Research Article |
Corresponding author: Manqiang Liu ( liumq@njau.edu.cn ) Academic editor: Wanda M. Weiner
© 2016 Daoyuan Yu, Qibao Yan, Manqiang Liu.
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.
Citation:
Yu D, Yan Q, Liu M (2016) New cave-dwelling species of Tomoceridae from China, with a study on the pattern of mesothoracic bothriotricha in Tomocerinae (Collembola, Entomobryomorpha). ZooKeys 574: 81-95. https://doi.org/10.3897/zookeys.574.7312
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Two new troglobitic species of Tomoceridae are described from Guizhou and Guangxi provinces, China. Tomocerus tiani sp. n. resembles Tomocerus kinoshitai Yosii, 1954, Tomocerus caecus Yu & Deharveng, 2015 and Tomocerus similis Chen & Ma, 1997 but differs from them mainly in the body colour, the cephalic chaetotaxy and the number of manubrial pseudopores. Monodontocerus cinereus sp. n. is similar to Monodontocerus mulunensis Yu, Deharveng & Zhang, 2014 but is different from the latter in the body colour, the length of antennae, the number of ungual teeth and the chaetotaxy on Abd. III and Abd. IV. Special remarks are made on the mesothoracic bothriotricha in Tomocerinae.
Tomocerus , Monodontocerus , troglobitic, Tomocerinae
Since the discovery of Tritomurus scutellatus Frauenfeld, 1854 in Slovenian caves, troglobitic Tomoceridae have frequently been reported in Asia, Europe and North America. Besides the troglobitic genera Tritomurus Frauenfeld, 1854 and Lethemurus Yosii, 1970, some other main groups of Tomoceridae have also been found with cave dwellers, and several genera, i.e. Monodontocerus Yosii, 1955, Plutomurus Yosii, 1956 and Aphaenomurus Yosii, 1956 have mainly or usually been found in caves. However, despite of several highly troglomorphic species, for example Tritomurus falcifer Cassagnau, 1958 and Tritomurus veles Lukić, Houssin and Deharveng, 2010 which are both eyeless and have very elongated antennae and claws, other cave tomocerids do not exhibit distinct troglomorphic characters: most of them have short to moderate antennae, normal or slightly elongated claws and a full set of 6+6 eyes for Tomocerinae, only the sizes of eyes are usually smaller than those of the edaphic species, and the tenent hair is often pointed as in other cave Collembola.
Four cave tomocerids have previously been reported in China, including Tomocerus caecus Yu & Deharveng, 2015, Monodontocerus absens Yu, Deharveng & Zhang, 2014 and Monodontocerus mulunensis Yu, Deharveng & Zhang, 2014 from Guangxi Province, and Monodontocerus trigrandis Yu, Deharveng & Zhang, 2014 from Hunan Province. The present paper reports two new species of Tomoceridae discovered in caves in the south-west karst regions of China. Both of the new species have small eyes and pointed tenent hair, but neither of them is highly troglomorphic. Tomocerus tiani sp. n. bears some unusual characters, including the single mesothoracic bothriotrichum, which lead to a comprehensive examination of the different genera of Tomocerinae.
Specimens were collected with aspirators and preserved in 99% ethanol. For detailed morphological study, specimens were cleared in Nesbitt’s fluid and mounted in Marc André II solution. For some specimens the furca, the ventral tube and the legs were cut off from the trunk and mounted separately for detailed observation. The slide-mounted specimens were studied using a Nikon Ni microscope. Photos were taken using Nikon DS-Fi1 cameras mounted respectively on Nikon SMZ1000 stereomicroscope and Nikon Ni microscope.
Ant. antennal segment
PAO postantennal organ
Th. thoracic segment
Abd. abdominal segment
Institutional acronyms
NJAU Nanjing Agricultural University, Nanjing, China
China, Guizhou Province: Zunyi, Suiyang County, Wenquan Town, Guihua Village, Hejiao Cave, inside cave, 7 November 2008, Mingyi Tian leg.
Holotype male (labelled 15cave15-1) and paratype juvenile (labelled 15cave15-2) on slide. Deposited in NJAU.
Species similar to Tomocerus kinoshitai Yosii, 1954, with short antennae, multi-furcated dental spines and apically curved mucro. Body length approximately 3.0 mm, with purplish grey pigment all over; antennae approximately half as long as body; eyes small; terminal hair of maxillary outer lobe with a small basal denticle; Th. II with only one bothriotrichum; tenent hairs pointed; unguis with two teeth, baso-internal ridges at approximately 1/2 distance from base; manubrium with 12–17 pseudopores on each side; dental spines formula as 4/2, II; dens dorsally with only a few feather-like chaetae; mucro without intermediate tooth. Cave-dwelling species.
Body length 2.9 mm. Body with uniform purplish grey pigment and unpigmented patches, appendages paler. Eye patches black (Fig.
