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A new species of the pseudoscorpion
genus Megachernes (Pseudoscorpiones:
Chernetidae) associated with a
threatened Sri Lankan rainforest
rodent, with a review of host
associations of Megachernes
Mark S. Harvey a b c d , Pamoda B. Ratnaweera e f , Preethi V.
Udagama e & Mayuri R. Wijesinghe e
a Department of Terrestrial Zoology, Western Australian Museum,
Locked Bag 49, Welshpool, DC, WA 6986, Australia
b Division of Invertebrate Zoology, American Museum of Natural
History, 79th Street @ Central Park West, New York, NY,
10024-5192, USA
c Department of Entomology, California Academy of Sciences,
Golden Gate Park, San Francisco, CA, 94103-3009, USA
d School of Animal Biology, University of Western Australia,
Crawley, WA, 6009, Australia
e Department of Zoology, University of Colombo, Cumaratunga
Munidasa Mawatha, Colombo 03, Sri Lanka
f Science and Technology Degree Programme, Uva Wellassa
University, Badulla, Sri Lanka
Version of record first published: 15 Nov 2012.
To cite this article: Mark S. Harvey, Pamoda B. Ratnaweera, Preethi V. Udagama & Mayuri R.
Wijesinghe (2012): A new species of the pseudoscorpion genus Megachernes (Pseudoscorpiones:
Chernetidae) associated with a threatened Sri Lankan rainforest rodent, with a review of host
associations of Megachernes , Journal of Natural History, 46:41-42, 2519-2535
To link to this article: http://dx.doi.org/10.1080/00222933.2012.707251
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Journal of Natural History
Vol. 46, Nos. 41–42, November 2012, 2519–2535
A new species of the pseudoscorpion genus Megachernes
(Pseudoscorpiones: Chernetidae) associated with a threatened
Sri Lankan rainforest rodent, with a review of host associations
of Megachernes
Mark S. Harveya,b,c,d*, Pamoda B. Ratnaweerae,f, Preethi V. Udagamae
and Mayuri R. Wijesinghee
aDepartment of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool
DC, WA 6986, Australia; bDivision of Invertebrate Zoology, American Museum of Natural
History, 79th Street @ Central Park West, New York, NY 10024-5192, USA; cDepartment of
Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94103-3009
USA; dSchool of Animal Biology, University of Western Australia, Crawley, WA 6009, Australia;
eDepartment of Zoology, University of Colombo, Cumaratunga Munidasa Mawatha, Colombo
03, Sri Lanka; fScience and Technology Degree Programme, Uva Wellassa University, Badulla,
Sri Lanka
(Received 14 March 2012; final version received 25 June 2012; printed 28 October 2012)
Most species of the Old World pseudoscorpion genus Megachernes are associated
with mammals, either within their nests or in their pelage, whereas others are
found in cave systems, usually in bat guano. The first Sri Lankan species of the
genus, Megachernes kanneliyensis, is described from the fur of an endemic, threat-
ened Sri Lankan mouse, Mus mayori pococki from the Kanneliya rainforest. The
species is compared with other species of the genus. Host associations of all named
and several unnamed species of Megachernes are reviewed, with two main habitat
types, those found in mammal nests and in their pelage, and those found in bat
guano. Based on similarities with Megachernes,Chiridiochernes is transferred to
Lamprochernetinae as the spermathecae are T-shaped, the sole diagnostic feature
of the subfamily.
Keywords: threatened species; Mus mayori pococki; rainforest; morphology;
taxonomy
Introduction
Mayor’s Mouse, Mus mayori (Thomas, 1915), is endemic to the rainforests of Sri
Lanka, with Mus mayori mayori found in the montane zone and Mus mayori pococki
Ellerman, 1947 found in the lowlands (Phillips 1980). It is listed as Vulnerable by the
IUCN because of deforestation of its native habitat (Goonatilake et al. 2008). Mus
mayori pococki is a medium-sized murid with a snout–vent length of approximately
90 mm (Phillips 1980; Ratnaweera et al. 2010). Previous studies on parasites occurring
on Mus mayori found the laelapid mite Laelaps atypicus Turk, 1950 (Laelapidae)
and the flea Stivalius phoberus Jordan and Rothschild, 1922 (Stivaliidae) (Turk 1950;
Phillips 1980; Crusz 1984). Recent studies on populations of Mus mayori pococki from
four rainforest reserves in Sri Lanka recorded several ectoparasitic or commensal
*Corresponding author. Email: mark.harvey@museum.wa.gov.au
ISSN 0022-2933 print/ISSN 1464-5262 online
© 2012 Taylor & Francis
http://dx.doi.org/10.1080/00222933.2012.707251
http://www.tandfonline.com
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2520 M.S. Harvey et al.
