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Australian Government’s Research Training

Program Journal of Medical Entomology, XX(X), 2021, 1–9 doi: 10.1093/jme/tjaa289 Morphology, Systematics, Evolution Research

Two New Species of Sucking Lice (: Phthiraptera: Hoplopleuridae) from Chestnut Mice,

Pseudomys gracilicaudatus and Pseudomys nanus Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021 (Rodentia: Muridae), in Australia

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

AADate 1GeneCology Research Centre, University of the Sunshine Coast, 90 Sippy Downs Drive, Sippy Downs, Queensland 4556, Australia, 2School of Science, Technology and Engineering, University of the Sunshine Coast, 90 Sippy Downs Drive, Sippy Downs, Queensland AAMonth 4556, Australia, 3Department of Biology, Georgia Southern University, Statesboro, GA 30458, and 4Corresponding author, e-mail: [email protected] AAYear Subject Editor: Richard Wilkerson Received 19 October 2020; Editorial decision 30 November 2020

Abstract We describe two new species of sucking lice in the genus Hoplopleura Enderlein, 1904 (Psocodea: Phthiraptera: Hoplopleuridae) from Australia: Hoplopleura gracilicaudatusa n. sp. from the eastern chestnut mouse Pseudomys gracilicaudatus (Gould) (Rodentia: Muridae), and Hoplopleura nanusa n. sp. from the western chestnut mouse Pseudomys nanus (Gould) (Rodentia: Muridae). Pseudomys Gray is the most speciose genus of rodents endemic to Australia with 24 species; however, only two Pseudomys species have been reported pre- viously to be hosts of sucking lice. The description of the new species in the present study doubles the number of species known to parasitize Pseudomys mice and increases the total number of sucking louse species known from endemic Australian rodents from 21 to 23. Pseudomys gracilicaudatus and P. nanus are closely related murines that diverged ~1 MYA with distinct and widely separated extant geographic distribu- tions. The two new Hoplopleura species described in the present study share some morphological characters and likely co-evolved and co-speciated with their chestnut mouse hosts.

Key words: Phthiraptera, Anoplura, Hoplopleura, new species, Old Endemic Australian rodents

Sixty-three species of endemic rodents in 14 genera are recognized chestnut mouse Pseudomys gracilicaudatus (Gould) and the western from Australia (Musser and Carleton 2005, Van Dyck and Strahan chestnut mouse Pseudomys nanus (Gould). These chestnut mice 2008). These rodents are divided into two groups based on their ar- inhabit grasslands, heathlands, open stands of eucalypt forest and rival time in Australia, the Old Endemics (5–8 million years ago) and woodland, and have a wide distribution range. The eastern chestnut the New Endemics (1–2 million years ago). The Old Endemics can mouse is patchily distributed along the eastern coast from northern be further divided into five informal divisions: Pseudomys, Uromys, Queensland to New South Wales as far south as Jervis Bay. The Hydromys, Xeromys, and Pogonomys (Lecompte et al. 2008, Cox western chestnut mouse occurs in northern Australia from Barrow and Hautier 2015). The Pseudomys division comprises eight genera Island in the west to the Sir Edward Pellew Group of Islands in the including Pseudomys Gray (Rodentia: Muridae), which is the most east (Breed and Ford 2007, Van Dyck and Strahan 2008). speciose genus of rodents in Australia with 24 species (Musser and The biodiversity of sucking lice on endemic rodents in Australia Carleton 2005, Van Dyck and Strahan 2008). All Pseudomys spe- has been understudied for decades. Twenty species of endemic cies are found only in Australia except for the little native mouse, sucking lice, all in the genus Hoplopleura Enderlein (Psocodea: Pseudomys delicatulus (Gould), which is also found in Papua New Phthiraptera: Hoplopleuridae), and the introduced spiny rat Guinea. Pseudomys species have a wide variety of body sizes, food louse, Polyplax spinulosa (Burmeister) (Psocodea: Phthiraptera: requirements, and habitats (Musser and Carleton 2005). Pseudomys Polyplacidae), have been reported previously from 23 species of mice are divided into seven subgroups based on morphological, endemic rodents in Australia (Johnson 1960, Kuhn and Ludwig molecular, and ecological evidence (Breed and Ford 2007). One of 1967, Kim 1972, Palma and Barker 1996, Weaver and Barton 2008, the subgroups, the chestnut mice, includes two species: the eastern Weaver 2017, Wang et al. 2018, Wang et al. 2020a, b). However, only

© The Author(s) 2021. 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 February 12, 2021 with fixed content and layout in compliance with Art. 8.1.3.2 ICZN. 2 Journal of Medical Entomology, 2021, Vol. XX, No. XX two Hoplopleura species have been described from Pseudomys mice: (Table 1). We collected 140 louse specimens in total, both adults and Hoplopleura calabyi Johnson from Pseudomys higginsi (Trouessart) nymphs, from these mice (Tables 1 and Table 2). Based on morpho- (Johnson 1960), and Hoplopleura gyomydis Kuhn and Ludwig from logical features, we identified these specimens as two new species of Pseudomys fumeus Brazenor (Kuhn and Ludwig 1967). In this study, Hoplopleura. we describe two new species of sucking lice from Pseudomys mice, the chestnut mice, P. gracilicaudatus and P. nanus.

