Opusc. Zool. Budapest, 2012, 43(1): 79–87

Draconizercon punctatus gen. et sp. nov., a peculiar zerconid (: : Zerconidae) from Taiwan

ZS. UJVÁRI*

Abstract. A new genus and species, Draconizercon punctatus gen. et sp. nov. is described from low and high elevation areas of Taiwan. The classification problems within the family Zerconidae are discussed.

Keywords. Zerconidae, new genus, new species, Taiwan, rainforest.

INTRODUCTION cerine. Preparates were examined using a light microscope; drawings were made with the aid of a erconid were thought to be distributed drawing tube. Mites are deposited in the Col- Z in the Holarctic region, inhabiting the cold lection of Soil Zoology of Hungarian Natural and temperate climate zones only, however se- History Museum (HNHM), and the National Mu- veral records of the last two years show that seum of Natural Sciences, Taichung, Taiwan Zerconidae were able to disperse southwards (NMNS), in 70% ethanol. The terminology of idi- through the high mountain zone, reaching and osomal setae follows Lindquist & Evans (1965), expanding beyond the Tropic of Cancer (Ujvári, with modifications for the caudal region as given 2010, 2011a, b, c; Ma et al., 2011). According to by Lindquist & Moraza (1998). The system of our present knowledge, altogether eight species of five genera are known from Taiwan, seven of the notation for dermal glands and lyrifissures follows species are endemic on the island, besides one Johnston & Moraza (1991). The description of genus (Rotundozercon Ujvári, 2011) and two sub- gnathosomal structures is in accordance with genera (Parazercon (Formosella) Ujvári, 2011 Ujvári (2011d). All measurements including scale and (Zercorientalia) Ujvári, 2011)) seem bars of the figures are given in micrometres. also to be endemic. The present paper contributes to the knowl- edge of the Zerconidae fauna of Taiwan reporting Draconizercon gen. nov. on a remarkable species which, because of the special character-combination necessitated estab- lishing a new genus. Diagnosis. Podonotal shield carapace-like, ex- panded anteroventrally and lateroventrally, setae It should be remarked that all previously re- j1 situated ventrally. The slit between peritrematal corded specimens were found between 1500– shields and dorsal shields not conspicuous, pos- 3100 meters a.s.l. and were missing from the low- terolateral tips of peritrematal shields expanded land rainforests of the island. However two spe- posteriorly. Peritremes of general size, expanded cimens of the species described herein were found to anterior half of coxae III, straight or slightly on 585 meters elevation and inhabit the latter bent. Setae r1 and r3 shifted ventrally to peri- vegetation type. trematal shields, both short, bristle-like. A single opening of glands gv2 present. Ventrianal shield MATERIAL AND METHODS bearing 19 setae, setae Zv1 absent. Setae z1 ab- sent. Glands gv3 situated posterolaterally to ad- Mites were extracted using Berlese funnels, anal setae. Third pair of opisthonotal pores as- then cleared with lactic acid and mounted in gly- sociated with the Z-series (gdZ4). Margin of opis-

______*Zsolt Ujvári, Department of Systematic Zoology and Ecology, Eötvös Loránd University, Budapest, H-1117 Pázmány P. sétány 1/c. E-mail: [email protected]

Ujvári: Draconizercon punctatus gen. et sp. nov. from Taiwan

thonotum generally with 7–8 pairs of R-setae. One female: Taiwan, Dabachinshan, 2120 m a.s.l., Opisthomarginal setae of distinctive shape, bent, from soil, 25 March 2004, leg. Huang, K-W apically tapering and serrate. (deposited in NMNS).

