European Journal of Taxonomy 176: 1-14

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2016 • Yu D. et al.

Research article

um:lsid:zoobank.org:pub:C94C7774-5176-460C-804D-A38520284D4B

Tomoceridae (Collembola, Entomobryomorpha) from the southern

Annamitic cordillera: redescription of Tomocerus ocreatus Denis, 1948

and description of a new species of Tomocerina Yosii, 1955

Daoyuan YU '•*, Le Cong MAN 2 & Louis DEHARVENG 3

1 Soil Ecology Lab, College of Resources and Environmental Sciences,

Nanjing Agricultural University, Nanjing 210095, China,

department of Biology, University of Natural Sciences,

Ho Chi Minh City National University, Vietnam.

3 Institut de Systematique, Evolution, Biodiversite, ISYEB - UMR 7205 - CNRS, MNHN, UPMC,

EPHE, Museum national d’Histoire naturelle, Sorbonne Universites,

57 rue Cuvier, CP 50, 75005 Paris, France.

* Corresponding author, dvvu [email protected] : [email protected]

2 Email: tnmletamljnh@va hoo. com. vn

3 Email: [email protected]

1 um:lsid:zoobank.org: author: 07321441-2070-4DB2-BC59-D3F1C8300BC3

2 urn:lsid:zoobank.org:author:0D7A5067-F6EF-456F-8B32-DlB61DCC4CDD

3 urn:lsid:zoobank.org:author:D8F5C679-C30C-442C-8621-D3B8EDB17EF7

Abstract. Two species of Tomoceridae were found near Dalat, southern Vietnam. Tomocerus ocreatus

Denis, 1948 is redescribed based on a neotype specimen. Previous records of Tomocerus ocreatus in

non-type localities are reevaluated. A new species Tomocerina annamitica sp. nov. is described. The

new species is mainly characterized by its small body size, pointed tenent hair, compound dental spines

and the absence of intermediate teeth on mucro.

Key words. Vietnam, Southeast Asia, Tomocerinae, taxonomy, neotype.

Yu D., Man L.C. & Deharveng L. 2016. Tomoceridae (Collembola, Entomobryomorpha) from the southern

Annamitic cordillera: redescription of Tomocerus ocreatus Denis, 1948 and description of a new species of

Tomocerina Yosii, 1955. European Journal of Taxonomy 176: 1-14. http://dx.doi.org/10.5852/ejt.2016.176

Introduction

The family Tomoceridae contains at least 169 known species (Bellinger et al. 1996-2015) distributed

worldwide. In Asia Tomoceridae are abundant in the north but have only a few reports in the south. Three

species were previously described from Vietnam: Tomocerus ocreatus Denis, 1948 from Dalat, Lam

Dong Province; Tomocerus nodentalis Nguyen, 1995 from Moc Chau, Son La Province; and Tomocerus

postantennalis Yu, Zhang & Deharveng, 2014 from Na Hang, Tuyen Quang Province. Tomocerus

ocreatus is characterized by the shape and arrangement of dental spines. Tomocerus nodentalis is very

European Journal of Taxonomy 176 : 1-14 ( 2016 )

peculiar for the absence of dental spines, which character is only observed in very early instars of other

species (Goto 1956; Uchida & Chiba 1958). Tomocernspostantennalis is a cave species with developed

postantennal organs in adult stage. To the present knowledge, Tomocerus nodentalis and Tomocerus

postantennalis are both endemic species, whereas Tomocerus ocreatus is known to be the most widely

distributed Asian species of Tomocerus Nicolet, 1842. Besides its type locality, Tomocerus ocreatus has

records from Himalaya (Yosii 1966), Japan (Yosii 1956, 1967, 1969, 1977; Chiba 1968), Korea (Yosii &

Lee 1963; Lee 1974, 1975), China (Stach 1964; Liu et al. 1998), northern Vietnam (Stach 1965), eastern

Russia (Sakhalin island) (Martynova 1977), etc. But intraspecific differences were often reported by

descriptions of non-type Tomocerus ocreatus. For instance, Chiba (1968) distinguished three varieties

from Japanese Tomocerus ocreatus according to the variation in the body size and relative length of

body parts. A molecular study (Zhang et al. 2014) also revealed “cryptic” diversity among Chinese

specimens previously identified as Tomocerus ocreatus.

