The Rhipiceridae of Taiwan
and Japan
(Insecta: Coleoptera: Dascilloidea)
Chi-Feng Lee1,*
,
Masataka Satô2, and Masahiro Sakai3
1Research Center for Biodiversity,
Academia Sinica, Taipei Taiwan 115, R.O.C.
2Dia Cuore 306, Kamegahora 3-1404, Midoriku,
Nagoya, 458-0804 Japan
3Entomological Laboratory, College of Agriculture, Ehime University,
3-5-7 Tarumi, Matsuyama,
Ehime Prefecture,
790-8566 Japan
*To whom
correspondence and reprint request should be addressed.
E-mail: cflee@gate.sinica.edu.tw
Tel: 886-2-27899525
Fax: 886-2-2785-8059
Received: 11 February
2004 Accepted: 20 April
2005 Published
online: 27 July 2005
Abstract Chi-Feng
Lee, Masataka Satô, and Masahiro Sakai (2005) The
Rhipiceridae of Taiwan and Japan
(Insecta: Coleoptera: Dascilloidea).
Zoological Studies 44(4): xxx-xxx. Species of the family Rhipiceridae in Taiwan and Japan are reviewed. Three
known species are regarded as valid:
Sandalus kani Sakai and Sakai
(1981), S. segnis Lewis (1887), and S. sauteri Emden (1924). Sandalus
takizawai Nakane (1985) is a junior synonym
of S. sauteri Emden. Sandalus taiwanicus Lee et al., sp. nov. and S.
sauteri lanyuensis Lee et al., ssp. nov. are
described and S. segnis Lewis (1887)
is recorded in Taiwan
for the 1st time.
Keywords: Taxonomy, Sandalus, New species, Cicada parasite
beetles
----------------------------------------------------------------------------------------------------------------
Introduction
The common name, “cicada parasite beetles”, of the Rhipiceridae
reflects the biology of Sandalus
larvae. According to observations
in North America (Elzinga 1977), the female of Sandalus niger
Knoch deposits enormous numbers of eggs (16,864 eggs; counted by Rings
1942) in holes or under bark of elms where cicadas possibly oviposit. Eggs are usually washed off by the 1st
rain and hatch into triungulin
larvae. A late-instar ectoparasitic
larva was discovered within the nymphal exuviae of a dead cicada which had
failed to emerge from its burrow
(Craighead 1921). Young (1956)
observed an abrupt abundance of S. niger adults in southern Indiana, and he
suggested that the emergence of this rhipicerid might have been connected with
the emergence of periodical cicadas (Magicicada
spp.) in the preceding year.
Regarding the Far Eastern species, Fukuda (1969) reported the
ovipositing behavior of S. segnis
Lewis in Aomori Prefecture,
Japan; he also
stated that the host cicada of this species is presumably Terpnosia
nigricosta (Motschulsky) or Tibicen
bihamatus (Motschulsky).
Rhipiceridae is a relatively small family, including 7 genera and
about 100 species (Lawrence 2005).
Only the genus Sandalus Knoch
occurs in East Asia. Four species of Sandalus have been reported from Taiwan and Japan: S. segnis Lewis (1887) recorded from the
main islands of Japan (Ohno
1995), S. kani Sakai
and Sakai (1981) from Amami-Ôshima
I., S.
takizawai Nakane (1985) from Ishigaki I., and S. sauteri Emden (1924) from Taiwan. Although Sakai
and Sakai
(1981) and Nakane (1985) mentioned some
diagnostic characters for
distinguishing species, some of them are neither reliable nor useful. This paper provides a comprehensive
study of the taxonomy of Sandalus
from Taiwan and Japan. Some diagnostic characters are proposed
through a comparison of all species.
----------------------------------------------------------------------------------------------------------------
MATERIALS AND METHODS
Specimens were examined with a Leica MZ95 stereoscopic microscope
under diffuse lighting. Genitalia
and antennae were examined with a Nikon Optiphot transmitted light microscope
and illustrated with the aid of a drawing tube. Illustrations of punctures on the
pronotum were made with Photoshop software.
Specimens examined are deposited in the following museums or
institutions (letter codes largely follow Arnett et al. 1993): BMNH, The
Natural History Museum, London, UK; BPBM, Bernice P. Bishop Museum, Honolulu,
HI, USA; DEI, Deutsches Entomologisches Institut im ZALF, Müncheberg, Germany;
EIHU, Hokkaido Univ., Sapporo, Hokkaido, Japan; EUMJ, Ehime University,
Matsuyama, Japan; HY, Collection of H. Yoshitomi, Hokkaido, Japan; MNHN, Muséum National d'Histoire Naturelle,
Paris, France; NCHU, National Chung Hsing Univ., Taichung, Taiwan; NMNS,
National Museum of Natural Science, Taichung, Taiwan; TARI, Taiwan Agricultural
Research Institute, Wufeng, Taichung, Taiwan; TFRI, Taiwan Forestry Research
Institute, Taipei, Taiwan.
