Bull. Natl. Mus. Nat. Sci., Ser. A, 41(3), pp. 155–162, August 21, 2015

Description of a New Land-hopper Genus, Kokuborchestia gen. nov.
and Redescription of K. kokuboi (Uéno, 1929) comb. nov.
(Crustacea, Amphipoda, Talitridae)
Hiroshi Morino1 and Hisashi Miyamoto2
1

Department of Zoology, National Museum of Nature and Science,
4–1–1 Amakubo, Tsukuba, Ibaraki 305–0005, Japan
E-mail: morino631@gmail. com
2
4–9 Kasumi, Maruoka-cho, Sakai, Fukui 910–0231, Japan
E-mail: dream_50000000690737@yahoo.co.jp(Received 20 May 2015; accepted 24 June 2015)

Abstract Kokuborchestia gen. nov. is described and the type species, K. kokuboi (Uéno, 1929)
comb. nov. is redescribed. This genus is characterized by elongate antenna 1 reaching mid-point ofpeduncular article 5 of antenna 2, 4- or 5- dentate lacinia on left mandible, maxilliped palp article
2 mediodistally lobate, palp article 4 reduced but distinct, gnathopod 1 (both sexes) deeply subchelate, pereopods cuspidactylate, pleopods with densely plumose setose peduncles and developedrami, uropod 1 outer ramus marginally robust setose. Several land-hopper genera from Japan andadjacent regions show similarity to this new genus in gnathopod 1 and maxilliped. However, elongate antenna 1, plumose setose peduncles and develop rami of pleopods, marginally robust-setoseouter ramus of uropod 1 are unique to the genus among them. The new genus also displays similarfeatures to Cochinorchestia Lowry and Peart, 2010 and “Parorchestia” gowerensis Bousﬁeld, 1976
from south oceans. Though, well-developed maxilliped palp 4 in Cochinorchestia and reduced
outer rami on pleopods in “P.” gowerensis are distinctive to each genus. This species occurs incoastal land habitats in restricted areas in the northwest Paciﬁc.
Key words : Crustacea, land-hoppers, Kokuborchestia kokuboi, new genus, new combination,
Talitridae.

Introduction
Orchestia kokuboi was described by Uéno
(1929) based on specimens secured on a hill of asmall island (Yu-no-shima) off Asamushi, northern tip of Honshu, Japan. After that, Iwasa(1939) described females of a talitrid species
under the name of O. kokuboi on the basis ofmaterial collected from Mt. Haruka, near Sapporo, Hokkaido. In the revision on the northPaciﬁc talitrids, Bousﬁeld (1982) proposed torestrict the concept of Orchestia to shore-dwelling and immediate coastal-terrestrial species of
the Atlantic-Mediterranean region, though he did
not indicate the generic allocation of Orchestia
kokuboi. Quite recently Lowry and Fanini (2013)

redeﬁned Orchestia further and conﬁned it tomarine supralittoral species, where they criticallyreviewed the world “Orchestia” species andnoted the possibility of “Orchestia” kokuboi
belonging to Paciforchestia Bousﬁeld, 1982.
Morino and Miyamoto (2015b) amended theconcept of Paciforchestia and alluded to the features of “Orchestia” kobukoi in comparison withPaciforchestia. Thus so far the generic positionof “Orchestia” kokuboi has not been formallyestablished. This is partly because the originaldescription was not adequate in modern sense
and also description (and ﬁgures) of female specimens given by Iwasa (1939) are not consistent
with the original description in essential characters. Unfortunately the type specimens of “O.”156