To clarify the true status and distribution of Tomocerus ocreatus , a redescription of its type specimen

is required, but the holotype and only specimen of the species has been lost in the Museum national

d’Histoire naturelle (Paris) where the Denis’ collection is currently stored. In a recent sample collected

from Dalat (Lam Dong Province) in 2008, we have found a specimen that matches well the original

description in morphology. We designate this specimen as neotype of Tomocerus ocreatus.

A new tomocerid species was found in the same sample as Tomocerus ocreatus. We assign the new

species to Tomocerina Yosii, 1955 because it has no toothlet on the outer basal mucronal tooth. To our

knowledge, Tomocerina annamitica sp. nov. is so far the southernmost record of the genus.

Materials and methods

Specimens were collected with aspirators or Berlese funnels. Photographs were taken under a Jenoptilc

ProgRes C10+ camera mounted on a Leica MZ 16 stereomicroscope. Specimens were cleared in lactic

acid and mounted in Hoyer’s solution (Krantz 1978). For some specimens the head, the furca and the

legs were cut off from the trunk and mounted separately for detailed observation. The slide-mounted

specimens were studied using a Leica DMLB microscope or a Nikon 80i microscope.

We follow Fjellberg (2007) for maxillary lamellae numbering, Yu et al. (2014) for the pattern of cephalic

dorsal chaetotaxy and Christiansen (1964) for body macrochaetotaxy. The description of the body

chaetotaxy refers to one side only since in most case it is symmetric. The exact morphology of each

chaeta was uncertain due to shedding. The dental spines formula follows that of Folsom (1913), in

which the dental spines are arranged from basal to distal, with a slash indicating the separation between

basal and medial subsegments and the Roman numerals referring to spines that are noticeably larger. If

not mentioned specially, all descriptions are based on fully developed individuals.

Abbreviations

Ant. = antennal segment

PAO = postantennal organ

Th. = thoracic segment

Abd. = abdominal segment

Institutional acronyms:

HCMCU = Ho Chi Minh City National University, Vietnam

MNHN = Museum national d’Histoire naturelle, Paris, France

NJAU = Nanjing Agricultural University, Nanjing, China

YU D. et al. , Tomoceridae from southern Annamitic cordillera

Results

Class Collembola Lubbock, 1873

Order Entomobryomorpha Borner, 1913

Superfamily Tomoceroidea Szeptycki, 1979

Family Tomoceridae Borner, 1913

Subfamily Tomocerinae Schaffer, 1896

Genus Tomocerus Nicolet, 1842

Diagnosis

Moderate to large sized Tomocerinae, usually longer than 3 mm; body colour pale to dark, some species

with distinct colour pattern; eyes at most 6+6; trochantero-femoral organ reduced to 1, 1 chaetae; dens

of furca basally without outer strong chaetae or inner large differentiated scales; shape of dental spines

from simple to compound among different species; mucro with two dorsal lamellae and two basal

teeth, outer basal tooth with corner toothlet. Commonest group of Tomocerinae in Eurasia Continent,

especially abundant in East Asia.

Tomocerus ocreatus Denis, 1948

Figs 1A, 2, 3

Diagnosis

Typical Tomocerus species with pale body colour, moderately long antennae and full set of 6+6 eyes.

Head without distinct PAO; Th. II with relatively reduced number of macrochaetae; tenent hair clavate,

moderately developed; unguis with 5-6 teeth; tenaculum unsealed, with numerous chaetae; manubrium

without dorsal scales and blunt prominent chaetae; dental spines compound with numerous moderate

sized denticles; no small dental spine between two distal large spines; mucro with 9-10 intermediate

teeth.

Neotype

VIETNAM: §, on slide. Collected in Hon Giao, Bi Doup massif, northeast of Dalat, Lam Dong Province,

108°42’53”E, 12°irH ,, N, alt. 1630 m, 12Jun. 2008, by Louis Deharveng & Anne Bedos (sample code

Vn08-150). Deposited in MNHN (specimen Vn08-150_Tol).

