2); all and Ludwig, 1967 (Psocodea: Phthiraptera: Hoplopleuridae) measurements are in micrometers (range followed by the mean). and H. melomydis, Weaver 2017 (Psocodea: Phthiraptera: Descriptive format and abbreviations of morphological features Hoplopleuridae), each of which is found on two endemic rodent spe- follow Kim et al. (1986) and Durden et al. (2018, 2019), with full cies: H. irritans on Rattus fuscipes (Waterhouse, 1839) (Rodentia: names of setae spelled out in full at first mention (Kim et al. 1986; Muridae) and Rattus lutreolus (Gray, 1841) (Rodentia: Muridae), Durden et al. 2018, 2019). We collected both adult and nymphal and H. melomydis on Melomys burtoni (Ramsay, 1887) (Rodentia: louse specimens, but we mounted only adult specimens on micro- Muridae) and Melomys capensis, Tate 1951 (Rodentia: Muridae) scope slides for detailed morphological examination. Following (Kuhn and Ludwig 1966, Weaver 2017). These 12 species are widely accepted museum preservation practices, the host rodents known from 8 of the 14 endemic rodent genera: Hydromys É. had been immersed in formalin prior to being stored in ethanol, so Geoffroy, 1804 (Rodentia: Muridae), Mastacomys Thomas, 1882 it was not possible to extract DNA from louse specimens for genetic (Rodentia: Muridae), Melomys Thomas, 1922 (Rodentia: Muridae), comparisons. Notomys Lesson, 1842 (Rodentia: Muridae), Pseudomys Gray, 1832 (Rodentia: Muridae), Uromys Peters, 1867 (Rodentia: Muridae), Zyzomys Thomas, 1909 (Rodentia: Muridae), and Results Rattus (Neumann 1909, Johnson 1960, Kuhn and Ludwig 1966, Hoplopleura Specimens Collected From Endemic Kim 1972, Weaver and Barton 2008, Weaver 2017, Wang et al. Rodents in Australia 2018). Until this study, sucking lice were unknown from the other We examined 162 endemic rodent specimens of eight spe- six endemic genera: Conilurus Ogilby, 1838 (Rodentia: Muridae), cies from six genera: Conilurus penicillatus (n = 22), Leggadina Leggadina Thomas, 1910 (Rodentia: Muridae), Leporillus Thomas, forresti (Thomas, 1906) (Rodentia: Muridae) (n = 45), Leggadina 1906 (Rodentia: Muridae), Mesembriomys Palmer, 1906 (Rodentia: lakedownensis Watts, 1976 (Rodentia: Muridae) (n = 36), Leporillus Muridae), Pogonomys Milne-Edwards, 1877 (Rodentia: Muridae), conditor (Sturt, 1848) (Rodentia: Muridae) (n = 5), Mesembriomys and Xeromys Thomas, 1889 (Rodentia: Muridae). In this study, we gouldii (Gray, 1843) (Rodentia: Muridae) (n = 10), Mesembriomys comprehensively examined rodent specimens in museums across macrurus (Peters, 1876) (Rodentia: Muridae) (n = 20), Pogonomys Australia for the presence of sucking lice. We describe eight new mollipilosus (n = 9), and Xeromys myoides (n = 15) (Table 1). We Hoplopleura species from the six genera of endemic rodents in collected 412 louse specimens in total, both adults and nymphs, from Australia from which sucking lice had not previously been recorded. these rodents (Table 2). The prevalence of lice on the rodents we examined ranged from 15% on M. macrurus to 20% on X. myoides, Materials and Methods 22.2% on P. mollipilosus, 27.3% on C. penicillatus, 30.6% on L. lakedownensis, 33.3 % on L. forresti, and 60% on L. conditor. Sample Collection For the rodent specimens that had lice, the abundance ranged from In order to investigate the biodiversity of sucking lice that parasi- 1.7 lice per rodent (average) on X. myoides to 5 on P. mollipilosus, tize native rodents in Australia, we examined ethanol-preserved 5.7 on M. macturus and L. lakedownensis, 8.6 on L. forresti, 12.3 rodent specimens in five museums across Australia (Queensland on C. penicillatus, 15 on L. conditor, and 23 on M. gouldii. Based Museum, QM; Museums Victoria, MV; Museum and Art Gallery of on the morphological features of adult lice, we identified these speci- the Northern Territory, MAGNT; South Australia Museum, SAM; mens as eight new species in the genus Hoplopleura. Western Australian Museum, WAM). As described in detail in Wang et al. (2020), individual rodent specimens were placed on a tray and the rodent pelage was gently and thoroughly combed with a comb Nomenclature designed for human head louse removal. The rodent was then trans- This paper and the nomenclatural act(s) it contains have been reg- ferred into a jar filled with 80% ethanol. The jar was sealed and was istered in Zoobank (www.zoobank.org), the official register of the gently shaken for ~1 min to dislodge lice from the host. The rodent International Commission on Zoological Nomenclature. The LSID was removed from the jar and the ethanol was filtered through a fine (Life Science Identifier) number of the publication is: urn:lsid:zoobank. mesh gauze. The filtered solid residue on the gauze was examined org:pub:8EE9E14C-1942-4B6B-9C5A-A25F90B3C265. Journal of Medical Entomology, 2020, Vol. XX, No. XX 3

Table 1. Collection of new species of Hoplopleura lice from murine rodent specimens in five museums across Australia

Rodent species (n = 8) Rodents sampled (n = 162) Rodents with Hoplopleura lice Hoplopleura lice collected

QMa MV WAM SAM MAGNT Adults Nymphs

Conilurus penicillatus -b - 4 7 11 6 24 50 Leggadina forresti 20 25 - - - 15 33 96 Leggadina lakedownensis 13 - 10 - 13 11 37 26 Leporillus conditor - 1 - 3 1 3 29 16 Mesembriomys gouldii 3 6 - - 1 3 26 43 Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 Mesembriomys macrurus - 1 12 4 3 3 10 7 Pogonomys mollipilosus 9 - - - - 2 6 4 Xeromys myoides 14 - - - 1 3 2 3 Total 59 33 26 14 30 46 167 245

aQM: Queensland Museum, Brisbane; MV: Museums Victoria, Melbourne; MAGNT: Museum and Art Gallery of the Northern Territory, Darwin; SAM: South Australian Museum, Adelaide; WAM: Western Australian Museum, Perth. bHyphen ‘-’ indicates no specimen was checked or collected.

Taxonomy pairs of StAS equal in length. Sternites 4, 6, 7 with 7 StAS. Sternites Family Hoplopleuridae Ewing, 1929 8–10 each with 3 pairs of StAS. Sternite 11 with 2 pairs of StAS and Genus Hoplopleura Enderlein, 1904 sternite 12 with 1 pair of StAS. One pair of ventral lateral abdominal Hoplopleura conilurudis Wang n. sp. setae (VLAS) adjacent to each of sternites 6, 7, 9, 11. (urn:lsid:zoobank.org:act:08920E3F-EED6-497E-AD8C- Paratergites: Paratergal plates (Fig. 1c) present on abdominal seg- 9D4AB70C7287). ments 1–8. All plates differentially sclerotized. Paratergal plate Male: [n = 6; Fig. 1a] Body length 911–1045 (983). I small and offset medially. Paratergal plates II–V each with 2 pos- Head: Head slightly longer than wide. Pre-antennal region short with terior lobes. Paratergal plate II with 1 small medial seta, 2 large pointed antero-lateral apex. Apical head setae (ApHS) 4, anterior posterior setae and acuminate posterior lobes. Paratergal plate III marginal head setae (AnMHS) 4. Dorsally, sutural head setae (SuHS) with 2 large setae and 2 serrated posterior lobes. Paratergal plates 4. Dorsal marginal head setae (DMHS) 4 on each side. DMHS not IV–VI each with 1 large seta posteriorly. Paratergal plates IV with aligned in row, first DMHS on top of postantennal angle, 2nd, 3rd, serrated posterior lobes. Paratergal plate V with 2 acuminate poste- and 4th DMHS offset medially. Small dorsal accessory head setae rior lobes. Paratergal plate VI with 1 acuminate posterior lobe dor- (DAcHS) 2, small dorsal anterior central head setae (DAnCHS) 2, sally. Paratergal plates VII, VIII each with 2 long setae and lacking small dorsal posterior central head setae (DPoCHS) 2, large dorsal posterior lobe. Paratergal plates I, II without spiracle. Spiracles on principal head setae (DPHS) 2. Ventrally, ventral principal head setae paratergal III–VII plates large in size, paratergal plate V spiracle (VPHS) 2. Antenna with five segments; first segment large, about as 20.5–30.7 (25.1) in diameter. Spiracle on paratergal plate VIII broad as long; second segment smaller, longer than wide; segments smaller than other paratergal spiracles. 3–5 about as wide as long. Distal seta on dorsal surface of antennal Genitalia (Fig. 1d): Subgenital plate with narrow anterolateral exten- segment 3 not sexually dimorphic. Antennal angle well developed. sion and 3 small lacunae on each side. Basal apodeme slightly longer Thorax: About as wide as long, with 1 dorsal principal thoracic seta than parameres. Parameres uniformly sclerotized, with pseudopenis (DPTS) on each side, DPTS length 108.5–156.6 (126.2). Thoracic tapering to a point extending slightly beyond apices of parameres. sternal plate shield-shaped with elongate anterior and posterior Female: [n = 3; Fig. 2a] Body length 1422–1447 (1434). processes, posterior process longer than anterior process (Fig. 1b). Head: Head slightly longer than wide, with 4 ApHS and 4 AnMHS. Forelegs small, with small acuminate claws; midlegs and hindlegs Pre-antennal region short with pointed antero-lateral apex. Dorsally, progressively larger with correspondingly more robust tibio-tarsal head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and claws, midlegs nearly two times longer than forelegs. 8 DMHS. Second, 3rd and 4th DMHS offset medially on each side. Abdomen: Wider than thorax. Dorsally, 1 tergite per segment, except Ventrally, head with 2 VPHS. Antenna with 5 segments; relative sizes for segment 3 with 2 tergites and segment 8 lacking tergite. Tergite 1 of segments as in male, antennal angle well developed. with 1 pair of small tergal abdominal setae (TeAS) posterolaterally. Thorax: About as wide as long, with 1 pair of DPTS dorsally, DPTS Tergite 2 with 2 pairs of TeAS posterolaterally, equal in length. length 110.5–113.6 (112.1). Thoracic sternal plate shield shaped Tergite 3 with 2 pairs of TeAS posterolaterally, with laterally placed with elongate anterior and posterior processes, posterior process seta on each side much longer. Tergites 4–6 each with 4 pairs of longer than anterior process (Fig. 2b). Forelegs small, with small TeAS, lateral pair of TeAS much longer on tergite 4. Tergite 7 with 7 acuminate claws; midlegs and hindlegs progressively larger with TeAS. Tergite 8 with 2 pairs of TeAS. Tergites 5–8 each with 1 pair correspondingly more robust tibio-tarsal claws, midlegs nearly two of dorsal lateral abdominal setae (DLAS). Ventrally, no sternites on times longer than forelegs. segments 1, 8. Segment 2 with 1 sternite. Segments 3–7 each with 2 Abdomen: Wider than thorax. Dorsally, 3 tergites per segment, except sternites, except for segment 4 with 3 sternites. Sternite 1 extended for segments 1, 2, 8 each with 1 tergite. All tergites narrow. Tergite 1 laterally to articulate with paratergal plate II and with 7 sternal ab- with 1 pair of TeAS posterolaterally. Tergite 2 with 2 pairs of TeAS dominal setae (StAS). Sternite 2 extended laterally to articulate with posterolaterally, all equal in length. Tergites 3, 4, 6, 15, 17 with 5 paratergal plate III and with 7 StAS, lateral 2 pairs long and stout; TeAS, respectively. Tergites 5, 7, 10, 12–14, and 16 each with 3 pairs medial StAS longer than 3rd pair StAS. Sternites 3, 5 each with 4 of TeAS. Tergites 8, 9, 11 each with 7 TeAS, respectively. Tergites 18, 4 Journal of Medical Entomology, 2020, Vol. XX, No. XX

Table 2. Murine rodent specimens (n = 46) from which new species of Hoplopleura sucking lice were collected

Host species Museum Collection Locality Hoplopleura Collection Museum accession No. spp. collected date

Conilurus penicillatus U6037 Angurugu, Groote Eylandt, NT 5 (5)a Unknown Museum and Art (13° 59′ 50.4″ S, 136° 28′ 34.8″ E) Gallery of the Northern Territory U5128 Melville Island, Tiwi Islands, NT 3 (12) 28-July-2000 Museum and Art Gallery of the Northern Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 Territory U5303 Araru Point,18 km from Araru Out- 1 (11) 23-June-2001 Museum and Art station, Cobourg Peninsula, NT. 11° Gallery of the 11′ S, 131° 53′ E Northern Territory M4071 Melville Island, NT. 11° 35′ S, 131° (1) 14-Aug.-1913 South Australian 10′ E Museum M54823 Wyndham-East Kimberley, Mitchell 8 (20) 01-Jan.-2003 Western Australian Plateau, WA. 14° 37′ S, 125° 52′ E Museum M54632 Wyndham-East Kimberley, Mitchell 7 (1) 20-May-2003 Western Australian Plateau, WA. 14° 35′ S, 125° 45′ E Museum Leggadina forresti JM12692 Blair Athol Coal Mine, QLD. 22° 42′ S, 21 (63) 02-Dec.-1997 Queensland Museum 147° 30′ E JM14858 Monkland Station via Alpha, QLD. 23° (4) 13-May-1998 Queensland Museum 23′ S, 146° 28′ E JM4486 Pelican Waterhole, Mt Leonard, QLD. (8) 17-April-1984 Queensland Museum 25° 42′ S, 140° 37′ E JM10077 Cannington Stn, 80 km S McKinlay, (2) 18-May-1993 Queensland Museum QLD. 21° 53′ S, 140° 55′ E JM11630 Blair Athol Coal Mine, QLD. 22° 42′ S, 1 (5) 22-Feb.-1997 Queensland Museum 147° 36′ E JM8816 LAKE YAMMA YAMMA,2 km EAST, (1) 12-Sept.-1991 Queensland Museum QLD. 26° 17′ S, 141° 37′ E C4530 Charlotte Waters, MacDonnell, NT. (2) Unknown Museums Victoria 25° 55′ S, 134° 55′ E C4527 Charlotte Waters, MacDonnell, NT. 1 Unknown Museums Victoria 25° 55′ S, 134° 55′ E C4517 Mulka, SA. 28° 21′ S, 138° 39′ E (4) Unknown Museums Victoria C4514 Mulka, SA. 28° 21′ S, 138° 39′ E 2 Unknown Museums Victoria C4518 Mulka, SA. 28° 21′ S, 138° 39′ E 4 (5) Unknown Museums Victoria C11331 Fowlers Gap, NSW. 31° 04′ S, 141° (2) 05-May-1973 Museums Victoria 42′ E C4513 Mulka, SA. 28° 21′ S, 138° 39′ E 2 09-Nov.-1931 Museums Victoria C4515 Mulka, SA. 28° 21′ S, 138° 39′ E 1 Unknown Museums Victoria C4519 Mulka, SA. 28° 21′ S, 138° 39′ E 1 Unknown Museums Victoria Leggadina JM12149 Clement SF, Rollingstone, ca. 5 km SE, 5 21-Oct.-1997 Queensland Museum lakedownensis Townsville, QLD. 19° 04′ S, 146° 25′ E JM11573 Undara NP, Archway Cave, Etheridge, (1) 20-Nov.-1996 Queensland Museum QLD. 18° 13′ S, 144° 34′ E JM15792 Unknown (1) Unknown Queensland Museum JM16574 Unknown 2 Unknown Queensland Museum U4198 Tjaynera Falls Area, Litchfield 1 (2) 08-Oct.-1995 Museum and Art National Park, NT. 13° 15′ S, Gallery of the 130° 44′ E Northern Territory M62366 Halls Creek, WA. 17° 36′ S, 128° 00′ E 17 (1) 01-Jan.-2006 Western Australian Museum M61252 Approx 9 km S Great Northern 8 (16) 31-Mar.-2012 Western Australian Highway Argyle, Wyndham-East Museum Kimberley, WA. 16° 34′ S, 128° 16′ E M64892 Roebourne, WA. 20° 48′ S, 116° 42′ E 1 Sept.-2006 Western Australian Museum M61814 Ashburton, WA. 22° 18′ S, 117° 30′ E 4 (2) Sept.-2005 Western Australian Museum Journal of Medical Entomology, 2020, Vol. XX, No. XX 5