Remarks. The new genus is similar to the ge- Measurements. Female. Length of idiosoma: nera Aleksozercon Petrova, 1978, Blaszakzercon 317–328 µm; width: 269–285 µm (n=6). Holo- Kemal & Koçak, 2009, Cosmozercon Błaszak, type: length: 323 µm; width: 280 µm. 1981, Krantzas Błaszak, 1981 and Sellnick, 1943 on the basis of the following fea- Dorsal side (Fig. 1). Podonotum with 22 pairs tures: carapace-like podonotal shield; two short of setae (j2–6, z2–6, s1–6, r2 and r4–5 inserted peritrematal setae; posteriorly expanded peritre- dorsally, setae j1 shifted ventrally, r1 and r3 in- matal shield; peritremes expanded to anterior half serted on peritrematal shields). Setae j3–5 and z5 of coxae III; third pair of podonotal glands smooth and needle-like, z4, z6 and s4–5 basally situated medial to the s-series; third pair of pilose. Among marginal setae s1, z2, s2, r2 and r4 opisthonotal glands associated to the Z-series. The serrate and pointed, others brush-like, plumose. six genera can be distinguished by the combina- Glands gds1 (po1) situated posterior to insertions tion of characters listed in Table 1. of s1; gdj4 (po2) positioned on line connecting j5 and z4, equidistantly; gds4 (po3) lying medial to Etymology. The name of the new genus is s5. Podonotal shield covered by reticulate pattern, composed of the Latin ’draco’, means dragon, posterior surface with a few small, alveolar pits. and the name Zercon. Opisthonotum with 22–23 pairs of setae (J1–5, Z1–5, S1–5, marginal R-series with 7–8 pairs of Gender. Male. setae). Setae J1–5 (Fig. 6) elongate, apically ta- Type species. Draconizercon punctatus sp. pering, pointed, provided with short pili, each nov. reaching far beyond bases of the following one in the series. Setae Z1–4 (Fig. 8) and S2 similar in Draconizercon punctatus sp. nov. shape to J-setae, but shorter; each reaching bases of the following one in the series. Setae Z5 Diagnosis. Central setae of podonotum smooth, straight, pointed, smooth or finely serrate. Setae submarginal and marginal setae pilose. Opistho- S3–5 (Fig. 7) elongate, brush-like, distally pilose, notal J-setae, Z1–4 and S2 elongate, apically expanding beyond margin of idiosoma. Marginal tapering, densely covered by short pili; Z5 setae (Fig. 9) bent, apically tapering and serrate. pointed, smooth or finely serrate; S3–5 brush-like, Size of opisthonotal setae and distances between apically plumose; marginal setae bent, serrate. their insertions as in Table 2. Glands gdZ1 (Po1) Glands gdZ4 (Po3) situated medially to Z4. Dor- anteromedial to Z1; gdS2 (Po2) on line connect- sal cavities small, well-sclerotized. The area bet- ing Z2 and S3; gdZ4 (Po3) medial to insertions of ween J-series covered by large alveolar pits. Z4; gdS5 (Po4) anterior to insertions of R7 (or R8, Sternal setae st2 situated in the central area of the if present). Marginal serration deep, several spines shield, near each other. can be found between setae Z5 (Fig. 5). The area between J-series covered by large alveolar pits, Material examined. Holotype. Taiwan, Hua- other parts of opisthonotal shield smooth. Dorsal lien County, Jhuosi Township, Wa-la-mi, 585 m cavities small, well-sclerotized, with finely undu- a.s.l., from leaf-litter, 05 May 2006, leg. Huang, late inner margin. Axes of cavities slightly con- K-W (deposited in HNHM). Paratypes. One fe- verging anteriorly. male: locality and date as for the holotype (de- Ventral side (Fig. 2). The slit between peritre- posited in HNHM). Three females: Taiwan, Tseu- matal and dorsal shields not conspicuous. Postero- hun lake, 1890 m a.s.l., from leaf-litter 21 October lateral tips of peritrematal shield reaching level of 2004, leg. Huang, K-W (deposited in HNHM). R6, surface of the shield covered by longitudinal

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Ujvári: Draconizercon punctatus gen. et sp. nov. from Taiwan

Figure 1. Dorsal view of Draconizercon punctatus sp. nov.

