European Journal of Taxonomy 226: 1-21

http://dx.doi.org/10.5852/ejt.2016.226

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This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2016 • Zhang F. et al.

Research article

urn:lsid:zoobank.org:pub:D85D7297-C5C3-4F0F-8B6E-7FA334723D07

Cave-dwelling Coecobrya from southern China with a survey of

clypeal chaetae in Entomobryoidea (Collembola)

Feng ZHANG '•*, Anne BEDOS 2 & Louis DEHARVENG 3

'Department of Entomology, College of Plant Protection, Nanjing Agricultural University,

1 Weigang, Nanjing 210095, P. R. China.

2 3 Institut de Systematique, Evolution, Biodiversite (ISYEB) UMR 7205 CNRS, MNHN, UPMC,

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

45 rue Buffon, CP50, F75005 Paris, France.

* Corresponding author: xtmtd.zf@gmail. com

23 Email: [email protected]

1 um:lsid:zoobank.org:author:612B65B3-C4B2-42BD-8A46-4DF70DB078BF

2 um:lsid:zoobank.org:author:CFD095B7-llC3-4A8A-9AA9-3682C0D75586

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

Abstract. Four new species of the genus Coecobrya , C. gejianbangi sp. nov., C. annulata sp. nov.,

C. ciliata sp. nov., and C. oculata sp. nov., are described from Guangxi caves as the representative of the

genus in China. Coecobrya oculata sp. nov. of the boneti- group has 1+1 eyes and a serrate outer edge of

the unguiculus. The other three species, devoid of eyes and with a tiny outer tooth on the unguiculus, are

assigned here to the tenebricosa- group, assuming that the large tooth on the unguiculus is transformed

into a tiny one in cave-obligate species. Clypeal chaetae in Entomobryoidea are systematically surveyed

for the first time, and are found to be well diversified at species level. They have a potential taxonomical

value in discriminating taxa of morphologically conserved groups.

Keywords. Entomobryidae, taxonomy, new species, Guangxi, clypeal chaetotaxy.

Zhang K, Bedos A. & Deharveng L. 2016. Cave-dwelling Coecobrya from southern China with a survey of clypeal

chaetae in Entomobryoidea (Collembola). European Journal of Taxonomy 226: 1-21. http://dx.doi.org/10.5852/

eit.2016.226

Introduction

The taxonomical history and diagnosis of the genus Coecobrya Yosii, 1956 have been recently reviewed

by Zhang et al. (2009). All species of the genus have polymacrochaetotic chaetotaxy, no labral papillae,

inverted intrusion on labral margin U-shaped, chaetae mel of labium always smooth, reduced eye

number (0 to 3 on each side), pigment reduced or absent, antennal apical bulb absent, falcate mucro

with a basal spine, and scales and dental spines absent. Later, Zhang et al. (2011a) divided the genus into

tenebricosa- and boneti-gxouys, blind and with a large outer tooth on unguiculus in the former and eyed

and with the outer edge of unguiculus smooth or serrate in the latter.

1

European Journal of Taxonomy 226: 1-21 (2016)

So far, 45 species of the genus have been reported in the world (Bellinger et al. 1996-2016). More

than Vs of them were recorded from cave or both cave and non-cave habitat, with Coecobrya draconis

Zhang & Dong, 2014 among species described from a Chinese cave. During biospeological expeditions

to Guangxi Province, South China, conducted during the last decade, many springtails, mainly

Entomobryidae, were discovered in caves (Deharveng et al. 2008). Among them were four Coecobrya

species new to science, which are described here in detail. The group assignation of these new species

is discussed. The taxonomical value of clypeal chaetae in Entomobryoidea is systematically presented

for the first time.

Material and methods

Specimens were mounted, after clearing in lactic acid, under a coverslip in Marc Andre II solution,

and were studied using a Leica DMLB microscope. Photographs were enhanced with Photoshop CS2/

PC (Adobe Inc.). The dorsal and ventral chaetotaxy of head and the Ant. Ill organ are described after

Chen & Christiansen (1993). Dorsal body chaetae are designated following Szeptycki (1979), Zhang

et al. (2011b) and Zhang & Deharveng (2015). The number of macrochaetae is given by half-tergite

in the descriptions. Material is deposited in the collections of the Department of Entomology, Nanjing

Agricultural University (NJAU), PR. China and Museum national d'Histoire naturelle (MNHN), Paris,

France.

Abbreviations

Ant. I-IV

Th. I-III

Abd. I-VI

mac

mic

ms

sens

Gr.

antennal segment I-IV

thoracic segment I-III

abdominal segment I-VI

macrochaeta, -ae

microchaeta, -ae

S-microchaeta, -ae (microsensillum, -a)

ordinary S-chaeta, -ae on terga group

group

Results

The following characters are shared by the four species included in this paper and are not repeated in the

descriptions: smooth spine-like mic at base of antennae (3 dorsal, 3 (2 in C. oculata ) ventral on Ant. I, 1

internal, 1 external and 1 ventral on Ant. II); Ant. IV apical bulb absent; labral papillae absent, inverted

intrusion on labral margin U-shaped; prelabral and labral chaetae 4/5, 5, 4, all smooth; five chaetae of

the first row of labrum subequal (Fig. 8A); subapical chaeta of maxillary outer lobe slightly larger than

the apical one; 3 smooth sublobal hairs on maxillary outer lobe; tip of lateral process of labial palp

reaching beyond apex of labial papilla; labial chaetae mel^ always smooth; inner outstanding tibiotarsal

mac 1-2, located at about 0.33 distance from base, ciliate and tapered; all tenent hairs pointed, subequal

to unguiculus in length; tenaculum with 4 + 4 teeth and one large striate chaeta; manubrium without

smooth or modified chaetae; mucro falcate with long basal spine, nearly reaching the tip of the apical

tooth.