Table 2. Continued

Host species Museum Collection Locality Hoplopleura Collection Museum accession No. spp. collected date M56949 East Pilbara, WA. 20° 48′ S, 119° 30′ E (1) May-2006 Western Australian Museum M56519 East Pilbara, WA. 22° 24′ S, 119° 48′ E (2) July-2004 Western Australian Museum Leporillus conditor M12291 West Franklin Island, SA. 32° 27′ S, 9 (11) 28-Oct.-1983 South Australian 133° 38′ E Museum

M12236 West Franklin Island, SA. 32° 27′ S, 19 (4) 29-May-1985 South Australian Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 133° 38′ E Museum M14053 West Franklin Island, SA. 32° 27′ S, 1 (1) 07-May-1984 South Australian 133° 39′ E Museum Mesembriomys gouldii DTC346 Lower Archer River, QLD. 13° 34′ S, 2 (1) 22-July-1933 Museums Victoria 142° 09′ E DTC349 Caledon Bay, NT. 12° 51′ S, 136° 33′ E 15 (33) Jan.-1943 Museums Victoria MG C Unknown 9 (5) Unknown Queensland Museum Mesembriomys M14148 Wyndham-East Kimberley, WA. 14° 36′ 1 Aug.-1987 South Australian macrurus S, 125° 48′ E Museum M21694 Wyndham-East Kimberley, WA. 14° 36′ 3 (2) Jan.-1982 Western Australian S, 125° 54′ E Museum M44975 Unknown 6 (5) Unknown Western Australian Museum Pogonomys JM10590 Shiptons Flat, QLD. 15° 48′ S, 145° 4 (4) Unknown Queensland Museum mollipilosus 00′ E JM14682 Shiptons Flat, QLD. 15° 48′ S, 145° 2 22-Oct.-1996 Queensland Museum 15′ E Xeromys myoides JM9480 Myora, N Stradbroke Is, QLD. 27° 27′ 1 (2) 21-Aug.-1992 Queensland Museum S, 153° 25′ E JM15078 Unknown (1) Unknown Queensland Museum U4744 Ramingining Area, Arafura Swamp, 1 28-July-1998 Museum and Art East Arnhem, NT. 12° 12′ S, 134° Gallery of the 59′ E Northern Terri- tory

NSW: New South Wales; QLD: Queensland; SA: South Australia; WA: Western Australia; NT: Northern Territory. Collection locality information is from https:// www.ala.org.au (access date: 2 April 2020). aNumber of adult specimens outside brackets, number of nymphal specimens inside brackets.

19 with 2 pairs of TeAS. One pair of DLAS adjacent to each of ter- of paratertal plate V 23.5–31.7 (27.6) in diameter. Spiracle on gites 6, 7, 9, 12, 15. Tergites 10, 13, 16 each with 2 pairs of adjacent paratergal plate VIII smaller than other paratergal spiracles. DLAS. Ventrally, 3 sternites per segment except for segment 2 with Genitalia (Fig. 2d): Subquadrangular subgenital plate tapering to 1 sternite, and segments 1, 8 each with no sternites. Sternite 1 with broadly rounded posterior apex, with 2 small mediolateral setae on 4 pairs of StAS and extended laterally to articulate with paratergal each side. Gonopods VIII, IX distinct; gonopods VIII with 3 pos- plate II. Sternite 2 extended laterally to articulate with paratergal terior setae subequal in size; gonopods IX with 3 posterior setae plate III and with 7 StAS, lateral 2 pairs long and stout, 3rd pair of of differing lengths, lateral seta longest and medial seta shortest. StAS short, medial StAS longer than 3rd pair. Sternites 3, 5, 8 each Vulvar fimbriae indistinct. Three small setae on each side medial to with 7 StAS. Sternites 4, 7, 10 each with 4 pairs of StAS. Sternites 6, gonopods IX. 9, 11–14 each with 3 pairs of StAS. Sternite 15 with 2 pairs of StAS. Type host. Conilurus penicillatus (Gould, 1842) (Rodentia: Sternite 16 with 4 pairs of StAS, lateral two pairs much longer than Muridae), brush-tailed rabbit rat. medial 2 pairs. One pair of VLAS adjacent to each of sternites 6, 8, 9, Type locality. Wyndham-East Kimberley, Mitchell Plateau, Western 11–13, and 16. Sternite 15 with 2 pairs of adjacent VLAS. Australia (14° 35′ S, 125° 45′ E). Paratergites: Paratergal plates (Fig. 2c) present on abdominal seg- Type material. Holotype male and allotype female ex Conilurus ments 1–8. All plates differentially sclerotized. Paratergal plate penicillatus (WAM, M54632): holotype, ♂, allotype, ♀. Paratypes: I small and offset medially. Paratergal plates II–V each with 2 poste- 3 ♂ and 1 ♀, same data as for the holotype. Museum accession num- rior lobes. Paratergal plate II with 1 small medial seta, 2 large pos- bers are linked to the ZooBank record. terior setae, and acuminate posterior lobes. Paratergal plate III with Additional material examined. 2 ♂ ex Conilurus penicillatus (WAM, 2 large posterior setae and 2 serrated posterior lobes. Paratergal M54832), Wyndham-East Kimberley, Mitchell Plateau, Western plates IV–VI each with 1 large seta postero-dorsally. Paratergal Australia (14° 35′ S, 125° 45′ E). 1 ♀ ex Conilurus penicillatus plates IV with serrated posterior lobes. Paratergal plate V with 2 (MAGNT, U6037), Angurugu, Groote Eylandt, NT (13° 59′ 50.4″ acuminate posterior lobes. Paratergal plate VI with 1 acuminate pos- S, 136° 28′ 34.8″ E). terior lobe postero-dorsally. Paratergal plates VII, VIII each with 2 Distribution. Australia: Western Australia. long setae and lacking posterior lobe. Paratergal plates I, II without Etymology. The species epithet is a noun in apposition referring to spiracle. Spiracles on paratergal III–VII plates large in size, spiracle the genus name of the host, Conilurus. 6 Journal of Medical Entomology, 2020, Vol. XX, No. XX Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020

Fig. 1. Hoplopleura conilurudis n. sp. Male. (a) Habitus (dorsal morphology to the left of the midline, ventral morphology to the right). (b) Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. Fig. 2. Hoplopleura conilurudis n. sp. Female. (a) Habitus (dorsal morphology to the left of the midline, ventral morphology to the right). (b) Thoracic sternal Remarks. The pre-antennal region of H. conilurudis n. sp. has plate. (c) Paratergal plates. (d) Genitalia. a pointed anterolateral apex which is different from that in H. irritans Kuhn and Ludwig, 1967, H. calabyi, Johnson, 1960 and H. uromydis Kuhn and Ludwig, 1967 (Johnson 1960, Kuhn V of H. conilurudis n. sp. is large in size (male 25.1, female 27.6), and Ludwig 1966). This region is rounded in H. irritans and and is larger than in H. gyomydis (7–9). Hoplopleura conilurudis H. calabyi and truncated in H. uromydis. The third antennal n. sp. has one posterior lobe dorsally on paratergal plate VI, segment of the male H. conilurudis is not sexually dimorphic, whereas H. zyzomydis, H. melomydis, H. notomydis, H. setosa, which is different from H. irritans. The DMHS of H. conilurudis H. uromydis, H. gyomydis, H. mastacomydis, and H. calabyi have n. sp. are not aligned, whereas in H. gyomydis Kuhn and two posterior lobes on this plate. Hoplopleura conilurudis does not Ludwig, 1967, H. mastacomydis Kuhn and Ludwig, 1967, and have posterior lobes on paratergal plate VII. However, H. setosa, H. melomydis, the DMHS are all aligned in a row (Kuhn and H. melomydis, H. zyzomydis, and H. mastacomydis have one pos- Ludwig 1966, Weaver 2017). The 2nd, 3rd, and 4th DMHS of terior lobe on this paratergal plate, whereas H. mastacomydis and H. conilurudis are offset medially, which is similar in H. uromydis, H. calabyi have two posterior lobes on this plate. Hoplopleura H. villosissima Wang, 2018 and H. irritans but different from conilurudis has two posterior setae on paratergal plate III, whereas H. uromydis, H. villosissima and H. irritans (Kuhn and Ludwig H. mastacomydis has one short and stout seta posteriorly. Males 1966, Wang et al. 2018). Hoplopleura conilurudis n. sp. has of H. conilurudis have one pair of DLAS on each segment of IV to four pairs of DMHS, which is different from H. zyzomydis VI; H. uromydis, H. calabyi, and H. mastacomydis have no DLAS Weaver, 2008 (two pairs), and H. notomydis Weaver, 2017 and on the abdomen. The female of H. conilurudis has two pairs of H. setosa Weaver, 2017 (three pairs) (Weaver and Barton 2008, DLAS on tergites 10, 13 and 16, and two pairs of VLAS on stern- Weaver 2017). The sternal plate of H. conilurudis n. sp. is shield ites 15, which differentiate it from all other Hoplopleura species shaped, which differentiates it from H. uromydis (less rounded in Australia. in shape). Hoplopleura conilurudis n. sp. has a small central seta on paratergal plate II, which is absent in H. pacifica Ewing, Hoplopleura leggadinadis Wang n. sp. 1924, H. bidentata (Neumann, 1909), H. cornata Kim, 1972, (urn:lsid:zoobank.org:act:0D839340-F044-49E4-A996- H. zyzomydis, and H. notomydis (Neumann 1909, Ewing 1924, D337CF663C33). Kim 1972). Paratergal plates IV–VI each have one large posterior Male: [n = 5; Fig. 3a] Body length 730–895 (803). seta in H. conilurudis n. sp. These three paratergal plates have Head: Head longer than wide, with 4 ApHS and 4 AnMHS. Pre- two posterior setae in H. uromydis, H. gyomydis, H. irritans, antennal region short, truncated anterolaterally. Antenna with 5 H. mastacomydis, and H. melomydis, but lack posterior setae in segments; 1st segment large, about as wide as long; 2nd segment H. notomydis and H. zyzomydis. The spiracle on paratergal plates smaller, much longer than wide; 3rd–5th segments about as long Journal of Medical Entomology, 2020, Vol. XX, No. XX 7 as wide; 4th segment slightly extended postero-apically; distal seta with 7 StAS. Sternite 11 with 2 pairs of StAS. Sternite 12 with 1 pair on dorsal surface of antennal segment 3 not sexually dimorphic. of StAS. One pair of VLAS adjacent to each of sternites 6, 8, 9, 11. Antennal angle not well developed. Dorsally, head with 4 SuHS, 2 Paratergites: Paratergal plates (Fig. 3c) present on abdominal seg- DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. Second, ments 1–8. All plates differentially sclerotized. Paratergal plate 3rd, and 4th DMHS offset medially on each side. Ventrally, head I small and offset medially. Paratergal plates II–VI each with 2 pos- with 2 VPHS. terior lobes. Paratergal plate II with 1 small medial seta, 2 large pos- Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS terior setae and acuminate posterior lobes. Paratergal plate III with length 72.7–92.1 (80.7). Thoracic sternal plate shield-shaped, 2 large posterior setae and 2 serrated posterior lobes. Paratergal with broadly rounded anterior apex and elongated posterior pro- plates IV–VI each with 1 large posterior seta and 2 posterior lobes.