Description

Body length 3.6 mm Ground colour uniformly yellowish white. Ant. III+IV, antero-ventral part of head,

coxae and tibiotarsi with dark pigment. Eye patches black. Scales brown (Fig. 1A).

Body densely clothed by scales and various types of chaetae. Scales of typical morphology of

Tomocerinae, with continuous longitudinal ridges on surface (Lubbock 1873). Ordinary chaetae of

different sizes. Microchaetae smooth and pointed. Macrochaetae and mesochaetae from slightly to

strongly ciliated, some slightly ciliated mesochaetae appearing to be smooth under optical microscope.

Most macrochaetae straight, rod-like and subcylindrical, some macrochaetae on posterior abdominal

segments long, curved and acuminate. Mesochaetae acuminate, shorter and thinner than macrochaetae.

S-chaetae subcylindrical, more hyaline than ordinary chaetae, as small as microchaetae except long ones

on Abd. IV. Dorso-inner chaetae on dens modified as strong pointed spines. Pseudopores as small circular

structures similar to chaetae sockets, distributed at least on Th. II to Abd. IV, coxae, and manubrium.

PAO not seen. Eyes 6+6. Ante nn ae nearly as long as body. Ante nn a length ratio as I:II:III+IV=

1.0:1.7:15.4. Ant. I and Ant. II dorsally scaled. Ant. III+IV unsealed. Prelabral and labral chaetae (labral

European Journal of Taxonomy 176 : 1-14 ( 2016 )

formula) 4/5, 5,4, the distal 4 chaetae stronger. Distal edge of labrum with four curved spine-like papillae

(Fig. 2A), and a well developed ventro-distal brush. Mandibular head asymmetrical, the left one with

4 teeth (including a basal rounded one) and the right one with 5, left molar plate distally with a tapered

tooth (Fig. 2B). Basal teeth of maxillary lamella 5 slightly longer than apical ones, without beard-like

appendage (Fig. 2C). Maxillary outer lobe with trifurcate palp, one basal chaeta and 4 sublobal hairs

(Fig. 2D). Both dorsal and ventral sides of head scaled. Cephalic dorsal macrochaetotaxy: anterior area:

2, 2; interocular area: 2, 6, central uneven macrochaeta absent; postocular area: 2+2; posterior area: 2.

Posterior margin of head with about 20+20 small chaetae (Fig. 2E). Mentum with 5 chaetae, submentum

with numerous chaetae.

Pattern of body chaetotaxy as in Fig. 2F. Bothriotricha 2, 1/0, 0, 1,2, 0, 0 on Th. II-Abd. VI, respectively,

as typical in Tomocerinae. Macrochaetae densely arranged along anterior margin of Th. II (not shown

in figure). Th. II with a row of macrochaetae behind anterior margin. Number of macrochaetae or large

mesochaetae in the posterior row as 3, 3/3, 3, 4, 2, 4 (3 dorsal+1 lateral) from Th. II to Abd. V. On Th.

II no macrochaeta near the pseudopore contrary to most other tomocerids; on Abd. Ill s-microchaeta

and accompanying microchaeta posterior to lateral macrochaeta; Abd. IV with one lateral macrochaeta

and numerous long s-chaetae; on Abd. V inner macrochaeta smaller than others in posterior row; Abd.

VI with numerous chaetae of moderate size. Most mesochaetae laterally and posteriorly on terga.

Pseudopores near the axis of terga, 1, 1/1, 1, 1, 1, 0, 0 from Th. II to Abd. VI.

Tegs with numerous ordinary chaetae. Trochantero-femoral organ with 1, 1 slender chaetae. Front,

middle and hind tibiotarsus dorsally with 1,1,3 long prominent chaetae, ventrally with 4-5, 5, 6 blunt

spine-lilce chaetae (Fig. 2G). Each tibiotarsus with a distal whorl of 11 chaetae, ventral 6 as ordinary

chaetae, dorsal 5 modified: tenent hair clavate, as long as inner edge of unguis; 2 accessory chaetae

extremely minute; 2 guard chaetae thin and long, slightly shorter than tenent hair (Fig. 3A, B). Unguis

Fig. 1 . Appearance of Tomocerus ocreatus Denis, 1948 and Tomocerina annamitica sp. nov. in alcohol.