2

ZHANG F. et al, Cave Coecobrya from China and clypeal chaetae

Class Collembola Lubbock, 1873

Order Entomonryomorpha Borner, 1913

Family Entomobryidae Tomdsvary, 1882

Genus Coecobrya Yosii, 1956

Coecobrya gejianbangi sp. nov.

urn:lsid:zoobank.org:act:FB0EB4BF-5950-4CFF-BC3A-A01347DB95C3

Figs 1A, 2, 3, 9B; Table 1

Diagnosis

Size large; Ant. IV annulated; 11 ciliate clypeal chaetae; labial chaetae and M l present; unguis

elongate and devoid of unpaired tooth; unguiculus truncate with a tiny tooth on outer edge; 9+9 mac

on Abd. I with al as mac; Abd. II with 4+4 central mac; Abd. IV with 4+4 central and 9+9 lateral mac.

Etymology

Named after Mr. Jianbang Ge, who provided help for our trip in the Mulun Natural Reserve.

Material examined

Holotype

CHINA: $ on slide, Guangxi, Huanjiang, Mulun Natural Reserve, Dong Qiao Dong, 25.181667° N,

108.008611° E, altitude 440 m, 5 Nov. 2009, Tian MY leg. (# CHIgx09-077). Deposited in NJAU.

Paratypes

CHINA: 2 $ ? on slides and 5 $ $ in alcohol, same data as holotype. One paratype deposited in NJAU

and others in MNHN.

Description

Body length up to 4.2 mm. Body colour white (Fig. 1A).

Antenna 2.5-2.7 times as long as cephalic diagonal. Antennal segments ratio as I : II : III : IV = 1 :

1.4-1.9 : 1.1-1.9: 3.6—4.3. Long smooth straight chaetae absent on antennae. Ant. II distally with 3

dorsal rod-like S-chaetae. Two internal S-chaetae of Ant. Ill organ slightly expanded (Fig. 2A). Ant. IV

distinctly annulated under light microscope but annulations not easily seen under stereoscope.

Eyes absent. Clypeal chaetae 11, all ciliate (Fig. 9B). Dorsal cephalic chaetotaxy with 5 sutural (S) and 4

mac in Gr. II (Fig. 2B). Lateral process of labial palp as thick as normal chaetae (Fig. 2C). Labial chaetae

as nynyrel^; r and X x as spiny mic; chaetae X, X 0 and X 4 as ciliate mic; chaeta X 3 absent (Fig. 2D).

Cephalic groove with 8 chaetae, anterior five smooth and others ciliate.

Trochanteral organ with 17-20 smooth spine-like chaetae; 8-9 in arms and 8-12 between them (Fig. 2E).

Partial inner differentiated tibiotarsal chaetae ciliate with ciliations not closely appressed to axis. Unguis

narrow with 2 subequal tiny inner teeth near base; unpaired tooth absent. Unguiculus truncate with a tiny

tooth on outer edge (Fig. 2F-G). Abd. IV 4.0-5.0 times as long as Abd. Ill along dorsal midline. Ventral

tube anteriorly with about 7 ciliate chaetae on each side (Fig. 2H); posteriorly with 16-18 chaetae, three

of them smooth (Fig. 21); each lateral flap with 7-8 smooth chaetae (Fig. 2J). Manubrial plaque with 2

pseudopores and 4 ciliate chaetae (Fig. 2K). Distal smooth part of dens 0.7-0.8 times as long as mucro.

Th. II with 3 medio-medial (ml, m2, m2i), 3(2) medio-lateral (m4, m4i, m4p) and 25-26 posterior

mac; p4-5 and p4i as mac; mac m4i rarely absent. Th. Ill with about 35 mac; mac m5i and a6i present;

p3p rarely present (Fig. 3A). Abd. I with 9 (al-3, m2^t, m2i, m4i, m4p) mac. Abd. II with 4 (a2, a3,

3

European Journal of Taxonomy 226 : 1-21 ( 2016 )

Fig. 1 . Habitus. A. Coecobrya gejianbangi sp. nov. B. Coecobrya annulata sp. nov. C. Coecobrya

ciliata sp. nov. D. Coecobrya oculata sp. nov. Scale bars: 500 pm.

4

ZHANG F. et al., Cave Coecobrya from China and clypeal chaetae

Fig. 2. Coecobrya gejianbangi sp. nov. A. Ant. Ill organ. B. Dorsal cephalic chaetotaxy. C. Lateral

process of labial palp. D. Chaetae on the ventral side of head. E. Trochanteral organ, ventral view.