cess (Fig. 3b). Forelegs small, with small acuminate claws; midlegs Paratergal plates VII, VIII each with 2 large posterior setae. Paratergal Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 and hindlegs progressively larger with correspondingly more robust plate VII with 1 posterior lobe dorsally, whereas paratergal plate tibio-tarsal claws. VIII lacks posterior lobe. Paratergal plates I, II without spiracles. Abdomen: Wider than thorax. Dorsally, 1 tergite per segment ex- Spiracle of paratergal plate V 11.3–19.4 (14.8) in diameter. Spiracle cept for segment 3 with 2 tergites. Tergite 1 with 1 pair of TeAS on paratergal plate VIII smaller than other paratergal spiracles. posterolaterally. Tergite 2 with 2 pairs of TeAS posterolaterally, equal Genitalia (Fig. 3d): Subgenital plate with narrow anterolateral ex- in length. Tergite 3 with 2 pairs of TeAS posterolaterally, lateral pair tension on each side and 4 lacunae; posterior lacunae slight larger very long, twice as long as medial pair. Tergites 4–8 each with 7 than anterior lacunae. Basal apodeme slightly longer than parameres. TeAS, TeAS length varies in tergite 4. One pair of DLAS adjacent to Parameres narrow, uniformly sclerotized, with pseudopenis tapering tergite 7. Ventrally, 2 sternites per segment except for segment 3 with to a point extending slightly beyond apices of parameres. 3 sternites, segments 2, 7, 8 each with 1 sternite, and segment 1 with Female: [n = 4; Fig. 4a] Body length 843–1317 (1007). no sternite. Sternite 1 extended laterally to articulate with paratergal Head: Head longer than wide, with 4 ApHS and 4 AnMHS. Pre- plate II and with 4 pairs of StAS, lateral two pairs stout. Sternite antennal region short, truncated anterolaterally. Antenna with 5 seg- 2 partially articulating with paratergal plate III and with 7 StAS, ments; relative sizes of segments as in male. Antennal angle weak. lateral 2 pairs long and stout, medial StAS slightly longer than 3rd Dorsally, head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 pair. Sternite 3 with 4 pairs of StAS, medial pair slightly short and DPHS, and 8 DMHS. Second, 3rd, and 4th DMHS offset medially narrow. Sternite 4 with 7 StAS, lateral two pairs slightly stouter and on each side. Ventrally, head with 2 VPHS. longer than others, medial StAS longer than 3rd lateral pair of StAS. Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS Sternites 5, 7, 9, 10 each with 4 pairs of StAS. Sternites 6, 8 each length 68.6–107.5 (88.6). Thoracic sternal plate shield shaped with broadly rounded anterior apex and elongate posterior process (Fig. 4b). Forelegs small, with small acuminate claws; midlegs and hindlegs progressively larger with correspondingly more robust tibio-tarsal claws. Abdomen: Wider than thorax. Dorsally, 3 tergites per segment except for segments 1, 2 each with 1 tergite and segment 8 with 2 tergites. All tergites narrow. Tergite 1 with 1 pair of TeAS posterolaterally. Tergites 2, 3 each with 2 pairs of TeAS posterolaterally, equal in length. Tergites 4, 6, 9, 12–16 each with 5 TeAS. Tergites 5, 17, 18 each with 2 pairs of TeAS. Tergites 7, 10 each with 7 TeAS. Tergites 8, 11 each with 3 pairs of TeAS. Tergites 16, 17 each with 2 pairs of TeAS. One pair of DLAS adjacent to each of tergites 9, 10, 13, 15, 17. Ventrally, 3 sternites per segment, except for segment 2 with 1 sternite, and segment 1, 8 with no sternite. Sternite 1 with 9 StAS, lateral pair stout and extended laterally to articulate with paratergal plate II. Sternite 2 extended laterally to articulate with paratergal plate III and with 7 StAS, lateral 2 pairs long and stout, 3rd pair of setae short, medial StAS slightly longer than 3rd pair. Sternites 3, 5, 11–14 each with 7 StAS. Sternites 4, 7–10 with 4 pairs of StAS, respectively. Sternites 6, 15 each with 3 pairs of StAS. Sternite 16 with 4 pairs of StAS, varying in size, lateral 2 pairs much longer than others. One pair of VLAS adjacent to each of sternites 8, 9, 11–13, 15, 16. Paratergites: Paratergal plates (Fig. 4c) present on abdominal segments 1–8. All plates differentially sclerotized. Paratergal plate I small and offset medially. Paratergal plates II–VI each with 2 posterior lobes. Paratergal plate II with 1 small medial seta, 2 large posterior setae, and acuminate posterior lobes. Paratergal plate III with 2 large posterior setae and 2 serrated posterior lobes. Paratergal plates IV–VI each with 1 large posterior seta. Paratergal plates VII, VIII

Fig. 3. Hoplopleura leggadinadis n. sp. Male. (a) Habitus (dorsal morphology each with 2 large posterior setae. Paratergal plate VII with 1 pos- to the left of the midline, ventral morphology to the right). (b) Thoracic sternal terior lobe dorsally, whereas paratergal plate VIII lacks posterior plate. (c) Paratergal plates. (d) Genitalia. lobes. Paratergal plates I, II without spiracle. Spiracle of paratergal 8 Journal of Medical Entomology, 2020, Vol. XX, No. XX plate V 16.4–26.6 (19.9) in diameter. Paratergal plate VIII with small H. gyomydis, H. mastacomydis, and H. melomydis, the DMHS are spiracle. all aligned in a row. The 2nd, 3rd, and 4th DMHS of H. leggadinadis Genitalia (Fig. 4d): Subtriangular subgenital plate with 5 small n. sp. are offset medially, whereas in H. uromydis, H. villosissima, mediolateral setae. Gonopods VIII, IX distinct; gonopods VIII with 3 and H. irritans, only the 2nd and 3rd DMHS offset medially. posterior setae subequal in size; gonopods IX with 3 posterior setae Hoplopleura leggadinadis n. sp. has four pairs of DMHS, which of differing lengths, lateral seta longest and medial seta shortest. is different from H. zyzomydis (two pairs), and H. notomydis and Vulvar fimbriae indistinct. Three small setae on each side medial to H. setosa (three pairs). The sternal plate of H. leggadinadis n. sp. is gonopods IX. shield-shaped with a broadly rounded anterior apex, differentiating Type host. Leggadina lakedownensis (Watts, 1976) (Rodentia: it from H. uromydis (less rounded in shape with a long anterior

Muridae), lakeland downs mouse. process). Hoplopleura leggadinadis n. sp. has a small central seta Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 Type locality. Approx. 9 km S Great Northern Highway Argyle, on paratergal plate II, which is absent in H. pacifica, H. bidentata, Wyndham-East Kimberley, Western Australia (16° 34′ S, 128° 16′ E). H. cornata, H. zyzomydis, and H. notomydis. Paratergal plates IV– Type material. Holotype male and allotype female ex Leggadina VI each have 1 large posterior seta in H. leggadinadis n. sp. These lakedownensis (WAM, M61252): holotype, ♂, allotype, ♀. Paratypes: three paratergal plates each have two posterior setae in H. uromydis, 2 ♂ and 2 ♀, same data as for the holotype. Museum accession num- H. gyomydis, H. irritans, H. mastacomydis, and H. melomydis, bers are linked to the ZooBank record. but no posterior setae in H. notomydis and H. zyzomydis. The Additional material examined. 2 ♂, 1 ♀ ex Leggadina lakedownensis paratergal plate VII of H. leggadinadis n. sp. has one posterior lobe (QM, JM12149), Clement SF, Rollingstone, ca. 5 km SE, Townsville, dorsally, in comparison to two posterior lobes in H. mastacomydis Queensland (19° 04′ S, 146° 25′ E). and H. calabyi, and no posterior lobe in H. cornata, H. gyomydis, Distribution. Australia: Queensland and Western Australia. H. irritans, H. uromydis, H. villosissima, H. bidentata, H. conilurudis, Etymology. The species epithet is a noun in apposition referring to and H. pacifica. Hoplopleura leggadinadis n. sp. has two posterior the genus name of the host, Leggadina. setae on paratergal plate III, whereas H. mastacomydis has one Remarks. The pre-antennal region of H. leggadinadis n. sp. is short, stout seta posteriorly. The spiracle on paratergal plate V of truncated anteriorly, which is different from that in H. irritans, H. leggadinadis n. sp. (male 14.8, female 19.9) is smaller than in H. calabyi, and H. bidentata. Anteriorly, the pre-antennal region is H. uromydis (34–37), H. bidentata (25–28), H. melomydis (male 25, rounded in H. irritans and H. calabyi, but narrow and pointed in female 25.8), and H. conilurudis (male 25.1, female 27.6), but larger H. bidentata. The third antennal segment of H. leggadinadis n. sp. is than in H. gyomydis (7–9). The VLAS of H. leggadinadis n. sp. are not sexually dimorphic, which differentiates it from H. irritans. The different from those of H. setosa, which has VLAS on small plates DMHS of H. leggadinadis n. sp. are not aligned in a row, whereas in next to the tergites. The male of H. leggadinadis n. sp. has one pair of DLAS/VLAS on abdominal segments 4–7; H. uromydis, H. calabyi, and H. mastacomydis have no DLAS on segments 4–7. The female of H. leggadinadis n. sp. has three sternites on segment 3–6, whereas females of H. bidentata have two sternites on the corresponding segments. Hoplopleura forrestima Wang n. sp. (urn:lsid:zoobank. org:act:64A59F63-CA08-4576-8157-5C666D45447B). Male: [n = 4; Fig. 5a] Body length 815–928 (856). Head: Head longer than wide, distinctly narrower in anterior half, with 4 ApHS and 4 AnMHS. Pre-antennal region short with round anterolateral margin. Antenna with 5 segments; 1st segment large, about as wide as long; 2nd segment smaller, longer than wide; 3rd–5th segments about as wide as long; distal seta on dorsal surface of antennal segment 3 not sexually dimorphic. Antennal angle not well developed. Dorsally, head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS; 2nd, 3rd, and 4th DMHS offset medially on each side. Ventrally, head with 2 VPHS. Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length 74.7–97.2 (85.5). Thoracic sternal plate shield shaped with short anterior process and elongate posterior process (Fig. 5b). Forelegs small, with small acuminate claws; midlegs and hindlegs progressively larger with correspondingly more robust tibio-tarsal claws. Abdomen: Wider than thorax. Dorsally, 1 tergite per segment, except for segment 3 with 2 tergites. Tergite 1 with 1 pair of TeAS posterolaterally. Tergite 2 with 2 pairs of TeAS posterolaterally, all equal in length. Tergite 3 with 2 pairs of TeAS posterolaterally, with TeAS much longer on each side. Tergite 4 with 9 TeAS, lateral 2 pairs longer than others. Tergite 5 with 5 pairs of TeAS. Tergites 6, 7 each with 4 pairs of TeAS. Tergite 8 with 7 TeAS. One pair of DLAS adjacent to each of tergites 5–8. Ventrally, 2 sternites per segment except Fig. 4. Hoplopleura leggadinadis n. sp. Female. (a) Habitus (dorsal mor- segment 4 which has 3 sternites, segment 2 with 1 sternite, segment phology to the left of the midline, ventral morphology to the right). (b) 1, 8 with no sternites. Sternite 1 extended laterally to articulate with Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. paratergal plate II and with 7 StAS. Sternite 2 extended laterally to Journal of Medical Entomology, 2020, Vol. XX, No. XX 9 Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020

Fig. 5. Hoplopleura forrestima n. sp. Male. (a) Habitus (dorsal morphology to the left of the midline, ventral morphology to the right). (b) Thoracic sternal Fig. 6. Hoplopleura forrestima n. sp. Female. (a) Habitus (dorsal morphology plate. (c) Paratergal plates. (d) Genitalia. to the left of the midline, ventral morphology to the right). (b) Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. articulate with paratergal plate III and with 7 StAS, lateral 2 pairs long and stout, 3rd pair setae short, medial StAS longer than 3rd pair. Sternite 3 with 4 pairs of StAS, equal in length. Sternites 4, 7 each with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS; 7 StAS. Sternites 5, 6 each with 9 StAS. Sternites 8–10 each with 3 2nd, 3rd, and 4th DMHS offset medially on each side. Ventrally, head pairs of StAS. Sternite 11 with 2 pairs of StAS. Sternite 12 with 1 pair with 2 VPHS. of TeAS. One pair of VLAS adjacent to each of sternites 6, 8, 10, 11. Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length Paratergites: Paratergal plates (Fig. 5c) present on abdominal seg- 93.1–102.4 (97.7). Thoracic sternal plate shield shaped with short ments 1–8. All plates differentially sclerotized. Paratergal plate anterior process and elongate posterior process (Fig. 6b). Forelegs I small and offset medially. Paratergal plates II–VI each with 2 pos- small, with small acuminate claws; midlegs and hindlegs progres- terior lobes. Paratergal plate II with 1 small medial seta, 2 large pos- sively larger with correspondingly more robust tibio-tarsal claws. terior setae, and acuminate posterior lobes. Paratergal plate III with Abdomen: Wider than thorax. Dorsally, 3 tergites per segment ex- 2 large setae and 2 serrated posterior lobes. Paratergal plates IV–VI cept for segments 1, 2, 8 each with 1 tergite. All tergites narrow. each with 1 large seta and 1 tiny seta. Paratergal plates VII, VIII each Tergite 1 with 1 pair of TeAS posterolaterally. Tergite 2 with 2 pairs with 2 large setae. Paratergal plate VI with 2 acuminate posterior of TeAS posterolaterally, equal in length. Tergites 3, 4, 6, 11, 15, 16 lobes. Paratergal plate VII with 1 posterior lobe dorsally. Paratergal each with 5 TeAS. Tergites 5, 7, 9, 10, 12–14 each with 3 pairs of plate VIII lacks posterior lobes. Paratergal plates I, II, VIII without TeAS. Tergite 8 with 7 TeAS. Tergites 17, 18 each with 2 pairs of spiracles. Spiracles on paratergal plates III–VII, medium in size, spi- TeAS. One pair of DLAS adjacent to each of tergites 9, 10, 12, 13, racle of paratergal plate V 13.3–17.4 (15.4) in diameter. 15, 16. Ventrally, 3 sternites on each segment except for segment 2 Genitalia (Fig. 5d): Subgenital plate with narrow anterolateral ex- with 1 sternite, and segments 1, 8 with no sternites. Sternite 1 extension on each side and 6 lacunae; 4 anterior lacunae slight smaller tended laterally to articulate with paratergal plate II and with 4 pairs than 2 posterior lacunae. Basal apodeme much longer than of StAS. Sternite 2 extended laterally to articulate with paratergal parameres. Parameres uniformly sclerotized, tapering posteriorly plate III and with 7 StAS, lateral 2 pairs long and stout, 3rd pair with pseudopenis tapering to point extending well beyond apices setae short, medial StAS longer than 3rd pair. Sternites 3, 9, 12, 15 of parameres. each with 3 pairs of StAS. Sternites 4–6, 8, 11, 14 each with 7 StAS. Female: [n = 5; Fig. 6a] Body length 973–1173 (1089). Sternites 7, 10, 13 each with 4 pairs of StAS. Sternite 16 with 4 pairs Head: Head longer than wide, distinctly narrower in anterior half, of StAS, lateral two pairs much longer than medial 2 pairs. One pair with 4 ApHS and 4 AnMHS. Pre-antennal region short, rounded of VLAS adjacent to each of sternites 8, 9, 11–13, 15, 16. anterolaterally. Antenna with 5 segments with relative sizes as in Paratergites: Paratergal plates (Fig. 6c) present on abdominal seg- male, distal seta on dorsal surface of antennal segment 3 not sexually ments 1–8. All plates differentially sclerotized. Paratergal plate dimorphic. Antennal angle not well developed. Dorsally, head with I small and offset medially. Paratergal plates II–VI each with 2 10 Journal of Medical Entomology, 2020, Vol. XX, No. XX posterior lobes. Paratergal plate II with 1 small medial seta, 2 large posterior setae, and acuminate posterior lobes. Paratergal plate III with 2 large setae and 2 serrated posterior lobes. Paratergal plates IV–VI each with 1 large seta and 1 tiny seta. Paratergal plates VII, VIII each with 2 large setae. Paratergal plate VI with 2 acuminate posterior lobes. Paratergal plate VII with 1 posterior lobe dorsally; paratergal plate VIII without posterior lobe. Paratergal plates I, II, VIII lacking spiracles. Spiracles on paratergal plates III–VII medium in size, spiracle on paratergal plate V 18.4–19.4 (19.1) in diameter.