Hiroshi Morino and Hisashi Miyamoto

kokuboi are not available (most probably lost), sothat in this paper, the specimens from a nearest
place to the type locality are redescribed, and a
new genus is erected to accept this species since
it displays features distinctive from other genera.
Methods
The general methodology follows Morino(2014). The specimens were dissected under astereomicroscope, and appendages and bodies
were depicted under a light microscope using adrawing tube. The illustrated appendages were
ﬁxed on slide mounts with Hoyerʼs medium or
kept in tubes with bodies. The body length measured from the tip of head to the tip of telson
along straightened dorsal margin. In the description of species, generic characters are basicallynot repeated. The specimens studied are lodged
in the collection of the National Museum of
Nature and Science, Tsukuba (NSMT), or in theprivate collection of Miyamoto.
Taxonomy
Family Talitridae
Kokuborchestia gen. nov.
[New Japanese name:
Kokubo-okatobimushi zoku]
Type species. Orchestia kokuboi Uéno, 1929
Diagnosis. Body size medium large, eyes
medium large. Antenna 1 elongate, exceeding
mid-point of peduncular article 5 of antenna 2;
peduncle subequal to ﬂagellum in length, peduncular article 3 longer than articles 1 and 2 each.
Antenna 2 in male not incrassate, ﬂagellum longer than peduncle. Mandible, left lacinia 4- or
5-dentate. Maxilliped, precoxa outer margin not
stepped, palp articles 2 and 3 broad, mediodistally lobate, article 4 reduced but distinct, apically positioned on article 3.
Gnathopod 1 in both sexes deeply subchelate,merus-propodus with pellucid lobe (weaker infemale), carpus and propodus with broad basedone; propodus lateral surface with a row of elongate robust setae. Gnathopod 2 in male, propodus
powerfully subchelate; in female, basis slightly
expanded anteriorly, propodus mitten-shaped.
Pereopods 3–7 cuspidactylate (bi-cuspate), dactyli locking robust setae lacking or weakly developed. Coxa of pereopod 4 as deep as wide. Coxaof pereopod 6, posterior lobe smoothly curved
anteroventrally. Pereopods 6 and 7 in male notincrassate. Coxal gills of pereopods 2 and 6
large, gill of pereopod 6 broad and truncated distally. Oostegites with simple-tipped setae middistally.
Pleopods, peduncles outer margin densely plumose-setose, with 2 retinacula; rami developed.
Uropod 1, peduncle distolateral robust seta
shorter than subdistal one; inner ramus with dorsomarginal and outer marginal robust setae; outerramus with marginal robust setae. Uropod 3,
peduncle dorsally expanded; ramus attenuate.
Telson lobe with lateral, laterodistal and distalrobust setae group (1–3 setae per group).
Remarks. Kokuborchestia gen. nov. is characterized by densely plumose-setose peduncles
on pleopods and distally truncated coxal gill on
pereopod 6. However, in other several charactersthe new genus shows similarity to several terrestrial or supralittoral genera worldwide. Deeply
subchelate gnathopod 1 (both sexes) with pellucid lobes on carpus-propodus (weak in female),maxilliped palp article 2 broad with distinct
mediodistal lobe, article 4 reduced but distinct,
and cuspidactylate pereopods in the new genus
are shared with the following genera in the North
Paciﬁc: Paciforchestia Bousﬁeld, 1982, from theNorth-east Paciﬁc, Pyatakovestia Morino andMiyamoto, 2015b, from the North-west Paciﬁc,Nipponorchesita Morino and Miyamoto, 2015a,from Japan, Bousﬁeldia Chou and Lee 1996,
from Taiwan and adjacent islands, andLanorchestia Miyamoto and Morino, 2010, from
Lanyu Island, Taiwan. Paciforchestia sense ofMorino and Miyamoto, 2015b is separable fromKokuborchstia by antenna 1 not reaching midpoint of peduncular article 5 of antenna 2, moderately reduced pleopod rami, 3 ＋ retinacula onKokuborchestia gen. nov.