F. Fore claw. G. Hind claw. H-J. Ventral tube. H. Anterior face. I. Posterior face. J. Lateral flap.

K. Manubrial plaque. Scale bars: A, C, E-G, K = 40 pm; B, D, H-J =100 pm.

5

European Journal of Taxonomy 226 : 1-21 ( 2016 )

m3, m3e) central and 1 (m5) lateral mac. Abd. Ill with 1 (m3) central and 4 (a7, am6, pm6, p6) lateral

mac; ms absent (Fig. 3B). Abd. IV with 4 central (I, M, B5, A6), 9 lateral mac (Fl-3, F3a, E2^1, E2p,

D3) and several additional large mesochaetae or small mac laterally (Fig. 3C). Abd. V with 3 sens (Fig.

3D). S-chaetae formula: 2+ms, 2/1+ms, 2, 2+ms, 2, 3. Uncertain number of longer S-like chaetae on

Abd. IV.

Ecology

Cave-restricted species.

Fig. 3. Coecobrya gejianbangi sp. nov. A. Thoracic chaetotaxy. B-D. Abdominal chaetotaxy. B. Abd.

I—III. C. Abd. IV. D. Abd. V. Scale bars: 200 pm.

6

ZHANG F. et al., Cave Coecobrya from China and clypeal chaetae

Table 1. Comparison of the five cave Coecobrya species from China (+: present;

c: ciliate; =: equal; j: unequal; approximately).

absent;

sm: smooth;

Characters

gejianbangi sp. nov.

annulata sp. nov.

ciliata sp. nov. oculata sp. nov. draconis

Maximum body length in mm

4.23

3.03

1.35

1.38

0.93

Antenna/head diagonal

2.50-2.72

2.14-2.77

1.4-1.8

1.7

1.44-1.80

Long smooth straight chaetae on antennae

-

+

-

Annulated Ant. IV

yes

weakly

no

Eyes

blind

1+1

blind

Clypeal chaetae

11c

9sm

15c

?c

7c

Dorsal cephalic chaetotaxy

sutural

5

4

5

Gr. II

4

4(5)

4

5(4)

4

Lateral process thicker than normal chaetae

no

yes

no

Chaetae on the ventral side of head

m ,

sm

-

r

sm, tiny

c

x,

sm, tiny

-

X 2

c

sm, tiny

c

X 3

-

c

-

X 4

c

sm, tiny

c

X

c

sm, tiny

c

H,

sm

c

sm

c

sm

c

groove

5sm, 3c

4sm, 4c

Ism, 6c

2sm, 4c

Ism, 6c

Spiny chaetae on trochanteral organ

17-20

15-23

9-16

9-13

10-13

Inner differentiated tibiotarsal chaetae

ciliate

smooth

ciliate

Unguis

paired teeth and distance from base

=,0-2

+ 0.4

+ 0.25

+ 0.4

+ 0.25

unpaired teeth

-

+

Unguiculus

shape

truncate

lanceolate

truncate or swollen

lanceolate

truncate

outer teeth

1, tiny

serrate

smooth

Ventral tube

anterior face on one side

~7c

7c

5-6c

4c

5c

posterior face

3sm, 13-15c

ll-14sm

2sm, 7-8c

6sm

2sm, 6c

lateral flap on one side

7-8sm

8-1Ism

5sm, l-2c

5sm, 0-1 c

5-6sm, 0-3 c

Ciliate chaetae on manubrial plaque

4

4(5)

4-5

3(2)

4

Mac ml + on Th. II

1

1(2)

2

1

Mac p4+ on Th. II

2

3

1(2)

2

Mac m6i on Th. Ill

+

-

+

-

Chaetotaxy of Abd. I

9+9

6+6

Mac a3 on Abd. II

+

-

Lateral mac on Abd. Ill

4+4

2+2

3+3

Ms on Abd. Ill

-

+

-

Chaetotaxy of Abd. IV

central mac

4+4

6+6

3+3

4+4

lateral mac

9+9

8+8

5+5

8+8

7+7

Remarks

Coecobrya gejianbangi sp. nov. is characterized by a very large size, distinctly annulated Ant. IV,

elongate unguis devoid of unpaired tooth, 11 ciliate clypeal chaetae, labial chaetae Xj and present,

mac al on Abd. I, mac a2 and a3 on Abd. II, mac a7 on lateral Abd. Ill, and 9+9 lateral mac on Abd. IV. It

is closest to another cave species, C. ciliata sp. nov., but differs from it in labial chaetae, and abdominal

chaetotaxy (Table 1).

7

European Journal of Taxonomy 226 : 1-21 ( 2016 )

Coecobryci annulata sp. nov.

um:lsid:zoobank.org:act:D7CC34A6-E891-46AA-8FBD-570CAB9C9449

Figs IB, 4, 5, 9C; Table 1

Diagnosis

Ant. IV weakly annulated; 9 smooth clypeal chaetae; tiny mic r, X, X 0 and X 4 on the ventral side of head;

partial inner differentiated tibiotarsal chaetae “smooth”; unguiculus lanceolate with a tiny tooth on outer

edge; 9+9 mac on Abd. Ill with 2+2 lateral mac; Abd. IV with 6+6 central and 8+8 lateral mac.

Etymology

Named after the weakly annulated Ant. IV (Latin word “annulata”).