Figure 1. Map showing distribution of Megachernes kanneliyensis ()andallother
Megachernes species ().
arthropods including mites of the genus Echinolaelaps Ewing, 1929 (Laelapidae), the
louse Polyplax serrata (Burmeister, 1839) (Hoplopleuridae), a species of the tick genus
Ixodes Latreille, 1795 (Ixodidae), and a pseudoscorpion of the genus Megachernes
Beier, 1932 (Ratnaweera et al. 2010). The pseudoscorpion is now named and described
in this paper.
Species of Megachernes occur widely throughout the eastern portion of the Old
World, with the 22 named species ranging from the Black Sea in the west to Japan
in the northeast and as far south as eastern Australia (Harvey 2011) (Figure 1).
Members of the genus are unusual among pseudoscorpions in that many appear to be
obligate commensals with mammals, occurring in their nests and living phoretically
in their pelage (e.g. Beier 1948; Martens 1975; Durden 1991), or occurring in caves
(e.g. Morikawa 1960). Despite records of Megachernes from other regions of Asia,
and only two records from India, both of Megachernes himalayensis (Ellingsen, 1914)
from the northern states of Uttarakhand (Ellingsen 1914) and Jammu and Kashmir
(Beier 1978) there had been no records from Sri Lanka. The discovery of specimens
of Megachernes in the fur of a Sri Lankan rodent allows us to record the genus from
Sri Lanka for the first time.
Materials and methods
The material examined here is lodged in the collections of the Western Australian
Museum, Perth (WAM). Terminology and mensuration mostly follow Chamberlin
(1931), with the exception of the nomenclature of the pedipalps and legs and with
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Journal of Natural History 2521
some minor modifications to the terminology of the trichobothria (Harvey 1992) and
chelicera (Judson 2007). The terminology for the mammalian hosts follows Wilson and
Reeder (2005).
The terms antaxial and paraxial have been recently proposed to describe the lat-
eral faces of pseudoscorpion appendages (Judson 2007) to replace the inadequate and
ambiguous terms anterior and posterior. This change is welcome but we here use the
terms prolateral and retrolateral as used in describing spider appendages (e.g. Forster
1967; Dondale and Redner 1978). When the appendage is (imaginarily) stretched at
90◦to the longitudinal axis of the body, the anterior-facing surface is the prolateral
face and the posterior-facing surface is the retrolateral face.
The specimens were collected and stored in 75% ethanol. They were studied using
a Leica DM2500 compound microscope and Leica MZ16A and M205C stereo micro-
scopes. The line drawings were prepared with the aid of a drawing tube attached
to the Leica DM2500 microscope, and the digital images were taken with a Leica
DFC 500 camera attached to the Leica MZ16A and M205C stereo microscopes.
Measurements were taken at the highest possible magnification using an ocular gratic-
ule. The material was studied using temporary slide mounts prepared by the dissection
of the pedipalps, one chelicera, leg I and leg IV from the body, immersion of the
larger portions in concentrated lactic acid at room temperature for several days, and
mounting them on microscope slides using 10-mm coverslips supported by small sec-
tions of nylon fishing line. After study the specimens were washed in distilled water
and returned to 75% ethanol with the dissected portions placed in 12 ×3-mm glass
genitalia microvials (BioQuip Products, Inc., Compton, CA, USA).
Family CHERNETIDAE Menge
Subfamily LAMPROCHERNETINAE Beier
Remarks
Harvey (1995) discussed the composition of the Lamprochernetinae, which was
restricted to those chernetids with T-shaped spermathecae. Although the genera
Allochernes Beier, 1932, Lamprochernes Tömösváry, 1882, Lasiochernes Beier, 1932,
Megachernes,Nudochernes Beier, 1935, Pselaphochernes Beier, 1932 and Wyochernes
Hoff, 1949 were included by Harvey (1995), it now appears that Wyochernes does not
have T-shaped spermathecae (Muchmore 1996) and is better placed in the Chernetinae,
despite the lack of a precise subfamilial definition (Harvey 1995).
