ore

JHR 45: |-14 (2015) JOURNAL COR. PA Peetresiewed open atontioureat
doi: 10.3897/JHR.45.5442 (Me) Hymenopter a

http://jhr.pensoft.net The insertional Society of ymenopterists. RESEARCH

Studies on Adelestini (Hymenoptera, Tenthredinidae),
particularly the long-tongued Nipponorhynchus
Takeuchi of Japan

David R. Smith!, Tikahiko Naito?

I Systematic Entomology Laboratory, Agricultural Research Service, U. S. Department of Agriculture, clo Na-
tional Museum of Natural History, Smithsonian Institution, PO. Box 37012, MRC 168, Washington, DC
20013-7012, USA 2. Aoyama-Minami 4-13-7, Himeji-shi, Hyogo Prefecture 671-2223, Japan

Corresponding author: David R. Smith (sawfly2@aol.com)

Academic editor: M. Yoder | Received 13 June 2015 | Accepted 12 August 2015 | Published 7 September 2015
http://zoobank.org/FA374BAD-F83E-4CBB-AEE2-69900D2C38FO

Citation: Smith DR, Naito T (2015) Studies on Adelestini (Hymenoptera, Tenthredinidae), particularly the long-
tongued Nipponorhynchus Takeuchi of Japan. Journal of Hymenoptera Research 45: 1-14. doi: 10.3897/JHR.45.5442

Abstract

Nipponorhynchus brevis Smith & Naito, sp. n., is described from Hokkaido, Japan. It is characterized by
shorter mouthparts than those of the other two species of the genus, VV. bimaculatus Naito and N. mirabilis
Takeuchi. The previously unknown female of N. bimaculatus is described. Larvae of N. bimaculatus and
N. mirabilis feed on Chrysosplenium macrostemon var. shiobarense (Saxifragaceae), and notes on the life
history are given. Nipponorhynchus is compared with the Nearctic Adelesta Ross, the only other genus of
Adelestini. A key to the genera and species of Adelestini is provided.

Keywords
Sawflies, Selandriinae, Adelesta, Chrysosplenium

Introduction

Nipponorhynchus was described by Takeuchi (1941) for a single unusual sawfly spe-
cies, N. mirabilis Takeuchi, 1941, in which the maxillolabial complex is extremely
elongated forming a proboscis-like structure. The mouthparts are so unique that Jervis
and Vilhelmsen (2000) proposed an additional type, “Type 8,” to the known seven

Copyright David R. Smith, Tikahiko Naito. This is an open access article distributed under the terms of the Creative Commons Attribution License
(CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

2 D.R. Smith & T. Naito / Journal of Hymenoptera Research 45: 1-14 (2015)

functional types of elongated proboscises previously recognized among Hymenoptera.
Type 8 was solely for Nipponorhynchus, and it differs from the other types by the
greatly elongated prementum and stipes, together forming a structure with two food
conduction channels.

A second species of Nipponorhynchus, N. bimaculatus, was added by Naito (1973),
which, among other structural features, the mouthparts are elongated, but not quite as
long as those of NV. mirabilis. Here, we describe a third species with elongated mouth-
parts, but much shorter than the two previously described species, thus showing a
gradient in length of the mouthparts from moderately to extremely long. Additionally,
we also describe the female of NV. bimaculatus for the first time.

Nipponorhynchus, known only from Japan, and the Nearctic Adelesta Ross are the
only members of what is recognized as the tribe Adelestini in the Selandriinae. Though
the two genera have a number of characters in common which distinguishes the tribe,
we prefer to keep the two genera separate. A brief history of the tribe and key to the
genera and species of the Adelestini is presented.