Material examined

Holotype

CHINA: $ on slide, Guangxi, Longzhou, Dong Liang Dong, 14 Apr. 2010, Louis Deharveng and Anne

Bedos leg. (# CHIgxl0-03). Deposited inNJAU.

Paratypes

CHINA: 2 $ $ on slides and 4 $ $ in alcohol, same data as holotype. One paratype deposited in NJAU

and others in MNHN.

Description

Body length up to 3.0 mm. Body colour white or with weak orange pigments (Fig. IB).

Antenna 2.1-2.8 times as long as cephalic diagonal. Antennal segments ratio as I : II : III : IV = 1 :

1.8-2.1 : 2.1-2.4: 2.7-3.4. Long smooth straight chaetae absent on antennae. Two internal S-chaetae of

Ant. Ill organ paddle-like (Fig. 4A). Ant. IV weakly annulated in large specimens.

Eyes absent. Clypeal chaetae 9, all smooth (Fig. 9C). Dorsal cephalic chaetotaxy with 5 sutural (S) and

4(5) mac in Gr. II (Fig. 4B). Papilla E with 4 guard chaetae; lateral process of labial palp as thick as

normal chaetae. Labial chaetae as mrel^; chaetae r, X, X 0 and X 4 as smooth tiny mic; chaeta X 3 absent

(Fig. 4C). Cephalic groove with 8 chaetae, anterior three and the fifth smooth and others ciliate.

Trochanteral organ with 15-23 smooth spine-like chaetae; 8-11 in arms and 7-12 between them (Fig. 4D).

Partial inner differentiated tibiotarsal chaetae “smooth” with ciliations closely or moderately appressed

to axis. Unguis with 3 inner teeth; 2 paired teeth unequal, outer one large. Unguiculus lanceolate with a

tiny tooth, sometimes inconspicuous, on its outer edge (Fig. 4E). Abd. IV 4.6-6.2 times as long as Abd.

Ill along dorsal midline. Ventral tube anteriorly with 7 ciliate chaetae on each side (Fig. 4F); posteriorly

with 11-14 smooth chaetae (Fig. 4G); each lateral flap with 8-11 smooth. Manubrial plaque with 2(1)

pseudopores and 4(5) ciliate chaetae. Distal smooth part of dens subequal to mucro in length (Fig. 4H).

Th. II with 3 medio-medial (ml, m2, m2i), 3 medio-lateral (m4, m4i, m4p) and 24-25 posterior mac; p5

as mac (Fig. 5A). Th. Ill with 27-28 mac; p5, m5i and a6i as mac (Fig. 5B). Abd. I with 6 (m2^l, m2i,

a3, m4p) mac. Abd. II with 3 (a2, m3, m3e) central and 1 (m5) lateral mac. Abd. Ill with 1 (m3) central

and 2 (pm6, p6) lateral mac (Fig. 5C). Abd. IV with 6 central (I, M, B4-5, A5-6), 8 lateral mac (Fl-3,

E2^1, E2p, D3) (Fig. 5D). Abd. V with 4 mac (m2, m3, m5, a5i) (Fig. 5E). S-chaetae formula: 2+ms,

2/1+ms, 2, 2+ms, 2, 3. Uncertain number of longer S-lilce chaetae on Abd. IV.

Ecology

Cave-restricted species with troglomorphic characters (long antennae, slightly modified claw, large size).

8

ZHANG F. et al., Cave Coecobrya from China and clypeal chaetae

Fig. 4. Coecobrya annulata sp. nov. A. Ant. Ill organ. B. Dorsal cephalic chaetotaxy. C. Chaetae on the

ventral side of head. D. Trochanteral organ, ventral view. E. Hind claw. F. Anterior face and lateral flap

of ventral tube. G. Posterior face of ventral tube. H. Mucro. Scale bars: A, C-E, H = 50 pm; B, F-G =

100 pm.

9

European Journal of Taxonomy 226 : 1-21 ( 2016 )

Remarks

Coecobrya annulata sp. nov. is characterized by annulated Ant. IV, two paddle-like internal S-chaetae

of Ant. Ill organ, tiny mic r, X, X 0 and X 4 on the ventral side of head, 9 smooth clypeal chaetae, an outer

tiny tooth on unguiculus (sometimes inconspicuous), 2+2 lateral mac on Abd. Ill, 6+6 central and 8+8

lateral mac on Abd. IV. It is also the only species having ms on Abd. Ill among Chinese cave Coecobrya

species. It is closest to C. oculata sp. nov. from which it differs in anophthalmy, smooth clypeal chaetae,

chaetae on the ventral side of head, “smooth” inner differentiated tibiotarsal chaetae, ventral tube and

dorsal chaetotaxy (Table 1).

Fig. 5. Coecobrya annulata sp. nov. A. Chaetotaxy of Th. II. B. Chaetotaxy of Th. III. C-E. Abdominal

chaetotaxy. C. Abd. I—III. D. Abd. IV. E. Abd. V. Scale bars: 200 pm.

10

ZHANG F. et al., Cave Coecobrya from China and clypeal chaetae

Coecobryci ciliata sp. nov.

urn: lsid: zoobanlc, org: act: 1E2353EF -E3B5 -4ED9- 8 A6 A-9 A43 71452F 3 7

Figs 1C, 6, 7, 9D; Table 1

Diagnosis

Small size; long smooth straight chaetae present on antennae; 15 ciliate clypeal chaetae; labial chaeta R

ciliate; postlabial chaetae H ( 4 ciliate; unguis elongate; unguiculus truncate or swollen with a tiny tooth

on outer edge; Abd. IV with 3+3 central and 5+5 lateral mac.