The differences between Megachernes and Lasiochernes are slight and rest on
the presence of a rounded protuberance on the posterolateral corner of coxa IV of
Megachernes, which is absent in Lasiochernes. Beier (1932) claimed that the relative
position of the tactile seta on legs III and IV differed between the genera, near the
middle of the segment in Megachernes and more distally placed in Lasiochernes,but
subsequent descriptions of other species of Lasiochernes suggest that a medial posi-
tion typical of Megachernes species also occurs (Beier 1949, 1961, 1963), casting doubt
on the utility of this feature. Other consistent differences may occur in the female
genitalia, but too few species of either genus have been examined so far to estab-
lish whether this feature is reliable. The spermathecal arms of Lasiochernes siculus
Beier, 1961, the only species described to date, are short (Callaini 1986), whereas those
of Megachernes are extremely elongated and T-shaped as described for Megachernes
pavlovskyi Redikorzev, 1949 (Vachon 1938, fig. 56k), Megachernes ochotonae Krumpál
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2522 M.S. Harvey et al.
and Keifer, 1982 (Krumpál and Kiefer 1982, fig. 7) and Megachernes tuberosus
Mahnert, 2009 (Mahnert 2009, fig. 25).
The southeast Asian chernetid genus Chiridiochernes Muchmore, 1972 is also asso-
ciated with rodents and has posterolaterally enlarged fourth coxae characteristic of
Megachernes.Chiridiochernes differs in numerous ways from Megachernes including
the lack of a tactile seta on tarsi III and IV, the lack of trichobothrium st, a prominent
rounded process on the proximal region of the prolateral face of the male pedipalpal
femur, flattened pedipalpal segments and the junction between the femur and patella of
legs I and II more like cheiridioids than cheliferoids (Muchmore 1972). The sole named
species of the genus, Chiridiochernes platypalpus Muchmore, 1972 from Sulawesi, is
only known from a single adult male (Muchmore 1972). Dr M. Judson (Muséum
National d’Histoire Naturelle, Paris) kindly informed us (in litt., 16 March 2012) that
females of an undescribed species of Chiridiochernes have T-shaped spermathecae and
therefore conform to current diagnoses of Lamprochernetinae, to which it is here
transferred.
Megachernes Beier
Megachernes Beier 1932: 128; Beier 1933: 518; Beier 1948: 476; Morikawa 1960: 144;
Harvey 1991: 599; Harvey 2011, without pagination.
Type species
Chernes grandis Beier, 1930, by original designation.
Diagnosis
Megachernes differs from other chernetid genera by the following combination of char-
acter states: posterolateral corner of coxa IV enlarged and rounded (Figures 3, 4B);
cheliceral rallum with three blades (Figure 4F); tarsus III and IV with long tactile seta
situated in middle of segment (Figure 4G); and female spermathecae very long and
T-shaped.
Megachernes kanneliyensis sp. nov.
(Figures 2–4)
Material examined
Holotype. Male, Kanneliya rainforest, Southern Province, Sri Lanka, 6◦09’ to 6◦18’ N,
80◦19’ to 80◦27’ E, 9 November 2006, from dorsal fur of Mus mayori pococki Ellerman,
P.B. Ratnaweera (WAM T121678).
Paratypes. One female (with attached brood-sac) (WAM T121679), two protonymphs
(WAM T121680), collected with holotype.
Etymology
This species is named after the type locality, Kanneliya rainforest.
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Journal of Natural History 2523
AB
C
D
Figure 2. Megachernes kanneliyensis sp. nov. (A) Holotype male, dorsal; (B) holotype male,
ventral; (C) paratype female, dorsal; (D) paratype female, ventral.
Diagnosis
Megachernes kanneliyensis is the only species of the genus in which males have been
described with long setae on the pedipalpal patella and chela, particularly on the pro-
lateral surfaces, but short setae on the femur (Figure 4C). Males of Megachernes with
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2524 M.S. Harvey et al.
Figure 3. Megachernes kanneliyensis sp. nov., paratype female, ventral showing detail of coxae
and brood-sac.