Materials and methods

Images for Figures 1-26 were acquired through an EntoVision micro-imaging system.
This system included a Leica M16 or Leica DRMB compound microscope with an
attached JVC KY-75U 3-CCD digital video camera or a GT-Vision Lw11057C-SCI
digital camera, that fed image data to a notebook or desktop computer. The program
Cartograph 6.6.0 was then used to merge an image series into a single in-focus image.
Figures 27, 28, 30, and 31 were taken by TN using a Nikon D200 digital camera with
AF Micro Nikon 200 mm lens. Figure 29 was taken by T. Saito using a Canon Power
Shot S5 IS digital camera with a Canon Zoom 12XIS 6.0—72.0 mm lens.
Abbreviations used are: NSMT, National Museum of Nature and Science, Ibaraki,
Japan; TN, collection of T. Naito, Himeji-shi, Hyogo Prefecture, Japan; USNM, Na-
tional Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

Results

Nipponorhynchus bimaculatus Naito
Figs 2, 6, 10, 14, 19, 23, 24, 27-29

Nipponorhynchus bimaculatus Naito, 1973: 95, fig. 1.

Description. Female: Length, 5.0 mm. Antenna and head black; anterior half of cly-
peus dark brown, labrum and upper surface of maxillolabial complex whitish; palpi
and undersurface of maxillolabial complex dark brown. Thorax black with tegula and
posterior corners of pronotum whitish. Legs black with apices of coxae, trochanters,

Studies on Adelestini (Hymenoptera, Tenthredinidae)... 3

‘

Figures 1-4. Head, front view. | Nipponorhynchus mirabilis 2 N. bimaculatus 3 N. brevis 4 Adelesta nova.

apical quarter of femora, tibiae except apical quarter with black stripe on inner sur-
face and basitarsomeres whitish. Abdomen black (terga 2 and 3 often with yellow
mark in male); center of apical tergum and cercus whitish. Wings hyaline; veins and
stigma black. Head very shiny, almost impunctate; inner orbits, gena and clypeus with
fine irregular punctures. Thorax and abdomen very shiny, sometimes terga 3—9 finely
reticulate; posterior margin of mesoscutellum with several large punctures.

4 D.R. Smith & T. Naito / Journal of Hymenoptera Research 45: 1-14 (2015)

Antenna length 1.7x head width, with scape and pedicel each longer than
broad; 3"! antennomere length 1.3x 4", 4" to 9* antennomeres gradually decreas-
ing in length. Clypeus slightly broadly, circularly emarginated, about 3x broader
than long. Malar space about equal to diameter of median ocellus. Lower interoc-
ular distance 1.3x eye height; eyes only slightly converging below. Maxillolabial
complex about 1.0—1.1x head width and about 2.2x eye height; maxillary palpus
about 0.6x length of proboscis, longer than distance between maxillary palpus
and labial palpus. Distances between eye and lateral ocellus, between lateral ocelli,
and between lateral ocellus and hind margin of head as 1.0:0.8:0.5. Interocellar
furrow represented by small pit. Postocellar furrow defined. Postocellar area 1.5x
broader than long. Posttergite distinct, length:width as1:4, triangular behind. Tar-
sal claws simple. Hind basitarsomere subequal to length of following 3 tarsomeres
combined. Sheath in lateral view straight above, rounded at apex and below; from
above, equally wide throughout. Lancet in Fig. 19; 10 or more annular spines in
two or three rows.

Male: Described by Naito (1973). Gentialia (Figs 23, 24), penis valve with valvi-
ceps almost round and apex somewhat flattened, dorsal spine slender.