SYSTEMATIC ACCOUNTS
Genus Sandalus Knoch, 1801
Diagnosis: Head elongate-quadrate, weakly
deflexed, inserted slightly into pronotum, and with prominent antennal tubercles. Eyes relatively large and visible. Antennae 11-segmented, flabellate from
segments 3 to 11 in
male (Fig. 3A),
serrate to pectinate from segments 3 to 9 in
female (Fig. 3B, C), 2 apical segments fused in females
of some species. Mentum triangular and
plate-like; submentum reduced; ligula small, conical, and densely setose;
palpus 3-segmented, 1st segment much smaller than others, 2nd and 3rd segments
subequal in length, apex of apical segment pointed, all segments covered with
fine hairs except for apex of apical
segment. Lacinia reduced; galea
well developed, cylindrical, setose or spinose; palpus 4-segmented, 1st segment small, other segments cylindrical
and with dense hairs except on apex. Mandibles large, apices strongly and
abruptly curved inward, unidentate.
Labrum small, bilobed, fused to head.
Pronotum densely punctate and setose, transverse, basally widened;
apical margin rounded; basal margin smooth or weakly crenulate, median lobe bidentate; base of pronotum distinctly narrower
than bases of elytra. Prosternal process small, connecting
with mesosternum. Anterior margin
of mesosternum with a forward process, posterior margin with a bilobed process. Scutellum small, rhomboid. Metasternum large, with transverse
suture parallel to posterior margin.
Elytra punctuate and pubescent,
with longitudinal and transverse carinae.
All legs similar in shape; 1st and 2nd coxae globular, hind coxae
transverse; trochanters small, but distinct;
femora with ventral groves for receiving tibiae; tibiae roughened, dorsal side
bearing various teeth, ventral surface smooth, apex with 2 long curved spurs;
tarsi 5-segmented, 1st 4 segments lobed, with paired, ventral, densely setose lobes; claws
simple; empodium club-shaped, bearing hairs near apex.
Abdomen with 5 ventrites, densely
pubescent. Male genitalia
trilobate, symmetrical; penis with dorsal and ventral lobes, ventral lobe more
slender than dorsal lobe, apex recurved; apices of parameres rounded, with
sparse hairs; basal piece short, transverse.
Notes: This genus has attracted the attention of many Japanese
entomologists. Ohno (1995) counted 104 papers on S. segnis Lewis and the allied species. In Taiwanese and Japanese species,
antenomeres 10 and 11 are separated, differing from the fused antenomeres in
American species.
Distribution: Nearctic, Neotropical, eastern Palaearctic, Oriental and
Afrotropical regions (25 species: Katovich 2002).
Sandalus kani Sakai and Sakai
(Figs. 1A, B, 2C, 4A,
5A)
Sandalus kani Sakai and Sakai 1981: 151; Nakane
1985: 33.
Type series: Holotype ♂ (Fig. 1A, B): “(Chuohrinodoh) Amami Kagoshima, 1st
July, 1980 Hikaru Kan leg. / Holotype Sandalus kani Sakai
et Sakai 1981
(red)” (EUMJ, examined).
Fig.
1.
Habitus and labels of Sandalus
species.
(A) Habitus of the holotype of S. kani; (B) labels of the holotype of S.
kani; (C) habitus of the syntype of S. sauteri sauteri; (D) labels of the
syntype of S. sauteri sauteri; (E) habitus of the paratype of S. takizawai; (F) labels of the
paratype of S. takizawai; (G) habitus of S.
segnis; (H) habitus of S. taiwanicus.
Description: Male: Length 10.8 mm, width 4.8 mm.
Body oblong, about 2.1 times longer than wide; coloration dark brown, but elytra brown; base of elytra and 1st to basal 1/2 of 3rd abdominal segments
yellowish-brown; head, pronotum, venter, and base of elytra clothed with
yellowish-brown pubescence, remainder of
elytra and last abdominal sternite clothed with blackish-brown pubescence.
Pronotum trapezoidal, slightly depressed on each side of middle at rear of anterior margin and anteriad posterior
margin, moderately angular near hind corner of lateral
margin (Fig. 2C); punctures of 2 sizes: large punctures much fewer and larger than small punctures (Fig. 4A).