A. Tomocerus ocreatus (lateral view). B. Tomocerina annamitica sp. nov. (lateral view). Scale bar: 1 mm.

YU D. et al ., Tomoceridae from southern Annamitic cordillera

slender, with baso-internal ridging visible in lateral view; lateral teeth pointed, of moderate size. Inner

edge of unguis with 5-6 teeth, the basal tooth smaller, the other subequal in size. Unguiculus about

half as long as unguis, its inner edge with 1 tooth. Pretarsal chaetae 1+1, much larger than tenent-hair

accessory chaetae (Fig. 3B).

Fig. 2. Tomocerus ocreatus Denis, 1948. A. Labrum (dorsal view). B. Mandibular head (dorsal view).

C. Lamella 5 of maxillary head (dorsal view). D. Maxillary outer lobe (ventral view). E. Cephalic

dorsal chaetotaxy (circles = sockets of chaetae, same as below; 1 = anterior area; 2 = interocular area;

3 = postocular area; 4 = posterior area). F. Dorsal chaetotaxy of Th. II-Abd. VI (circle with a slash =

pseudopore, same as below). G. Right tibiotarsi (lateral view). Scale bars: A, B, G = 100 pm; C = 10 pm;

D = 50 pm; E, F = 500 pm.

European Journal of Taxonomy 176 : 1-14 ( 2016 )

Ventral tube with scales on both anterior and posterior faces, lateral flap unsealed, anterior face with

about 15 chaetae on each side, posterior face with about 55 chaetae, each lateral flap with 45M7 chaetae.

Rami of tenaculum with 4+4 teeth, anterior face with 13 small chaetae and without scale (Fig. 3C).

Furcaratio manubrium: dens: mucro=2.9-3.1:4.1M.4:1.0. Manubrium ventrally scaled, without chaetae,

laterally with large round scales and 11 chaetae, proximal 2 chaetae small and slender, distal 9 distinctly

stronger and serrated; dorsal scales absent; each dorsal chaetal stripe with about 90 chaetae in different

sizes, without blunt chaetae; pseudopores 11-12 on each side (Fig. 3D); external corner chaeta as large

as small mesochaetae in chaetal stripe (Fig. 3E). Dens basally without inner modified scale or outer

Fig. 3. Tomocerus ocreatus Denis, 1948. A. Diagram of tibiotarsal distal chaetae (distal view; t = tenent

hair; a = accessory chaeta; g = guard chaeta; p = pretarsal chaeta, same as below). B. Hind claw (lateral

view). C. Tenaculum (anterior view). D. Left side of manubrium (dorsal view). E. Disto-extemal corner

of manubrium (dorsal view). F. Basal and middle subsegments of dens with dental spines (dorsal view).

G. Mucro (dorso-inner view). Scale bars: B, C, E, G = 50 pm; D, F = 100 pm.

YU D. et al., Tomoceridae from southern Annamitic cordillera

strong chaetae. Dental spines formula as 3/4, II, distal spine strongest, about 0.1 times as long as dens;

all spines with numerous denticles of moderate size (Fig. 3F). Dens dorsally with ordinary chaetae and

feather-like chaetae as typical in Tomocerinae, ventrally with only scales. Mucro elongated, bearing

numerous smooth chaetae with elongated sockets; both basal teeth with proximal lamellae, the outer

tooth with a toothlet; apical and subapical tooth subequal; structure of dorsal lamellae of Tomocerus

type, two dorsal lamellae running from subapical tooth, outer lamella ending in inner basal tooth, inner

lamella ending freely at base of mucro; outer lamella with 9-10 subequal intermediate teeth (Fig. 3G).

Remarks

Denis (1948) provided accurate description for Tomocerus ocreatus. For instance, he described clearly

the dental spines as “covered with secondary spines, more similar to scales than denticles”, and the

figure showed that those scale-like denticles were of moderate size and covered at least basal half of

each spine, on which account we rejected some records of non-type Tomocerus ocreatus (see below).

But naturally, some later revealed characters were not mentioned by him. For instance, chaetotaxy had

not been used for taxonomy in Tomocerinae before Yosii (1956). The limit of original description is a

main reason for the misidentification of Tomocerus ocreatus in some subsequent records, and finally

turned the species into a complex including a number of closely related forms from Vietnam to eastern

Russia. To overcome the deep confusion about this species, a redescription of type specimen is necessary.