Specimens examined. JAPAN: Honshu, Bicchuzawa, Bato, Tochigi Pref.
15.IV.2004, K. Katayama (12); same locality, 23.1V.2005, T. Naito (24); same
locality, 24—-30.IV.2005, S. Ibuki (42), same locality, 23, 29.IV.2005, A. Shinohara
(22); same locality, 16-17.IV.2012, S. Ibuki (32); same locality, 29.11.2012, S. Ibuki
(emerged from Chrysosplenium macrostemon Maxim var. shiobarense (Franch. Hara) )
(22); Akazai-keikoku, Hyogo Pref. 20.V.1999, T. Okushima (12); Onzui-keikoku,
Hyogo Pref., 21.1V.2008, T. Naito (12); Mt. Ooginosen, Hyogo Pref., 22.V.2006,
T. Naito (19, 1 3); same locality, 7.V.2012, T. Naito; Mt. Hachibuseyama, Hyogo
Pref., 5.V.2013, T. Naito (12); Honshu, Oodaigahara, Nara Pref., 7.VI.1976, T. Nai-
to(1 3 paratype); Honshu, Bicchuzawa, Bato, Tochigi Pref., 23.1V.2005, T. Naito (2
©, 1 2); Hyogo Pref., Onzui-keikoka, 700 m, 21.IV.2008, T. Naito (1 9).

Distribution. Japan (Honshu).

Host and biological notes. TN observed adult habits on the host plant in April
2005 at Bicchuuzawa, Tochigi Pref., Japan. Later, Saito and Ibuki (2010) succeeded in
rearing this sawfly. Adults appear in early spring and fly on the flowers of the host plant,

Chrysosplenium macrostemon Maxim var. shiobarense (Franch. Hara) (Saxifragaceae).
They insert the long proboscis into the flower and absorb the nectar (Fig. 27). Females
cut the seed capsule of the host plant with the sawlike ovipositor and lay their eggs into
the capsule (Fig. 28). Larvae mainly eat young seeds and bracts (Fig. 29). Mature larvae
fall to the ground, enter it, and make a fragile cocoon of soil particles held together by
saliva (similar to NV. mirabilis, Fig. 31). They overwinter in the prepupal stage.

The life cycle of N. mirabilis is quite similar to N. bimaculatus, and the host plant
is the same for both species. The two species coexist in Honshu, Japan, where UN.
mirabilis appears somewhat earlier than NV. bimaculatus. The body of the larva is light
brown in both species, but the head is light brown in N. bimaculatus (Fig. 29), whereas
it is black in N. mirabilis (Fig. 30) (Saito and Ibuki 2010).

Studies on Adelestini (Hymenoptera, Tenthredinidae)...

Figures 5-8. Head, lateral view. 5 Nipponorhynchus mirabilis 6 N. bimaculatus 7 N. brevis 8 Adelesta nova.

6 D.R. Smith & T. Naito / Journal of Hymenoptera Research 45: 1-14 (2015)

Nipponorhynchus brevis Smith & Naito, sp. n.
http://zoobank.org/937B73BC-65A3-4007-82D5-0942A3A95EC6
Figss5; 7s cltls, Lo.e17Z, 20225526

Description. Female: Length 4.3 mm. Antenna and head black; anterior half of cl-
ypeus light brown, labrum and upper surface of maxillolabial complex white; palpi and
undersurface of maxillolabial complex black. Thorax black with tegula and posterior
corners of pronotum white. Legs black with apices of coxae, trochanters, apical quarter
of femora, most of tibiae except about apical quarter with black stripe on inner surface;
tarsi mostly black with inner surface of first two tarsomeres paler to white. Abdomen
black; apical tergum and cercus white. Wings hyaline; veins and stigma black. Head
shiny but roughened with scattered fine punctures, denser on genae, postocellar area,
clypeus, and above toruli. Thorax and abdomen shiny; posterior margin of mesoscutel-
lum with several large punctures.