Elytra elongate, about 1.6 times as long as wide, each with 3 distinct
longitudinal carinae; intervals finely punctured. Genitalia:
parameres like those of S. sauteri;
apex of ventral lobe recurved like that of
S. taiwanicus, but recurved area and teeth smaller (Fig. 5A).
Female: Similar to male, but larger (16.5
mm); coloration blackish; elytral pubescence yellowish-brown;
elytral punctures smaller; elytral carinae distinct, a number of cross-carinae
present on elytra; antennae similar to those of S. segnis.
Diagnosis: This species is similar to
S. sauteri but has a greater number of large-sized pronotal punctures and
differs by the fine elytral punctures.
Distribution: Japan
(Amami-Ôshima I.).
Fig.
2.
Pronota. (A) Sandalus
taiwanicus; (B) S. sauteri sauteri; (C) S. kani.
Fig.
3. Antenna. (A) Sandalus
sauteri sauteri, male; (B) S. segnis, female; (C) S. sauteri sauteri, female.
Fig. 4. Pronotal punctures. (A) Sandalus
kani; (B) S. sauteri
sauteri; (C) S. sauteri lanyuensis; (D) S.
segnis; (E) S.
taiwanicus.
Fig.
5. Male genitalia and apices of penises, ventral
view. (A) Sandalus kani, apex of
penis; (B) S. sauteri sauteri, apex
of penis; (C) S. segnis, apex of penis; (D) S. taiwanicus, apex of
penis; (E) S. sauteri sauteri,
genitalia; (F) S. taiwanicus, genitalia; (G) S. segnis, genitalia.
Sandalus
sauteri sauteri Emden
(Figs. 1C,
D, 2B, 3C, 4B, 5B, E)
Sandalus sauteri Emden 1924: 28; Miwa 1928: 374.
Sandalus takizawai Nakane 1985: 35. syn. nov.
Type series: Lectotype ♂, herewith designated (Fig. 1C, D): “Hoozan, Formosa, H. Sauter IX 10 / coll. DEI
Müncheberg / Sandalus sauteri n. sp.
v.Emden det. 1923 / Syntype (red)” (DEI, examined); number of syntypes
uncertain.
Synonym: Holotype ♂ of Sandalus
takizawai: “Omotodake, Ishgaki Is., Ryukyus, Japan, 6.VII.1974, H. Takizawa
leg.”; paratype ♂ (Fig. 1E, F): “Omotodake, Ishigaki Is., 30.V.1973, K Sugino
leg. / Sandalus takizawai n s. Det T. Nakane (pink) / PARATYPE (red)” (all in
EIHU, paratype examined).
Description: Male: Length 12.1~14.4 mm, width 4.9~6.7 mm.
Body oblong, about 2.1~2.4 times longer wide; coloration blackish-brown,
but antennae brown, elytra brown or blackish-brown, elytral bases pale; head,
pronotum, venter, and elytral bases clothed with yellowish-brown pubescence, remainder of elytra with blackish-brown
pubescence. Pronotum trapezoidal,
slightly depressed on each side of middle at rear of
anterior margin and anteriad posterior margin, moderately angular near hind
corner of lateral margin, hind corner lobed (Fig. 2B); punctures on surface of
2 sizes: large punctures fewer and much larger
than small punctures (Fig. 4B). Elytra elongate, about 1.8 times as long
as wide, each provided with 4 distinct
longitudinal carinae, 5th carina indistinct in some individuals; intervals
prominently punctured. Genitalia (Fig. 5E): parameres abruptly
narrowed at apical 1/4, forming long, slender apices; apex of ventral lobe
open, ring-like, with lateral, upcurved teeth (Fig. 5B).
Female: Similar to male, but larger (20.0
mm); elytral pubescence yellowish-brown; elytral carinae more
prominent, elytral punctures tiny; antennal segments 3~11 (Fig 3C) pectinate, progressively longer
toward segment 8, then progressively shortened toward apical segment; apical
segment dilated
Diagnosis. See diagnosis of S. kani.
Material examined: 1 ♂: “C. Taiwan, Kukuan 730 m,
Taichung Hsien 20-22.VI.1978. KSLin” (TARI); 1 ♂, 1 ♀: “Nakasuji,
Ishigaki-jima, 26-IV-2001, T. Fukaishi” (NMNS); 9 ♂♂, 2 ♀♀: “Taiwan (Formosa), Puli (Hori), June~July,
1953~1954, Native Collector” (BPBM).
Distribution: Taiwan and Japan (Ishigaki
I.).