Because the holotype and unique type specimen of Tomocerus ocreatus was lost, we propose to set up a

neotype for the species from specimens of the type locality.

Denis (1948) described Tomocerus ocreatus on one specimen, from “plateau du Lang Biang, 2400 m.

alt., foret tropicale”. It is not the Lang Bian peak itself that reaches 2400 m but the Chu Yang Sin

massif north of the Lang Bian (= Dalat) plateau, the Lang Bian peak being only 2197 m in altitude.

The Chu Yang Sin massif was and still is of very difficult access, and was certainly not the place where

the collector, Dawydoff, operated. There are two main patches of subtropical forests around Dalat: a

small one on the Lang Bian peak, and a much larger one 25 km northeast on the Bi Doup massif, which

reaches 2287 m and is less easy to access. In any case, it is most likely on the slopes of one of these

massifs that Tomocerus ocreatus was collected. On this account, we propose to designate a specimen

that we obtained from litter sample in the Bi Doup mountain as the neotype.

The neotype is highly identical with the original description in almost all described characters, including

the body colour, the length of antennae, the structure of claws and the morphology and arrangement of

dental spines. The only difference is that in the neotype specimen the numbers of ungual teeth, dental

spines and mucronal teeth are slightly larger than in the original one. It could happen in Tomoceridae

that within the same species larger individuals have a few more teeth and dental spines than smaller

ones, so the slight difference mentioned above can be treated as intraspecific considering the neotype

specimen is slightly larger than the lost holotype (3.6 mm versus 3.3 mm).

Regarding previous non-type records of Tomocerus ocreatus , Stach (1964, 1965) redescribed two different

forms of Tomocerus ocreatus from southeastern China and northern Vietnam, respectively, and also claimed

that the Japanese record of “Tomocerus minor ” by Uchida (1953) was actually Tomocerus ocreatus ; Yosii

described a Japanese species Tomocerus kawamurai (Yosii, 1954) and then synonymized it with Tomocerus

ocreatus (Yosii 1956, 1967); Chiba (1968) recognized three forms in Japanese Tomocerus ocreatus ; Lee

(1975) made descriptive notes on Korean specimens; Martynova (1977) recorded Tomocerus ocreatus in

Sakhalin and made some descriptive notes on it. These redescriptions had subtle to considerable differences

to each other, and were never identical to the original description. The Chinese “ ocreatus ” recorded in

Hangzhou (“Hangchow”), Zhejiang Province by Stach (1964) has brownish body colour and distinctly

short antenna about half the length of body; the northern Vietnamese record from Lao Cai Province by

the same author (Stach 1965) bears a small dental spine between two large distal spines, and the denticles

European Journal of Taxonomy 176 : 1-14 ( 2016 )

on spines are finer than those in the original description, therefore it is more similar to Tomocerus folsomi

Denis, 1929 from Yunnan Province, China; the Japanese “ ocreatus ” described by Yosii (1967) has dorsal

scales and 2+2 blunt principal chaetae on manubrium which are absent in the true Tomocerus ocreatus ; other

records (Chiba 1968; Lee 1975; Martynova 1977) all have dental spines with one or two whorls of crown¬

like denticles near the base which is not the morphology of the true ocreatus form. But these remarkable

differences were treated as intraspecific variations when other characters showed high similarity. Inferred

from the present morphological review and the molecular study on Chinese ocreatus complex (Zhang et al.

2014), the aforementioned non-type “ ocreatus ” may each represent an independent species in the complex,

and so far the true Tomocerus ocreatus is known as only endemic to southern Annamitic range in Vietnam.

Genus Tomocerina Yosii, 1955

Diagnosis

Small to moderate sized Tomocerinae, usually less than 2 mm in length; body colour pale to light grey,

seldom dark; antennae usually much shorter than body; eyes at most 6+6; trochantero-femoral organ

with 1, 1 or more chaetae; dens of furca basally without outer strong chaetae or inner large differentiated

scales; shape of dental spines from simple to compound; mucro with two dorsal lamellae and two basal

teeth, outer basal tooth without comer toothlet. Widely distributed in Northern Hemisphere.