Materials and Methods Family Hoplopleuridae Ewing, 1929

Sample Collections Genus Hoplopleura Enderlein, 1904 Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021 Sucking lice were collected from ethanol-preserved rodent specimens in Queensland Museum, Australian Museum and Museum and Art Hoplopleura gracilicaudatusa Wang n. sp. Gallery of the Northern Territory. As described in detail previously (Wang et al. 2018, Wang et al. 2020a, b), individual rodent speci- ZooBank Registration mens were thoroughly combed with a delousing comb. Each rodent To comply with the regulations set out in the article 8.5 was then put into a jar filled with 80% ethanol and gently shaken of the amended 2012 version of the International Code of for ~1 min to dislodge lice from the pelage. The ethanol wash was fil- Zoological Nomenclature (ICZN 2012), details of the new tered through a gauze on a fine mesh and solid contents on the gauze species have been submitted to ZooBank. The Life Science Identifier were examined under a dissecting microscope (Nikon SMZ800N). (LSID) of the article is urn:lsid:zoobank.org:pub:175ADCBE- Lice were collected and kept in 2-ml screw-cap vials filled with eth- 4F4A-40A5-BFDA-368352CAEC8F. The LSID for the new name anol, one vial per host specimen. Louse specimens were stored in a Hoplopleura gracilicaudatusa is urn:lsid:zoobank.org:act:9D7F9D1B- −20℃ freezer until further examination. 9700-4F29-A02F-8FC8DA78DB63. Male: [Based on 3 specimens; Fig. 1a] Body length 934–950 (943). Head: Head as long as wide with 4 apical head setae (ApHS) and 4 Morphological Examination anterior marginal head setae (AnMHS). Pre-antennal region short Louse specimens were mounted on microscope slides with Canada with rounded anterior margin. Antenna with five segments; 1st seg- balsam as described in detail in Wang et al. (2018, 2020a, b). Intact ment large, about as wide as long; 2nd segment narrower, much louse specimens with minimal gut contents were selected for slide longer than wide; 3rd–5th segments about as long as wide; distal mounting. The specimens were cleared and softened in KOH (20%), seta on dorsal surface of antennal segment 3 not sexually dimor- and then washed in distilled water and acetic acid solution (10%). phic. Antennal angle well developed. Dorsally, head with 4 sutural After that, the specimens were stained in acid fuchsin (1%), and head setae (SuHS), 2 small dorsal accessory head setae (DAcHS), 2 dehydrated in ethanol in different concentrations (40%, 70%, and small dorsal anterior central head setae (DAnCHS), 2 small dorsal 100%). Specimens were then cleared in pure clove oil and mounted posterior central head setae (DPoCHS), 2 large dorsal principal head on slides using Canada balsam. Specimens mounted on slides were setae (DPHS) and 8 dorsal marginal head setae (DMHS). DMHS not examined and measured with a photomicroscope (Nikon ECLIPSE aligned in row, 3rd DMHS offset medially. Ventrally, head with 2 Ts 2); all measurements were in micrometers (range followed by the ventral principal head setae (VPHS). mean). Descriptive format and abbreviations of morphological fea- Thorax: Wider than long, with 1 pair of dorsal principal thoracic tures follow Kim et al. (1986) and Durden et al. (2018, 2019) with setae (DPTS), DPTS length 83.6–92 (84.5). Thoracic sternal plate names of setae spelled out in full at first mention. We collected both shield-shaped with short anterior process and elongate posterior adult and nymphal louse specimens; we checked all adult specimens process (Fig. 1b). Mesothoracic spiracle 13.5–16.2 (14.5) in di- and mounted selected male and female adult specimens on micro- ameter. Forelegs small, with small acuminate claws; midlegs and scopic slides for detailed morphological examination. hindlegs progressively larger with correspondingly more robust tibio-tarsal claws. Results Abdomen: Wider than thorax. Dorsally, 1 tergite per segment, except for segment 3 with 2 tergites. All tergites broad. Tergite 1 with 1 pair Hoplopleura Specimens Collected From Endemic small tergal abdominal setae (TeAS) posterolaterally. Tergite 2 with Rodents in Australia 2 pairs of TeAS posterolaterally, lateral pair slightly shorter. Tergite 3 We examined 30 endemic rodent specimens of the two chestnut with 2 pairs of TeAS posterolaterally, with lateral setae much longer. mouse species: P. gracilicaudatus (n = 18) and P. nanus (n = 12) Tergite 4 with 9 TeAS, 2nd pair of TeAS longer than others, medial

Table 1. Specimens of Hoplopleura lice collected from Pseudomys gracilicaudatus and Pseudomys nanus in Australia

Rodent species Rodents sampled Rodents with Hoplopleura lice Hoplopleura lice collected

QM1 AM MAGNT Adults Nymphs

Pseudomys gracilicaudatus 13 5 -2 7 14 28 Pseudomys nanus - - 12 5 66 32 Total 13 5 12 12 80 60

Notes: 1QM: Queensland Museum, Brisbane; AM: Australian Museum, Sydney; MAGNT: Museum and Art Gallery of the Northern Territory, Darwin. 2hyphen “-” indicates no specimen was checked or collected. Journal of Medical Entomology, 2021, Vol. XX, No. XX 3

Table 2. Pseudomys specimens (n = 12) from which new species of Hoplopleura lice were collected in Australia

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

Pseudomys JM16940 Callide Mine via Biloela, QLD. (6)a 12-Apr-2005 Queensland gracilicaudatus 24°19′23″S, 150°36′00″E Museum JM19212 Lamington NP, Duck Creek Road, 1 (2) 21-Jun-2010 Queensland QLD. 28°12′08.1″S, 153°05′59.8″E Museum JM19213 Burke Development Road, 100m 4 (13) 16-Jul-2008 Queensland from, 1.7km E of Chillagoe, QLD. Museum

M36335 Myall Lakes National Park, NSW. 4 26-Jun-1999 Australian Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021 32°25′S, 152°22′E Museum M36337 Myall Lakes National Park, NSW. 1 (1) 25-Jan-1990 Australian 32°25′S, 152°22′E Museum M36338 Myall Lakes National Park, NSW. 32° 1 (5) 29-Nov-1994 Australian 25’ 55’’ S, 152° 22’ 05’’ E Museum M36339 Myall Lakes National Park, NSW. 3 (1) 25-Jun-1995 Australian 32°25′S, 152°22′E Museum Pseudomys nanus U5222 Victoria River, Gregory National 9 (1) 22-Mar-2001 Museum and Park, NT. 15° 35’ 22.2’’ S, 131° 07’ Art Gallery of 03.6’’ E the Northern Territory U5900 Tanami Desert, NT. 18°56′S, (2) 1-Jul-1983 Museum and 131°22′E Art Gallery of the Northern Territory U3508 Coomarie Springs, Tanami Desert, 54 (22) 5-May-1982 Museum and NT. 19°42′S, 129°47′E Art Gallery of the Northern Territory U4610 Unknown. 2 (7) Unknown Museum and Art Gallery of the Northern Territory U4350 Bullita Area, Gregory National Park, 1 29-Apr-1994 Museum and NT. 16°07′S, 130°25′E Art Gallery of the Northern Territory

Note: anumber of adult specimens outside brackets, number of nymphal specimens inside brackets. NSW: New South Wales; QLD: Queensland; NT: Northern Territory. Collection locality information is from https://www.ala.org.au (access date: 20 July 2020).