Paratergal plate VIII lacking spiracle. Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 Genitalia (Fig. 6d): Subtriangular subgenital plate with 5 small mediolateral setae. Gonopods VIII, IX distinct; gonopods VIII with 3 posterior setae subequal in size; gonopods IX with 3 posterior setae of differing lengths, lateral seta longest and medial seta shortest. Vulvar fimbriae indistinct. Three small setae on each side medial to gonopods IX. Type host. Leggadina forresti (Thomas, 1906) (Rodentia: Muridae), forrest’s mouse. Type locality. Blair Athol Coal Mine, Queensland, Australia (22° 42′ S, 147° 30′ E). Type material. Holotype male and allotype female ex Leggadina forresti (QM, JM 12692): holotype, ♂, allotype, ♀. Paratypes: 1 ♂ and 2 ♀, same data as for the holotype. Museum accession numbers are linked to the ZooBank record. Additional material examined. 2 ♂ and 1 ♀ ex Leggadina forresti (MV, C4518), Mulka, South Australia, (28° 21′ S, 138° 39′ E); 1 ♀ ex Leggadina forresti (MV, C4514), Mulka, South Australia (28° 21′ S, 138° 39′ E). Fig. 7. Hoplopleura leporilludis n. sp. Male. (a) Habitus (dorsal morphology Distribution. Australia: Queensland and South Australia. to the left of the midline, ventral morphology to the right). (b) Antennae of Etymology. The species epithet is a noun in apposition referring to male and female. (c) Thoracic sternal plate. (d) Paratergal plates. (e) Genitalia. the specific name of the host, Leggadina forresti. Remarks. The pre-antennal region of H. forrestima n. sp. is pair of DLAS adjacent to abdominal segments IV–VI and one pair of rounded anteriorly, which is different from that in H. uromydis, VLAS adjacent to segments IV–VII, whereas males of H. uromydis, H. mastacomydis, H. leggadinadis, and H. bidentata. The anterior H. calabyi, and H. mastacomydis have no DLAS on these segments. pre-antennal region is truncate in H. uromydis and H. leggadinadis, The female of H. forrestima n. sp. has three sternites on segments less rounded in H. mastacomydis, and narrow and prominent in 3–6, whereas H. bidentata has two sternites on the corresponding H. bidentata. The third antennal segment of the male of H. forrestima segments. The female of H. forrestima n. sp. has 12 pairs of DLAS, n. sp. is not sexually dimorphic, which is different from H. irritans. whereas H. leggadinadis has eight pairs. The DMHS of H. forrestima n. sp. are not aligned in a row, whereas in H. gyomydis and H. mastacomydis, the DMHS are all aligned in a row. The sternal plate of H. forrestima n. sp. is shield shaped Hoplopleura leporilludis Wang n. sp. with a small anterior and a longer posterior process, whereas in (urn:lsid:zoobank.org:act:52062691-ADD3-42DD-99A1- H. uromydis, it is less rounded in shape and has long anterior and 35A5B7A39F8D). posterior processes. Hoplopleura forrestima n. sp. has a small central Male: [n = 5; Fig. 7a] Body length 861–877 (874). seta on paratergal plate II, which is absent in H. pacifica, H. bidentata, Head: Head slightly longer than wide, with 4 ApHS and 4 AnMHS. H. cornata, H. zyzomydis, and H. notomydis. Paratergal plates Pre-antennal region short, truncated anteriorly. Antenna with 5 seg- IV–VI of H. forrestima n. sp. each have one large posterior and ments. Male distal setae on 3rd antennal segment different from fe- one tiny posterior seta. Paratergal plates IV–VI of H. villosissima, male (Fig. 7b): distal setae stout and long in male, but narrow and H. setosa, H. conilurudis, and H. leggadinadis have one large poste- short in female. Antennal angle well developed. Dorsally, head with rior seta but lack posterior setae on H. notomydis and H. zyzomydis. 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. Paratergal plate VII of H. forrestima n.sp. has one posterior lobe DMHS aligned in row and offset medially. Ventrally, head with dorsally, whereas in H. mastacomydis and H. calabyi this plate has 2 VPHS. two posterior lobes. Hoplopleura cornata, H. gyomydis, H. irritans, Thorax: Wider than long, with one pair of DPTS dorsally, DPTS H. uromydis, H. villosissima, H. bidentata, H. conilurudis, H. pa- length 93.1–97.2 (98.1). Thoracic sternal plate sub-hexagonal with cifica, H. mastacomydis, and H. calabyi all lack a posterior lobe slightly elongated anterior and posterior processes (Fig. 7c). Forelegs on paratergal plate VII. Hoplopleura forrestima n. sp. has two pos- small, with small acuminate claws; midlegs and hindlegs progres- terior setae on paratergal plate III, whereas H. mastacomydis has sively larger with correspondingly more robust tibio-tarsal claws, one short, stout seta. The spiracle diameter on paratergal plate V midlegs almost two times longer than forelegs. of H. forrestima n. sp. (male 15.4, female 19.1) is smaller than in Abdomen: Wider than thorax. Dorsally, 1 tergite per segment ex- H. uromydis (34–37), H. bidentata (25–28), H. melomydis (male 25, cept for segment 3 with 2 tergites. All tergites broad and wide female 25.8), and H. conilurudis (male 25.1, female 27.6), but larger with no adjacent DLAS. Tergite 1 with 1 pair of long, narrow than in H. gyomydis (7–9). The male of H. forrestima n. sp. has one TeAS posterolaterally. Tergite 2 with 2 pairs of narrow TeAS Journal of Medical Entomology, 2020, Vol. XX, No. XX 11 posterolaterally, all equal in length. Tergite 3 with 2 pairs of narrow Genitalia (Fig. 7e): Subgenital plate with narrow anterolateral exten- setae posterolaterally, lateral pair very long, nearly twice as long as sion on each side. Basal apodeme about twice as long as parameres. medial pair. Tergite 4 with 7 TeAS, lateral pair of TeAS long and Parameres uniformly sclerotized, with broad pseudopenis tapering stout, second lateral pair of TeAS longer than lateral pair, other TeAS to broad apex extending slightly beyond apices of parameres. on tergite 4 very short. Tergites 5–7 each with 7 narrow TeAS. Middle Female: [n = 5; Fig. 8a] Body length 1146–1276 (1215). TeAS on tergite 5 short. Tergite 8 with 5 narrow TeAS, with middle Head: Head slightly longer than wide, with 4 ApHS and 4 AnMHS. TeAS shorter. Tegite 9 with 1 pair of TeAS. Ventrally, 2 sternites per Pre-antennal region short, truncated anteriorly. Antenna with 5 segment except for segment 4 with 3 sternites, segment 2 with 1 ster- segments. Distal seta on 3rd segment of antenna narrow and short nite, but segments 1, 8 without sternites. All sternites broad and wide (Fig. 7b). Antennal angle well developed. Dorsally, head with 4 SuHS,

with no adjacent VLAS. Sternite 1 with 8 StAS, gradually increasing 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. DMHS Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 in length laterally to medially. Sternite 1 extended laterally to ar- aligned in row and offset medially. Ventrally, head with 2 VPHS. ticulate with paratergal plate II. Sternite 2 with 7 StAS, lateral 2 Thorax: Wider than long, with 1 DPTS on each side, DPTS length pairs longer and stouter, 3rd pair of StAS shorter than medial StAS. 86–100.3 (95.2). Thoracic sternal plate sub-hexagonal with elongate Sternite 3 with 7 StAS, equal in length. Sternites 4, 6, 8, 10 each with anterior and posterior processes (Fig. 8b). Forelegs small, with small 5 StAS. Sternites 5, 7, 9 each with 3 pairs of StAS. Sternite 11 with 2 acuminate claws; midlegs and hindlegs progressively larger with pairs of StAS. Sternite 12 with 1 pair of StAS. correspondingly more robust tibio-tarsal claws, midlegs almost two Paratergites: Paratergal plates (Fig. 7d) present on abdominal seg- times longer than forelegs. ments 1–8. All plates differentially sclerotized. Paratergal plate Abdomen: Wider than thorax. Dorsally, 3 tergites per segment ex- I small and offset medially. Paratergal plates II–V each with 2 pos- cept for segments 1, 2, 8 each with 1 tergite. All tergites broad and terior acuminate lobes. Paratergal plate II with 1 small medial seta wide with no adjacent DLAS. Tergite 1 with 1 pair of long TeAS and 2 large posterior setae. Paratergal plate III with 2 large setae. posterolaterally. Tergite 2 with 2 pairs of TeAS posterolaterally, Paratergal plates IV–VI each with 1 large seta dorsally. Paratergal equal in length. Tergite 3 with 2 pairs of TeAS posterolaterally. plates VII, VIII each with 2 long setae and lacking posterior lobe. Tergites 4, 6, 7, 9 10, 12, 13, 16 each with 5 TeAS. Tergites 5, 8, Paratergal plates I, II without spiracles. Spiracle on paratergal plate 11, 14, 15, 17, 18 each with 2 pairs of TeAS. Ventrally, 3 sternites V 16.4–24.6 (20.5) in diameter. Spiracle on paratergal plate VIII in per segment, except for segment 2 with 1 sternite, and segments 1, small size. 8 no sternites. Sternite 1 with 4 pairs of StAS and extended laterally to articulate with paratergal plate II. Sternite 2 with 7 setae, lateral 2 pairs long and stout, middle StAS longer than 3rd pair of StAS. Sternites 3, 4, 7, 9, 10, 12, 13, 15 each with 3 pairs of StAS, equal in length. Sternites 5, 8, 11, 14 each with 5 StAS. Sternite 6 with 7 StAS. Sternite 16 with 4 pairs of StAS, of varying lengths, lateral 2 pairs of StAS longest. One pair of VLAS adjacent to sternite 15. Paratergites: Paratergal plates (Fig. 8c) present on abdominal segments 1–8. All plates differentially sclerotized. Paratergal plate I small and offset medially. Paratergal plates II–V each with 2 posterior acuminate lobes. Paratergal plate II with 1 small medial seta and 2 large posterior setae. Paratergal plate III with 2 large setae. Paratergal plates IV–VI each with 1 large seta dorsally. Paratergal plates VII, VIII each with 2 long setae and lacking posterior lobe. Paratergal plates I, II without spiracles. Spiracle on paratergal plate V 18.4–22.5 (22.5) in diameter. Spiracle on paratergal plate VIII small. Genitalia (Fig. 8d): Subtriangular subgenital plate with 4 small mediolateral setae. Gonopods VIII, IX distinct; gonopods VIII with 3 posterior setae subequal in size; gonopods IX with 3 posterior setae of differing lengths, lateral seta longest and medial seta shortest. Vulvar fimbriae indistinct. Three small setae on each side medial to gonopods IX. Type host. Leporillus conditor (Sturt, 1848) (Rodentia: Muridae), greater stick-nest rat. Type locality. West Franklin Island, South Australia, Australia (32° 27′ S, 133° 38′). Type material. Holotype male and allotype female ex Leporillus conditor (SAM, M12236): holotype, ♂, allotype, ♀. Paratypes: 2 ♂ and 3 ♀, same data as for the holotype. Museum accession numbers are linked to the ZooBank record. Additional material examined. 2 ♂ and 1 ♀ ex Leporillus conditor (SAM, M12291). West Franklin Island, South Australia (32° 27′ S, 133° 38′).

Fig. 8. Hoplopleura leporilludis n. sp. Female. (a) Habitus (dorsal morphology Distribution. Australia: South Australia. to the left of the midline, ventral morphology to the right). (b) Thoracic sternal Etymology. The species epithet is a noun in apposition referring to plate. (c) Paratergal plates. (d) Genitalia. the genus name of the host, Leporillus. 12 Journal of Medical Entomology, 2020, Vol. XX, No. XX

Remarks. The pre-antennal region of H. leporilludis n. sp. is trun- 25, female 25.8), and H. conilurudis (male 25.1, female 27.6) but cated anterolaterally, which differentiates it from H. irritans, larger than in H. gyomydis (7–9). Hoplopleura leporilludis n. sp. H. calabyi, H. forrestima n. sp., and H. bidentata. The anterolat- lacks posterior lobes on paratergal plate VII. Hoplopleura setosa, eral pre-antennal region is rounded in H. irritans, H. calabyi, and H. melomydis, and H. zyzomydis have one posterior lobe on this H. forrestima n. sp., and narrow and pointed in H. bidentata. The paratergal plate, whereas H. mastacomydis, H. leggadinadis, and third antennal segment of H. leporilludis is sexually dimorphic, H. calabyi have two posterior lobes on this plate. Hoplopleura which is seen in H. irritans but not in other Hoplopleura species leporilludis n. sp. has two posterior setae on paratergal plate III, in Australia. The DMHS of H. leporilludis n. sp. are aligned in a whereas H. mastacomydis has one short, stout seta posteriorly. row and offset medially, whereas in H. uromydis, H. villosissima, Hoplopleura leporilludis n. sp. does not have DLAS or VLAS, except