Etymology

Named after the ciliate chaetae on the ventral side of head.

Material examined

Holotype

CHINA: $ on slide, Guangxi, Huanjiang, Muhin, Mashan Dong, 19 May 2007, Louis Deharveng, Anne

Bedos and Youbang Li leg. (# CHIgx07-19-01). Deposited in NJAU.

Paratype

CHINA: $ on slide and same data as holotype. Deposited in MNHN.

Additional material

CHINA: 1 $ on slide and 4 $ 5 in alcohol, Guangxi, Huanjiang, Muhin, Dong Zai Dong, 20 May 2007,

L. Deharveng et al. leg. (# CHIgx07-20-01). Deposited in MNHN.

Description

Body length up to 1.35 mm. Ground colour pale yellow in alcohol (Fig. 1C).

Antenna 1.4-1.8 times as long as cephalic diagonal. Antennal segments ratio as I : II : III : IV = 1 :

1.5-2.2 : 1.1-1.9: 2.5-3.0. Ant. I ventrally with long smooth straight chaetae. Two internal S-chaetae of

Ant. Ill organ slightly expanded (Fig. 6A). Ant. IV not annulated.

Eyes absent. Clypeal chaetae 15, all ciliate (Fig. 9D). Dorsal cephalic chaetotaxy with 5 sutural mac and

4 mac in Gr. II (Fig. 6B). Lateral process of labial palp thicker than normal chaetae (Fig. 6C). Labial

base as mReljf,, all smooth except R; R/m = 0.5; chaetae H^, X and ciliate (Fig. 6D). Cephalic

groove with 7 chaetae, the anterior one (G^ smooth and others ciliate.

Trochanteral organ with 9-16 smooth spine-like chaetae; 7-8 in arms and 3-6 between them (Fig. 6E).

Inner differentiated tibiotarsal chaetae ciliate with ciliations not closely appressed to axis, most distal

one smooth on hind claw. Ungual basal paired teeth unequal with outer one larger, 20% distance from

the base; distal unpaired tooth tiny, 25% distance from the base. Unguiculus truncate or swollen with

a tiny tooth, sometimes inconspicuous, on outer edge (Fig. 6F). Abd. IV 3.1-3.5 times as long as Abd.

Ill along dorsal midline. Ventral tube anteriorly with 5-6 ciliate chaetae on each side, one of them

much larger than others (Fig. 6G); posteriorly with 2 apical smooth and 7-8 ciliate chaetae (Fig. 6H);

lateral flap with 5 smooth and 1-2 ciliate chaetae. Manubrial plaque with 2 pseudopores and 4-5 ciliate

chaetae. Distal smooth part of dens 0.7 times as long as mucro (Fig. 61).

Th. II with 3(4) (ml, m2, m2i) medio-median, 3 (m4, m4p, m4i) medio-lateral and about 20 posterior

mac; p4, p4i and p5 as mac; mac mli and p4i rarely present. Th. Ill with about 31 mac; m5i, a6i and p5

as mac (Fig. 6J). Abd. I with 6 (m2i, m2, m3, a3, m4 and m4p) mac. Abd. II with 3 (a2, m3, m3e) central

and 1 (m5) lateral mac. Abd. Ill with 1 (m3) central and 3 (am6, pm6, p6) lateral mac (Fig. 7A). Abd.

11

European Journal of Taxonomy 226 : 1-21 ( 2016 )

Fig. 6. Coecobrya ciliata sp. nov. A. Ant. Ill organ. B. Dorsal cephalic chaetotaxy. C. Lateral process

of labial palp. D. Chaetae on the ventral side of head. E. Trochanteral organ, ventral view. F. Hind claw.

G. Anterior face of ventral tube ad lateral flap. H. Posterior face of ventral tube. I. Mucro. J. Thoracic

chaetotaxy. Scale bars: A, C-I = 20 pm; B, J = 100 pm.

12

ZHANG F. et al.. Cave Coecobrya from China and clypeal chaetae

IV with 3 (A6, M, B5) central, 5 (FI, E2^t, E2p) lateral mac (Fig. 7B). Abd. V with 3 sens (Fig. 1C).

S-chaetae formula: 2+ms, 2/1+ms, 2, 2+ms, 2, 3. Uncertain number of longer S-like chaetae on Abd. IV.

Ecology

Cave-restricted species, moderately troglomorphic (slightly modified claw).

Remarks

Ciliate chaetae H ] 4 on the ventral side of head in C. ciliata sp. nov. are observed in Coecobrya for the

second time since C. draconis. Small size, 15 ciliate clypeal chaetae, long smooth straight chaetae on

antennae, elongate unguis, truncate or swollen unguiculus, tiny outer tooth on unguiculus (sometimes

inconspicuous), and 5+5 lateral mac on Abd. IV are also characteristic. It is closest to C. draconis but

differs from it in long smooth straight chaetae on antennae and chaetotaxy of Abd. IV (Table 1).