Antennal length 1.3x head width, with scape and pedicel each longer than broad;
3" antennomere slightly longer than 4", 4° to 9" antennomeres gradually decreasing
in length. Clypeus slightly broadly, circularly emarginated, about 3x broader than
long. Malar space about equal to diameter of median ocellus. Lower interocular dis-
tance 1.3x eye height; eyes only slightly converging below. Maxillolabial complex
about 0.6x head width and about 1.3x eye height; maxillary palpus subequal to length
of proboscis, longer than distance between maxillary palpus and labial palpus. Dis-
tances between eye and lateral ocellus, between lateral ocelli, and between lateral ocel-
lus and hind margin of head as 1.0:0.8:0.5. Postocellar area 1.5x broader than long.
Posttergite distinct, long, rounded behind. Tarsal claws simple. Hind basitarsomere
subequal to length of following 3 tarsomeres combined. Pulvilli minute on tarsomeres
1-4. Sheath in lateral view straight above, rounded at apex and below (Fig. 17); from
above, equally wide throughout. Lancet (Fig. 20) with about 10 spines on basal two
annuli and single row of about 6 spines on remaining annuli.

Male: Length 4.1 mm. Color and structure similar to female. Genitalia (Figs 25,
26) penis valve with valviceps elongate and rounded at apex, dorsal spine broad.

Type material. Holotype female “Japan: Hokkaido, Usubetsu, Sapporo, Ishi-
kari, 42.9348°N, 141.1206°E, 29.1V—24.V.2012, small stream, Mal. trap, N. Kuhara”
(NSMT). Paratypes: 4 2, 5 4, same data as holotype (NSMT, TN, USNM).

Distribution. Japan (Hokkaido).

Etymology. From the Latin brevis, meaning short, referring to the shorter mouth-
parts compared to the other species.

Comments. The new species is separated from both N. mirabilis and N. bimacu-
latus by its shorter mouthparts. In NV. mirabilis, the mouthparts are about 1.3x head
width and 3.0x eye height (Fig.1) and in N. bimaculatus, the mouthparts are about
1.0-1.1x head width and 2.2x eye height (Fig. 2). In Adelesta nova, the mouthparts
are about 0.3x head width and 0.4x eye height (Fig. 4). It is also separated from UN.
mirabilis by the long maxillary palpus, which is longer than the distance between the
maxillary palpus and labial palpus, and the longer more triangular posttergite, and

Studies on Adelestini (Hymenoptera, Tenthredinidae)... .

y ©
v/.

Figures 9-16. 9-12 Head, dorsal view. 13-16 Apex of mesonotum showing posttergite 9, 13 Nippono-
rhynchus mirabilis \0, 14 N. bimaculatus \\, 15 N. brevis 12, 16 Adelesta nova.

from NV. bimaculatus by the finely punctate head, which is very shiny in NV. bimacula-
tus. The female lancet and male genitalia appear close to N. bimaculatus (Figs. 19, 20,
23-26). In N. brevis, the lancet has the basal annuli more slanted and the annuli have
fewer spines, and the male penis valve is more slender and has a broader dorsal spine.
All females and males of NV. brevis studied are of the same length.

This is the first record of Nipponorhynchus from Hokkaido. The host plant is

unknown.

8 D.R. Smith & T. Naito / Journal of Hymenoptera Research 45: I-14 (2015)

Figures 17-20. 17 Nipponorhynchus brevis, lateral view 18-20 Female lancets 18 N. mirabilis 19 N.
bimaculatus 20 N. brevis.

Studies on Adelestini (Hymenoptera, Tenthredinidae)... 9

Figures 21-26. Male genitalia. 21, 23, 25 Genital capsule, ventral view 22, 24, 26 Penis valve, lateral
view 21, 22 N. mirabilis 23, 24 N. bimaculatus 25, 26 N. brevis.

Nipponorhynchus mirabilis Takeuchi
Biggrlsnd ye), Wo,altOea2ib, 2 203 0 1

Nipponorhynchus mirabilis Vakeuchi, 1941: 233; Takeuchi and Tokunaga 1941: figs
1-7; Malaise 1963: 169, 213, figs 138-140; Naito 1973: 96, fig. 2; Jervis and
Vilhelmsen 2000: 134, fig. 5 (mouthparts).