Tomocerus tiani sp. n. and Monodontocerus cinereus sp. n. A and B, Tomocerus tiani sp. n. A appearance in alcohol (lateral view) B dorsal chaetae on dens (dorsal view, arrows pointing to feather-like chaetae) C Monodontocerus cinereus sp. n. C appearance in alcohol (lateral view). Scale bars: 1000 μm (A, C), 50 μm (B).
Antennae approximately half length of body. Length ratio of antenna segments as I:II:III+IV = 1.0:1.9–2.0:9.6–9.7. Only dorsal side of Ant. I and Ant. II scaled, Ant. III+IV unscaled. PAO not seen. Eyes 6+6, relatively small. Labral formula as 4/5, 5, 4. Distal edge of labrum with four curved spine-like papillae. Mandibular heads asymmetrical, the left one with four teeth and the right one with five, left molar plate distally with a tapered tooth (Fig.
Tomocerus tiani sp. n. A mandibular heads (dorsal view) B palp of left maxillary outer lobe (dorsal view); C cephalic dorsal chaetotaxy (dorsal view, circle: socket of chaeta, same as below) D dorsal chaetotaxy of Th. II–Abd. V (dorsal view, circle with a slash: pseudopore, wavy line: bothriotricha, same as below) E trochantero-femoral organ (inner view) F hind tibiotarsus (lateral view, showing spine-like inner chaetae) G hind claw (lateral view, t: tenent hair, a: accessory chaeta, g: guard chaeta, p: pretarsal chaeta, same as below).
Pattern of body chaetotaxy as in Fig.
Trochantero-femoral organ with 1, 1 small slender chaetae (Fig.
Anterior face of ventral tube with scales, posterior face and lateral flaps unscaled, anterior face with ca. 25 chaetae on each side, posterior face with ca. 30 chaetae, each lateral flap with ca. 15 chaetae. Rami of tenaculum with 4+4 teeth, anterior face with one chaeta and without scale (Fig.
Tomocerus tiani sp. n. A tenaculum (anterior view) B right side of manubrium (dorsal view, showing prominent dorsal chaetae, T-shaped mark: socket of scale, same as below) C disto-external corner of manubrium (dorsal view) D basal and middle subsegments of dens (dorsal view, showing dental spines and prominent dorso-basal chaeta) E feather-like dental chaeta (lateral view) F mucro (lateral view).
Named after the collector Prof. Mingyi Tian.
Tomocerus tiani sp. n. is similar to Tomocerus caecus Yu & Deharveng, 2015, Tomocerus kinoshitai (materials from Changbai Mountain, China) and Tomocerus similis Chen & Ma, 1997 (type materials) in the length of antennae, the general pattern of chaetotaxy on Th. II, the number and position of spine-like tibiotarsal inner chaetae, the size of external corner chaeta on the manubrium, the type and general arrangement of dental spines and the shape of mucro, but it is clearly different from the first species in having eyes and pigment, and is different from the other two species mainly in the body colour, the cephalic chaetotaxy, the sharply pointed tibiotarsal strong inner chaetae and tenent hair, and the more slender unguiculus; besides, with similar body size, Tomocerus tiani sp. n. has more manubrial pseudopores than the three known species. The small basal denticle of the terminal hair of maxillary outer lobe is so far unique to Tomocerus tiani sp. n. and is useful for diagnosis if dissected and exposed carefully. The baso-internal ridges of unguis are located at approximately 1/2 distance from the base in Tomocerus tiani sp. n., whereas in most other species the distance between the ridges and the base is only 1/3 or less. The dorsal dental chaetae in Tomocerus tiani sp. n. is also characteristic, that the dense stripes of feather-like chaetae in most other tomocerids are almost replaced by ordinary chaetae and swollen serrated chaetae, leaving only a few feather-like ones. Similar condition was also reported in Tomocerus kinoshitai and Tomocerus caecus that some spine-like chaetae are present on dens (
The juvenile specimen is almost identical to the adult in most characters, including the macrochaetotaxy, the number of teeth on claws and the dental spines formula. However, some characters on manubrium are distinctly different between juvenile and adult. In the juvenile specimen, there are ca. 80 dorsal chaetae on each side of manubrium, the number of pseudopores is only 4–5 on each side, and the external corner chaeta is as large as a mesochaeta in the dorsal chaetal stripe. These differences provide interesting information in the postembryonic development of manubrium in Tomocerinae, and indicate these characters are not suitable for the identification of immature specimens at different instars.
China, Guangxi Province: Hechi, Duan County, Chengjiang Township, Ganwan Village, Nongsi Cave, 23°56'24"N, 108°10'12"E, alt. ca. 270 m, inside cave, 25 July 2015, Jujian Chen, Xinhui Wang and Mingruo Tang leg.
Holotype male (labelled 15cave6-1) and two paratypes female (labelled 15cave6-2 and -3) on slides, one paratype (labelled 15cave6) in alcohol. Deposited in NJAU.