Tomocerina annamitica sp. nov.

um:lsid:zoobank.org:act:9AD947EC-5A05-4385-9A92-8B0DC590A94D

Figs IB, 4, 5

Diagnosis

Tomocerina species with typical body size and pigmentation; antennae about 0.8 times as long as body,

relatively long for Tomocerina ; Th. II with only one bothriotrichum; tibiotarsi with multiple strong

chaetae; tenet hair small and pointed; unguis with 1-2 inner teeth; dental spines compound with denticles;

only one distal spine distinctly larger; mucro without intermediate tooth.

Etymology

Specific name derived from its type locality: the southern Annamitic range.

Type material

Holotype

VIETNAM: §, on slide. Collected in Hon Giao, Bi Doup massif, northeast of Dalat, Lam Dong Province,

108 o 42’53”E, 12°irH”N, alt. 1630 m, mixed forest litter, 12 Jun. 2008, by Louis Deharveng & Anne

Bedos (sample code Vn08-150), deposited in MNHN (specimen Vn08-150_Tal).

Paratypes

VIETNAM: 2 §§, on slides, same data as holotype, 1 in HCMCU (specimen Vn08-150_Ta2) and 1 in

NJAU (specimen Vn08-15 0_Ta3).

Other material examined

VIETNAM: 5 specimens on slides and 28 specimens in alcohol, collected in the Hon Ba Nature Reserve,

Kanh Hoa Province, located 25 km east-southeast of Hon Giao and 25 km south-southwest of Nha

Trang, 108°56 , 55 ,, -108°58 , 51 ,, E, 12°06 , 35 ,, -12°07 , 09 ,, N, mixed forest litter, Nov. 2013, by Louis

Deharveng & Anne Bedos: alt. 850 m, Vnl3-072 (3 in alcohol); 890 m, Vnl3-126 (2 in alcohol), Vnl3-

135 (4 in alcohol); 1050 m, Vnl3-054 (2 §§, on slides), Vnl3-057 (1 S and 1 $, on sildes), Vnl3-114

(1 on slide, sex undetermined), Vnl3-120 (8 in alcohol); 1350 m, Vnl3-039 (5 in alcohol); 1500 m,

Vnl3-231 (6 in alcohol).

YU D. et al ., Tomoceridae from southern Annamitic cordillera

Description

Body length 1.6-1.8 mm. Body colour light grey, antennae light purple, eye patches black, scales brown

(Fig. IB). Clothing of Tomocerinae type, similar to that of Tomocerus ocreatus.

PAO not seen. Eyes 6+6. Antenna about 0.75-0.90 times as long as body. Antenna length ratio LILIII:

IV=1.0:1.3-1.5:5.5-6.0:2.2-2.5, Ant. I and Ant. II dorsally scaled. Ant. Ill and Ant. IV unsealed. Labral

formula 4/5, 5,4, the distal 4 chaetae stronger. Distal edge of labrum with four curved spine-like papillae,

ventro-distal brush well developed. Mandibular head asymmetrical, the left one with 4 teeth (including a

Fig. 4. Tomocerina annamitica sp. nov. A. Apical half of mandibular head (dorsal view). B. Lamella 5

of maxillary head (dorsal view). C. Maxillary outer lobe (ventral view). D. Cephalic dorsal chaetotaxy.

E. Dorsal chaetotaxy of Th. II-Abd. VI. F. Left tibiotarsi (lateral view, b = blunt inner chaeta). Scale bars:

A, C = 50 pm; B = 10 pm; D, E = 250 pm; F = 100 pm.

European Journal of Taxonomy 176 : 1-14 ( 2016 )

basal rounded one) and the right one with 5, left molar plate distally with tapered tooth (Fig. 4A). Basal

teeth of maxillary lamella 5 elongated without beard-like appendage (Fig. 4B). Maxillary outer lobe

with trifurcate pulp, basal chaeta and 4 sublobal hairs (Fig. 4C). Cephalic dorsal chaetotaxy: anterior

area: 2, 4; interocular area: 2, 6, central chaeta absent; postocular area: 2+2; posterior area: 2; posterior

margin: about 20+20 small chaetae of different sizes (Fig. 4D). Both dorsal and ventral side of head

scaled. Mentum with 5 chaetae, submentum with numerous chaetae.