Diagnosis. Head roughened, with punctures. Maxillolabial process about 1.4x head
width and 3x eye height (Figs 1, 5). Malar space longer than diameter of median ocel-
lus. Posttergite short, narrow, of equal width throughout (Fig. 13). Lancet with single
row of annular spines, not more than 4 or 5 spines on basal annuli (Fig. 18). Male
genitalia (Figs 21, 22) with valviceps of penis valve round.

Specimens examined. JAPAN: Mt. Ooginosen, Hyogo Pref., 900 m, 22.V.2006,
T. Naito (3 9, 1); Hyogo Pref., Akazai-keikoku, 600 m, 21.IV.2008, T. Naito (2 3).

Distribution. Japan (Honshu).

10 D.R. Smith & T. Naito / Journal of Hymenoptera Research 45: 1-14 (2015)

Host and biological notes. The host and life history are similar to those of J.
bimaculatus. The larva (Fig. 31) also feeds on Chrysosplenium macrostemon Maxim vat.
shiobarense, but N. mirabilis occurs somewhat earlier in the season, and the head of the
larva is black (Fig. 30). See notes under NV. bimaculatus.

Adelestini

Ross (1937) first proposed the family-group name Adelestinae, including only Ade-
lesta described at the same time, with the single species A. nova (Norton, 1867). He
differentiated the subfamily by the absence of the epicnemium, the posttergite not
differentiated, propleurae with the mesal margins narrow and pointed, the distal anal
cell of the forewing almost as long as the proximal anal cell, and 11-m joining Rs near
separation from R in the hind wing.

Benson (1938) did not agree with Ross, indicating that the characters Ross used
did not justify a separate category because of fluctuating morphological characters in
the Selandriinae. However, his treatment as “?Adelestini” left the door open to accept-
ing it as a possible tribe.

Takeuchi (1941) described Nipponorhynchus and Takeuchi and Tokunaga (1941)
were the first to recognize that characters of Adelesta almost perfectly coincided with
those of Nipponorhynchus. They placed both genera in Adelestini, one of three tribes in
the Selandriinae. Their conclusion was based on Ross’ (1937) description of Adelesta
and not on comparison of actual specimens. The main character for the tribe was the
absence of an epicnemium on the anterior part of the mesopleuron.

Ross (1951) and Smith (1969, 1979) continued to recognize the tribe Adelestini,
mainly characterized by the subserrate antennae, forewing with an anal crossvein, long
distal anal cell and constriction of the basal anal cell of the forewing, absence of an
epicnemium, and indistinctly defined posttergite on the mesoscutellum. Both authors
based their concept only on Adelesta.

Malaise (1963) did not recognize tribes, but keyed Adelesta and Nipponorhynchus to
the same couplet, separated from other genera by the scutellar appendage (posttergite) not
differentiated or very narrow, front wings with short anal crossvein at about the middle
of the anal cell, labrum narrow and triangular, and antennae filiform or, slightly serrated.
He recognized Adelesta as having a very short perpendicular anal crossvein in the forewing
and antennae stout with the pedicel about as long as wide and Nipponorhynchus with an
oblique anal crossvein in the forewing and antenna slender with pedicel longer than wide.
In addition, though not mentioned for Adelesta, he gave the elongated mouthparts and
broadly emarginated propodeum as additional characters for Nipponohynchus.

Naito (1975) proposed a phylogeny of some related genera in Selandriinae and
concluded that Nipponothynchus and Adelesta belong to a distinct clade. However, he
did not assign tribal names.