long setae on the patella usually also have long setae on the femur as reported for
M. afghanicus Beier, 1959, M. barbatus Beier, 1951, M. glandulosus Mahnert, 2009,
M. limatus Hoff and Parrack, M. loebli Schawaller, 1991, M. monstrosus Beier, 1966,
M. trautneri Schawaller, 1994 and M. tuberosus Mahnert, 2009 (Beier 1951, 1959, 1966,
1967; Hoff and Parrack 1958; Schawaller 1991, 1994; Mahnert 2009). Other species for
which males have been described have short setae as in the majority of females. These
include M. grandis (Beier, 1930), M. mongolicus (Redikorzev, 1934), M. soriciola Beier,
1974, M. titanius Beier, 1951 and M. vietnamensis Beier, 1967 (Beier 1930, 1951, 1967,
1974; Redikorzev 1934). Of the species for which males have not been fully described
or illustrated, M. kanneliyensis is substantially larger than M. crinitus Beier 1948,
M. himalayensis (Ellingsen 1914), M. ochotonae Krumpál and Kiefer 1982,
M. papuanus Beier 1948, M. pavlovskyi Redikorzev 1949, M. penicillatus Beier 1948
and M. queenslandicus Beier 1948, which have a female pedipalpal femur length
less than 1.30 mm (Beier 1932, 1948; Redikorzev 1949; Krumpál and Kiefer 1982;
Dashdamirov 2005), compared with 1.44 mm for M. kanneliyensis. Of those species
of a comparable size, M. kanneliyensis differs as follows: the pedipalpal segments are
less slender than in M. philippinus Beier, 1966 (Beier 1966); and the prolateral face of
the chelal hand is only slightly curved in M. kanneliyensis, but is noticeably curved in
M. ryugadensis Morikawa, 1954 (Morikawa 1954).
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Journal of Natural History 2525
Figure 4. Megachernes kanneliyensis sp. nov., paratype female, unless stated otherwise.
(A) Carapace, dorsal; (B) coxae III and IV, ventral; (C) right pedipalp, male holotype, dorsal;
(D) right pedipalp, dorsal; (E) right chelicera, dorsal; (F) right rallum; (G) left leg IV, lateral;
(H) left leg I, lateral; (I) left chela, lateral; (J) left chela, protonymph, dorsal. Scale bars =1.0 mm
(A,C,D,G,H),0.5mm(B,I),0.2mm(E,J),0.1mm(F).
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2526 M.S. Harvey et al.
Description – Adults
Pedipalps, cephalothorax, abdomen and legs uniformly pale yellow (Figures 2, 3).
Vestitural setae generally short and inconspicuous, lightly dentate; most sternal setae
acicular. Pleural membrane longitudinally wrinkled striate for entire length, without
setae.
Pedipalps. Robust (Figures 2A–D, 4C,D), with trochanter 1.45 (♂), 1.55 (♀), femur
2.48 (♂), 2.62 (♀), patella 2.14 (♂), 2.56 (♀), chela (with pedicel) 2.78 (♂), 3.41 (♀),
chela (without pedicel) 2.54 (♂), 3.08 (♀), hand (without pedicel) 1.26 (♂), 1.44 (♀)
times longer than wide; movable finger 1.12 (♂), 1.14 (♀) times length of hand. All
pedipalpal surfaces finely granulate. Fixed finger with c.60 (♂), c.64 (♀) marginal teeth,
movable finger with c.64 (♂), c.70 (♀) marginal teeth; accessory teeth present in dis-
tal half of fingers, fixed finger with six (♂), eight (♀) retrolateral and 12 (♂), seven
(♀) prolateral teeth; movable finger with six (♂), seven (♀) retrolateral and 10 (♂), 6
(♀) prolateral teeth; fingers not gaping when closed; chelal gland pores not observable.
Pedipalpal setae: male with setae on trochanter and femur small and inconspicuous, on
patella and chela long and slightly curved, especially on prolateral margin (Figure 4C);
female with all setae small and inconspicuous (Figure 4D). Fixed finger with eight tri-
chobothria, movable chelal finger with four trichobothria (Figure 4C,D,I); eb closer
to esb than to est;est closer to et than to esb;ib and ist inserted basally; isb closer
to ist than to ib;sb closer to bthan to st;st closer to tthan to sb. Venom appara-
tus only present in movable finger with nodus ramosus terminating between tand st
(Figure 4I).
Chelicerae. Hand with seven (♂,♀) setae, including the usual five setae and two
additional bs setae, all except ls and is dentate; movable finger with one acuminate
seta (Figure 4E). Galea long and stout with seven (♂), nine (♀) prominent dis-
tal rami (Figure 4E). Rallum (Figure 4F) composed of three blades; each dentate
along anterior margin. Serrula exterior with 26 (♂), 20 (♀) lamellae. Lamina exterior
present.