Figure 2. Ventral view of Draconizercon punctatus sp. nov.

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Ujvári: Draconizercon punctatus gen. et sp. nov. from Taiwan

Figures 3–10. Different structures of Draconizercon punctatus sp. nov. female: 3 = chelicera, 4 = sternal shield of paratype, 5 = posteriormost setae with marginal spines, 6 = seta J1, 7 = seta S5, 8 = seta Z3, 9 = seta R2, 10 = epistome. sutures. Peritrematal setae r1 and r3 short, smooth Gnathosoma. Situation of hypostomal and sub- and pointed. Peritremes straight, with a dilation capitular setae typical for the family. Setae h1–2 near stigmata. Sternal shield (Fig. 4) well sclero- similar in appearance, elongate, needle-like. Setae tized, 45 µm long and 32 µm wide at the level of h3 shorter than h1–2, needle-like, h4 somewhat setae st2, with fine, reticulate ornamentation. longer than previous setae, serrate. Corniculi Setae st2 situated extraordinarily near each other. horn-like, internal malae with a pair of bifurcate Ventrianal setae short, smooth and needle-like, anterocentral branches and with serrate margins. setae Zv1 absent. Setae Jv3–4 1.5 times longer, Fixed digit of chelicerae (Fig. 3) with 5 teeth, adanal setae 2 times longer, postanal seta 3 times movable digit with 4 teeth. Epistome (Fig. 10) of longer than anterior ventrianal setae. Setae Jv5 Prozercon-type. similar in shape and length to marginal setae of opisthonotum. Anal valves with vestigial euanal Male and immature stages unknown. setae. Glands gv3 situated posterolateral to adanal setae. Anterior surface of ventrianal shield cover- Etymology. The name of the new species refers ed by reticulate pattern, posterior surface smooth. to the punctuate opisthonotal ornamentation.

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Ujvári: Draconizercon punctatus gen. et sp. nov. from Taiwan

DISCUSSION 1976, Mesozercon Błaszak, 1976 and Prozercon Sellnick, 1943). The species belonging to this The generic classification of Zerconidae suf- type are generally small. The other type has not fers from several problems. With the appearance been defined yet, but the podonotal shield of these of more and more exotic and special species it has zerconids is not extended ventrally and they become obvious that some of the previously used usually have longer legs in proportion to the body characters are not suitable for generic delimitation length. Latter type has large (Zercon C. L. Koch, or some of them have to be handled with care. 1836) and small (Zeronella Willmann, 1953) The newest results show that most of the cha- representatives as well. Draconizercon gen. nov. racters previously kept discrete (e.g. shape of belongs to the first group possessing carapace-like peritrematal setae and shields) have several inter- podonotal shield. mediate stages, others show intraspecific variati- on, which makes them difficult to apply. The case The second character chosen was the number of genus Prozercon Sellnick 1943 serves as a and general appearance of setae shifted to peritre- perfect example: due to the intensive research of matal shields. Those genera which possess three the last decades (e.g. Mašán & Fenďa, 2004; pairs of setae on peritrematal shields are most Urhan, 2008; Ujvári, 2011e) the genus became the probably closely related to each other, constitute- second largest among Zerconidae, with more than ing a separate evolutionary lineage within the 50 species. With the discovery of several in- family. The unique, hipertrichous Syskenozercon teresting species the diagnosis of the genus was Athias-Henriot, 1977 represents also a separate, changed from time to time, some restrictions have probably the most ancient linage of Zerconidae. been erased, new observations have been added, However the majority of zerconid mites possess but it is almost sure that neither the latest two pairs of peritrematal setae (r1 and r3). The diagnosis given is perfect (Ujvári, 2011e). shape of peritrematal setae shows a large degree of variation and has to be handled with care Another problem is, however the new diag- (Ujvári, 2010, 2011e). Setae r1 are always re- nosis of Prozercon is in accordance with the latest latively short, and apart from some exceptions observations, most of the genera of the family (e.g. Echinozercon Błaszak, 1976) are always have not been dealt with. It is obvious that the shorter than or as long as setae r3. To reduce the whole family badly needs an overall revision. mutual character stages of r1 and r3 I considered only the relative size of them: both r1 and r3 short Because of these problems I had much trouble or r3 significantly longer than r1. Draconizercon with the insertion of the new species described gen. nov. belongs to the group possessing two above into the existing Zerconidae system there- short peritrematal setae. fore, it was necessary to establish a new genus Draconizercon gen. nov. to accommodate it. As the third character I have chosen the shape of peritrematal shields. Nowadays we recognize I tried to choose the most reliable characters several different stages: in some groups these during the classification of the new species. There shields end truncate, in most of the cases posteri- is a well-recognizable character regarding the orly separated from podonotal shields (e.g. Zercon shape of the body which is not easy to define. C. L. Koch, 1836); in other cases the postero- There are at least two different types of Zerco- lateral tips of these shields are expanded pos- nidae on the basis of the body shape. One of these teriorly, in a different degree (e.g. Prozercon possesses a carapace-like podonotal shield, which Sellnick, 1943). Rarely can we see special scle- is extended anteroventrally and lateroventrally rotized areas next to the ventrianal shield which (Lindquist & Moraza, 1998) and have relatively are usually called ventrilateral shields. The short legs in proportion to the body length (e.g. originof these sclerotized areas is not clear, but Aspar Halašková, 1977, Echinozercon Błaszak, presumably is different in different groups. In Ma-