Pattern of body chaetotaxy as Fig. 4E. 1, 1/0, 0, 1, 2, 0, 0 bothriotricha on Th. II-Abd. VI. Macrochaetae

densely along anterior margin of Th. II (not shown in figure). Th. II with a row of macrochaetae behind

anterior margin. Number of macro- or large mesochaetae in the posterior row as 3, 3, 3, 3, 3, 5, 4 (3

dorsal+1 lateral) from Th. II to Abd. V. On Th. II pseudopore near posterior macrochaeta of the 3 central

ones; Th. Ill with anterior macrochaeta; on Abd. Ill s-microchaeta posterior to lateral macrochaetae;

Abd. IV with antero-lateral macrochaeta and numerous long s-chaetae, 4 inner mesochaetae in posterior

Fig. 5. Tomocerina annamitica sp. nov. A. Diagram oftibiotarsal distal chaetae (distal view). B. Tibiotarsal

distal chaetae (dorsal view). C. Hind claw (lateral view). D. Tenaculum (anterior view). E. Teft side of

manubrium (dorsal view). F. Disto-external corner of manubrium (dorsal view). G. Basal and middle

subsegments of dens with dental spines (dorsal view, 1 = ventral side of spine, showing only distal

serrations; 2 = dorsal side of spine, showing fine longitudinal ribs; s = small inner scale). H. Mucro (inner

view). Scale bars: B-D = 20 pm; E-H = 50 pm.

YU D. et al., Tomoceridae from southern Annamitic cordillera

Table 1 . Discrimination between Tomocerina annamitica sp.

Hou & Li, 1999.

nov. and Tomocerina purpurithora Liu,

Species

Number of tibiotarsal

strong inner chaetae

Number of chaetae

on tenaculum

Dental spines

formula

Number of intermediate

teeth on mucro

T. annamitica sp. nov.

5-7, 4, 5-6

4/3,1

T. purpurithora

0, 0,1

5/7, 1

5-7

row prominent but smaller than outer macrochaeta; on Abd. VI dorsal flap with 2+2 and lateral flap with

3 macrochaetae. Most mesochaetae laterally and posteriorly on terga. Pseudopores near the axis of terga,

1, 1/1, 1, 1, 1, 0, 0 from Th. II-Abd. VI.

Legs with numerous ordinary chaetae. Trochantero-femoral organ with 1, 1 chaetae. Tibiotarsi dorsally with

1, 1, 3 prominent chaetae from front to hind leg, ventrally with 5-7, 4, 5-6 strong chaetae, only 2, 1, 2 of

them blunt (Fig. 4F). Each tibiotarsus with a distal whorl of 11 chaetae, the dorsal 5 modified: tenent hair

pointed, short and thin; 2 accessory chaetae about as long as but thicker than tenent hair, and longer than

pretarsal chaetae; 2 outer guard chaetae strong, as long as inner edge of unguis (Fig. 5A-C). Scales absent

from front and mid tibiotarsi but present on hind tibiotarsus. Unguis slender, lateral teeth of moderate size,

internal ridging present. Inner edge of unguis with a small basal and 0-1 larger more distal teeth. Unguiculus

about half as long as unguis, without tooth on inner or outer edge. Pretarsus with 1+1 chaetae (Fig. 5C).

Ventral tube anteriorly with a few small scales, each side with 20-25 chaetae; posteriorly unsealed, with

about 40 chaetae; lateral flap unsealed, each side with about 25 chaetae. Tenaculum with 4+4 teeth,

unsealed, anterior face with 1 chaeta about as long as rami (Fig. 5D). Ratio manubrium: dens: mucro=

2.0-2.2:2.9-3.2:1.0. Manubrium ventrally with only scales, laterally with scales and 8 chaetae, proximal

2 chaetae small and slender, distal 6 chaetae distinctly stronger; dorsal scales absent; each dorsal chaetal

stripe with about 70 chaetae in different sizes, without blunt chaetae; pseudopores 5-6 on each side

(Fig. 5E); external corner chaeta as large as small chaetae in chaetal stripe (Fig. 5F). Dental spines formula

4/3,1, a small pointed scale present ventrally to basal spines; all spines with several large denticles near

base, superficial texture different between dorsal and ventral sides: dorsally with only fine longitudinal

ribs, ventrally with longitudinal ribs and distal tiny serrations (Fig. 5G). The largest spine about 0.1 times

as long as dens. Dens dorsally with feather-like chaetae between ordinary chaetae. Stripe of feather-like

chaetae starting from centre of middle subsegment of dens, ending a short distance to the apex of dens.