TeAS shortest. Tergite 5 with 9 TeAS, 3rd pair and medial TeAS short. and VIII without spiracle. Spiracles on paratergal III–VII plates small Tergite 6 with 9 TeAS, lateral 2 pairs longer than others. Tergite 7 in size, paratergal plate V spiracle 12.5–13.5 (13.2) in diameter. with 9 TeAS, lateral pair longer and stouter than others. Tergite 8 Genitalia: (Fig. 1d)Subgenital plate with narrow anterolateral ex- with 7 TeAS. One pair of dorsal lateral abdominal setae (DLAS) ad- tension on each side, with 2 large and 2 small lacunae; posterior jacent to each of tergites 6–8. Ventrally, 2 sternites per segment ex- lacuna larger than anterior lacuna. Basal apodeme longer than cept segment 4 with 3 sternites, segment 2 with 1 sternite, segments parameres and with lateral acuminate process on each side posteri- 1 and 8 with no sternites. Sternite 1 extended laterally to articulate orly. Parameres uniformly sclerotized, with pseudopenis tapering to with paratergal plate II and with 4 pairs of sternal abdominal setae a point extending beyond apices of parameres. (StAS), equal in length. Sternite 2 extended laterally to articulate with Female: [Based on 3 specimens; Fig. 2a] Body length paratergal plate III and with 7 StAS, lateral 2 pairs long and stout, 1194–1218 (1206). 3rd pair setae short, medial StAS longer than 3rd pair. Sternite 3 with Head: Head slightly longer than wide with 4 ApHS and 4 AnMHS. 9 StAS. Sternites 4, 5, 7 and 9 each with 4 pairs of StAS. Sternites 6, Pre-antennal region short with rounded anterior. Antenna with 5 seg- 8 and 10 each with 7 StAS. Sternite 11 with 2 pairs of StAS. Sternite ments; 1st segment large, about as wide as long; 2nd segment narrower, 12 with 1 pair of StAS. One pair of ventral lateral abdominal setae much longer than wide; 3rd–5th segments about as long as wide; 4th (VLAS) adjacent to each of sternites 6, 8, 10 and 11. segment slightly extended postero-apically; distal seta on dorsal sur- Paratergites: Paratergal plates (Fig. 1c) present on abdominal seg- face of antennal segment 3 not sexually dimorphic. Antennal angle ments 1–8. All plates differentially sclerotized. Paratergal plate I small well developed. Dorsally, head with 4 SuHS, 2 DAcHS, 2 DAnCHS, and offset medially. Paratergal plate II with 1 tiny central seta, 2 pos- 2 DPoCHS, 2 DPHS and 8 DMHS. DMHS not aligned in row, 3rd terior setae and 2 acuminate posterior lobes. Paratergal plate III with DMHS offset medially. Ventrally, head with 2 VPHS. 2 large setae and 2 serrated posterior lobes. Paratergal plates IV–VI Thorax: Wider than long, with 1 pair of DPTS, DPTS length 93.5– each with 1 large seta and 2 serrated posterior lobes. Paratergal plate 107 (98.5). Thoracic sternal plate shield-shaped with short ante- VII with 2 large setae and 1 dorsal posterior lobe. Paratergal plate rior process and elongate posterior process (Fig. 2b). Mesothoracic VIII with 2 long setae and no posterior lobe. Paratergal plates I, II, spiracle 14.8–17.5 (15.8) in diameter. Forelegs small, with small 4 Journal of Medical Entomology, 2021, Vol. XX, No. XX Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021