Fig. 7. Abdominal chaetotaxy of Coecobrya ciliata sp. nov. A. Abd. I—III. B. Abd. IV. C. Abd. V. Scale

bars: 50 pm.

13

European Journal of Taxonomy 226: 1-21 (2016)

Coecobrya oculata sp. nov.

um:lsid:zoobank.org:act:ABEEC710-582D-4381-9D81-83C2D5948912

Figs ID, 8; Table 1

Diagnosis

Small size; eyes 1+1; clypeal chaetae ciliate; chaeta R, H p H 0 , X, X 0 and X 4 ciliate on the ventral side

of head; unguiculus lanceolate with outer edge serrate; Abd. IV with 4+4 central and 8+8 lateral mac.

Etymology

Named after the presence of eyes.

Material examined

Holotype

CHINA: $ on slide, Guangxi, Shanglin, Longshan, Jinlun Dong, 23.45239° N, 108.74154° E, 19 Mar.

2005, Louis Deharveng and Anne Bedos leg. (# CHIgx05-137). Deposited in NJAU.

Paratypes

CHINA: $ on slide and 5 in alcohol, same data as holotype; 3 $ $ on slides and 4 in alcohol, Guangxi,

Shanglin, Longshan, Ganlin Dong, 23.45932° N, 108.73436° E, 18 Mar. 2005, Louis Deharveng and

Anne Bedos leg. (# CHIgx05-136). Deposited in MNHN.

Description

Body length up to 1.38 mm. Body colour white (Fig. ID).

Antenna 1.6-1.7 times as long as cephalic diagonal. Antennal segments ratio as I : II : III : IV = 1:

1.5-1.8: 1.2—1.6: 2.6-2.7. Long, smooth straight chaetae absent on antennae. Ant. II distally with 1 small

rod-like S-chaeta. Ant. Ill organ with 2 rods. Ant. IV not annulated.

Eyes 1+1. Clypeal chaetae ciliate but their arrangement not clearly seen. Dorsal cephalic chaetotaxy

with 4 sutural chaetae and 5(4) mac in Gr. II (Fig. 8B). Lateral process of labial palp straight, thicker

than normal chaetae (Fig. 8C). Labial base as mRel^, chaeta R ciliate and 0.45-0.60 length of M;

chaetae H p H 0 , X, X 0 and X 4 ciliate; chaeta X 3 absent (Fig. 8D). Cephalic groove with 6 chaetae, the first

and the third smooth and others ciliate.

Trochanteral organ with 9-13 smooth spine-like chaetae; 7-9 in arms and 2M between them (Fig. 8E).

Inner differentiated tibiotarsal chaetae ciliate with ciliations not closely appressed to axis. Ungual basal

paired teeth unequal, outer one large. Unguiculus lanceolate with outer edge serrate (Fig. 8F). Abd. IV

3.(Ml.4 times as long as Abd. Ill along dorsal midline. Ventral tube anteriorly with 4 ciliate chaetae on

each side and two of them much larger (Fig. 8G); posteriorly with 6 smooth chaetae (Fig. 8H); each

lateral flap with 5 smooth chaetae and 1 additional ciliate chaeta rarely present. Manubrial plaque with

2 pseudopores and 3(2) ciliate chaetae. Distal smooth part of dens slightly longer than mucro (Fig. 81).

Th. II with 4 (ml, mli, m2, m2i) medio-median, 3 (m4, m4i, m4p) medio-lateral and 21-23 posterior

mac. Th. Ill with about 29 mac; m5i, a6i, p5, m6e, m6i as mac; a4i2 and a6i sometimes absent in young

specimens (Fig. 8J). Abd. I with 6 (m2M, m2i, a3, m4p) mac. Abd. II with 3 (a2, m3, m3e) central and

1 (m5) lateral mac. Abd. Ill with 1 (m3) central and 3 (am6, pm6, p6) lateral mac (Fig. 8K). Abd. IV

with 4 central (A6, B4-5, M), 8 lateral mac; F2 and F3 as mac (Fig. 8L). Abd. V with 3 sens (Fig. 8M).

S-chaetae formula: 2+ms, 2/1+ms, 2, 2+ms, 2, 3. About 6 long S-like chaetae on Abd. IV.

14

ZHANG F. et al., Cave Coecobrya from China and clypeal chaetae

Fig. 8. Coecobrya oculata sp. nov. A. Labrum. B. Dorsal cephalic chaetotaxy. C. Lateral process of

labial palp. D. Chaetae on the ventral side of head. E. Trochanteral organ, ventral view. F. Hind claw.

G. Anterior face of ventral tube and lateral flap. H. Posterior face of ventral tube. I. Mucro. J. Thoracic

chaetotaxy. K-M, Abdominal chaetotaxy. K. Abd. I—III. L. Abd. IV. M. Abd. V. Scale bars: A, C-I =

20 pm; B, J-M =100 pm.

15

European Journal of Taxonomy 226: 1-21 (2016)

Ecology

Cave-restricted species, non-troglomorphic.

Remarks

Coecobrya oculata sp. nov. is characterized by small size, 1+1 eyes, ciliate chaetae R, H p H 0 , X,

and X 4 on the ventral side of head, serrate outer edge of unguiculus, absence of ms on Abd. Ill and 4+4

central mac on Abd. IV. See Table 1 for comparison with other Chinese cave species.