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Ujvári: Draconizercon punctatus gen. et sp. nov. from Taiwan

crozercon Błaszak, 1975 it seems to be an expan- near r4 in former genera) suggests that Draconi- sion of the ventrianal shield, in Rotundozercon zercon gen. nov. does not belong to this group Ujvári, 2011 it seems if the dorsal sclerotization (besides Draconizercon gen. nov. possesses would shifted ventrally, but the ventrilateral epistome of Prozercon-type, while the other East shields in Krantzas Błaszak, 1981 may have been Asian genera have epistome of Parazercon-type). detached from the peritrematal shields. To reduce the number of stages of peritrematal shields’ Close phylogenetic relationship between Dra- shape I considered only the relative ending of it: conizercon gen. nov. and some Nearctic genera can also be excluded by considering the situation posteriorly truncate or expanded (including of the third pair of opisthonotal pores. There is an Krantzas as well, with its special ventrilateral endemic Nearctic group of genera possessing the shields). Draconizercon gen. nov. belongs to the third pair of opisthonotal glands in association group possessing posteriorly expanded peritrema- with the J-series (e. g. Bakeras Błaszak, 1984, tal shields. Blaszakiella Sikora & Skoracki, 2008 and Mic- rozercon Błaszak, 1975). This unique phenome- The fourth major character chosen was the size non can not be observed on any Palearctic spe- of peritremes. Three stages can be recognized. cies, the third pair of opisthonotal glands of latter The majority of species possess peritremes reach- are always associated to the Z-series. ing the anterior margin of coxae III. Some groups have reduced peritremes reaching only the poste- Applying these restrictions only five genera rior margin of coxae III (Skeironozercon Halaš- remained as a possibility to classify the new ková, 1977, Syskenozercon Athias-Henriot, 1977 Taiwanese species with. Two of these (Prozercon and Zerconella Willmann, 1953), other groups possess elongate peritremes reaching level of Sellnick, 1943 and Aleksozercon Petrova, 1978) coxae II (Carpathozercon Balan, 1991 and Echi- are distributed in the Western Palearctic region, nozercon Błaszak, 1976). No matter which of the other three (Blaszakzercon Kemal et Koçak, these three stages is the ancient one, it seems to be 2009, Cosmozercon Błaszak, 1981 and Krantzas sure that each derived stages are formed in- Błaszak, 1981) are Nearctic genera. If we con- dependently in different groups, as a result of sider the special shape of opisthomarginal setae as convergent evolution. Draconizercon gen. nov. a mark of phylogenetic relationship, the new belongs to the largest group possessing peritremes species can only be related to Cosmozercon reaching anterior margin of coxae III. Błaszak, 1981 and Krantzas Błaszak, 1981. This is also supported by the distribution pattern of With the discovery of more and more new these genera: the eastern distribution border of species in the Nearctic and Eastern Palearctic Prozercon Sellnick, 1943 lies in Mongolia (Bła- regions the significance of dorsal adenotaxy is szak, 1979), the single species of Aleksozercon becoming increasingly conspicuous in systematics Petrova, 1978 is known only from the north- of Zerconidae. The special position of proper eastern shores of the Black Sea (Petrova, 1978), dorsal pores most probably reflects phylogenetic while Blaszakzercon Kemal & Koçak, 2009 relationship reliably, which is also supported by seems to be endemic in the Appalachian Moun- the distribution pattern of the morphologically tains (Ujvári, 2012). Therefore, apparently there is similar genera. I chose the position of the third no sign of possible biogeographical connection pair of podonotal and the third pair of opistho- notal glands. There is a group of genera in East between the former genera and the new Tai- Asia the members of which (Aquilonozercon wanese species as well. On the contrary, Cos- Halašková, 1979, Eurozercon Halašková, 1979, mozercon Błaszak, 1981 and Krantzas Błaszak, Kaikiozercon Halašková, 1979, Koreozercon Ha- 1981 are distributed on the Pacific Coast of North lašková, 1979 and Mesozercon Błaszak, 1975) are America. Asia and Western North America were very similar to the newly established genus connected by Trans-Beringian land bridges seve- Draconizercon gen. nov., but the position of the ral times from the Mid Creataceous, which served third podonotal glands (situated lateral to s-series, as an important dispersal route for different ani-