Fig. 1. Hoplopleura gracilicaudatusa n. sp. Male. (a) Habitus (dorsal mor- phology to the left of the midline, ventral morphology to the right). (b) Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. Morphological fea- tures are labeled following Kim et al. (1986) as: 1 - apical head setae (ApHS); Fig. 2. Hoplopleura gracilicaudatusa n. sp. Female. (a) Habitus (dorsal mor- 2 - anterior marginal head setae (AnMHS); 3 - distal seta on dorsal surface of phology to the left of the midline, ventral morphology to the right). (b) antennal segment 3; 4 - sutural head setae (SuHS); 5 - small dorsal accessory Thoracic sternal plate. (c) Paratergal plates. (d) Genitalia. Morphological head setae (DAcHS), 6 - small dorsal anterior central head setae (DAnCHS), features of female genitalia are labeled following Kim et al. (1986) as: U - 7 - small dorsal posterior central head setae (DPoCHS), 8 - large dorsal prin- subgenital plate; V - gonopods VIII; W - gonopods IX. cipal head setae (DPHS); 9 - dorsal marginal head setae (DMHS); 10 - ven- tral principal head setae (VPHS); 11 - dorsal principal thoracic setae (DPTS); 12 - tergal abdominal setae (TeAS); 13 - sternal abdominal setae (StAS); pairs of StAS. Sternite 16 with 4 pairs of StAS, varying in size, lateral 14 - dorsal lateral abdominal setae (DLAS); 15 - ventral lateral abdominal 2 pairs much longer than others. One pair of VLAS adjacent to each setae (VLAS); A - mesothoracic spiracle; B - thoracic sternal plate; T - tergite; S - sternite; U - subgenital plate; BA - basal apodeme; PM - parameres; PP of sternites 8, 9, 11–13, 15 and 16. - pseudopenis. Paratergites: Paratergal plates (Fig. 2c) present on abdominal seg- ments 1–8. All plates differentially sclerotized. Paratergal plate acuminate claws; midlegs and hindlegs progressively larger with cor- I small and offset medially. Paratergal plate II with 1 tiny central respondingly more robust tibio-tarsal claws. seta, 2 posterior setae and 2 acuminate posterior lobes. Paratergal Abdomen: Wider than thorax. Dorsally, 3 tergites per segment, ex- plate III with 2 large setae and 2 serrated posterior lobes. Paratergal cept for segments 1, 2 and 8 each with 1 tergite. All tergites narrow. plates IV–VI each with 1 large seta and 2 serrated posterior lobes. Tergite 1 with 1 pair of short TeAS posterolaterally. Tergite 2 with Paratergal plate VII with 2 long setae and 1 dorsal posterior lobe. 2 pairs of TeAS posterolaterally, lateral pair slightly shorter. Tergite Paratergal plate VIII with 2 long setae and with no posterior lobe. 3 with 2 pairs of TeAS posterolaterally, equal in length. Tergites 4, Paratergal plates I, II, and VIII without spiracles. Spiracles on 7, 8, 10, 12 and 13 each with 7 TeAS. Tergites 5, 6, 11, 14 and 15 paratergal plates III–VII small, paratergal plate V spiracle 12.8–14 each with 3 pairs of TeAS. Tergite 9 with 4 pairs of TeAS. Tergite 16 (13.5) in diameter. with 5 TeAS. Tergites 17 and 18 with 2 pairs of TeAS. One pair of Genitalia: Subtriangular subgenital plate with 4 small mediolateral DLAS adjacent to each of tergites 9, 10, 12, 13, 15 and 16. Ventrally, setae (1 specimen with 5 small mediolateral setae) (Fig. 2d). 3 sternites per segment, except for segment 4 with 4 sternites, seg- Gonopods VIII and IX distinct; gonopods VIII with 3 posterior setae ment 7 with 2 sternites, segment 2 with 1 sternite, segments 1 and subequal in size; gonopods IX with 3 posterior setae of differing 8 with no sternites. Sternite 1 extended laterally to articulate with lengths, lateral seta longest and medial seta shortest. Vulvar fimbriae paratergal plate II and with 4 pairs of StAS. Sternite 2 extended lat- indistinct. Three small setae on each side medial to gonopods IX. erally to articulate with paratergal plate III and with 7 StAS, lateral 2 Type Host: Pseudomys gracilicaudatus (Gould, 1845) (Rodentia: pairs long and stout, 3rd pair short, medial StAS slightly longer than Muridae), eastern chestnut mouse. 3rd pair. Sternites 3, 6, 9, 10, 12 and 13 each with 4 pairs of StAS. Type Locality: Burke Development Road, 100m from, 1.7 km E of Sternites 4, 5, 7, 8, 11 and 14 each with 7 StAS. Sternite 15 with 3 Chillagoe, Queensland, Australia. Journal of Medical Entomology, 2021, Vol. XX, No. XX 5