Cephalothorax. Carapace (Figures 2A,C, 4A) 1.09 (♂), 0.99 (♀) times longer than
wide, broadest posteriorly; unicolored; eyes and eye-spots completely absent; setae
small and inconspicuous, with numerous setae, including about six (♂), five (♀)setae
near anterior margin, and c.59 (♂), c.62 (♀) setae on posterior disk; posterior half
with two deeply incised transverse furrows, anterior furrow crosses the carapace at
c.0.56 (♂), c.0.57 (♀) of carapace length, posterior furrow crosses at c.0.80 (♂), c.0.84
(♀) of carapace length; completely granulate with exception of transverse furrows.
Manducatory process with one long distal and several other sub-distal setae, and
with short, curved sub-oral seta; remainder of maxilla with numerous setae. Coxae
(Figures 2B,D, 3, 4B) with numerous small setae over entire ventral surface, with
longer setae on posterior margin of coxae IV; posterior corners of coxae IV rounded
and slightly protruding but not particularly enlarged.
Abdomen. Tergites I–X and sternites V–X divided (Figure 2). Tergal chaetotaxy, ♂: 44:
49: 53: 66: 68: 64: 63: 65: 48: 39: 17: 2; ♀: 62: 60: 72: 76: 74: 72: 70: 68: 54: 52: 24: 2; setae
usually restricted to posterior and lateral tergal margins. Sternal chaetotaxy, ♂: 70: (4)
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Journal of Natural History 2527
61 (4): (2) 32 (2): 58: 66: 62: 60: 53: 45: 14: 2; ♀: 40: (2) 14 (2): (4) 42 (4): 66: 72: 82: 76:
72: 48: 14: 2.
Genital region. Male not studied in detail; female genitalia with anterior genital oper-
culum with setae arranged in inverted-U, some long most short; spermathecae not
observable. Brood-sac with c.24 embryos (Figure 3).
Legs. Legs I and II with a strongly oblique junction between femur and patella
(Figure 4H). Leg IV (Figure 4G) with femur +patella 3.82 (♂), 3.72 (♀) times longer
than broad. Tibiae of legs III and IV without tactile setae. Tarsi of legs III and IV with
long tactile seta (Figure 4G), TS =0.59 (♂), 0.54 (♀). Tarsi with slightly raised slit
sensillum. Sub-terminal tarsal setae arcuate, not dentate. Tarsal claws simple; arolium
shorter than claws.
Dimensions (mm).Male holotype: body length 4.2. Pedipalps: trochanter 0.94/0.65,
femur 1.54/0.62, patella 1.56/0.73, chela (with pedicel) 2.47/0.89, chela (without pedi-
cel) 2.26, hand length (without pedicel) 1.12, movable finger length 1.25. Chelicera
0.51/0.27, movable finger length 0.36. Carapace 1.72/1.58. Leg I: femur 0.48/0.30,
patella 0.82/0.29, tibia 0.68/0.20, tarsus 0.53/0.16. Leg IV: femur +patella 1.45/0.38,
tibia 1.16/0.23, tarsus 0.70/0.19.
Female paratype. Body length 4.2. Pedipalps: trochanter 0.79/0.51, femur 1.44/0.55,
patella 1.36/0.53, chela (with pedicel) 2.08/0.61, chela (without pedicel) 1.88, hand
length (without pedicel) 0.88, movable finger length 1.00. Chelicera 0.39/0.23, movable
finger length 0.30. Carapace 1.52/1.54. Leg I: femur 0.42/0.27, patella 0.75/0.25, tibia
0.61/0.19, tarsus 0.48/0.14. Leg IV: femur +patella 1.34/0.36, tibia 1.04/0.22, tarsus
0.65/0.16.
Description – protonymph
Uniformly very pale yellow, nearly white. Vestitural setae generally short and incon-
spicuous, lightly dentate. Pleural membrane longitudinally striate for entire length,
without setae.
Pedipalps. Robust, with trochanter 1.60, femur 2.44, patella 2.12, chela (with pedicel)
3.35, chela (without pedicel) 3.10, hand (without pedicel) 1.65 times longer than wide;
movable finger 0.97 times the hand (with pedicel). All surfaces of pedipalps finely gran-
ulate with exception of fingers. Fixed finger with three trichobothria, movable chelal
finger with one trichobothrium (Figure 4J); eb and ist situated basally; et situated
subdistally; tmedially. Venom apparatus only present in movable finger with nodus
ramosus terminating slightly distal to t.
Chelicerae. With four setae on hand; movable finger without seta. Galea long and stout
with seven prominent distal rami. Rallum composed of three blades.
Cephalothorax. Carapace: 0.91 times longer than wide, broadest posteriorly; eyes and
eye-spots completely absent; setae small and inconspicuous, with five setae on anterior
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2528 M.S. Harvey et al.
margin, and six setae on posterior margin; shallow median furrow present, posterior
furrow absent.