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Ujvári: Draconizercon punctatus gen. et sp. nov. from Taiwan

mal groups (Sanmartín et al. 2001). Besides, with Acknowledgements. Dr. Kun-Wei Huang is gratefully the drop of sea level several other land bridges acknowledged for lending mesostigmatid mite specimens may have emerged in the Northern Pacific region, from Taiwan. especially through the recent island arcs of the Pacific Plate (Aleutian Islands, Kuril Islands, REFERENCES Japanese Islands etc.), resulting in other possible dispersal routes. These paleogeographic connec- BŁASZAK, C. (1979): Systematic studies on the family tions might explain the relationship between the Zerconidae IV. Asian Zerconidae (Acari, Mesosti- mata). Acta Zoologica Cracoviensia, 24: 3–112. fauna of the Western Nearctic and the Eastern Palearctic and support the hypothesis of the com- JOHNSTON, D. E. & MORAZA, M. L. (1991): The mon origin of Draconizercon gen. nov., Cosmo- idiosomal adenotaxy and poroidotaxy of Zerconi- dae (Mesostigmata: Zerconina). In: DUSBÁBEK, F. zercon and Krantzas. & BUKVA, V. (eds): Modern Acarology. Vol. 2., Academia, Prague, pp. 349–356. The presence or absence of adgenital glands gv2 is applied with a great emphasis, and (so far) LINDQUIST, E. E. & EVANS, G. O. (1965): Taxonomic concepts in the Ascidae, with a modified setal seems to be a very useful and stable character in nomenclature for the idiosoma of the Gamasina classification of Zerconidae. Both Cosmozercon (Acarina: Mesostigmata). Memoirs of the Entomo- Błaszak, 1981 and Krantzas Błaszak, 1981 lack logical Society of Canada, 47: 1–64. these glands, this is the main reason why I de- LINDQUIST, E. E. & MORAZA, M. L. (1998): Observa- cided to establish a new genus for the new Tai- tions on homologies of idiosomal setae in Zerconi- wanese species. dae (Acari: Mesostigmata), with modified notation for some posterior body setae. Acarologia, 39: The presence or absence of ventrianal setae 203–226. Zv1 is a character I did not take into account with MA, L. M., HO, C. C. & WANG, S. C. (2011): One new great relevance. It seems that the presence of Zv1 species of Zerconidae and one new recorded spe- is a plesiomorphic stage within Zerconidae. Dur- cies of Blattisocidae from Taiwan (Acari: Meso- ing the evolutionary history, many groups might stigmata). Formosan Entomologist, 31: 157–165. lost these setae independently from each other, as a result of convergent evolution. This can be MAŠÁN, P. & FENĎA, P. (2004): Zerconid mites of Slovakia (Acari, Mesostigmata, Zerconidae). NOI proven by examples selected from genus Zercon Press, Bratislava, 238 pp. C. L. Koch, 1836. There are different, closely re- lated groups (from