H. irritans, H. leggadinadis, and H. conilurudis they are not aligned for the segment 7 of female, which has 1 pair of VLAS. Hoplopleura Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 in a row. Hoplopleura leporilludis n. sp. has four pairs of DMHS, gyomydis, H. irritans, H. melomydis, H. forrestima, H. leggadinadis, which is different from H. zyzomydis (two pairs), and H. notomydis and H. villosissma have both DLAS and VLAS on segments 4–7; and H. setosa (three pairs). The sternal plate of H. leporilludis however, H. calabyi and H. zyzomydis have VLAS on those seg- n. sp. is sub-hexagonal; however, the sternal plate is shield shaped ments, whereas H. setosa has VLAS on small plates next to the in H. gyomydis, H. irritans, H. mastacomydis, H. villosissima, tergites. The female of H. leporilludis n. sp. has three sternites on H. zyzomydis, H. melomydis, H. conilurudis, and H. forrestima. segments 3–7, whereas H. bidentata has two sternites on the corre- Hoploplura leporilludis n. sp. has a small central seta on paratergal sponding segments. plate II, which is not present in H. pacifica, H. bidentata, H. cornata, H. zyzomydis, and H. notomydis. Paratergal plates IV–VI have one large posterior seta in H. leporilludis n. sp. These three paratergal Hoplopleura mesembriomydis Wang n. sp. plates have two posterior setae in H. uromydis, H. gyomydis, (urn:lsid:zoobank.org:act:4E420990-E8D0-4BE1-9B65- H. irritans, H. mastacomydis, H. melomydis, and H. forrestima BBCED51FE7D6). n. sp., but lack posterior setae in H. notomydis and H. zyzomydis. Male: [n = 3; Fig. 9a] Body length 1011–1232 (1067). Paratergal plate VI has two posterior lobes in H. leporilludis n. sp. Head: Head longer than wide, with 4 ApHS and 4 AnMHS. Pre- but only one in H. villosissima and H. conilurudis. The spiracle antennal region long with pointed anterolateral apex. Antenna with on paratergal plate V of H. leporilludis n. sp. is smaller than in 5 segments, distal seta on dorsal surface of antennal segment 3 not H. uromydis (34–37), H. bidentata (25–28), H. melomydis (male sexually dimorphic. Antennal angle not well developed. Dorsally, head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. DMHS not aligned in row, 2nd, 3rd, and 4th DMHS offset medially. Ventrally, head with 2 VPHS. Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length 80.9–124.96 (102.3). Thoracic sternal plate shield shaped with elongated anterior and posterior processes (Fig. 9b). Forelegs small, with small acuminate claws; midlegs and hindlegs progressively larger with correspondingly more robust tibio-tarsal claws. Abdomen: Wider than thorax. Dorsally, 1 tergite per segment except for segment 3 with 2 tergites. Tergite 1 with 1 pair of TeAS posterolaterally. Tergite 2 with 2 pairs of TeAS posterolaterally, lateral pair shorter. Tergite 3 with 2 pairs of TeAS posterolaterally, lateral pair very long, twice as long as medial pair. Tergites 4–6 each with 5 TeAS, middle TeAS shorter than others on tergite 4. Tergites 7, 8 each with 3 pairs of TeAS. One pair of DLAS adjacent to each of tergites 5–8. Ventrally, 2 sternites per segment except for segment 3 with 3 sternites, and segments 2, 7, 8 each with 1 sternite. Sternite 1 extended laterally to articulate with paratergal plate II and with 4 pairs of StAS. Sternite 2 partially articulating with paratergal plate III and with 7 StAS, lateral 2 pairs long and stout, 3rd pair of setae short, middle StAS longer than 3rd pair. Sternites 3, 7 each with 3 pairs of StAS. Sternites 4–6 and 8–10 each with 5 StAS, sternite 11 with 2 pairs of StAS, sternite 12 with 1 pair of StAS. One pair of VLAS adjacent to each of sternites 8, 10, 11. Paratergites: Paratergal plates (Fig. 9c) present on abdominal segments 1–8. All plates differentially sclerotized. Paratergal plate I small and offset medially. Paratergal plate II with 1 small medial seta, 2 large posterior setae and acuminate posterior lobes. Paratergal plate III with 2 large setae and 2 serrated posterior lobes. Paratergal plates IV–VI each with 2 large setae, equal in length. Paratergal plates IV–VI with 2 serrated posterior lobes. Paratergal

Fig. 9. Hoplopleura mesembriomydis n. sp. Male. (a) Habitus (dorsal mor- plate VII with pointed posterior lobe dorsally. Paratergal plates VII, phology to the left of the midline, ventral morphology to the right). (b) VIII each with 2 long setae. Paratergal plate VIII lacking posterior Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. lobe. Paratergal plates I, II with no spiracle. Spiracles on paratergal Journal of Medical Entomology, 2020, Vol. XX, No. XX 13 plates III–VII large in size, spiracle on paratergal plate V 25.6–34.8 to each of tergites 10, 12, 13, 15, 16. Ventrally, 3 sternites per (31.7) in diameter. Spiracle on paratergal plate VIII much smaller in segment except for segment 2 with 1 sternite and segments 1, 8 size than those on other plates. with no sternites. Sternite 1 extended laterally to articulate with Genitalia (Fig. 9d): Subgenital plate with narrow anterolateral ex- paratergal plate II and with 4 pairs of StAS. Sternite 2 extended tension on each side and 3 lacunae; 2 anterior lacunae smaller than laterally to articulate with paratergal plate III and with 7 StAS, posterior lacuna. Basal apodeme slightly longer than parameres. lateral 2 pairs long and stout, 3rd pair of setae short, medial StAS Parameres uniformly sclerotized, with pseudopenis tapering to point longer than 3rd pair. Sternites 3, 5, 8 each with 7 StAS; sternites extending slightly beyond apices of parameres. 4, 7, 9, 10, 16 each with 4 pairs of StAS; sternites 6, 11–14 each Female: [n = 3; Fig. 10a] Body length 1354–1509 (1460). with 3 pairs of StAS; sternite 15 with 2 pairs of StAS. Sternite 16

Head: Head longer than wide, with 4 ApHS and 4 AnMHS. Pre- with 4 pairs of StAS varying in size, lateral 2 pairs much longer Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 antennal region short with rounded anterior apex. Antenna with 5 than medial 2 pairs. One pair of VLAS adjacent to each of stern- segments, distal seta on dorsal surface of antennal segment 3 not sex- ites 10, 12, 13, 15, 16. ually dimorphic. Antennal angle weak. Dorsally, head with 4 SuHS, Paratergites: Paratergal plates (Fig. 10c) present on abdominal 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. Second, segments 1–8. All plates differentially sclerotized. Paratergal plate 3rd, and 4th DMHS offset medially on each side. Ventrally, head I small and offset medially. Paratergal plates II–VI each with 2 pos- with 2 VPHS. terior lobes. Paratergal plate II with 1 small medial seta, 2 large Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length posterior setae, and acuminate posterior lobes. Paratergal plate III 124.9–140.2 (130.7). Thoracic sternal plate shield shaped with elon- with 2 large setae and 2 serrated posterior lobes. Paratergal plates gate anterior and posterior processes (Fig. 10b). Forelegs small, with IV–VI each with 2 large setae, equal in length. Posterior lobes on small acuminate claws; midlegs and hindlegs progressively larger paratergal plates IV–VI serrated. Paratergal plate VII with pointed with correspondingly more robust tibio-tarsal claws. posterior lobe dorsally. Paratergal plates VII, VIII each with 2 long Abdomen: Wider than thorax. Dorsally, 3 tergites per segment setae. Paratergal plate VIII lacking posterior lobe. Paratergal plates I, except for segments 1, 2, 8 with 1 tergite each; all tergites narrow. II with no spiracle. Spiracle on paratergal plate V 35.8–43 (38.4) in Tergite 1 with 1 pair of TeAS posterolaterally. Tergite 2 with 2 diameter. Spiracle on paratergal plate VIII much smaller in size than pairs of TeAS posterolaterally, lateral pair shorter than other pair. those on other plates. Tergites 3, 5, 17, 18 each with 2 pairs of TeAS. Tergites 4, 6, 8, Genitalia (Fig. 10d): Subtriangular subgenital plate with 5 small 15 each with 5 TeAS. Tergites 7, 9, 11–14, and 16 with 3 pairs mediolateral setae. Gonopods VIII, IX distinct; gonopods VIII of TeAS. Tergite 10 with 7 TeAS. One pair of DLAS adjacent with 3 posterior setae with medial seta shorter than 2 lateral setae; gonopods IX with 3 posterior setae of differing lengths, lateral seta longest and medial seta shortest. Vulvar fimbriae indistinct. Three small setae on each side medial to gonopods IX. Type host. Mesembriomys gouldii (Gray, 1843) (Rodentia: Muridae), black-footed tree rat. Type locality. Caledon Bay, Northern Territory, Australia (12° 51′ S, 136° 33′ E). Type material. Holotype male and allotype female ex Mesembriomys gouldii (MV, DTC349): holotype, ♂, allotype, ♀. Paratypes: 2 ♂ and 2 ♀, same data as for the holotype. Museum accession numbers are linked to the ZooBank record. Distribution. Australia: Northern Territory. Etymology. The species epithet is a noun in apposition referring to the name of the host genus, Mesembriomys. Remarks. The pre-antennal region of H. mesembriomydis n. sp. is pointed anterolaterally, which is different from H. irritans, H. calabyi, and H. uromydis. The anterior pre-antennal region is rounded in H. irritans and H. calabyi, and truncated in H. uromydis. The third antennal segment of H. mesembriomydis n. sp. is not sexually dimorphic, which differentiates it from H. irritans and H. leporilludis. The DMHS are not aligned in a row in H. mesembriomydis n. sp. but are aligned in H. gyomydis, H. mastacomydis, H. melomydis, and H. leporilludis. The sternal plate of H. mesembriomydis n. sp. is shield shaped, different from H. uromydis (less rounded) and H. leporilludis (sub-hexagonal). Paratergal plate II of H. mesembriomydis n. sp. has a small central seta, which is not present in H. pacifica, H. bidentata, H. cornata, H. zyzomydis, or H. notomydis. Paratergal plates IV–VI of H. mesembriomydis n. sp. have two posterior setae and two pointed posterior lobes. These three paratergal plates all have one large seta and one small/minute posterior setae in H. uromydis, H. gyomydis, H. irritans, H. mastacomydis, H. forrestima, and H. melomydis, and Fig. 10. Hoplopleura mesembriomydis n. sp. Female. (a) Habitus (dorsal morphology to the left of the midline, ventral morphology to the right). (b) have one posterior seta in H. setosa, H. villosissimus, H. conilurudis, Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. H. leggadinadis, and H. leporilludis, but lack posterior setae in 14 Journal of Medical Entomology, 2020, Vol. XX, No. XX

H. notomydis and H. zyzomydis. The spiracle on paratergal plate Male: [n = 3; Fig. 11a] Body length 880–1125 (965). V of H. mesembriomydis n. sp. is larger than in H. irritans (12– Head: Head longer than wide, with 4 ApHS and 4 AnMHS. Pre- 16), H. mastacomydis (11–14), H. conilurudis (male 25.1, female antennal region long and rounded anterolaterally. Antenna with 5 27.6), H. leporilludis (male 20.5, female 22.5), H. villosissima segments, distal seta on dorsal surface of antennal segment 3 not sex- (male 18.4, female 20), H. zyzomydis (male 15.8, female 17.6), ually dimorphic. Antennal angle well developed. Dorsally, head with H. notomydis, H. setosa, H. forrestima (male 15.4, female 19.1), 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. H. leggadinadis (male 14.8, female 19.9), H. gyomydis (7–9), and DMHS all aligned in row. Ventrally, head with 2 VPHS. H. calabyi. Paratergal plate VII of H. mesembriomydis n. sp. has one Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length posterior lobe dorsally. Hoplopleura mastacomydis and H. calabyi 101.6–123.5 (115.7). Thoracic sternal plate broadly rounded anteri-

both have two posterior lobes on this plate, whereas H. cornata, orly with elongated posterior process (Fig. 11b). Forelegs small, with Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 H. gyomydis, H. irritans, H. uromydis, H. villosissima, H. bidentata, small acuminate claws; midlegs and hindlegs progressively larger H. pacifica, H. conilurudis, and H. leporilludis lack posterior lobes with correspondingly more robust tibio-tarsal claws. on paratergal plate VII. Hoploplura mesembriomydis n. sp. has two Abdomen: Wider than thorax. Dorsally, 1 tergite per segment ex- posterior setae on paratergal plate III, which differentiate it from cept for segment 3 with 2 tergites. Tergite 1 with 1 pair of TeAS H. mastacomydis, which has one short, stout seta posteriorly. The posterolaterally. Tergite 2 with 2 pairs of TeAS posterolaterally, all male of H. mesembriomydis n. sp. has both DLAS and VLAS on equal in length. Tergite 3 with 2 pairs of TeAS posterolaterally, lat- segments 4–7. Hoplopleura calabyi and H. zyzomydis, however, eral pair distinctly longer that medial pair. Tergite 4 with 7 TeAS, have only one VLAS on the corresponding segments, whereas lateral pair of setae very long. Tergites 5–8 each with 9 TeAS. One H. uromydis, H. mastacomydis, H. notomydis, and H. leporilludis pair of DLAS adjacent to tergite 8. Ventrally, 2 sternites per segment have no VLAS. In comparison with H. mesembriomydis n. sp., except for segment 3 with 3 sternites, segment 2 with 1 sternite, and H. setosa has VLAS only on the small plates next to tergites on segment 1 with no sternites. Sternite 1 extended laterally to artic- those segments. The female of H. mesembriomydis n. sp. has three ulate with paratergal plate II and with 4 pairs of StAS. Sternite 2 sternites on abdominal segments 3–6, whereas H. bidentata has two extended laterally to articulate with paratergal plate III and with 7 sternites on the corresponding segments. StAS, lateral 2 pairs long and stout, medial StAS longer than 3rd pair of StAS. Sternites 3, 6, 8 each with 3 pairs of StAS. Sternites 4, 5, 7, Hoplopleura macrurusa Wang n. sp. 9, 10 each with 7 StAS. Lateral 3rd pair of StAS short on sternite (urn:lsid:zoobank.org:act:5886C27A-24AC-4FA8-A915- 4. Sternite 11 with 2 pairs of StAS. Sternite 12 with 1 pair of StAS. B82C4D362002). One pair of VLAS adjacent to each of sternites 6, 8, 9, 11.