Abdomen. Tergites I–X and sternites II–X divided. Tergal chaetotaxy: 6: 6: 6: 6: 6: 6:
6: 6: 6: 6: T2T: 2. Sternal chaetotaxy: 0: (0) 2 (0): (1) 4 (1): 7: 6: 6: 6: 6: 6: 4: 2.
Legs. Tarsi of legs III and IV with long tactile seta, situated medially.
Dimensions (mm).Body length 1.56. Pedipalps: trochanter 0.24/0.15, femur 0.39/0.16,
patella 0.36/0.17, chela (with pedicel) 0.67/0.20, chela (without pedicel) 0.62, hand
length (without pedicel) 0.33, movable finger length 0.32. Carapace 0.56/0.61.
Remarks
Megachernes kanneliyensis is the first species of Megachernes recorded from Sri Lanka
and the first pseudoscorpion recorded from the pelage of a Sri Lankan mammal. The
occurrence of adult and nymphal specimens of M. kanneliyensis in the fur of Mus
mayori pococki strongly suggests that the pseudoscorpions are either breeding in the
pelage or in their nests and are being transported phoretically by the mouse. Species
of Megachernes do not appear to be restricted to individual host species (Table 1).
Host associations of Megachernes
Species of Megachernes are closely associated with other animals, either in the nests
or fur of small mammals, nests of birds and bumblebees, or in bat guano within
caves (Tables 1, 2). The following species have been collected most frequently or exclu-
sively in the nests or pelage of small mammals: M. barbatus,M. crinitus,M. grandis,
M. himalayensis,M. kanneliyensis,M. limatus,M. monstrosus,M. ochotonae,M. peni-
cillatus,M. philippinus,M. queenslandicus,M. soricicola,M. titanius,M. trautneri
and M. vietnamensis, as well as three undescribed species from Papua New Guinea
and Australia (Table 1). This assemblage may also include M. loebli, specimens of
which were sifted from leaf litter at the foot of a wall (Schawaller 1991), a habitat
that Schawaller (1991) suggested is frequently occupied by murids and that a close
association with a small mammal was likely.
Three other species, M. afghanicus,M. pavlovskyi and M. ryugadensis, have been
collected from guano in caves (Table 1). Although the origin of the guano is rarely
stated, deposits of bat faeces are the likely source in all cases. Two of these species have
also been recorded from other habitats. Megachernes pavlovskyi has been taken from
within a barn, under stones, in pigeon guano, forest litter, plant roots and rodent nests,
and the subspecies M. ryugadensis myophilus differs from the other two subspecies
(M. ryugadensis ryugadensis and M. ryugadensis naikaiensis) by its close associa-
tion with other animals, having been collected on small mammals (Rattus norvegicus
and Mogera wogura) and from nests of bumblebees. Of the remaining three species
(Table 1), M. glandulosus and M. tuberosus have been collected in caves without any
mention of whether they were taken in guano (Mahnert 2009), and M. mongolicus
lacks any habitat data (Redikorzev 1934).
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Journal of Natural History 2529
Table 1. Habitat data and hosts recorded for species of Megachernes
Taxon Country Host and habitat References
M. afghanicus Beier, 1959 Afghanistan from guano in cave (Beier 1959, 1960)
M. barbatus Beier, 1951 Vietnam from Niviventer tenaster (given as Rattus niviventer champa),
Maxomys surifer (given as R. rajah koratis)andR. sladeni
sladeni
(Beier 1967)
Vietnam taken from fur of various species of rats (Judson 1997)
M. crinitus Beier, 1948 Indonesia on Maxomys bartelsii (given as Rattus bartelsi); from Hylomys
suillus;fromRattus lepturus
(Beier 1948)
M. glandulosus Mahnert, 2009 China in cave (Mahnert 2009)
M. grandis (Beier, 1930) Philippines from Apomys insignis,A. sp., Bullimus bagobus (given as Apomys
bagobus and Rattus bagobus), Cyanopterus brachyotis,Maxomys
panglima (given as Rattus palawanensis)andRattus sp.