external morphological point PETROVA, A. D. (1978): A new genus and species of of view) within Zercon, the members of which peculiar zerconid-mites (Parasitiformes, Gamasoi- differ by the presence of Zv1. These hypothetical dea, Zerconidae) with entire dorsal shield. Revue groups represent different branches within the d’Entomologie de l’URSS, 57: 218–220. evolutionary tree of Zercon. Presumably these SANMARTÍN, I, ENGHOFF, H. & RONQUIST, F. (2001): groups diverged much earlier than the separation Patterns of dispersal, vicariance and diversi- inside a group has taken place by losing Zv1. fication in the Holarctic. Biological Journal of the Therefore losing setae Zv1 can be considered as a Linnean Society, 73: 345–390. frequent phenomenon, which usually occurs in- UJVÁRI, ZS. (2010): Zerconella Willmann, 1953, a dependently during speciation processes in dif- forgotten group of Zerconidae (Acari, Mesostigma- ferent groups of Zerconidae. Hence it does not ta). Zootaxa, 2558: 33–47. seem to be a good character for generic deli- UJVÁRI, ZS. (2011a): New Zerconid mites (Acari: mitation. Excluding species with lacking Zv1 Mesostigmata: Zerconidae) from Taiwan. Zoologi- from a genus usually possessing Zv1 may results cal Studies, 50 (1): 87–102. a morphological classification which does not UJVÁRI, ZS. (2011b): First records of Zerconidae correspond to the evolutionary history, puts (Acari: Mesostigmata) south of the Tropic of proper, closely related species into separate ge- Cancer, Mexico, with description of five new spe- nera, while other, more distant relatives would cies. International Journal of Acarology, 37 (3): remain in the same genus. 201–215.

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UJVÁRI, ZS. (2011c): A new subgenus and two new UJVÁRI, ZS. (2012): Review of the American genera species of Zercon C. L. Koch, 1836 (Acari: Zerco- Aspar Halašková, 1977 and Blaszakzercon Kemal nidae) from Southeast Asia. Zootaxa, 2995: 45–54. et Koçak, 2009 (Acari, Mesostigmata, Zerconidae) UJVÁRI, ZS. (2011d): Comparative study on the with description of three new species. Acta Zoo- taxonomic relevance of gnathosomal structures in logica Academiae Scientiarum Hungaricae, in the family Zerconidae (Acari: Mesostigmata). press. Opuscula Zoologica Budapest, 42 (1): 75–93. URHAN, R. (2008): Contributions to the genus Pro- UJVÁRI, ZS. (2011e): Six new species of Prozercon zercon Sellnick, 1943 (Acari: Zerconidae) from Sellnick, 1943 (Acari, Mesostigmata, Zerconidae) Turkey, with the description of two new species from Greece, with remarks on the genus. Zootaxa, and a key to species. Zoology in the Middle East, 2785: 1–31. 45: 97–104.

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