Type Material: Holotype male and allotype female ex Pseudomys H. pogonomydis (male 13.5, female 15.4) and H. xeromydis (male gracilicaudatus (Queensland Museum, JM19213): holotype, ♂ (QM 23.5, female 26.6). The female of H. gracilicaudatusa n. sp. has three T250530), allotype, ♀ (QM T250531), same data as for the holo- sternites on abdominal segments 3–6, whereas H. bidentata has two type. Paratypes: 1 ♂ (QM T250532) same data as for the holotype. sternites on these segments. Museum accession numbers are linked to the ZooBank record. Additional Material Examined: 1 ♂ (AM K.541484 ) and 2 ♀ (AM Hoplopleura nanusa Wang n. sp. K.541482, AM K.541483) ex Pseudomys gracilicaudatus (AM ZooBank Registration M36335), Myall Lakes National Park, NSW. 32°25′S, 152°22′E. The LSID for the new name Hoplopleura nanusa is urn:lsid:zoobank. Distribution. Australia: New South Wales and Queensland. org:act:CE5AB370-DEC1-4C81-8FB8-5DAD6233CF80. Etymology: The species epithet is a noun in apposition referring to Male: [Based on 3 specimens; Fig. 3a] Body length 715–873 (734.9). Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021 the species name of the host, gracilicaudatus. Head: Slightly longer than wide, with 4 ApHS and 4 AnMHS. Pre- Remarks: The preantennal region of H. gracilicaudatusa n. sp. is antennal region short with pointed antero-lateral apex. Antenna slightly rounded anterolaterally, which is different from H. uromydis with five segments; 1st segment large, about as wide as long; 2nd Kuhn and Ludwig, 1967, H. leporilludis Wang, 2020, H. conilurudis segment narrower, much longer than wide; 3rd–5th segments about Wang, 2020, H. leggadinadis Wang, 2020, H. mastacomydis Kuhn and as long as wide; distal seta on dorsal surface of antennal segment Ludwig, 1967, H. bidentata (Neumann, 1909), H. mesembriomydis 3 not sexually dimorphic. Antennal angle well developed. Dorsally, Wang, 2020 and H. xeromydis Wang, 2020 (Neumann 1909, Kuhn head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS and and Ludwig 1967, Wang et al. 2020b). The anterior preantennal re- 8 DMHS. Second and 3rd DMHS offset medially. Ventrally, head gion is truncated in H. uromydis, H. leporilludis, H. conilurudis, and with 2 VPHS. H. leggadinadis; it is less rounded in H. mastacomuydis, and narrow Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length and pointed in H. bidentata, H. mesembriomydis and H. xeromydis. 95–98 (96.3). Thoracic sternal plate shield shaped with elongate The third antennal segment of H. gracilicaudatusa n. sp. is not anterior and posterior processes, posterior process longer than an- sexually dimorphic, which is different from H. irritans Kuhn terior process (Fig. 3b). Mesothoracic spiracle 11.1–13.5 (12.6) in and Ludwig, 1967 and H. leporilludis (Kuhn and Ludwig 1967). diameter. Forelegs small, with small acuminate claws; midlegs and The 3rd DMHS of H. gracilicaudatusa n. sp. are offset medially, hindlegs progressively larger with correspondingly more robust whereas all DMHS are aligned in a row in H. gyomydis Kuhn and tibio-tarsal claws. Ludwig, 1967, H. mastacomydis, H. leporilludis and H. melomydis Abdomen: Wider than thorax. Dorsally, with 8 tergites. Tergite 1 Weaver, 2017 (Kuhn and Ludwig 1967, Weaver 2017). Hoploplura with 1 pair TeAS posterolaterally. Tergite 2 with 2 pair of TeAS gracilicaudatusa n. sp. has four DMHS on each side; H. zyzomydis posterolaterally, all TeAS equal in length. Tergite 3 with 2 pairs of Weaver, 2008 has only two DMHS on each side, whereas H. notomydis Weaver, 2017 and H. setosa Weaver, 2017 each have three DMHS on each side (Weaver and Barton 2008, Weaver 2017). The sternal plate of H. gracilicaudatusa n. sp. is shield shaped which differentiates it from H. uromydis (less rounded) and H. leporilludis (subhexagonal). Paratergal plate II of H. gracilicaudatusa n. sp. has a tiny central seta, which is absent in H. bidentata, H. cornata Kim, 1972, H. zyzomydis and H. notomydis (Kim 1972). Paratergal plates IV–VI of H. gracilicaudatusa n. sp. each have one large posterior seta. Two posterior setae are adjacent to these plates in H. uromydis, H. gyomydis, H. irritans, H. mastacomydis, H. forrestima Wang, 2020, H. melomydis, H. notomydis and H. mesembriomydis, whereas no posterior seta is adjacent to these plates in H. zyzomydis (Wang et al. 2020b). Paratergal plate VI has two serrated posterior lobes, which differentiates it from H. villosissima Wang, 2018, which has two pointed posterior lobes on paratergal plate VI (Wang et al. 2018). Paratergal plate VII of H. gracilicaudatusa n. sp. has one pos- terior lobe dorsally, compared to no posterior lobe on that plate in H. cornata, H. gyomydis, H. irritans, H. uromydis, H. villosissima, H. bidentata, H. conilurudis, H. leporilludis, H. macrurusa Wang, 2020, H. pogonomydis Wang, 2020 and H. xeromydis, and two pos- terior lobes in H. mastacomydis and female H. calabyi Johnson 1960 (Johnson 1960, Wang et al. 2020b). Hoplopleura gracilicaudatusa n. sp. has two setae on paratergal plate III, whereas H. mastacomydis has one short, stout seta posteriorly. The spiracle on paratergal plate V of H. gracilicaudatusa n. sp. is small (male 13.2, female 13.5), smaller than in H. irritans (12–16), H. leporilludis (male 20.5, fe- male 22.5), H. villosissima (male 18.4, female 20), H. zyzomydis (male 15.8, female 17.6), H. forrestima (male 15.4, female 19.1), H. leggadinadis (male 14.8, female 19.9), H. conilurudis (male