Fig. 11. Hoplopleura macrurusa n. sp. Male. (a) Habitus (dorsal morphology Fig. 12. Hoplopleura macrurusa n. sp. Female. (a) Habitus (dorsal mor- to the left of the midline, ventral morphology to the right). (b) Thoracic sternal phology to the left of the midline, ventral morphology to the right). (b) plate. (c) Paratergal plates. (d) Genitalia. Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. Journal of Medical Entomology, 2020, Vol. XX, No. XX 15

Paratergites: Paratergal plates (Fig. 11c) present on abdominal seg- gonopods IX with 3 posterior setae of differing lengths, lateral seta ments 1–8. All plates differentially sclerotized. Paratergal plate I small longest and medial seta shortest. Vulvar fimbriae indistinct. Three and offset medially. Paratergal plate II with 1 small medial seta, 2 small setae on each side medial to gonopods IX. large posterior setae and acuminate posterior lobes. Paratergal plate Type host. Mesembriomys macrurus (Peters, 1876) (Rodentia: III with 2 large setae and 2 acuminate posterior lobes. Paratergal Muridae), golden-backed tree rat. plates IV–VI each with 1 large seta posteroventrally. Paratergal Type locality. Wyndham-East Kimberley, Western Australia, plates IV, V each with 2 serrated posterior lobes. Paratergal plate VI Australia (14° 36′ S, 125° 54′ E). with 1 acuminate posterior lobe dorsally. Paratergal plates VII, VIII Type material. Holotype male and allotype female ex Mesembriomys each with 2 long setae and lacking posterior lobes. Paratergal plates macrurus (WAM, M21694): holotype, ♂, allotype, ♀. Museum ac-

I, II with no spiracle. Spiracles on paratergal plates III–VII large. cession numbers are linked to the ZooBank record. Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 Spiracle on paratergal plate V 18.5–22.3 (20.9) in diameter. Spiracle Additional material examined. 2 ♂ and 1 ♀ ex Mesembriomys on paratergal plate VIII much smaller than those on other plates. macrurus (WAM, M44975), location unknown. Genitalia (Fig. 11d): Subgenital plate with narrow anterolat- Distribution. Australia: Western Australia. eral extension and 1 lacuna. Basal apodeme almost twice as long Etymology. The species epithet is a noun in apposition referring to as parameres. Parameres uniformly sclerotized, with pseudopenis the name of the host species, Mesembriomys macrurus. tapering to point extending slightly beyond apices of parameres. Remarks. The pre-antennal region of H. macrurusa n. sp. is Female: [n = 2; Fig. 12a] Body length 1320–1452 (1386). rounded anteriorly, which differs from H. uromydis, H. leporilludis, Head: Head longer than wide, with 4 ApHS and 4 AnMHS. Pre- H. conilurudis, H. bidentata, and H. mesembriomydis. The anterior antennal region long and rounded anterolaterally. Antenna with 5 pre-antennal region is truncated in H. uromydis, H. leporilludis, segments, distal seta on dorsal surface of antennal segment 3 not sex- and H. conilurudis, and narrow and pointed in H. bidentata and ually dimorphic. Antennal angle well developed. Dorsally, head with H. mesembriomydis. The third antennal segment of H. macrurusa 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. n. sp. is not sexually dimorphic, which differs from H. irritans and DMHS all aligned in row. Ventrally, head with 2 VPHS. H. leporilludis. The DMHS are aligned in a row in H. macrurusa n. sp. Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length but not in H. uromydis, H. villosissima, H. bidentata, H. irritans, 115.5–135.6 (125.5). Thoracic sternal plate broadly rounded anteri- H. zyzomydis, H. conilurudis, H. forrestima, H. leggadinadis, or orly and with elongated posterior process (Fig. 12b). Forelegs small, H. mesembriomydis. The sternal plate of H. macrurusa n. sp. is with small acuminate claws; midlegs and hindlegs progressively rounded anteriorly, which differs from that of H. uromydis (less larger with correspondingly more robust tibio-tarsal claws. rounded) and H. leporilludis (sub-hexagonal). Paratergal plate II Abdomen: Wider than thorax. Dorsally, 3 tergites per segment except of H. macrurusa n. sp. has a small central seta, which is not pre- for segments 1, 2, 8 with 1 tergite each. Tergite 1 with 1 pair of TeAS sent in H. pacifica, H. bidentata, H. cornata, H. zyzomydis, or posterolaterally. Tergite 2 with 2 pairs of TeAS posterolaterally, equal H. notomydis. Paratergal plates IV–VI each have one large poste- in length. Tergite 3 with 2 pairs of TeAS posterolaterally, medial pair rior seta in H. macrurusa n. sp. These three paratergal plates each twice as long as lateral pair. Tergites 4, 7, 9, 10, 15 each with 3 pairs have two posterior setae in H. uromydis, H. gyomydis, H. irritans, of TeAS. Tergites 5, 8, 14, 17, 18 each with 2 pairs of TeAS. Tergites H. mastacomydis, H. forrestima, H. melomydis, H. notomydis, 6, 11, 13, 16 each with 5 TeAS. Tergite 12 with 7 TeAS. One pair and H. mesembriomydis, but no posterior seta in H. zyzomydis. of DLAS adjacent to each of tergites 10, 12, 13, 15, 16. Ventrally, 3 Hoplopleura macrurusa n. sp. does not have a posterior lobe on sternites per segment except for segment 3 with 4 sternites, segments paratergal plate VII, but H. forrestima, H. leggadinadis, H. setosa, 2, 7 each with 1 sternite, and segments 1, 8 each with no sternites. H. melomydis, H. zyzomydis, and H. mesembriomydis all have one Sternite 1 extended laterally to articulate with paratergal plate II and posterior lobe, and H. mastacomydis and H. calabyi all have two with 4 pairs of StAS. Sternite 2 extended laterally to articulate with posterior lobes on paratergal plate VII. Hoplopleura macrurusa paratergal plate III and with 7 StAS, lateral 2 pairs long and stout, n. sp. has two posterior setae on paratergal plate III, whereas 3rd pair of StAS short, medial StAS twice as long as 3rd pair of StAS. H. mastacomydis has one short, stout posterior seta on that plate. Sternites 3, 6, 7, 9, 10, 12, 13, 15 each with 3 pairs of StAS. Sternites The spiracle on paratergal plate V of H. macrurusa n. sp. is me- 4, 5, 8 each with 7 setae. Third pair of StAS on sternite 5 short. dium in size (male 20.9, female 22.7), smaller than in H. uromydis Sternites 11, 14 each with 5 StAS. Sternite 16 with 4 pairs of StAS (34–37), H. bidentate (25–28), H. melomydis (male 25, female 25.8), varying in size, lateral 2 pairs of StAS long, medial 2 pairs very small. and H. mesembriomydis (male 31.7, female 38.4), but larger than in One pair of VLAS adjacent to each of sternites 8, 10, 13, 15, 16. H. gyomydis (7–9). The posterior lobes on paratergal plates III–VII Paratergites: Paratergal plates (Fig. 12c) present on abdominal seg- of H. macrurusa n. sp. are pointed and unilobate, which is distinct ments 1–8. All plates differentially sclerotized. Paratergal plate from other Australian Hoplopleura species, except for H. setosa and I small and offset medially. Paratergal plates II–V each with 2 pos- M. mesembriomydis. The male of H. macrurusa n. sp. only has one terior lobes. Paratergal plate II with 1 small medial seta, 2 large pos- pair of DLAS on segment 7, which differentiates it from all other terior setae and acuminate posterior lobes. Paratergal plate III with Hoplopleura species in Australia. The female of H. macrurusa n. sp. 2 large posterior setae and 2 acuminate posterior lobes. Paratergal has three sternites on segment 3–6, whereas H. bidentata has two plates IV–VI each with 1 large seta posteroventrally. Paratergal plates sternites on these segments. IV, V each with 2 serrated posterior lobes. Paratergal plate VI with 1 acuminate posterior lobe dorsally. Paratergal plates VII, VIII each with 2 long posterior setae and lacking posterior lobes. Paratergal Hoplopleura pogonomydis Wang n. sp. plates I, II with no spiracle. Spiracle on paratergal plate V 20.2–25.3 (urn:lsid:zoobank.org:act:297BA919-4610-4D5D-BA2F- (22.7) in diameter. Spiracle on paratergal plate VIII small in size. 19CC924EEDBE). Genitalia (Fig. 12d): Genitalia with subquadrangular subgenital Male: [n = 2; Fig. 13a] Body length 735–816 (775.5). plate with 2 small mediolateral setae on each side. Gonopods VIII, Head: Head slightly longer than wide, with 4 ApHS and 4 AnMHS. IX distinct; gonopods VIII with 3 posterior setae subequal in size; Pre-antennal region short, rounded anterolaterally. Antenna with 5 16 Journal of Medical Entomology, 2020, Vol. XX, No. XX segments, distal seta on dorsal surface of antennal segment 3 not sex- Paratergites: Paratergal plates (Fig. 13c) present on abdominal ually dimorphic. Antennal angle well developed. Dorsally, head with segments 1–8. All plates differentially sclerotized. Paratergal plate 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. I small and offset medially. Paratergal plates II–V each with 2 poste- Second, 3rd, and 4th DMHS offset medially on each side. Ventrally, rior lobes. Paratergal plate II with 1 small medial seta, 2 large poste- head with 2 VPHS. rior setae and 2 acuminate posterior lobes. Paratergal plate III with 2 Thorax: Wider than long, with 1 pair of DPTS dorsally, short in large posterior setae and 2 serrated posterior lobes. Paratergal plates length, DPTS length 67.6–81.9 (74.7). Thoracic sternal plate IV–VI each with 1 large seta posterodorsally. Paratergal plates IV–VI broadly rounded anteriorly with elongate posterior process each with 2 serrated posterior lobes. Paratergal plate VII with 2 long (Fig. 13b). Forelegs small, with small acuminate claws; midlegs and posterior setae and 1 posterior lobe dorsally. Paratergal plate VIII

hindlegs progressively larger with correspondingly more robust with 2 long posterior setae but no posterior lobe. Paratergal plates Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 tibio-tarsal claws. I, II, VIII with no spiracle. Spiracle on paratergal plate V 13.3–13.7 Abdomen: Wider than thorax. Dorsally, 1 tergite per segment except (13.5) in diameter. for segment 3 with 2 tergites, and segments 1, 8 each with no tergites. Genitalia (Fig. 13d): Subgenital plate with narrow anterolateral Tergite 1 with 1 pair of TeAS posterolaterally. Tergite 2 with 2 pairs of extension and 2 lacunae on each side; anterior lacunae larger than TeAS posterolaterally, equal in length. Tergite 3 with 2 pairs of TeAS posterior lacunae. Basal apodeme slightly longer than parameres. posterolaterally, lateral pair long. Tergite 4 with 4 pairs of TeAS, lat- Parameres uniformly sclerotized, with pseudopenis tapering to eral pair of setae longer than others. Tergites 5–7 each with 9 TeAS. pointed apex extending beyond apices of parameres. Tergite 8 with 4 pairs of TeAS. One pair of DLAS adjacent to each of Female: [n = 2; Fig. 14a] Body length 906–1048 (977). tergites 5–8. Ventrally, 2 sternites per segment except for segment 3 Head: Head about as long as wide, with 4 ApHS and 4 AnMHS. Pre- with 3 sternites, segment 2 with 1 sternite, and segments 1, 8 each with antennal region very short, rounded anterolaterally. Antenna with no sternites. Sternite 1 extended laterally to articulate with paratergal 5 segments. Antennal angle well developed. Dorsally, head with 4 plate II and with 4 pairs of StAS. Sternite 2 extended laterally to ar- SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and 8 DMHS. ticulate with paratergal plate III and with 7 StAS, lateral 2 pairs long Second, 3rd, and 4th DMHS offset medially on each side. Ventrally, and stout. Sternite 3 with 4 pairs of StAS, lateral two pairs longer than head with 2 VPHS. others. Sternites 4, 6, 8–10 each with 7 StAS. Sternites 5, 7 each with 4 Thorax: Wider than long, with 1 pair of DPTS dorsally, short in length, pairs of StAS. Sternite 11 with 2 pairs of StAS. Sternite 12 with 1 pair DPTS length 78.9–91.1 (85). Thoracic sternal plate broadly rounded of StAS. One pair of VLAS adjacent to each of sternites 6, 7, 9–11. anteriorly with elongate posterior process (Fig. 14b). Forelegs small,

Fig. 13. Hoplopleura pogonomydis n. sp. Male. (a) Habitus (dorsal mor- Fig. 14. Hoplopleura pogonomydis n. sp. Female. (a) Habitus (dorsal morphology to the left of the midline, ventral morphology to the right). (b) phology to the left of the midline, ventral morphology to the right). (b) Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. Journal of Medical Entomology, 2020, Vol. XX, No. XX 17 with small acuminate claws; midlegs and hindlegs progressively H. notomydis, H. setosa, H. gyomydis, H. mastacomydis, larger with correspondingly more robust tibio-tarsal claws. H. leporilludis, and H. melomydis. The DMHS are aligned in Abdomen: Wider than thorax. Dorsally, 3 tergites per segment, ex- a row in H. gyomydis, H. mastacomydis, H. leporilludis, and cept for segments 1, 2 with 1 tergite each. Tergite 1 with 1 pair of H. melomydis. In H. uromydis, H. villosissima, and H. irritans, the TeAS posterolaterally. Tergite 2 with 2 pairs of TeAS posterolaterally, 2nd and 3rd DMHS are offset medially. Hoplopleura zyzomydis has equal in length. Tergite 3 with 2 pairs of TeAS posterolaterally, lat- two pairs of DMHS, whereas H. notomydis and H. setosa, have eral pair longer than others. Tergites 4, 11, 14, 15, 17 each with 7 three pairs of DMHS. The sternal plate of H. pogonomydis n. sp. TeAS. Tergites 5–10, 12, 13, 16, 18 each with 3 pairs of TeAS. Tergite is broadly rounded anteriorly; in H. uromydis and H. leporilludis, 19 with 2 pairs of TeAS; tergite 20 with 1 pair of TeAS; tergite 10 the sternal plate is less rounded anteriorly. Paratergal plate II of