Fig. 9. Clypeal chaetae. A. Diagram. B. Coecobrya gejianbangi sp. nov. C. Coecobrya annulata sp. nov.

D. Coecobrya ciliata sp. nov. E. Coecobrya brevis. F. Coecobrya pani. Figures of the latter two species

after Xu et al. (2012). Scale bars: 40 pm.

16

ZHANG F. et al., Cave Coecobrya from China and clypeal chaetae

Discussion

Modification of the unguiculus in cave species and redefinition of the tenebricosa- group

Most blind Coecobrya species have a large outer tooth on unguiculus and can be assigned to the

tenebricosa- group (Zhang et al. 2011a). Among the five cave species from China, C. oculata sp. nov.

lacks troglomorphic features and is similar to non-cave members of the boneti- group. The four other

species exhibit more or less marked troglomorphic traits, e.g., elongated unguis with reduction of inner

ungual teeth which is displaced towards unguis base when present in C. gejianbangi sp. nov., C. ciliata

sp. nov. and C. draconis , or two paddle-like internal S-chaetae on Ant. Ill organ in C. annulata sp. nov.

Among these four blind species, the outer tooth on unguiculus is extremely tiny or absent (Figs 2F, 4E,

6F) instead of being large in typical species of the tenebricosa- group. A tiny outer tooth of unguiculus

is not exceptional in Entomobryidae, and often present in cave Pseudosinella Schaffer, 1897 species

(Christiansen 1961). The link between cave life and reduction of unguiculus outer tooth in species

of remotely related genera suggest evolutionary convergence of the character under similar selective

pressure, though the functional basis of this evolution is obscure. Smooth outer edge in C. draconis

might be a further reduction from a tiny tooth. Under the assumption of tiny tooth modified from large

one, we place the five blind cave Coecobrya species in the tenebricosa- group, that is re-defined for

those species devoid of eyes. Size and number of outer teeth on unguiculus are not the key character

separating the two groups.

Clypeal chaetae in Entomobryoidea

Clypeal area locates between frontal and prelabral areas, always with several chaetae. Its details in

Entomobryoidea are difficult to observe for its peculiar position, being hard to expose during slide

preparation, or covered by antennae. Clypeal chaetae are usually overlooked by collembologists, but

were widely used by Yoshii and Suhardjono (Yoshii 1982, 1983, 1985, 1987, 1988, 1989, 1990a, 1990b;

Yoshii & Suhardjono 1989, 1992). Zhang (2013) also applied clypeal chaetae in distinguishing species

in Sinella Brook, 1882 and Coecobrya.

To easily study clypeal chaetae, we divide this area into prefrontal and facial part (after Yoshii &

Suhardjono 1992, modified), with a few lateral chaetae (Fig. 9A; Table 2). Lateral chaetae ( L ] and L 0 ),

ciliate or smooth and usually larger than clypeal chaetae, are external to prelabral and clypeal chaetae.

Prefrontal chaetae are 3 arranged in a row, usually all smooth or ciliate; middle chaeta is designated as

pf 0 , lateral ones as pf r Chaeta pf 0 is sometimes different from p^ in morphology, e.g., in Alio sc opus

tetracanthus (Borner, 1906) and in Coecobrya edenticulata (Handschin, 1926) (both after Yoshii &

Suhardjono 1989).

Facial chaetotaxy is highly diversified in the number and arrangement of its chaetae, even in several

groups morphologically conserved. Three families have a basic pattern of two chaetae arranged in a

row. In lepidocyrtids where tergal chaetotaxy and many characters are quite conserved, number of facial

chaetae varies from 2 to 13 in the recorded species. For the species with the same number of facial

chaetae, they often have different arrangement and size. For example, both Coecobrya pani Xu, Yu &

Zhang, 2012 and Coecobrya gejianbangi sp. nov. have 8 facial chaetae, the former with 2,2, 2, 2 smooth

chaetae arranged in four rows and two chaetae of the third row tiny (Fig. 9F), the latter with 1, 3, 2, 2

ciliate chaetae and two median ones of them much smaller than others (Fig. 9B).

Clypeal chaetae in Entomobryoidea don’t exhibit obvious differentiation at the familial level, but are

diversified at the species level. It could be a powerful tool to help to discriminate species particularly in

morphologically conserved groups, e.g., lepidocyrtids. To easily observe this character in a specimen on

slide, it is suggested adjusting the specimen to the proper pose during slide preparation.

17

Table 2 (continued on next page). Clypeal chaetae in Entomobryoidea (sm = smooth; c = ciliate; * = original observations, different from the literature).

European Journal of Taxonomy 226: 1-21 (2016)

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19

European Journal of Taxonomy 226: 1-21 (2016)

Acknowledgments

We thank the staff of the natural reserves of Guangxi for having organized the field trips. Tian Mingyi

and Li Youbang provided specimens as well as invaluable help in the field. The “Guangxi Integrated

Forestry Development and Biodiversity Conservation Project”, a GEF-World Bank project, funded

biological cave surveys in the province. Liu Jin (World Bank) and Tony Whitten (FFI) greatly facilitated

our work all through the duration of the project. The Museum national d’Histoire naturelle, Paris (France)

provided a grant to the first author for working at the MNHN.