Sandalus
sauteri lanyuensis Lee et al., ssp. nov.
(Fig. 4C)
Holotype ♂: “TAIWAN
Orchid I.
(Botel Tobago) 4-8.III.1991 C.K.
Starr and H.Y. Wang / NMNS ENT 1164-93 (NMNS).
Paratypes: 1 ♂, same collecting data for holotype / NMNS ENT 1164-97; 1 ♂: “Taiwan: Taitung, Lanyu I., Tienshih,
Apr. 2003, J.Y. Liou leg.” (NCHU).
Description: Male: Length 9.6~11.0 mm, width 4.3~4.9 mm. Body oblong, about 2.1~2.2 times longer than
wide; coloration blackish, but elytra brown
and pale basally; head, pronotum, venter, and elytral bases clothed with
yellowish-brown pubescence, remainder of
elytra with blackish-brown pubescence.
Pronotum trapezoidal, slightly depressed on each side of midline behind
anterior margin, irregularly depressed, weakly angular near hind corner of lateral
margin; punctures of 2 sizes: large punctures fewer than small punctures (Fig. 4C). Elytra elongate, about 1.7 times longer
than wide, each with 4 distinct longitudinal carinae; intervals prominently
punctured. Genitalia: similar to nominate subspecies.
Female: Unknown.
Diagnosis: This new subspecies differs from the nominate subspecies by the
smaller “large-sized punctures” on pronotum, and weak angular processes near
the hind corners of the pronotum.
Distribution: Taiwan (Orchid I.
[Lanyu in Chinese]).
Sandalus segnis Lewis
(Figs. 1G, 3B,
4D, 5C, G)
Sandalus segnis Lewis 1887: 316; Miwa
1928: 374; Nakane 1985: 33. (see Ohno 1995 for additional references)
Sandalus semitestaceus Pic 1906: 1,
synonymized by Jakobson (1913)
Type series: Lectotype ♂ : “CO Type (yellow letters, circle label) / Sandalus
segnis Lewis DET. Miwa / 17”;
one paralectotype ♀, same data with holotype except 17 replaced with 16, TARI,
herewith designated.
Synonym: Holotype ♂ of Sandalus
semitestaceus: “Japon: Kioto / type (hand writing) / Type (printed, black
ink in red card) / Sandalus semitestaceus
Pic (hand writing)” (MNHN, examined).
Description: Male: Length 12.9 mm, width 5.4 mm. Body oblong, about 2.4 times longer than wide;
coloration blackish-brown, elytra blackish or yellowish; head, pronotum,
venter, and elytral bases clothed with yellowish-brown pubescence, remainder of elytra with blackish-brown
pubescence. Pronotum trapezoidal,
lightly depressed on each side of middle at rear of
anterior margin; moderately angular near hind corner of lateral margin;
punctures subequal in size (Fig.
4D). Elytra elongate, about 2.0
times as long as wide, each with 4 indistinct longitudinal carinae; intervals
prominently punctured. Genitalia (Fig. 5G): parameres parallel, slightly narrowed near
apices; apex of ventral lobe depressed, with a pair of acute upcurved processes
(Fig. 5C).
Female: Similar to male, but larger (length 18.1~19.2 mm); elytral pubescence yellowish-brown; elytral
carinae indistinct; elytral punctures tiny; antennae similar to those of S. sauteri sauteri, but antennal rami
shorter (Fig. 3B).
Diagnosis: It is characterized by the equal sized pronotal punctures and the
indistinct elytral carinae.
Material examined: 1 ♂ (Fig. 1G): “(SHIKOKU) Mt. Takanawa Ehime Pref. 29-VI, 1980 Y. SEIYAMA
leg.” (EUMJ); 2 ♀♀, same collector and locality, but with different dates:
“26-VI, 1979”; and “27-VI, 1978” (EUMJ); 1 ♂: “Iyo (Ehime Pref.),
Japan, 201116”; 1♂: Houheikyo, Sapporo-shi, Hokkaido,
29-VI-2000, H.Yoshitomi (HY); 1♀: Kanehachi-toge, Hokkaido, 20-VI-2001, H.Yoshitomi (HY); 1 ♂: “Fushan A2, Ilan
Co., TAIWAN AN, III-(8-22)-2004, S. S. Lu / 00089428 (barcode)” (TFRI).
Distribution: Japan (Hokkaido, Honshu, Shikoku, and Kyushu) and Taiwan (new record).
Sandalus taiwanicus Lee et al., sp. nov.
(Figs. 1H, 2A, 4E,
5D, F)
Holotype ♂: Taiwan:
“Pingtung, Dahanshan, 26-IV-2003, K.-S. Hsu” (NMNS).