(Beier 1966)
Indonesia from Maxomys musschenbroekii and Niviventer fulvescens (given
as Rattus bukit treubi)
(Beier 1948; Durden
1986)
Malaysia from Maxomys rajah (given as Rattus rajah) (Beier 1964)
M. himalayensis (Ellingsen,
1914)
China from Niviventer confucianus (given as Echimys confucianus) (Beier 1948) (as
M. sinensis Beier,
1932)
India from conifer leaf litter (Beier 1978; Schawaller
1988)
Nepal from Apodemus gurkha,Apodemus sylvaticus,Mus musculus,
Niviventer fulvescens (given as Rattus fulvescens)andRattus
pyctoris (given as Rattus rattoides)
(Beier 1974; Martens
1975; Schawaller 1983)
M. kanneliyensis sp. nov. Sri Lanka from pelage of Mus mayori pococki this study
M. limatus Hoff and Parrack,
1958
Papua New
Guinea
from Melomys sp., Pogonomys sp., Rattus niobe,Rattus nitidus
(given as Rattus ruber trimitius∗), and from rat
(Hoff and Parrack 1958;
Beier 1965; Durden
1991)
M. limatus crassus Beier, 1965 Indonesia from unidentified rodents [given as Rattus sp. by Beier (1965)] and
from Paramelomys cf. platyops [given as Pseudocheirus sp. by
Beier (1965)]
(Beier 1965; Tenorio and
Muchmore 1982)
(Continued)
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2530 M.S. Harvey et al.
Table 1. (Continued)
Taxon Country Host and habitat References
M. loebli Schawaller, 1991 Nepal from forest litter (Schawaller 1991)
M. mongolicus (Redikorzev,
1934)
Mongolia habitat not stated (Redikorzev 1934)
M. monstrosus Beier, 1966 Philippines from nest of Maxomys panglima (given as Rattus palawanensis by
Beier (1966)
(Beier 1966)
M. ochotonae Krumpál and
Kiefer, 1982
Mongolia under stones in hole of Ochotona dauurica;innestofPasser sp. (Krumpál and Kiefer
1982)
M. papuanus Beier, 1948 Papua New
Guinea
from Paramelomys levipes (given as Melomys levipes), Melomys
sp., Mus sp., Rattus niobe, from unidentified rats and Rainbow
Lory (Trichoglossus haematododus)
(Beier 1948, 1965; Hoff
and Parrack 1958;
Durden 1991; Judson
1997)
M. pavlovskyi Redikorzev, 1949 Afghanistan from guano in cave (Beier 1959)
Azerbaijan in cave; in bat guano (Rhinolophus mehelyi,Myotis blythi,
Miniopterus scheibersii)
(Krumpál 1986; as M.
caucasicus Krumpál
1986; Schawaller and
Dashdamirov 1988)
Kyrgyzstan in bat guano; and in pigeon guano (Schawaller 1986)
Pakistan roots of Aesculus and nest of rodent; and under stones at the edge
of a stream
(Dashdamirov 2005)
Russia Taxus –Buxus woods with Fagus (Schawaller and
Dashdamirov 1988)
Turkmenistan in cracks of a livestock barn (Redikorzev 1949)
Turkmenistan in guano (Dashdamirov and
Schawaller 1995)
M. penicillatus Beier, 1948 Australia from Phascogale sp. (Beier 1948; Judson
1997)
M. philippinus Beier, 1966 Philippines from Rattus sp. (Beier 1966; Tenorio and
Muchmore 1982)
(Continued)
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Journal of Natural History 2531
Table 1. (Continued)
M. queenslandicus Beier, 1948 Australia from Rattus fuscipes assimilis,R. fuscipes and grey scrub rat (Beier 1948; Judson
1997)
M. ryugadensis Morikawa,
1954
Japan in bat guano in caves (Morikawa 1954, 1960)
M. ryugadensis myophilus
Morikawa, 1960
Japan from nest of Bombus diversus and B. ardens;onMogera wogura;
on Rattus norvegicus
(Morikawa 1960)
M. ryugadensis naikaiensis
Morikawa, 1957
Japan from bat guano in caves (Morikawa 1957, 1960)
M. soricicola Beier, 1974 Nepal from Soriculus nigrescens (Beier 1974; Martens
1975)
M. titanius Beier, 1951 Vietnam from Niviventer tenaster (given as Rattus niviventer champa),
Maxomys surifer (given as R. rajah koratis), R. sladeni sladeni;
and “from the fur of various species of rats”
(Beier 1967; Judson
1997)
M. trautneri Schawaller, 1994 Thailand from small mammal nest (Schawaller 1994)
M. tuberosus Mahnert, 2009 China in cave (Mahnert 2009)
M. vietnamensis Beier, 1967 Vietnam from Niviventer fulvescens and Rattus andamanensis (originally
reported as Rattus niviventer and R. sladeni sladeni)
(Beier 1967; Tenorio and
Muchmore 1982)
M. sp. indet. Papua New
Guinea
from nests of Micromurexia habbema (given as Antechinus
habbema)andPhascomurexia naso (given as Antechinus naso)
(Woolley 1989)
M. sp. nov. (Papua New
Guinea)
Papua New
Guinea
from Rattus niobe,Melomys sp. and Murexechinus melanurus
(given as Antechinus wilhelmina)
(Durden 1991)
M. sp. nov. (Australia) Australia from Vespadelus regulus (given as Eptesicus regulus)and
Falsistrellus tasmaniensis
(Harvey and Parnaby
1993)
M. sp. nov. (Australia) Australia from nest box of Leadbeater’s Possum (Gymnobelideus
leadbeateri)
(Harvey and Parnaby
1993)
∗The host name “Rattus ruber trimitius” recorded by Beier (1965) appears to be a lapsus. Rattus ruber is now a synonym of R. nitidus (Wilson and
Reeder, 2005) and “trimitius” may refer to the trichozoan blood parasite Trimitus.