25.1, female 27.6), H. uromydis (34–37), H. bidentata (25–28), Fig. 3. Hoplopleura nanusa n. sp. Male. (a) Habitus (dorsal morphology to H. melomydis (male 25, female 25.8), H. macrurusa (male 20.9, the left of the midline, ventral morphology to the right). (b) Thoracic sternal female 22.7) and H. mesembriomydis (male 31.7, female 38.4) plate. (c) Paratergal plates. (d) Genitalia. 6 Journal of Medical Entomology, 2021, Vol. XX, No. XX

TeAS posterolaterally, with setae much longer on each side. Tergite 4 with 5 pairs of TeAS, all TeAS narrow and short, equal in length. Tergite 5 with 9 TeAS, 2nd pair longer and stouter than others. Tergites 6 and 7 each with 9 TeAS, lateral 2 pairs longer and stouter than others. Tergite 8 with 7 TeAS, lateral pair longer than others. One pair of DLAS adjacent to tergites 5–8. Ventrally, with 10 stern- ites. Two sternites per segment except segment 4 with 3 sternites, segment 2 with 1 sternite, and segments 1 and 8 with no sternites. Segment 1 extended laterally to articulate with paratergal plate II and with 4 pairs of StAS, equal in length. Sternite 2 extended later- Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021 ally to articulate with paratergal plate III and with 7 StAS, lateral 2 pairs long and stout, 3rd pair short, medial StAS longer than 3rd pair. Sternites 3, 5, 7, 8 and 9 each with 4 pairs of StAS. Sternites 4, 6 and 10 each with 7 StAS. Sternite 11 with 2 pairs of StAS. Sternite 12 with 1 pair StAS. One pair of VLAS adjacent to each of sternites 6, 8, 10 and 11. Paratergites: Paratergal plates (Fig. 3c) present on abdominal seg- ments 1–8. All plates differentially sclerotized. Paratergal plate I small and offset medially. Paratergal plate II with 1 small me- dial seta, 2 large posterior setae and 2 acuminate posterior lobes. Paratergal plate III with 2 large posterior setae and 2 serrated pos- terior lobes. Paratergal plates IV–VI each with 1 large seta postero- ventrally, 1 tiny seta postero-dorsally and 2 serrated posterior lobes. Paratergal plates VII and VIII each with 2 long setae and lacking posterior lobe. Paratergal plates I and II without spiracles. Spiracles on paratergal III–VII plates small in size, spiracle of paratertal plate V 10–12 (11.3) in diameter. Spiracle on paratergal plate VIII smaller than other paratergal spiracles. Genitalia: (Fig. 3d)Subgenital plate with narrow anterolateral and mediolateral extensions on each side, with 1 large and 2 small lacunae; Fig. 4. Hoplopleura nanusa n. sp. Female. (a) Habitus (dorsal morphology to posterior lacuna larger than anterior lacunae. Basal apodeme longer the left of the midline, ventral morphology to the right). (b) Thoracic sternal than parameres and with lateral acuminate process on each side pos- plate. (c) Paratergal plates. (d) Genitalia. teriorly. Parameres uniformly sclerotized, with pseudopenis tapering to point extending beyond apices of parameres. 8 with no sternites. Sternite 1 extended laterally to articulate with Female: [Based on 3 specimens; Fig. 4a] Body length paratergal plate II and with 4 pairs of StAS. Sternite 2 extended lat- 1110–1240 (1165). erally to articulate with paratergal plate III and with 7 StAS, lateral Head: Head slightly longer than wide, with 4 ApHS and 4 AnMHS. 2 pairs long and stout, 3rd pair of StAS short, medial StAS longer Pre-antennal region short with pointed antero-lateral apex. Antenna than 3rd pair. Sternite 3 with 2 pairs of StAS, posterolateral pair with five segments; 1st segment large, about as wide as long; 2nd stout. Sternite 4 with 4 pairs of StAS. Sternite 5 with 7 StAS, lateral segment narrower, much longer than wide; 3rd–5th segments about two pairs long and stout, 3rd pair short, medial StAS longer than as long as wide; distal seta on dorsal surface of antennal segment 3rd pair. Sternite 6 with 3 pairs of StAS, equal in length. Sternite 3 not sexually dimorphic. Antennal angle well developed. Dorsally, 7 with 4 pairs of StAS, lateral two pairs stout and long. Sternite 8 head with 4 SuHS, 2 DacHS, 2 DAnCHS, 2 DPoCHS, 2 DPHS and with 7 StAS, 3rd pair short and narrow. Sternite 9 with 3 pairs of 8 DMHS. Second and 3rd DMHS offset medially. Ventrally, head StAS. Sternite 10 with 7 StAS. Sternite 11 with 7 StAS, medial StAS with 2 VPHS. short. Sternites 12–14 each with 7 StAS. Sternite 15 with 3 pairs Thorax: Wider than long, with 1 pair of DPTS dorsally, DPTS length of StAS. Sternite 16 with 4 pairs of StAS, lateral two pairs much 104–112 (107.6). Thoracic sternal plate shield shaped with elongate longer than medial 2 pairs. One pair of VLAS adjacent to each of anterior and posterior processes, posterior process longer than an- sternites 8, 9, 11–13, 15 and 16. terior process (Fig. 4b). Mesothoracic spiracle 12.5–14.2 (13.7) in Paratergites: Paratergal plates (Fig. 4c) present on abdominal seg- diameter. Forelegs small, with small acuminate claws; midlegs and ments 1–8. All plates differentially sclerotized. Paratergal plate hindlegs progressively larger with correspondingly more robust I small and offset medially. Paratergal plate II with 1 small me- tibio-tarsal claws. dial seta, 2 large posterior setae and 2 acuminate posterior lobes. Abdomen: Wider than thorax. Dorsally, 18 tergites; 3 tergites per Paratergal plate III with 2 large posterior setae and 2 serrated pos- segment, except for segments 1, 2 and 8 each with 1 tergite. Tergites terior lobes. Paratergal plates IV–VI each with 1 large seta postero- 2–4 broad. Tergite 1 with 1 pair of TeAS posterolaterally. Tergite dorsally, 1 tiny seta postero-dorsally and 2 serrated posterior lobes. 2 with 2 pairs of long TeAS posterolaterally, lateral pair shorter. Paratergal plates VII and VIII each with 2 long setae and lacking Tergite 3 with 2 pairs of long TeAS, equal in length. Tergites 4, 7, posterior lobe. Paratergal plates I and II without spiracles. Spiracles 9 and 10 each with 7 TeAS. Tergites 5, 6, 8 and 11–15 each with 3 on paratergal III–VII plates small in size, spiracle of paratertal plate pairs of TeAS. Tergite 16 with 5 TeAS. Tergites 17 and 18 each with V 11.5–13.2 (12.4) in diameter. Spiracle on paratergal plate VIII 2 pairs of TeAS. One pair of DLAS adjacent to each of tergites 6, smaller than other paratergal spiracles. 7, 9, 10, 12, 13, 15 and 16. Ventrally, 16 sternites; 3 sternites per Genitalia: (Fig. 4d) Subtriangular subgenital plate with large ante- segment except for segment 2 with 1 sternite, and segments 1 and rior indentation on each side, tapering to broadly rounded posterior Journal of Medical Entomology, 2021, Vol. XX, No. XX 7

apex, with 4 small mediolateral setae. Gonopods VIII and 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: Pseudomys nanus (Gould, 1858) (Rodentia: Muridae), Western chestnut mouse. Type Locality: Coomarie Springs, Tanami Desert, Northern Territory,