with 4 pairs of TeAS. One pair of DLAS adjacent to each of tergites H. pogonomydis n. sp. has a small central seta, which is not pre- Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 11–18. Ventrally, 3 sternites per abdominal segment except for seg- sent in H. pacifica, H. bidentata, H. cornata, H. zyzomydis, or ment 2 with 1 sternite, and segments 1, 8 each with no sternites. H. notomydis. Paratergal plates IV to VI have one large posterior Sternite 1 extended laterally to articulate with paratergal plate II and seta dorsally in H. pogonomydis n. sp. These three paratergal plates with 4 pairs of StAS, lateral 2 pairs of StAS long and stout. Sternites have two posterior setae in H. uromydis, H. gyomydis, H. irritans, 2, 8, 10, 11, 13 each with 7 setae. Sternites 3–5, 7, 9, 12, 14, 16 each H. mastacomydis, H. forrestima, H. melomydis, H. notomydis, and with 4 pairs of StAS. StAS on sternite 16 varying in size. Sternite 6 H. mesembriomydis, but lack posterior setae in H. notomydis and with 9 StAS. Sternite 15 with 3 pairs of StAS. One pair of VLAS ad- H. zyzomydis. A dorsal posterior lobe is present on paratergal plate jacent to each of sternites 8–16. VII in H. pogonomydis n. sp. but not in H. cornata, H. gyomydis, Paratergites: Paratergal plates (Fig. 14c) present on abdominal H. irritans, H. uromydis, H. villosissima, H. bidentata, H. pa- segments 1–8. All plates differentially sclerotized. Paratergal plate cifica, H. conilurudis, H. leporilludis, or H. macrurusa. However, I small and offset medially. Paratergal plates II–V each with 2 poste- H. mastacomydis and H. calabyi have two posterior lobes on the rior lobes. Paratergal plate II with 1 small medial seta, 2 large poste- corresponding plate. Hoplopleura pogonomydis n. sp. has two rior setae and 2 acuminate posterior lobes. Paratergal plate III with 2 posterior setae on paratergal plate III, whereas H. mastacomydis large posterior setae and 2 serrated posterior lobes. Paratergal plates has one short, stout seta posteriorly on that plate. The spiracle on IV–VI each with 1 large seta posterodorsally and 2 serrated posterior paratergal plate V of H. pogonomydis n. sp. is small in size (male lobes. Paratergal plate VII with 2 long posterior setae and 1 posterior 13.5, female 15.4), smaller than in H. villosissima (male 18.4, fe- lobe dorsally. Paratergal plate VIII with 2 long posterior setae but no male 20), H. zyzomydis (male 15.8, female 17.6), H. conilurudis posterior lobes. Paratergal plates I, II, VIII with no spiracle. Spiracles (male 25.1, female 27.6), H. forrestima (male 15.4, female 19.1), on paratergal plates III–VII small in size. Spiracle on paratergal plate H. leggadinadis (male 14.8, female 19.9), H. uromydis (34–37), V 14.3–16.7 (15.4) in diameter. H. bidentata (25–28), H. melomydis (male 25, female 25.8), Genitalia (Fig. 7d): Genitalia with subtriangular subgenital plate H. leporilludis (male 20.5, female 22.5), H. macrurusa (male 20.9, with 2 small mediolateral setae on each side. Gonopods VIII, IX female 22.7), and H. mesembriomydis (male 31.7, female 38.4). The distinct; gonopods VIII with 3 posterior setae subequal in size; male of H. pogonomydis n. sp. has one pair of DLAS on abdominal gonopods IX with 3 posterior setae of differing lengths, lateral seta segments 4–7; H. macrurusa, however, has only one pair of DLAS on longest and medial seta shortest. Vulvar fimbriae indistinct. Three segment 7. The male of H. pogonomydis n. sp. has one pair of VLAS small setae on each side medial to gonopods IX. adjacent to the anterior and posterior sternites of segments 5 and Type host. Pogonomys mollipilosus (Peters and Doria, 7 but has no VLAS on segment 6, which is different from all other 1881) (Rodentia: Muridae), prehensile-tailed rat. Hoplopleura species in Australia. The female H. pogonomydis n. sp. Type locality. Shiptons Flat, Queensland, Australia (15° 48′ S, 145° has three sternites on segments 3–6, whereas H. bidentata has two 00′ E). sternites on those segments. Type material. Holotype male and allotype female ex Pogonomys mollipilosus (QM, JM10590): holotype, ♂, allotype, ♀. Paratype: 1 ♀, same data as for the holotype. Museum accession numbers are Hoplopleura xeromydis Wang n. sp. linked to the ZooBank record. (urn:lsid:zoobank.org:act:57899858-7D28-4116-B19E- Additional material examined. 1 ♂ ex Pogonomys mollipilosus 3639B4658D2C). (QM, JM14682), Shiptons Flat, Queensland (15° 48′ S, 145° 00′ E). Male: [n = 1; Fig. 15a] Body length 845. Distribution. Australia: Queensland. Head: Head about as long as wide, with 4 ApHS and 4 AnMHS. Etymology. The species epithet is a noun in apposition referring to Pre-antennal region long, slightly pointed anterolaterally. Antenna the name of the host genus, Pogonomys. with 5 segments, distal seta on dorsal surface of antennal segment Remarks. The pre-antennal region of H. pogonomydis n. sp. 3 not sexually dimorphic. Antennal angle well developed. Dorsally, is rounded anteriorly, which is different from H. uromydis, head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS, and H. leporilludis, H. conilurudis, H. leggadinadis, H. mastacomuydis, 8 DMHS. Second, 3rd and 4th DMHS offset medially on each side. H. bidentata, and H. mesembriomydis. The anterior pre-antennal Ventrally, head with 2 VPHS. region is truncated in H. uromydis, H. leporilludis, H. conilurudis, Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length and H. leggadinadis; it is less rounded in H. mastacomuydis, and 73. Thoracic sternal plate shield shaped, broadly rounded anteriorly, narrow and pointed in H. bidentata and H. mesembriomydis. The and with elongated posterior process (Fig. 16b). Forelegs small, with third antennal segment of H. pogonomydis n. sp. is not sexually di- small acuminate claws; midlegs and hindlegs progressively larger morphic, unlike in H. irritans and H. leporilludis. The DMHS of with correspondingly more robust tibio-tarsal claws. H. pogonomydis n. sp. are not aligned in a row. The most anterior Abdomen: Wider than thorax. Dorsally, 1 tergite per abdominal seg- DMHS is at the top of the postantennal angle, and the other DMHS ment except for segment 3 with 2 tergites. Tergite 1 with 1 pair of are offset medially in H. pogonomydis n. sp., which differentiates TeAS posterolaterally, short in length. Tergite 2 with 2 pairs of TeAS it from H. uromydis, H. villosissima, H. irritans, H. zyzomydis, posterolaterally, lateral pair longer than other TeAS. Tergite 3 with 18 Journal of Medical Entomology, 2020, Vol. XX, No. XX

2 pairs of TeAS posterolaterally, lateral pair very long. Tergites 4–6 23.5 in diameter. Spiracle on paratergal plate VIII distinctly smaller each with 9 TeAS, posterolateral 2 pairs of TeAS longest; other TeAS than spiracles on plates III–VII. pairs varying in length with middle TeAS shortest. Tergite 7 with 4 Genitalia (Fig. 15d): Subgenital plate with narrow anterolateral ex- pairs of TeAS, posterolateral pair long and stout, other 3 pairs short tension on each side and 2 lacunae. Basal apodeme slightly longer and equal in length. Tergite 8 with 3 pairs of TeAS. One pair of than parameres. Parameres uniformly sclerotized with pseudopenis DLAS adjacent to tergite 8. Ventrally, 2 sternites per segment except tapering to point slightly beyond apices of parameres. for segments 2, 8 with 1 sternite each, and segment 1 with no stern- Female: [n = 1; Fig. 16a] Body length 1055. ites. Sternite 1 extended laterally to articulate with paratergal plate Head: Head about as long as wide, with 4 ApHS and 4 AnMHS. II and with 4 pairs of StAS, posterolateral two pairs long and stout. Pre-antennal region long, slightly pointed anterolaterally. Antenna

Sternite 2 extended laterally to articulate with paratergal plate III with 5 segments, distal seta on dorsal surface of antennal segment Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 and with 7 StAS, posterolateral 2 pairs long and stout, medial StAS 3 not sexually dimorphic. Antennal angle well developed. Dorsally, longer than 3rd pair of StAS on this sternite. Sternite 3 with 4 pairs head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS and 8 of StAS, posterolateral 2 pairs long and stout. Sternites 4, 6, 8 each DMHS. Second, 3rd and 4th DMHS shifting medially on each side. with 7 StAS, posterolateral 2 pairs StAS long and stout. Sternites 5, 7 Ventrally, head with 2 VPHS. each with 4 pairs of StAS. Sternites 9, 10 each with 3 pairs of StAS. Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length Sternite 11 with 2 pairs of StAS; sternite 12 with 1 pair of StAS. One 90.1. Thoracic sternal plate shield-shaped, broadly rounded anteri- pair of VLAS adjacent to each of sternites 6, 8, 9, 11. orly and with elongated posterior process. Forelegs small, with small Paratergites: Paratergal plates (Fig. 8c) present on abdominal seg- acuminate claws; midlegs and hindlegs progressively larger with cor- ments 1–8. All plates differentially sclerotized. Paratergal plate respondingly more robust tibio-tarsal claws, midlegs nearly twice as I small and offset medially. Paratergal plates II–VI each with 2 poste- long as forelegs. rior lobes. Paratergal plate II with 1 small medial seta, 2 large poste- Abdomen: Wider than thorax. Dorsally, 3 tergites per segment ex- rior setae, and 2 acuminate posterior lobes. Paratergal plate III with cept for segments 1, 2, 8 each with 1 tergite. All tergites narrow. 2 large posterior setae and 2 serrated posterior lobes. Paratergal Tergite 1 with 1 pair of TeAS posterolaterally. Tergite 2 with 2 pairs plates IV–VI each with 1 large seta posterodorsally, paratergal plates IV, V each with 2 serrated posterior lobes, paratergal plate VI with 2 serrated undivided acuminate posterior lobes. Paratergal plates VII, VIII each with 2 long posterior setae but no posterior lobes. Paratergal plates I, II with no spiracle. Spiracle on paratergal plate V

Fig. 15. Hoplopleura xeromydis n. sp. Male. (a) Habitus (dorsal morphology Fig. 16. Hoplopleura xeromydis n. sp. Female. (a) Habitus (dorsal mor- to the left of the midline, ventral morphology to the right). (b) Thoracic sternal phology to the left of the midline, ventral morphology to the right). (b) plate. (c) Paratergal plates. (d) Genitalia. Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. Journal of Medical Entomology, 2020, Vol. XX, No. XX 19 of TeAS posterolaterally, equal in length. Tergite 3 with 2 pairs of DMHS on each side; H. zyzomydis has only two DMHS on each long TeAS posterolaterally, equal in length. Tergite 4 with 7 TeAS, side, whereas H. notomydis and H. setosa each have three DMHS 2nd pair of lateral TeAS long. Tergites 5, 7, 10, 13, 15 each with 3 on each side. The sternal plate of H. xeromydis n. sp. is shield pairs of TeAS. Tergites 6, 8, 9, 12, 14 each with 7 TeAS. Tergites 11, shaped which differentiates it from H. uromydis (less rounded) and 17, 18 each with 5 TeAS. Tergite 16 with 2 pairs of TeAS. One pair H. leporilludis (sub hexagonal). Paratergal plate II of H. xeromydis of DLAS adjacent to each of tergites 9, 13, 15. Ventrally, 3 sternites n. sp. has a small central seta, which is not present in H. pacifica, per segment except for segment 2 with 1 sternite, segment 7 with H. bidentata, H. cornata, H. zyzomydis, or H. notomydis. Paratergal 2 sternites and segments 1, 8 each with no sternites. Sternite 1 ex- plates IV–VI of H. xeromydis n. sp. each have one large posterior tended laterally to articulate with paratergal plate II and with 4 pairs seta. Two posterior setae are adjacent to these plates in H. uromydis,

of StAS. Sternite 2 extended laterally to articulate with paratergal H. gyomydis, H. irritans, H. mastacomydis, H. forrestima, Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 plate III and with 7 StAS, lateral 2 pairs long and stout, middle StAS H. melomydis, H. notomydis, and H. mesembriomydis, whereas no longer than 3rd pair of StAS. Sternites 3–9, 12, 14 each with 3 pairs posterior seta is adjacent to these plates in H. zyzomydis. Paratergal of StAS. Sternites 10, 11, 13 each with 7 StAS. Sternite 15 with 4 plate VII of H. xeromydis n. sp. has no posterior lobe, compared pairs of StAS, varying in size, lateral two pairs much longer than with one posterior lobe on that plate in H. forrestima, H. setosa, other pairs. One pair of VLAS adjacent to each of sternites 8, 11, 15. H. melomydis, H. zyzomydis, H. mastacomydis, H. leggadinadis, and Paratergites: Paratergal plates (Fig. 8c) present on abdominal seg- H. mesembriomydis, and two posterior lobes in H. mastacomydis ments 1–8. All plates differentially sclerotized. Paratergal plate and H. calabyi. Hoplopleura xeromydis n. sp. has two setae on I small and offset medially. Paratergal plates II–VI each with 2 poste- paratergal plate III, whereas H. mastacomydis has one short, stout rior lobes. Paratergal plate II with 1 small medial seta, 2 large poste- seta posteriorly. The spiracle on paratergal plate V of H. xeromydis rior setae, and 2 acuminate posterior lobes. Paratergal plate III with n. sp. is large (male 23.5, female 26.6), larger than those in H. irritans 2 large posterior setae and 2 serrated posterior lobes. Paratergal (12–16), H. mastacomydis (11–14), H. leporilludis (male 20.5, fe- plates IV–VI each with 1 large seta posterodorsally, paratergal plates male 22.5), H. villosissima (male 18.4, female 20), H. zyzomydis IV, V each with 2 serrated posterior lobes, paratergal plate VI with (male 15.8, female 17.6), H. forrestima (male 15.4, female 19.1), 2 serrated undivided acuminate posterior lobes. Paratergal plates H. leggadinadis (male 14.8, female 19.9), H. gyomydis (7–9), and VII, VIII each with 2 long posterior setae but no posterior lobe. H. pogonomydis (male 13.5, female 15.4). The male H. xeromydis Paratergal plates I, II with no spiracle. Spiracles on paratergal plates n. sp. has one DLAS only on segment 7, unlike all other Hoplopleura III–VII large in size. Spiracle on paratergal plate V 26.6 in diameter. species in Australia except for H. macrurusa. The female of No spiracle on paratergal plates I, II. Spiracle on plate VIII distinctly H. xeromydis n. sp. has three sternites on abdominal segments 3–6, smaller than spiracles on plates III–VII. whereas H. bidentata has two sternites on these segments. Genitalia (Fig. 16d): Genitalia with subtriangular subgenital plate with broadly rounded anterolateral and posterior margins and 3 Discussion small mediolateral setae on each side. Gonopods VIII, IX distinct; gonopods VIII with 3 posterior setae subequal in size; gonopods IX We describe eight new species of sucking lice belonging to the genus with 3 posterior setae of differing lengths, lateral seta longest and Hoplopleura from six genera of Old Endemic rodents in Australia: medial seta shortest. Vulvar fimbriae indistinct. Three small setae on Conilurus, Leggadina, Leporillus, Mesembriomys, Pogonomys, each side medial to gonopods IX. and Xeromys. The new species of sucking lice, H. conilurudis, Type host. Xeromys myoides (Thomas, 1889) (Rodentia: Muridae), H. leggadinadis, H. forrestima, H. leporilludis, H. mesembriomydis, false water rat. H. macrurusa, H. pogonomydis, and H. xeromydis, are similar in Type locality. Ramingining Area, Arafura Swamp, East Arnhem, overall morphology to each other and with the 12 Hoplopleura Northern Territory, Australia (12° 12′ S, 134° 59′ E). species described previously from endemic rodents in Australia. Type material. Holotype male ex Xeromys myoides (MAGNT, However, these eight new species are distinct from each other and U4744): holotype, ♂. Museum accession number is linked to the from other previously described species based on a combination of ZooBank record. detailed key morphological characters, which allow us to provide Additional material examined. 1 ♀ ex Xeromys myoides (QM, an updated identification key for the Australian Hoplopleura species JM9480), Myora, N Stradbroke Is, Queensland, Australia (27° 27′ (Table 3). S, 153° 25′ E). We collected only two adult louse specimens from the false water Distribution. Australia: Northern Territory and Queensland. rat, Xeromys myoides: a male from a rat specimen collected in the Etymology. The species epithet is a noun in apposition referring to East Arnhem, Northern Territory, and a female from a rat specimen the name of the host genus, Xeromys. collected on North Stradbroke Island, Queensland (Table 2). The Remarks. The pre-antennal region of H. xeromydis n. sp. is slightly false water rat is a vulnerable species with only 34 specimens stored pointed anterolaterally. However, H. uromydis, H. leporilludis, in museums across Australia (from https://www.ala.org.au). These H. conilurudis, and H. leggadinadis have a truncated pre-antennal re- two louse specimens from different localities (>3,000 km apart) gion anterolaterally, whereas in H. irritans, H. calabyi, H. macrurusa, share six morphological characters (Figs. 15 and 16): 1) the head and H. pogonomydis, it is rounded. The third antennal segment of pre-antennal region slightly pointed anterolaterally; 2) the position H. xeromydis n. sp. is not sexually dimorphic, which is different of DMHS on each side of head not aligned in a row with the 2nd, from H. irritans and H. leporilludis. The DMHS of H. xeromydis 3rd, and 4th DMHS offset medially; 3) the thoracic sternal plate n. sp. are not aligned in a row. The most anterior DMHS is on the top shield-shaped, broadly rounded anteriorly with elongated posterior of the postantennal angle and the 2nd, 3rd, and 4th DMHS are offset process; 4) only one posterior seta each on paratergal plates IV–VI; medially in H. xeromydis n. sp. The DMHS are aligned in a row in 5) the posterior lobes are serrated on paratergal plates IV and V; 6) H. gyomydis, H. mastacomydis, H. leporilludis, and H. melomydis. the posterior lobe is pointed on paratergal plate VI; and 7) no poste- In H. uromydis, H. villosissima and H. irritans, only the 2nd and rior lobes on paratergal plates VII–VIII. We describe these two spe- 3rd DMHS are offset medially. Hoploplura xeromydis n. sp. has four cimens as the same species, Hoplopleura xeromydis, based on these 20 Journal of Medical Entomology, 2020, Vol. XX, No. XX