pleopod peduncle, peduncle of uropod 1 withelongate distolateral robust seta, etc.; Pyatakovestia is by stepped outer margin on maxillipedprecoxa, strongly reduced pleopod rami, elongate
telson, etc.; Nipponorchestia is by antenna 1 notreaching mid-point of peduncular article 5 ofantenna 2, weak pellucid lobe on female gnathopod 1, moderately reduced pleopod rami, anduropod 1 with elongate distolateral robust seta on
peduncle and marginally bare outer ramus, etc.
The new genus is closer to Bousﬁeldia in having
developed pleopod rami and also to Lanorchestia
in having elongate antenna 1 (reaching end of
peduncular article 5 of antenna 2). However,both genera are separable from the new genus byuropod 1 with elongate distolateral robust seta
and marginally bare outer ramus. In addition,
Bousﬁeldia is distinguished from Kokuborchestia
in having antenna 1 not reaching mid-point ofpeduncular article 5 of antenna 2, female gnathopod 1 with weak or lacking pellucid lobe; Lanorchesita is in having moderately reduced pleopodrami.
Kokuborchestia also shows similarity to a few
taxa from the southern oceans. Cochinorchestia
Lowry and Peart 2010, from southern India andMoçanbique, is allied to the genus in having
elongate antenna 1, maxilliped palp article 2 withmediodistal lobe, and deeply subchelate (to chelate) gnathopod 1 (merus-propodus with pellucidlobe), pleopods well developed, uropod 1 outerramus marginally robust setose. However, welldeveloped palp article 4 of maxilliped, telsonwith apical setae (lacking lateral robust setae) of
Cochinorchestia differentiate this genus fromKokuborchestia.
“Parorchestia” gowerensis Bousﬁeld, 1976,
from Lord Howe Island, off Australia, displays
several similar characters to the new genus in
antenna 1 (exceeding beyond end of peduncular
article 4 of antenna 2), maxilliped palp articles 2
mediodistally lobate and article 4 reduced but
distinct, merus-propodus of male gnathopod 1with pellucid lobe, pleopods peduncle plumose
setose (though not dense), outer ramus of uropod
1 with marginal robust setae. However, strongly157

slender and elongate antenna 2 and pereopods,
female gnathopod 1 with weak pellucid lobe, distally attenuate coxal gill of pereopod 6, fewer
plumose setae on pleopod peduncles, reduced
outer ramus of pleopods, and elongate distolateral robust seta on uropod 1 are features distinctive to this species. The cusps of pereopod dactyli are not known for “P.” gowarensis. The
generic allocation of this species is better toawait the study of types.
Etymology. The generic name is a combination of the speciﬁc name of the type species andOrchestia.
Kokuborchestia kokuboi (Uéno, 1929) comb. nov.
[Japanese name: Kokubo-okatobimushi](Figs. 1–3)

Orchestia kokuboi Uéno, 1929: 7, text-ﬁg. 1; Iwasa, 1939:
266, text-ﬁg. 9, pl. XII; Bulycheva, 1957: 163, ﬁg. 59.
“Orchestia” kokuboi: Morino, 2015: 1087, 1076 (ﬁg. 2).

Material examined. HOKKAIDO: 2 males, 1
ovig. female, 2 females (NSMT-Cr 24097);
Kumoishi, Kumaishi-cho, Hiyama; 31 Jul. 1984;
Otaka, A. collect. 9 males, 3 ovig. females, 13
females (NSMT-Cr 24099); Tomarigawa, Mt.
Obira, Shimamaki-gun; 8 Aug. 1986; Udagawa,T. collect. Male 13.0 mm (NSMT-Cr 24092),
female 14.4 mm (NSMT-Cr 24093), 3 males, 2
females, 1 juvenile (NSMT-Cr 24094); Fukushima, Matsumae-gun; 29 Sept. 1979; Kato, H.
collect. 1 male, 1 female, 3 juveniles (NSMT-Cr
24095); Shiriuchi River, Matsumae-gun; 21 Sept.
1980; Ando, A. collect. Male 11.0 mm (NSMT-Cr
24101), 2 males, 2 females, 1 unsexed (NSMTCr 24102); Mt. Hakodate, Hakodate; 26 May
1994; Tsubokura, T. collect.
AOMORI: 1 male, 1 ovig. female, 2 females(NSMT-Cr 24089); Wakinozawa (Cryptomeria
forest near sea-shore), Shimokita-gun; 4 Sept.
1974; Morino, H. collect. 2 males, 6 females, 1juvenile (NSMT-Cr 24098); Wakinozawa,
Shimokita-gun; 13 Dec. 1986; Yanagihashi collect. 3 males, 1 ovig. female, 1 female (NSMT-158