Ludwig, 1967 Ludwig, Australia (19°42′S, 129°47′E). H. nanusa n. sp. n. sp. H. gracilicaudatusa 3 Kuhn & H. gyomydis 2 Johnson, 1960 Johnson, H. calabyi Species Type Material: Holotype male and allotype female ex Pseudomys Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021 nanus (Museum and Art Gallery of the Northern Territory, U3508): holotype, ♂ (NTM I010693), allotype, ♀ (NTM I010694), same data as for the holotype. Paratypes: 1 ♂ (NTM I010695) and 1 ♀ (NTM I010696), same data as for the holotype. Museum accession num- bers are linked to the ZooBank record. Additional Material Examined: 1 ♂ (NTM I010697), 1 ♀ (NTM I010698), ex Pseudomys nanus (NT U5222), location Victoria River, Gregory National Park, Northern Territory, Australia (15°35′22.2″S, 131°07′03.6″E). Distribution. Australia: Northern Territory. Etymology: The species epithet is a noun in apposition referring to the species name of the host, nanus. Remarks: The preantennal region of H. nanusa n. sp. is pointed anterolaterally. Conversely, H. uromydis, H. leporilludis, H. conilurudis, and H. leggadinadis have a truncated preantennal re- gion anterolaterally, whereas in H. irritans, H. calabyi, H. macrurusa, H. pogonomydis, and H. gracilicaudatusa, it is rounded. The third antennal segment of H. nanusa n. sp. is not sexually dimorphic, which is different from H. irritans and H. leporilludis. The 2nd and 3rd DMHS of H. nanusa n. sp. are offset medially, whereas the DMHS are aligned in a row in H. gyomydis, H. mastacomydis, H. leporilludis, and H. melomydis. Hoploplura nanusa n. sp. has four DMHS on each side; H. zyzomydis has only two DMHS on each side, whereas H. notomydis and H. setosa each have three DMHS on each side. The sternal plate of H. nanusa n. sp. is shield shaped which differentiates it from H. uromydis (less rounded) and H. leporilludis (subhexagonal). Paratergal plate II of H. nanusa n. sp. has a small central seta, which is not present in H. bidentata, H. cornata, H. zyzomydis, and H. notomydis. Paratergal plates IV–VI of H. nanusa n. sp. each have one large and one tiny pos- terior seta. Paratergal plates IV-VI of H. villosissima, H. setosa, H. conilurudis, H. leggadinadis, H. leporilludis, H. macrurusa, H. pongonomydis, H. xeromydis, and H. gracilicaudatusa each have 1 large posterior seta only, whereas no posterior seta is adjacent to these plates in H. zyzomydis. Paratergal plate VII of H. nanusa n. sp. has no posterior lobe, compared to one posterior lobe on that plate in H. forrestima, H. setosa, H. melomydis, H. zyzomydis, H. mastacomydis, H. leggadinadis, H. mesembriomydis, and H. gracilicaudatusa, and two posterior lobes in H. mastacomydis and female H. calabyi. Hoplopleura nanusa n. sp. has two setae on paratergal plate III, whereas H. mastacomydis has one short, stout seta posteriorly. The spiracle on paratergal plate V of H. nanusa n. sp. is small (male 11.3, female 12.4), smaller than those in H. irritans (12–16), H. mastacomydis (11–14), H. leporilludis (male 20.5, fe- Pseudomys nanus . VIII with 1 small spiracle; on Pseudomys lobes; paratergal plate . gracilicaudatus VIII with no spiracle; on Pseudomys paratergal plate male 22.5), H. villosissima (male 18.4, female 20), H. zyzomydis Second and 3rd DMHS offset medially; paratergal plates IV to VI each with 1 large and 1 tiny posterior seta; paratergal plate VII without posterior VI each with 1 large and tiny posterior seta; paratergal plate Second and 3rd DMHS offset medially; paratergal plates IV to Third DMHS offset medially; paratergal plates IV to VI each with 1 posterior seta dorsally; paratergal plate VII with 1 posterior lobe dorsally; VI each with 1 posterior seta dorsally; paratergal plate Third DMHS offset medially; paratergal plates IV to Pseudomys gyomys. not on Pseudomys DMHS not aligned in a row, Pseudomys gyomys. on Pseudomys DMHS all aligned in a row, Pseudomys higginsi. not on Pseudomys VII not sexually dimorphic, Thoracic sternal plate not sub-hexagonal; posterior lobes of paratergal Pseudomys higginsi. on Pseudomys VII sexually dimorphic, Thoracic sternal plate sub-hexagonal; posterior lobes of paratergal Key characters (male 15.8, female 17.6), H. forrestima (male 15.4, female 19.1), H. leggadinadis (male 14.8, female 19.9), H. conilurudis (male 25.1, female 27.6), H. uromydis (34–37), H. bidentata (25–28),

rodents in Australia rodents in lice in the genus Hopopleura parasitizing Pseudomys for the identification of sucking Key H. melomydis (male 25, female 25.8), H. macrurusa (male 20.9,

female 22.7) and H. mesembriomydis (male 31.7, female 38.4) H. pogonomydis (male 13.5, female 15.4), and H. xeromydis (male

3 2 1 Table 3. Table No. 23.5, female 26.6). The female of H. nanusa n. sp. has three sternites 8 Journal of Medical Entomology, 2021, Vol. XX, No. XX Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021