Table 3. Key for the identification of sucking lice in the genusHopopleura parasitizing native rodents in Australia

No. Key characters Species

1 Distal seta on dorsal surface of antennal segment 3 markedly sexually dimorphic; tergites and sternites narrow. 2 Distal seta on dorsal surface of antennal segment 3 not, or only slightly, sexually dimorphic. 3 2 Abdominal segments with DLAS and VLAS; tergites and sternites narrow; on Rattus lutreolus and Rattus H. irritans Kuhn & fuscipes. Ludwig, 1967 Abdominal segments without DLAS and VLAS, except 1 pair of VLAS present on segment 7 in female; tergites H. leporilludis n. sp. and sternites wide; on Leporillus conditor. 3 Head with 8 DMHS in both ♂ and ♀. 6 Head with DMHS number less than 8. 4 Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 4 Head with same number DMHS in both ♂ and ♀. 5 Head with 6 DMHS in ♂ and 8 DMHS in ♀; on Notomys alexis. H. notomydis Weaver 2017 5 Head with 4 DMHS in both ♂ and ♀; paratergites of segments 4 to 6 with no posterior setae; on Zyzomys H. zyzomydis argurus. Weaver and Barton 2008 Head with 6 DMHS in both ♂ and ♀; abdominal segments 4 to 7 each with VLAS on small plates next to the H. setosa Weaver tergites; on Notomys alexis. 2017 6 Thoracic sternal plate without an elongated anterior process. 7 Thoracic sternal plate with an elongated anterior and posterior process. 8 7 Abdominal segments with 1 pair of DLAS in ♂; on Mesembriomys macrurus. H. macrurus n. sp. Abdominal segments with 4 pairs of DLAS in ♂; on Pogonomys mollipilosus. H. pogonomydis n. sp. 8 Paratergite of abdominal segment 2 without central setae. 9 Paratergite of abdominal segment 2 with 1 pair of tiny central setae. 10 9 Abdominal segments 4 to 6 with 2 sternites in ♀; on Hydromys chrysogaster. H. bidentata (Neu- mann 1909) Abdominal segments 4 to 6 with 3 sternites in ♀; on Rattus sordidus. H. cornata Kim 1972 10 Paratergite of abdominal segment 8 without distinct posterior lobe. 11 Paratergite of abdominal segment 8 with distinct posterior lobe; paratergite of segment 3 with 1 short apical H. mastacomydis seta; on Mastacomys fuscus. Kuhn & Ludwig, 1967 11 Abdominal spiracles large, diameter greater than 0.030 mm. 12 Abdominal spiracles small, diameter less than 0.030 mm. 13 12 Paratergites of abdominal segments 4 to 6 each with 2 large posterior setae and 2 pointed posterior lobes; on H. mesembriomydis Mesembriomys gouldii. n. sp. Paratergites of abdominal segments 4 to 6 each with 1 large and 1 tiny posterior seta and 2 slightly bilobate H. uromydis Kuhn posterior lobes; on Uromys caudimaculatus. & Ludwig, 1967 13 Thoracic sternal plate heart shaped and with reduced posterior process; DMHS aligned in a row; on H. gyomydis Kuhn Pseudomys gyomys. & Ludwig, 1967 Thoracic sternal plate not heart shaped with elongate anterior and posterior process. 14 14 Paratergite of abdominal segments 6 with 2 posterior lobes. 15 Paratergite of abdominal segments 6 with 1 posterior lobe. 19 15 Paratergites of abdominal segments 4 and 5 each with 2 serrated post4erior lobes, segment 6 with 2 pointed H. xeromydis n. sp. posterior lobes; paratergites of abominal segments 4 to 6 each with 1 large posterior seta dorsally; only segment 8 with 1 pair of DLAS in ♂ on Xeromys myoides. Paratergites of abdominal segments 4 and 5 each with 2 serrated posterior lobes. 16 16 Paratergites of abdominal segments 4 to 6 each with 1 large posterior seta; paratergite of abdominal segment 7 H. leggadinadis with 1 posterior lobe dorsally. DMHS not aligned in a row; on Leggadina lakedownensis. n. sp. Paratergites of abdominal segments 4 to 6 each with 1 large and 1 tiny posterior seta. 17 17 No DLAS on abdominal segments in ♂; thoracic sternal plate less round with elongate anterior and posterior H. calabyi Johnson process; paratergite of abdominal segment 7 with 1 ventral posterior lobe; paratergites of abdominal seg- 1960 ments 4 to 6 with 1 large and 1 tiny posterior seta; on Pseudomys higginsi. DLAS present on abdominal segments in ♂. 18 18 Paratergite of abdominal segment 7 with 1 posterior lobe dorsally; paratergites of abdominal segments 4 to 6 H. melomydis each with 2 serrated posterior lobes; paratergites of abdominal segments 4 to 6 each with 1 large and 1 tiny Weaver 2017 posterior seta; DLAS only present on abdominal segment 6 in ♂; DMHS all aligned in a row; on Melomys burtoni and Melomys capensis. Paratergite of abdominal segment 7 with no posterior lobes; paratergites of abdominal segment 4 and 5 H. forrestima n. sp. each with 2 serrated posterior lobes; paratergite of abdominal segment 6 with 2 pointed posterior lobes; paratergites of abdominal segments 4 to 6 each with 1 large and 1 tiny posterior seta. DMHS not aligned in a row; on Leggadina forresti. 19 Paratergite of abdominal segment 5 with 2 pointed posterior lobes; paratergite of abdominal segment 6 with 1 H. conilurudis n. sp. pointed posterior lobe dorsally; paratergites of abdominal segments 4 to 6 each with 1 large posterior seta; 2 pairs of DLAS adjacent to tergites 10, 13, 16 and 2 pairs of VLAS adjacent to sternite 15 in female; on Conilurus penicillatus. Paratergites of abdominal segments 4 and 5 each with 2 divided posterior lobes; paratergite of abdominal segment H. villosissima 6 with 1 pointed posterior lobe dorsally; paratergite of abdominal segments 4 to 6 each with 1 large posterior Wang 2018 seta; DLAS present on abdominal segments 5 to 8 in ♂ DMHS not aligned in a row; on Rattus villosissimus. Journal of Medical Entomology, 2020, Vol. XX, No. XX 21 shared morphological characters. We designated the male louse spec- 3) tergites 5–8 each with one pair of adjacent DLAS in male imen as the holotype and the female louse specimen as additional H. mesembriomydis, whereas only tergite 8 has a pair of adjacent material, not as the allotype, due to the fact that the male and the DLAS in male H. macrurusa; 4) VLAS adjacent to sternites 8, 10, female specimens were collected from two different rat specimens in and 11 in male H. mesembriomydis but adjacent to sternites 6, different localities. As stated earlier, DNA for genetic comparisons 8, 9, and 11 in male H. macrurusa; 5) DLAS adjacent to tergites could not be recovered from these specimens due to previous expo- 10, 12, 13, 15, and 16 in female H. mesembriomydis but adjacent sure of the host rodents to formalin. to tergites 10, 12, 13, 15, 16, and 17 in female H. macrurusa; Among the eight Hoplopleura species described in the present 6) VLAS adjacent to sternites 10, 12, 13, 15, and 16 in female study, H. leggadinadis and H. forrestima are morphologically most H. mesembriomydis but adjacent to sternites 8, 10, 13, 15, and

similar to each other but still have sufficient differences that allow 16 in female H. macrurusa; 7) paratergal plates IV–VI each with Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa206/5962924 by guest on 13 November 2020 us to recognize them as separate species (Figs. 3–6): 1) the poste- two equally long posterior setae in H. mesembriomydis but only rior abdomen is wide and blunt in H. leggadinadis but narrow and one seta in H. macrurusa; 8) paratergal plate VI with two pos- pointed in H. forrestima; 2) males of H. leggadinadis have one pair terior lobes in H. mesembriomydis but one acuminate posterior of DLAS, whereas males of H. forrestima have four pairs of DLAS; lobe dorsally in H. macrurusa; and 9) spiracle on paratergal plate 3) females of H. leggadinadis have five pairs of DLAS, whereas fe- V in H. mesembriomydis larger (31.7 for male, 38.4 for female) males of H. forrestima have six pairs of DLAS; 4) H. leggadinadis than in H. macrurusa (male 20.9, female 22.7).The current study has one large posterior seta on paratergal plates IV–VI, whereas increases the number of Australian Hoplopleura species from 12 H. forrestima has one large seta and one tiny seta on these paratergal to 20 and extends the records of sucking lice to all of the 14 plates; and 5) H. leggadinadis has a small spiracle on paratergal genera of endemic rodents in Australia. Overall, the Australian plate VIII, whereas H. forrestima has no spiracle on this paratergal Hoplopleura species show a higher host specificity than those plate. The hosts of these two lice, Leggadina lakedownensis and in other parts of the world. Eighteen of the 20 Australian Leggadina forresti, diverged 0.4 MYA (Rowe et al. 2016) and differ Hoplopleura species have been found only on a single host spe- in ecology and distribution (Lee 1995, Van Dyck and Strahan 2008) cies; the two exceptions, H. irritans and H. melomydis, each par- (Fig. 17), which explains the differences in morphology between asitize two host species (Kuhn and Ludwig 1966, Weaver 2017). H. leggadinadis and H. forrestima, assuming these two species of Worldwide, ~54% of the 140 Hoplopleura species recorded are lice have co-speciated and co-evolved with their hosts. found on a single host species (Durden and Musser 1994b, Light Hoplopleura mesembriomydis and H. macrurusa are also sim- et al. 2010). The high host specificity of Australian Hoplopleura ilar to each other in overall morphology but have more obvious species could be the result of louse-rodent co-speciation and differences than the pair of lice from Leggadina spp. These two co-evolution, which should be investigated in future studies when lice are found on Mesembriomys gouldii and M. macrurus re- more samples are available. Our results show that the presence spectively, which diverged 2.39 MYA (Smissen and Rowe 2018). of sucking lice on endemic Australian rodents is much more di- The differences between H. mesembriomydis and H. macrurusa verse than previously recorded. The 20 Australian Hoplopleura are (Figs. 9–12): 1) dorsal marginal head seta (DMHS) not species are found on 21 endemic rodent species; it is unknown aligned in a row in H. mesembriomydis but aligned in a row whether or not the other 42 species of Australian endemic ro- in H. macrurusa; 2) thoracic sternal plate elongated both an- dents (including 10 extinct species) are/were hosts to sucking teriorly and posteriorly into processes in H. mesembriomydis lice except that two of the 42 species—Pseudomys occidentalis but elongated only posteriorly into a process in H. macrurusa; Tate, 1951 (Rodentia: Muridae) and Rattus tunneyi Thomas,

Fig. 17. Geographical distribution of eight rodent species in Australia (in green) and locations of the rodents from which Hoplopleura species were collected in this study (red triangles) (rodent distribution adapted from https://ala.org.au). 22 Journal of Medical Entomology, 2020, Vol. XX, No. XX

1904 (Rodentia: Muridae) are hosts to the introduced spiny rat Kim, K. C. 1972. A new species of Hoplopleura (Anoplura: Hoplopleuridae) louse, Polyplax spinulosa (Wang et al. 2020). If co-speciation and from an Australian rat. Pac. Insects 14: 675–678. co-evolution are indeed a common pattern between Australian Kim, K. C., and H. W. Ludwig. 1978. The family classification of the Anoplura. endemic rodents and their sucking lice, we can expect more spe- Syst. Entomol. 3: 249–285. Kim, K. C., H. D. Pratt, and C. J. Stojanovich. 1986. The sucking lice of North cies of sucking lice to be discovered in Australia as ectoparasites America. An illustrated manual for identification, pp. 241. University of more endemic rodent species are studied. Park, Pennsylvania State University Press, University Park, PA. Kuhn, H. J., and H. W. Ludwig. 1966. Sucking lice of the genus Hoplopleura (Anoplura: Insecta) from Australian Muridae. J. Nat. Acknowledgments Hist. 13: 657–674.

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