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2532 M.S. Harvey et al.
Table 2. Host animals recorded for Megachernes species, excluding associations with mammal
guano (see Table 1).
Phylum Chordata
Class Mammalia
Order Dasyuromorpha
Micromurexia habbema (Tate and Archbold, 1941)
Murexechinus melanurus (Thomas, 1899)
Phascomurexia naso (Jentink, 1911)
Phascogale sp.
Order Diprotodontia
Gymnobelideus leadbeateri McCoy, 1867
Order Soricomorpha
Mogera wogura (Temminck, 1842)
Soriculus nigrescens (Gray, 1842)
Order Erinaceomorpha
Hylomys suillus Müller, 1840
Order Lagomorpha
Ochotona dauurica (Pallas, 1776)
Order Rodentia
Apodemus gurkha Thomas, 1924
Apodemus sylvaticus (Linnaeus, 1758)
Apomys insignis Mearns, 1905
Apomys sp.
Bullimus bagobus Mearns, 1905
Maxomys bartelsii (Jentink, 1910)
Maxomys musschenbroekii (Jentink, 1878)
Maxomys panglima (Robinson, 1921)
Maxomys rajah (Thomas, 1894)
Maxomys surifer (Miller, 1900)
Melomys sp.
Mus mayori pococki Ellerman, 1947
Mus musculus Linnaeus, 1758
Mus sp.
Niviventer confucianus (Milne-Edwards, 1871)
Niviventer fulvescens (Gray, 1847)
Niviventer lepturus (Jentink, 1879)
Niviventer tenaster Thomas, 1916
Paramelomys levipes (Thomas, 1897)
Paramelomys cf. platyops (Thomas, 1906)
Pogonomys sp.
Rattus andamanensis (Blyth, 1860)
Rattus fuscipes (Waterhouse, 1839)
Rattus fuscipes assimilis Gould, 1858
Rattus niobe (Thomas, 1906)
Rattus nitidus (Hodgson, 1845)
Rattus norvegicus (Berkenhout, 1769)
Rattus pyctoris (Hodgson, 1845)
Rattus sp.
(Continued)
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Journal of Natural History 2533
Table 2. (Continued).
Order Chiroptera
Cyanopterus brachyotis (Müller, 1838)
Vespadelus regulus (Thomas, 1906)
Falsistrellus tasmaniensis (Gould, 1858)
Class Aves
Order Psittaciformes
Trichoglossus haematododus (Linnaeus, 1771)
Order Passeriformes
Passer sp.
Phylum Arthropoda
Class Hexapoda
Order Hymenoptera
Bombus ardens Smith, 1879
Bombus diversus Smith, 1869
Acknowledgements
We gratefully acknowledge the assistance of the National Science Foundation, Sri Lanka (grant
number: RG/2004/B2) and the University of Colombo for funding this project; the Forest
Department and the Department of Wildlife Conservation, Sri Lanka, for granting permission
to carry out studies on small mammals in the forest reserves of Sri Lanka; Mr P.M. Weerawarna
for field assistance and his support which extended throughout the study; Dr M. Burger
(Western Australian Museum) who kindly provided some of the digital images; Dr M. Judson
(Muséum National d’Histoire Naturelle, Paris) for providing information on Chiridiochernes;
and Dr Judson and three anonymous reviewers for the comments on the manuscript.
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