Typical Monodontocerus species with multi-furcated dental spines and single mucronal basal tooth. Body length approximately 4.0 mm, with light grey pigment all over; antennae slightly shorter than body; eyes small; chaetotaxy typical for the genus; tenent hair pointed; unguis with 2–4 teeth; manubrium with 28–35 pseudopores on each side; dental spines formula as 4, II/6, I, 3, I or 5, I/6, I, 3, I; mucro with 3–4 intermediate teeth. Cave-dwelling species.
Body length 3.9–4.1 mm. Body colour uniformly light grey with unpigmented patches, appendages paler. Eye patches black. (Fig.
Antennae approximately 0.7–0.9 times as long as body. Length ratio of antenna segments as I:II:III:IV = 1.0:1.7–2.0:10.5–11.0:1.8. Only dorsal side of Ant. I and Ant. II scaled, Ant. III and Ant. IV unscaled. PAO not seen. Eyes 6+6, relatively small. Mouthparts typical for Tomocerinae. Labral formula as 4/5, 5, 4. Distal edge of labrum with four curved spine-like papillae. Mandibular heads asymmetrical, the left one with four teeth and the right one with five, left molar plate distally with a tapered tooth (Fig.
Monodontocerus cinereus sp. n. A mandibular heads (dorsal view) B maxillary lamella 5 (dorsal view) C cephalic dorsal chaetotaxy (dorsal view) D dorsal chaetotaxy of Th. II–Abd. V (dorsal view) E trochantero-femoral organ (inner view) F hind tibiotarsus (lateral view, showing spine-like inner chaetae) G middle claw (lateral view).
Pattern of body chaetotaxy as in Fig.
Trochantero-femoral organ with 1, 1 small slender chaetae (Fig.
Ventral tube with scales on both anterior and posterior faces, lateral flaps unscaled, anterior face with ca. 70 chaetae on each side, posterior face with ca. 160 chaetae, each lateral flap with ca. 100 chaetae. Rami of tenaculum with 4+4 teeth, anterior face with one chaeta and without scale (Fig.
Named for its light grey body colour, from the Latin cinereus, meaning ash-coloured.
Within the genus, Monodontocerus cinereus sp. n. is more similar to Monodontocerus mulunensis Yu, Deharveng & Zhang, 2014 in the cephalic chaetotaxy, the pointed tenent hair and the number of mucronal intermediate teeth, but can be distinguished from the latter by having longer antennae, fewer teeth on unguis and more macrochaetae on Abd. III and Abd. IV. In alcohol the new species can be identified from other known species of Monodontocerus by the grey body colour.
Tomocerus tiani sp. n. has only one bothriotrichum on Th. II, while there are two in most other Tomocerus species observed previously. The pattern of bothriotricha is significant for the taxonomy and phylogeny of Collembola (
There are two main obstacles to determining the pattern of mesothoracic bothriotricha in Tomocerinae. Firstly, these long, thin and ciliated chaetae, as well as the macrochaetae, are easily lost during specimen collection and slide preparation, leaving only the sockets. The sockets of bothriotricha are usually characteristic for their round shape and small size, but sometimes can still be confused with the sockets of macro- or mesochaetae. Secondly, in case of two bothriotricha present they are arranged transversely, and the outer one is usually near the lateral margin of the tergum, thus could possibly be omitted when the margin is wrinkled or damaged. To avoid those disadvantages, we examined the specimens with almost intact coating of chaetae, most of which are pre-molting specimens with new chaetae under the old cuticle.
In the observed species, there are three main patterns of mesothoracic bothriotricha, which appear to be relevant to the generic division except for several species of Tomocerus and Tomocerina. Pattern A: all species of Tomocerus ocreatus complex (
Dorsal view of the left antero-lateral corner of mesonotum (arrows directing anterior). A Tomocerus sp. (ocreatus complex) from China (b: bothriotricha, same as below) B Tomocerus similis from China (mb: macrochaeta at the location of outer bothriotricha, same as below) C Tomocerina annamitica from Vietnam D Tomocerus laxalamella from China (with bothriotricha-like macrochaeta two times magnified) E Pogonognathellus sp. from France. Scale bars: 50 μm (A, B, C, E), 100 μm (D).
This study has revealed several distinct patterns of mesothoracic bothriotricha in Tomocerinae, and has proved their taxonomic importance. However, since our study covered mostly Asian edaphic species, we have probably not exhausted the variability of this character among Tomocerinae. The exact pattern in Monodontocerus has not been successfully determined because of the lack of specimen with intact bothriotricha, though the sockets indicate pattern A more possible. For Pogonognathellus more examination is required since several species were described having two mesothoracic bothriotricha (
We sincerely thank Prof. Mingyi Tian and his team in South China Agricultural University who collected and provided the specimens of the new species, and Dr. Louis Deharveng in Museum National d’Histoire Naturelle (Paris) who transferred the specimens to us. This study 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).