Paratypes: 2 ♂♂ (Fig. 1H): “Nan Feng Shan, near Liu Kui, S. Taiwan, 18.IV.1986, Col. K. Baba” (EUMJ); 1♂, same locality as holotype, 25.III.2005,
leg. S.-J. Lyu (DEI); 1♀: “S. TAIWAN: KaoShiung Co. Mid-elevation Exp. Station
near Teng Zhih 27/V-22/VI/2005 alt. ca 1600m
by FIT collr. C.-L. Li” (NMNS).
Description: Male: Length 13.4~17.7 mm, width 6.0~7.1mm. Body oblong, about 2.2~2.5 times longer than
wide; coloration blackish-brown; body clothed with yellowish-brown
pubescence. Pronotum trapezoidal,
slightly depressed on each side of middle at rear of
anterior margin and deeply depressed anteriad posterior margin; without angular process near hind corner of
lateral margin (Fig. 2A);
punctures of 2 sizes: large punctures much fewer and much larger than small punctures (Fig. 4E). Elytra
elongate, about 1.8 times longer than wide, each provided with four distinct longitudinal carinae, intervals finely
punctured. Genitalia (Fig. 5F):
parameres parallel, internal margins abruptly depressed distally, forming
short, slender apices; apex recurved, with 1 pair of teeth (Fig. 5D).
Female: Similar to male, but larger (length 19.2 mm); elytral carinae indistinct; elytral
punctures tiny; antennae similar to those of S. sauteri sauteri.
Diagnosis: Similar to S. kani Sakai and Sakai
with small elytral punctures, but characterized by lacking angular processes
near the hind corners of the pronotum and having fewer large-sized pronotal
punctures.
Distribution: Taiwan.
----------------------------------------------------------------------------------------------------------------
DISCUSSION
A number of characters have previously been proposed as being
diagnostic characters. Miwa (1928)
indicated that S. sauteri sauteri was
recognized by 4 longitudinal carinae on each elytron and a median longitudinal
depression on the pronotum.
Actually, most species have 3 or 4 distinct elytral carinae, and others
are distal and too faint to observe.
The latter character is shared among all species
of Japan and Taiwan. Sakai and
Sakai (1981) mentioned a number of characters for distinguishing
S. segnis and S. kani. Most of them
are too variable except for the punctures on
the pronotum which are reliable. In addition to those characters, the elytral punctures, the pronotal shapes, and the male genitalic structures are also useful
in differentiating the species. It
should be noted that body sizes and coloration are so variable that it is impossible to use these characters to
identify species. Variation in
these characters are likely related to the sizes and species of hosts.
Key to males of Sandalus
species from Taiwan and Japan
1. Elytra finely punctate …..…………................…………………………………. 2
- Elytra bearing prominent punctures ……. .…………………………………….. 3
2.
Pronotum furnished with an angular process near corner, and bearing many
large-sized punctures (Fig. 4A) …......………...…………… S. kani Sakai
and Sakai
- Pronotum without an angular
process near corner (Fig. 2A), and bearing fewer
large-sized punctures …..………………...............
S. taiwanicus Lee et al., sp. nov.
3.
Pronotum with
punctures of 1 size (Fig. 4D); elytral carinae
indistinct ……………………........................................................................
S. segnis Lewis
- Pronotum bearing different
sizes of punctures (Fig. 4B, C); elytral carinae
distinct
..................................................................................................................
4
4.
Large-sized punctures on
pronotum smaller, about 2 times longer than small-sized punctures in diameter
(Fig.4C)
..................................................................................
..................................................................
S. sauteri
lanyuensis Lee et al., ssp. nov.
-
Large-sized punctures on pronotum larger, about 3 times longer than small-sized
punctures in diameter (Fig. 4B) ........................... S. sauteri sauteri Emden
Acknowledgments: We are
indebted to John F. Lawrence for providing valuable information. We also thank Yu-Long Lin and Chun-Lin
Li (Taipei), Kuo-Sheng Hsu (Pingtung), Sheng-Jhih Lyu (Kaoshiung), Mei-Ling
Chan (NMNS), Shen-Shan Lu (TFRI), Masahiro Ôhara (EIHU), Hsien-Tzung Shih
(TARI), Man-Miao Yang (NCHU), Hiroyuki Yoshitomi (Sapporo) and Lothar Zerche (DEI) for loans of type specimens and unidentified
materials. We also thank Ming-Luen
Jeng who examined types at the Paris
museum (MNHN).
----------------------------------------------------------------------------------------------------------------
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