Hiroshi Morino and Hisashi Miyamoto

Cr 24091); Shimofuro, Shimokita-gun; 12 Aug.
1979; Kato, H. collect. 1 male, 4 females(NSMT-Cr 24090); Mimmaya, Tsugaru-gun; 26
Sept. 1979; Kato, H. collect. 5 males, 2 females(NSMT-Cr 24100); Juniko Lakes (forest near
Ketoba pond), Nishitsugaru-gun; 16 Nov. 1991;
Otaka, A. collect. 5 males, 2 females (NSMT-Cr
24106); Juniko Lakes (near Ketoba pond);
Nishitsugaru-gun; 24 Apr. 1999; Otaka, A. collect. Male 13.9 mm (NSMT-Cr 24103), male
16.0 mm (NSMT-Cr 24123), female 15.9 mm(NSMT-Cr 24104), female 12.5 mm (NSMT-Cr
24122), 5 males, 9 females, 2 juveniles (NSMTCr 24105); hill behind Asamushi Marine Station(Tohoku University), Asamushi; 7 Oct. 1977;
Morino, H. collect. Male 20 mm, female 16 mm(Miyamoto collection); Asamushi Marine Station(Tohoku University); 8 Jul. 1978; Miyamoto H.
collect. 23 males, 32 females (Miyamoto collec-tion); Yu-no-shima Island (forest ﬂoor), Asamushi; 8 Jul. 1978; Miyamoto, H. collect. 1
female (NSMT-Cr 24096); Asamushi; 16 Jun.
1983; Nunomura, N. collect. 3 males, 5 females(Miyamoto collection); Azigasawa (forest on
slope); Nishitsugaru-gun; 8 Aug. 1978; Miyamoto, H. collect. 3 males, 3 females (Miyamotocollection); Obata (Cryptomeria forest, upstreamof Nakamura River), Azigasawa, Nishitsugarugun; 8 Aug. 1978; Miyamoto, H. collect.
Description of male (NSMT-Cr 24103,
13.9 mm). Eyes medium large, subround.
Antenna 1 (Fig. 1B), peduncular article 3 longest, ﬂagellum as long as peduncle, with 7 articles. Antenna 2 (Fig. 1A), peduncular article 5
subequal to articles 3 and 4 combined, ﬂagellum
longer than peduncle, with 17 articles. Upper lip(Fig. 1C) lacking robust setae. Mandible (Fig.
1E), left lacinia 4-dentate. Maxilliped (Fig.

Fig. 1. Kokuborchestia kokuboi (Uéno, 1929) comb. nov. Male, 13.9 mm (NSMT-Cr 24103). Asamushi, Japan.
—A, habitus, lateral view (after Morino, 2015); B, antenna 1; C, upper lip; D, lower lip; E, left mandible; F,
distal part of right mandible; G, maxilliped; H, palp article 4 of maxilliped; I, maxilla 1; J, maxilla 2. Scale: A,
2.50 mm; B, 1.43 mm; C–G, I–J, 0.42 mm; H, 0.14 mm.

Kokuborchestia gen. nov.

1G–H), palp article 4 reduced, distinct, positioned apically. Other mouth parts (Fig. 1D, F,
I–J), same as those of other talitrid species.
Gnathopod 1 (Figs. 1A, 2A), carpus ca. 1.2
times as long as propodus, propodus lateral surface with row of 6 submarginal elongate setae,
palmar margin vertical, exceeding dactylus. Gnathopod 2 (Figs. 1A, 2B), propodus palmar margin smooth, slightly shorter than posterior margin, dactylus attenuate apically. Pereopod 4
shorter than pereopod 3 (Fig. 1A), propodi ofboth pereopods lacking locking robust setae (Fig.
2F–G), dactylus of pereopod 4 pinched. Pereopods 5–7, propodi locking robust setae minute(Fig. 2H–J). Coxal gill of pereopod 2 (Fig. 2K),
lobed at middle, those of pereopods 3–5 (Fig.
2 L–N) smaller, convoluted, coxal gill of pereo-159

pod 6 (Fig. 2O) as large as that of pereopod 2,
distal half broad and truncated distally.
Pleonite side plates (Fig. 2Q), posterodistal
corner angulate, with several setules on posterior
margin, lacking marginal pits. Pleopods 1–3 (Fig.
2R–T), peduncles with facial ﬁne robust setae,
and with dense plumose-setae on whole outermargin, bearing 2 retinacula; rami developed,(0.92, 0.88, 0.89 times as long as respective
peduncle, with about 13 articles.
Uropod 1 (Fig. 3D), inner ramus with 3 dorsomarginal, 3 outer marginal robust setae, outerramus with 3 marginal robust setae. Uropod 2(Fig. 3E), both rami subequal in length, with
marginal and apical robust setae. Uropod 3 (Fig.
3F), peduncle with 3 robust setae, rami shorter
than peduncle (ca. 0.6 times), with marginal andFig. 2. Kokuborchestia kokuboi (Uéno, 1929) comb. nov. A–B, F–T, male, 13.9 mm (NSMT-Cr 24103); C–E,
female, 15.9 mm (NSMT-Cr 24104). Asamushi, Japan.—A, distal articles of gnathopod 1; B, distal articles ofgnathopod 2; C, gnathopod 1; D, distal articles of gnathopod 1; E, gnathopod 2; F–J, distal articles of pereopods 3–7; K–O, coxal gills of pereopods 2–7; P, coxal plate of pereopod 6; Q, pleonite side plates; R–T, pleopods 1–3 with enlarged retinacula. Scale: A, D, 0.94 mm; B–C, E, K–O, 1.61 mm; F–J, 0.48 mm; P–Q,
1.43 mm; R–T, 0.71 mm.