References

Bellinger P.F., Christiansen K.A. & Janssens F. 1996-2016. Checklist of the Collembola of the World

[online]. Available from http://www.collembola.org [accessed 12 Mar. 2016]

Chen J -X. & Christiansen K.A. 1993. The genus Sinella with special reference to Sinellas. s. (Collembola:

Entomobryidae) of China. Orient Insects 27: 1-54. http://dx.doi.org/10.1080/003Q5316.1993.10432236

Christiansen K. 1961. Convergence and parallelism in cave Entomobryinae. Evolution 15: 288-301.

http://dx.doi.org/10.2307/24Q6229

Deharveng L., Brehier F., Bedos A., Tian M.Y., Li Y.B., Zhang F., Qin W.G. & Tan X.F. 2008. Mulun

and surrounding karsts (Guangxi) host the richest cave fauna of China. Subterranean Biology 6: 75-79.

Ding Y.H. & Zhang F. 2015. Contribution to the eyed Sinella from China: two new species and additional

reports on nine known species (Collembola: Entomobryidae). Zootaxa 3973: 474M90. http://dx.doi.

org/10.11646/zootaxa.3973.3.4

Szeptycki A. 1979. Morpho-systematic studies on Collembola. IV. Chaetotaxy of the Entomobryidae

and its phylo gene deal significance. Polska Akademia Nauk, Zakiad Zoologii Systematycznej i

Doswiadczalnej, Panstwowe Wydawnictwo Naukowe, Warszawa, Krakow.

Xu G.-L., Yu D.-Y. & Zhang F. 2012. Two new species of Coecobrya (Collembola: Entomobryidae:

Entomobryinae) from China, with a key to the Chinese specie of the genus. Zootaxa 3399: 61-68.

Yoshii R. 1982. Lepidocyrtid Collembola of Sabah. Entomological Report from the Sabah Forest

Research Centre 5: 1—47.

Yoshii R. 1983. Entomological report from the Sabah forest research centre. No. 7. Studies on Paronellid

Collembola of East Asia. Japan International Cooperation Agency 1-28.

Yoshii R. 1985. Notes on paronellid Collembola of southeast Asia. Contributions from the biological

laboratory Kyoto University 27: 73-90.

Yoshii R. 1987. Notes on some cyphoderid Collembola of the tropical Asia. Contributions from the

biological laboratory Kyoto University 27: 121-136.

Yoshii R. 1988. Paronellid Collembola from caves of Central and South America collected by P. Strinati.

Revue Suisse de Zoologie 95: 449-459. Available from http://biodiversitvlibrarv.org/page/41228721

[accessed 2 Aug. 2016]

Yoshii R. 1989. Finding of Lepidosinella armata Handschin from east Java. In: Dallai R. (eds) Third

International Seminar of Apterygota: 89-91. University of Siena, Siena.

Yoshii R. 1990a. Miscellaneous notes on the Collembola of Macaronesia. Contributions from the

biological laboratory Kyoto University 27: 535-540.

Yoshii R. 1990b. Miscellaneous notes on the Collembola of caves. I. Sinella spinidentata Yosii, 1942

and Sinella coeca (Schott, 1897). Annals of the Speleological Research Institute of Japan (Iwaizumi) 8:

1 - 6 .

20

ZHANG F. et al., Cave Coecobrya from China and clypeal chaetae

Yoshii R. & Suhardjono Y.R. 1989. Notes on the Collembolan Fauna of Indonesia and its vicinities.

I. Miscellaneous notes, with special references to Seirini and Lepidocyrtini. Acta Zoologica Asiae

Orientalis 1: 23-90.

Yoshii R. & Suhardjono Y.R. 1992. Notes on the Collembolan Fauna of Indonesia and its vicinities. Ill:

Collembola of Timor Island. Acta Zoologica Asiae Orientalis 2: 75-96.

Zhang F. 2013. Five new eyed species of Sinella (Collembola: Entomobryidae) from China, with a key

to the eyed species of the genus. Zootaxa 3736: 549-568. http: //dx. doi. org/10.11646/zootaxa.3736.5.7

Zhang F. & Deharveng L. 2015. Systematic revision of Entomobryidae (Collembola) by integrating

molecular and new morphological evidence. Zoologica Scripta 44: 298-311. http://dx.doi.org/10.llll/

zsc.12100

Zhang F., Deharveng L. & Chen, J.-X. 2009. New species and rediagnosis of Coecobrya (Collembola:

Entomobryidae), with a key to the species of the genus. Journal of Natural History 43: 2597-2615.

http://dx.doi.org/10.1080/0022293090324397Q

Zhang F., Man L.C. & Deharveng L. 2011a. A review of the boneti-grouy of the genus Coecobrya

(Collembola: Entomobryidae). Zootaxa 2748: 61-68.

ZhangF., YuD.-Y. & Xu G.-L. 2011b. Transformational homology ofthe tergal setae duringpostembryonic

development in the Sinella-Coecobrya group (Collembola: Entomobryidae). Contributions to Zoology

80: 213-230.

Manuscript received: 22 January 2016

Manuscript accepted: 5 April 2016

Published on: 1 September 2016

Topic editor: Gavin Broad

Desk editor: Kristiaan Hoedemakers

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; Naturalis Biodiversity Center, Leiden, the Netherlands.

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