Sinonerva, with the single species S. albipes, was described by Wei (1998). He
compared Sinonerva with Adelesta and Nipponorhynchus and mentioned that Sinonerva
differs from the two by the bidentate mandibles, second segment of the antenna longer

Studies on Adelestini (Hymenoptera, Tenthredinidae)... 11

“all

Figures 27-31. Nipponorhynchus spp. and the host plant, Chrysosplenium macrostemon vat. shiobarense.
27 N. bimaculatus, adult drinking honey of the host plant 28 NV. bimaculatus adult female ovipositing into
a seed capsule of the host plant. 29 NV. dimaculatus larva eating young seeds of host plant 30 NV. mirabilis,

larva eating young seeds of host plant 31 NV. mirabilis pupa in cocoon in soil.

than wide, developed epicnemium and posttergite, tarsal claws with a long inner tooth
and small basal lobe, and the hind wing with cell Rs remote from vein Sc+R and the
anal cell shortly petiolate. Later, Wei and Nie (1998) placed Sinonerva in the Ade-
lestini with Adelesta and Nipponorhynchus. We have not seen Sinonerva, but it differs
from Adelesta and Nipponorhynchus in several ways. The latter two genera have triden-
tate mandibles, antenna with flagellum gradually reduced towards apex, epicnemium
absent, tarsal claws simple, and cell Rs in hind wing elongate and very near or touching
vein Sc+R. Thus, we do not include Sinonerva in Adelestini.

12 D.R. Smith & T. Naito / Journal of Hymenoptera Research 45: 1-14 (2015)

We continue to recognize the Adelestini as a category in Selandriinae, separated
from other Selandriinae by the presence of a short almost perpendicular anal cell in
the forewing near the center of the anal cell, absence of an epicneium, an elongated
maxillolabial process (the long proboscis), tridentate mandibles, lateral halves of first
tergum widely separated on the meson thus exposing a broad membranous area, and
simple tarsal claws. ‘The indistinct posttergite, as mentioned by Ross (1937), Malaise
(1963), and Smith (1969), is present only in Adelesta; it is well developed in Nippono-
rhynchus. Though not mentioned by previous authors, Adelesta does have somewhat
elongated mouthparts, though not nearly as long as those in Nipponorhynchus. Though
we see a gradient from the short proboscis in Adelesta nova to the unusually elongated
proboscis in Nipponorhynchus mirabilis, and Adelesta and Nipponorhynchus share many
characters in common, they can still be distinguished from each other by the shape of
the posttergite, length of the mouthparts, antennae, placement of the anal crossvein
in the forewing, and the shape of clypeus, and we choose to continue to keep them
separate. [he two genera and species can be separated by the following key.

Key to genera and species of Adelestini

i Posttergite very narrow, of equal width throughout (Fig. 16); mouthparts
short, less than eye length and about 0.3x head width (Figs 4, 8); apical labial
palpomeres enlarged, palpi appear capitate; anal cell of forewing perpendicu-
lar, placed basally of the middle of the anal cell; antenna stout, antennomeres
4—9 at most 2x longer than broad; clypeus about 2x broader than long; east-
Seis NO Pole AITTeRICA a see. oeans cn PRN ee cee, Soeeer 2 ne reson Adelesta nova

= Posttergite distinct, long, longer at center than at sides (Figs 13-15); mouth-
parts exceptionally developed, half or more head width and longer than eye
height (Figs 1— 3, 5—7); labial palpomeres about equal in size; anal cell of
forewing slightly slanted, placed at center of anal cell; antenna slender, anten-
nomeres 4—9 more than 2x longer than broad; clypeus about 3x broader than

GH la pany NEP PON ONIN CIES a. wéionsemeroniemertpnsmees vineReseruebiosmaes opraidiainwet lena 2
92 Maxillolabial complex short, about 0.6x head width and about 1.3x eye
height (Figs 3, 7); maxillary palpus long, longer than distance between maxil-
Fary-palous-and: labial Palos a 0h hekaeassarcdeenemmaraeslbara ts taetene bap N. brevis
= Maxillolabial complex longer, 1.1 to 1.4x head width and 2x or more eye
height (Figs 1, 2, 5, 6); length of maxillary palpus various «0.0... cee eee 3
3 Posttergite long, triangular, longer at center than at sides (Fig. 14); head shiny;

maxillolabial process subequal to head width and about 2x eye height (Figs 2,
G6); malar space about equal to diameter of median ocellus.....N. bimaculatus
2 Posttergite short, narrow, of equal width throughout (Fig. 13); head roughened,
with punctures; maxillolabial complex about 1.4x head width and 3x eye height
(Figs 1, 5); malar space longer than diameter of median ocellus......N. mirabilis