160

Hiroshi Morino and Hisashi MiyamotoFig. 3. Kokuborchestia kokuboi (Uéno, 1929) comb.
nov. A–B, female, 15.9 mm (NSMT-Cr 24104);
C, female, 12.5 mm (NSMT-Cr 24122); D–G,
male, 13.9 mm (NSMT-Cr 24103). Asamushi,
Japan. A–B, oostegites of pereopods 2 and 5; C,oostegite of pereopod 2; D–F, uropods 1–3; G,
telson. Scale: A–E, 1.43 mm; F–G, 0.56 mm.

apical robust setae. Telson (Fig. 3G) with dorsal
longitudinal suture, bearing 5–6 robust setae per
lobe.
Female (sexual characters NSMT-Cr 24104,
15.9 mm; NSMT-Cr 24122, 12.5 mm). Gnathopod 1 (Fig. 2C–D), pellucid lobes on merus-propodus distinct but weaker than of those of male,
carpus ca. 1.7 times as long as propodus, withrow of 5 submarginal elongate setae on lateralsurface, palmar margin vertical, slightly exceeding dactylus. Gnathopod 2 (Fig. 2E) basis anteroproximally slightly expanded, merus and carpus
with dome-shaped pellucid lobe, propodus lateralsurface with rows of facial and submarginal

setae. Oostegite of pereopod 2 (Fig. 3A, C) with
25, that of pereopod 5 (Fig. 3B) with 14 simpletipped setae.
Distribution. Kokuborchestia kokuboi has
been collected from coastal forests (see also
Tsubokura et al. 1998), occasionally from inland
forests, in southwestern Hokkaido, and northern
tip of Honshu (Fig. 4). Most sites face theTsugaru Strait. Gongalsky et al. (2014) recorded
this species from broad-leaved forests inKunashir Island, Kuril Islands.
Remarks. The present material displays no
distinct inconsistencies with the original description. Females from Mt. Haruka, near Sapporo,
Hokkaido, of which Iwasa (1939) gave illustration, exhibit same features as types and the present material, except 5-dentate left lacinia on
mandible and marginally bare peduncles on pleopods. Close examination of additional material athand reveals that a specimen from Mt. Hakodate,
Hokkaido (NSMT-Cr 24101) bears 5-dentate
lacinia. Also in some specimens the plumose
setae on the pleopod peduncles tightly adhered tothe peduncles and were hard to recognize. Thus,
it is concluded that Kokuborchestia kokuboi has
4-, occasionally 5-, dentate left lacinia. And it ishighly probable that Iwasa (1939) has failed torecognize the plumose setae on pleopods.
In Japan, this species occurs on a restricted
area on the coasts of Honshu and Hokkaido, facing the Tsugaru Strait. The strait is known as
Blakiston Line, a zoogeographical barrier line forterrestrial animals between Honshu andHokkaido. Geologically the strait is estimated tohave been bridged during the glacial periods, 10
to several 10 thousands years before. Twohypotheses are possible to explain the pattern of
this distribution: due to dispersal by rafting
between Hokkaido and Honshu, or reﬂecting theancient pattern since the time of land bridge.
Recently Gongalsky et al. (2014) reported thisspecies as a component of soil fauna characteristic to broad-leaved forests in Kunashir Island,
Kuril, Russia. It is possible that future studies
expand the distributional range in these areas. In
any case, K. kokuboi exhibits spotted-pattern ofKokuborchestia gen. nov.