Fig. 5. Geographical distribution of Pseudomys gracilicaudatus, Pseudomys nanus, Pseudomy fumeus and Pseudomys higginsi in Australia (in green in online version or gray in black and white version) and locations of the rodents from which Hoplopleura species were collected in this study (red triangles in online version or black triangles in black and white version) (rodent distribution adapted from https://ala.org.au). on abdominal segments 3–6, whereas H. bidentata has two sternites except for the introduced Polyplax spinulosa, which was found on on these segments. P. occidentalis recently (Wang et al. 2020a). The four Hoplopleura louse species from Pseudomys rodents show clear morphological differences from each other: 1) DMHS are not aligned in a row in H. gracilicaudatusa and H. nanusa (Figs. 1a, 2a, 3a, and 4a), but are Discussion aligned in a row in H. gyomydis (Kuhn and Ludwig 1967); 2) tho- racic sternal plate is shield shaped with elongate anterior and poste- We describe two new Hoplopleura species from chestnut mice, rior in H. gracilicaudatusa and H. nanusa (Figs. 1b, 2b, 3b, 4b) but one from P. gracilicaudatus and one from P. nanus, in this study. is heart shaped in H. gyomydis, and sub-hexagonal in H. calabyi These new species double the number of sucking louse species re- (Johnson 1960); 3) DLAS adjacent to tergites are present in male corded from Pseudomys species and bring the total number of H. gyomydis, male H. gracilicaudatusa and male H. nanusa but are Hoplopleura louse species known from endemic Australian rodents absent in male H. calabyi; 4) DLAS adjacent to tergites VII-VIII are from 20 to 22. Pseudomys is the most speciose genus of rodents present in male H. gracilicaudatusa and male H. nanusa but are ab- in Australia with 24 species (Musser and Carleton 2005, Van Dyck sent in male H. gyomydis; 5) paratergal plates IV-VI each have one and Strahan 2008). Pseudomys species vary widely in distribution, large posterior seta in H. gracilicaudatusa (Figs. 1c, 2c), whereas habitat, body size and food requirements (Musser and Carleton their corresponding plate has one large and one tiny posterior seta in 2005). The first sucking louse species described from Pseudomys H. gyomydis, H. calabyi and H. nanusa (Figs. 3c, 4c); 6) paratergal mice was H. calabyi from the long-tailed mouse, P. higginsi, fol- plate VI has two pointed posterior lobes in H. gyomydis but has lowed by H. gyomydis from P. fumeus (Johnson 1960, Kuhn and two serrated posterior lobes in H. gracilicaudatusa, H. nanusa, and Ludwig 1967). Prior to the present study, no sucking louse spe- H. calabyi; 7) paratergal plate VII has one long and one short pos- cies had been reported from Pseudomys mice for more than 50 yr, terior lobes in female H. calabyi but has one short posterior lobe Journal of Medical Entomology, 2021, Vol. XX, No. XX 9 dorsally in female H. gracilicaudatusa and no posterior lobes in fe- Durden, L. A., C. Robinson, J. A. Cook, B. S. McLean, B. Nyamsuren, and male H. nanusa and female H. gyomydis; and 8) paratergal plate S. E. Greiman. 2019. A New Species of Sucking Louse from the Long- VII has one short posterior lobe dorsally in male H. calabyi and tailed Ground Squirrel, Urocitellus undulatus, from Mongolia, with a male H. gracilicaudatusa but no posterior lobes in male H. nanusa Key to Species, and a Review of Host Associations and Geographical Distributions of Members of the Genus Linognathoides (Psocodea: and male H. gyomydis. Based on these morphological differences, Anoplura: Polyplacidae). J. Parasitol. 105: 469–479. we provide a dichotomous key for the identification of the four ICZN. 2012. International Commission on Zoological Nomenclature: Hoplopleura species from Pseudomys mice (Table 3). Amendment of articles 8, 9, 10, 21 and 78 of the International Code of Overall, H. gracilicaudatusa and H. nanusa share more mor- Zoological Nomenclature to expand and refine methods of publication. phological characters (characters 1–4 and 6 above) to each Bull. Zool. Nomencl. 69: 161–169. other than to H. calabyi or H. gyomydis. The similarity between Johnson, P. T. 1960. A new species of Hoplopleura from Australia (Anoplura: Downloaded from https://academic.oup.com/jme/advance-article/doi/10.1093/jme/tjaa289/6133203 by guest on 13 February 2021 H. gracilicaudatusa and H. nanusa is likely the result of co-evolution Hoplopleuridae). Proc. Entomol. Soc. Wash. 62: 111–113. and co-speciation with their host mice. Pseudomys gracilicaudatus Kim, K. C. 1972. A new species of Hoplopleura (Anoplura: Hoplopleuridae) and P. nanus are the most closely related sister species among the from an Australian rat. Pac. . 14: 675–678. 24 Pseudomys species and they diverged approximately one million Kim, K. C., H. D. Pratt, and C. J. Stojanovich. 1986. The sucking lice of North America. An illustrated manual for identification. Pennsylvania State years ago (MYA), which is much more recent than their divergences University Press, Pennsylvania, USA. from P. fumeus (3.47 MYA) and P. higginsi (3.75 MYA) (Smissen Kuhn, H. J., and H. W. Ludwig. 1967. Sucking lice of the genus Hoplopleura and Rowe 2018). Pseudomys gracilicaudatus and P. nanus also (Anoplura: Insecta) from Australian Muridae. J. Nat. Hist. 13: overlap in their distribution along the northern coast of Australia 657–674. but are distant from P. fumeus and P. higginsi, which are distributed Lecompte, E., K. Aplin, C. Denys, F. Catzeflis, M. Chades, and P. Chevret. in the southern parts of Australia (Fig. 5). Given the higher host spec- 2008. Phylogeny and biogeography of African Murinae based on mito- ificity observed among Australian Hoplopleura species than those chondrial and nuclear gene sequences, with a new tribal classification of in other parts of the world (Wang et al. 2020b), co-evolution and the subfamily. BMC Evol. Biol. 8: 199. co-speciation with rodent hosts are likely a common mechanism for Musser, G. G., and M. D. Carleton. 2005. Superfamily Muroidea. pp. 894– the speciation of Hoplopleura species in Australia as observed in the 1531. In: D. E. Wilson, D. M. Reeder, (eds.), Mammal species of the world: a taxonomic and geographic reference. 2nd ed. Johns Hopkins University current study. We expect future studies will provide more evidence Press, Baltimore. on the speciation mechanisms of sucking lice in Australia and will Neumann, L. G. 1909. Notes sur les Pédiculidés. Arch. Parasitol. 13: 497–538. reveal more species of sucking lice from Pseudomys mice. Palma R. L., and S. C. Barker. 1996. Phthiraptera. pp. 81–247. In: A. Wells, (ed.), Zoological catalogue of Australia. Vol. 26. , Phthiraptera, Thysanoptera. CSIRO Publishing, Melbourne. Acknowledgments Smissen, P. J., and K. C. Rowe. 2018. Repeated biome transitions in the evolu- We thank Heather Janetzki (Queensland Museum, Brisbane, Australia), Sandy tion of Australian rodents. Mol. Phylogenet. Evol. 128: 182–191. Ingleby and Derek Smith (Australian Museum, Sydney, Australia), Gavin Dally Van Dyck, S., and R. Strahan (editors). 2008. The Mammals of Australia. 3rd (Museum and Art Gallery of the Northern Territory, Darwin, Australia) for edition. New Holland Publishers, Sydney. 887pp. providing access to rodent specimens in museums; Yalun Dong (University Wang, W., H. J. Weaver, F. Song, L. A. Durden, and R. Shao. 2018. A new species of the Sunshine Coast, Australia) for assisting with sample collection; Allison of sucking louse Hoplopleura villosissima n. sp. (Psocodea: Phthiraptera: Morrison and Darren Morrow (University of the Sunshine Coast, Australia) Hoplopleuridae) and a new host record of the spiny rat louse Polyplax for providing field work support. We thank the reviewers for comments that spinulosa Burmeister, 1839 (Psocodea: Phthiraptera: Polyplacidae) from have improved this manuscript greatly. This study was funded by Australian the long-haired rat Rattus villosissimus Waite (Rodentia: Muridae) in Government’s Research Training Program Scholarship to WW and the Australia. Parasit. Vectors. 11: 476. Australian Biological Resources Study (ABRS, RF217-51). Wang, W., L. A. Durden, and R. Shao. 2020a. Rapid host expansion of an introduced parasite, the spiny rat louse Polyplax spinulosa (Psocodea: Phthiraptera: Polyplacidae), among endemic rodents in Australia. Parasit. 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A new species of sucking louse Polyplacidae) From the Gray Mouse Lemur, Microcebus murinus (Primates: (Phthiraptera: Anoplura) from Australia, and a key to the Australian spe- Cheirogaleidae), in Madagascar. J. Med. Entomol. 55: 910–914. cies of Hoplopleura. Zootaxa. 1679: 55–62.