Studies on Adelestini (Hymenoptera, Tenthredinidae)... 13

Acknowledgments

We thank T. Saito for allowing use of the photograph of the larva of N. bimaculatus
(Fig. 29) and E. Talamas, Systematic Entomology Laboratory, USDA, Washington,
DC, for help with some of the images. Mention of trade names or products in the pub-
lication is solely for the purpose of providing specific information and does not imply
recommendation or endorsement of the USDA. The USDA is an equal opportunity
provider and employer.

References

Benson RB (1938) On the classification of sawflies (Hymenoptera: Symphyta). Transactions of
the Royal Entomological Society of London 87: 353-384.

Jervis M, Vilhelmsen L (2000) Mouthpart evolution in adults of the basal, “symphytan”, hy-
menopteran lineages. Biological Journal of the Linnean Society 70: 121-146.

Malaise R (1963) Hymenoptera Tenthredinoidea, subfamily Selandriinae, key to genera of the
world. Entomologisk Tijdskrift 84: 159-216.

Naito T (1973) A new species of Nipponorhynchus Vakeuchi from Japan (Hymenoptera: Ten-
thredinidae. Kontyd 41: 95-96.

Naito T (1975) Phylogeny and distribution of five related genera, Eriocampidea, Hemitaxonus,
Pseutohemitaxonus, Nipponorhynchus and Adelesta, referred to the Selandriinae (Hymenop-
tera, Tenthredinidae). Kontyti 43: 330-342.

Norton E (1867) Catalog of the described Tenthredinidae and Uroceridae of North America.
Transactions of the American Entomological Society 1: 193-280.

Ross HH (1937) A generic classification of the Nearctic sawflies (Hymenoptera: Symphyta).
Illinois Biological Monographs 34, 173 pp.

Ross HH (1951) Tenthredinidae. In: Muesebeck CFW, Krombein KV, Townes HK (Eds)
Hymenoptera of America North of Mexico, Synoptic Catalog. United States Department
of Agriculture, Agriculture Monograph 2, 22-82.

Saito T, Ibuki S (2010) [Biological notes on the sawfly genus Nipponorhynchus at Bicchuzawa,
Nakagwa-cho, and Yamaguchi, Nikko-shi, in Tochigi Pref.]. Insekuto 61: 101-107. [In
Japanese]

Smith DR (1969) Nearctic sawflies II. Selandriinae: Adults (Hymenoptera: Tenthredinidae).
United States Department of Agriculture, Technical Bulletin No. 1398, 48 pp., 10 pls.

Smith DR (1979) Symphyta. In: Krombein KV, Hurd PD Jr., Smith DR, Burks BD (Eds)
Catalog of Hymenoptera in America north of Mexico, Volume 1, Symphyta and Apocrita
(Parasitica). Smithsonian Institution Press, Washington, D.C., 3-137.

Takeuchi T (1941) A systematic study on the suborder Symphyta of the Japanese Empire (IV).
Tenthredo 3: 230-274.

Takeuchi K, Tokunaga M (1941) Morphological study of head and mouthparts of a remarkable
saw-fly, Nipponorhynchus mirabilis Takeuchi. Tenthredo 3: 275-291.

14 D.R. Smith & T. Naito / Journal of Hymenoptera Research 45: 1-14 (2015)

Wei M (1998) A new genus and species of Selandriidae of China (Hymenoptera: Tenthredino-
morpha). Journal of Central South Forestry University 18: 1-3.

Wei M, Nie H (1998) Generic list of Tenthredinoidea s. str. (Hymenoptera) in new system-
atic arrangement with synonyms and distribution data. Journal of Central South Forestry

University 18: 23-31.