161

Fig. 4. Distribution of Kokuborchestia kokuboi (Uéno, 1929) comb. nov. in the Japan Archipelago. Star marks
are from Iwasa (1939) and Gongalsky et al. (2014).

geographical distribution and peculiar morphological features, which might suggest that theendemicity of this species is of paleoendemic.
Acknowledgments
We sincerely thank Mr. Kato, H., Dr. Otaka, A.
and colleagues who kindly forwarded their sam-ples for this study. Dr. Komatsu, H. of the
National Museum of Nature and Science has
supported us in preparing the manuscript, to who
we are grateful. We also thank Dr. Lowry, J. K.
for careful reading and valuable comment on an
earlier draft.

162

Hiroshi Morino and Hisashi Miyamoto

References
Bousﬁeld, E. L. 1976. A new terrestrial amphipod fromLord Howe Island. Records of the Australian Museum,30: 118–122.
Bousﬁeld, E. L. 1982. The amphipod superfamily Talitroidea in the northeastern Paciﬁc region. 1. Family
Talitridae: systematics and distributional ecology. Publications in Biological Oceanography, 11: i–vii, 1–72.
Bulycheva, A. I. 1957. Morskie bloxi morej SSSR isopredelʼnyx vod (Amphipoda - Talitroidea). Opredeliteli po Faune SSSR, Akademiya Nauk SSSR, 65:
1–185. (In Russian.)Chou, W.-H. and J.-D. Lee 1996. A new terrestrial amphipod (Crustacea) from a subtropical forest in Taiwan,
with description of a new genus. Bulletin of the
National Museum of Natural Science, 8: 43–55.
Gongalsky, K. B., D. M. Kuzunetsova, A. D. Elagin, S. A.
Malyavin and A. S. Zaitsev. 2014. Soil macrofauna of
the south of Kunashir Island (Kuril Islands, Russia).
Doklady Biological Sciences, 457: 240–243.
Iwasa, M. 1939. Japanese Talitridae. Journal of the Faculty of Science, Hokkaido Imperial University, Series
6, Zoology, 6: 255–296, pls. 9–18.
Lowry, J. K. and L. Fanini 2013. Substrate dependent talitrid amphipods from fragmented beaches on the northcoast of Crete (Crustacea, Amphipoda, Talitridae),including a redeﬁnition of the genus Orchestia anddescriptions of Orchestia xylino sp. nov. and Cryptorchestia gen. nov. Zootaxa, 3709: 201–229.
Lowry, J. K. and R. Peart 2010. The genus Microrchestia(Amphipoda: Talitridae) in eastern Australia. Zootaxa

2349: 21–38.
Miyamoto, H. and H. Morino 2010. Taxonomic studies on
the Talitridae (Crustacea: Amphipoda) of Taiwan. IV.
The genera Paciforchestia and Lanorchestia gen. nov.
Species Diversity, 15: 155–167.
Morino, H. 2014. A new land-hopper genus, Mizuhorchesita, from Japan (Crustacea, Amphipoda, Talitridae). Bulletin of the National Museum of Nature andScience, Series A (Zoology), 40: 117–127.
Morino, H. 2015. Amphipoda. In Aoki, J. (ed.): Pictorial
Keys to Soil Animals of Japan, 2nd ed, pp. 1069–1089.
Tokai University Press, Hadano. (In Japanese.)Morino, H. and H. Miyamoto 2015a. A new land-hopper
genus, Nipponorchestia, with two new species fromJapan (Crustacea, Amphipoda, Talitridae). Bulletin of
the National Museum of Nature and Science, Series A(Zoology), 41: 1–13.
Morino, H. and H. Miyamoto 2015b. Redeﬁnition ofPaciforchestia Bousﬁeld, 1982 and description of
Pyatakovestia gen. nov. (Crustacea, Amphipoda,
Talitridae). Bulletin of the National Museum of Natureand Science, Series A (Zoology), 41: 105–121.
Tsubokura, T., S. Kikuchi and S. Nakano 1998. Distribution pattern, gill area relationships, and ﬁne structuralcharacteristics of the gill epithelia in supralittoral andterrestrial talitrid amphipods. Journal of Crustacean
Biology, 18: 625–642.
Uéno, M. 1929. A new terrestrial amphipod Orchestia
kokuboi, sp. nov. from Asamushi. Science Reports of
the Tôhoku Imperial University, Fourth series, Biology,
4: 7–9.