Ventral side of dens covered by scales, several chaetae present apically. Mucro elongated and multi-

setaceous, with two dorsal lamellae; both basal teeth with proximal lamellae, outer tooth without toothlet;

apical tooth elongated, stronger than subapical tooth; intermediate teeth absent (Fig. 5H).

Remarks

Among Tomocerina , the new species is similar to those of the minuta group in its small body size, grey

body colour and shape of mucro, but is different from them mainly in the chaetotaxy and the shape of

dental spines. It is similar to Tomocerina purpurithora Liu, Hou & Li, 1999 from Sichuan Province,

China in the shape of dental spines, but is different from the latter mainly in the cephalic chaetotaxy,

the number of tibiotarsal strong inner chaetae, the number of chaetae on tenaculum, the dental spines

formula and the number of teeth on mucro (Table 1). Tomocerina annamitica sp. nov. is the southernmost

distributed species of Tomocerina , a genus mostly diversified in northern temperate to cold temperate

regions. However, so far Tomocerina is poorly defined by only a minute character: the absence of corner

toothlet on outer basal tooth of mucro. In this respect, the presence of only 1+1 bothriotricha on Th. II

of Tomocerina annamitica sp. nov. might provide an interesting character, while most other species of

Tomocerinae have 2+2. Investigation on this character in different groups of Tomocerinae is in progress

to assess its phylogenetic value.

European Journal of Taxonomy 176 : 1-14 ( 2016 )

Discussion

Tomoceridae are mostly distributed in the northern temperate areas, with a diversity peak in eastern

Palearctic: Japan (Yosii 1967), Korea (Lee 1975), Russian Far-East (Martynova 1977) and China

(Ma 2004). The diversity rapidly decreases towards southern regions, but this family is still present

in the caves or at high altitude of the subtropics (southern China and Vietnam, Denis 1929; Yu et al.

2014; Yu & Deharveng 2015; Nepal, Yosii 1966), or even at high altitude in tropical regions (Sumatra,

Oudemans 1891; Vietnam, Denis 1948). Before this study, all three species of Tomoceridae known from

Vietnam belong to Tomocerus. The fourth species added here, Tomocerina annamitica sp. nov., belongs

to another genus which had so far a single species distributed south of 30°N ( Tomocerina simplex Yosii,

1966 from Nepal). Unexpectedly, the new species was frequent in litter and soil as low as 1000 m of

elevation. Besides Tomocerus ocreatus and Tomocerina annamitica sp. nov., at least one other species is

probably present in our collections from southern Annamitic Cordillera, confirming its status of southern

extension for Palaearctic fauna of springtails (Deharveng & Bedos 2000).

Acknowledgements

We thank Dr. Ann e Bedos from MNHN Paris who helped to collect specimens in Tang Bian, Bi Doup

and Hon Ba, Truong Quang Tam and Nguyen Tran Vy from Institute of Tropical Biology of Ho Chi Minh

City who organized the field trip and the staff of the Bi Doup-Nui Ba National Park. Field work in Hon

Ba was carried out in the frame of a project with the Nature Reserve, and supported by a French grant

of the Agence Nationale de la Recherche under the TabEx ANR-10-TABX-0003-BCDiv. Laboratory

work in NJAU was supported by the National Natural Science Foundation of China (41501056), the

Fundamental Research Funds for the Central Universities (KJQN201668) and the Scientific Grant for

Post-doctors of Jiangsu Province (1402054C).

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Manuscript received: 17 October 2015

Manuscript accepted: 6 November 2015

Published on: 19 February 2016

Topic editor: Koen Martens

Desk editor: Charlotte Thionois

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, Denmark.