Acari: Oribatida) of Georgia

New Carabodidae (Acari: Oribatida) of Georgia

Maka Murvanidze & Gerd Weigmann

New samples from different regions of Georgia contribute to the knowledge of Carabodidae species in this region. Austrocarabodes foliaceisetus georgiensis subsp. n. and Carabodes djaparidzae sp. n., are described as new taxa for the Georgian fauna. A regional form of the Nearctic Carabodes granulatus Banks, 1895 is described. It is the first Palaearctic record of the species, collected in ��litter and soil of Carpinus-Fagus forest in Western Georgia�. A. f. georgiensis subsp. n. occurs in a semi-desert region of Eastern Georgia, C. djaparidzae sp. n. was found in a Carpinus forest in Eastern Georgia. The characters of the species are discussed and compared with those of similar species in these genera. M. Murvanidze *, Georgia LEPL Institute of Zoology, Chavchavadze av. 31, 0079 Tiblisi, Georgia. [email protected] G. Weigmann, Institute for Zoology, Freie Universitaet Berlin, Koenigin Luise Str. 1-3, D-14195 Berlin, Germany. [email protected]

Introduction Material and methods The Carabodidae is a large and globally distri- The material was collected by soil samples; the mites buted family of oribatid mites with about 34 gen- were extracted by use of modified Berlese-funnels. era (Mahunka 1986, 1987) and many species. Three The animals were stored in ethanol until preparation genera of Carabodidae have been recorded from and were studied after clearance in lactic acid in open Georgia: Austrocarabodes Hammer, 1966, Carabodes hollow-ground microscope slides covered partly with C. L. Koch, 1835 and Odontocepheus Berlese, 1913, a cover slip, which allows to view the animal in dif- with 17 species in total (see Shtanchaeva 2001). ferent positions. Permanent slide preparations are Research carried out recently in different regions of mounted in modified Berlese-fluid. Georgia provided additional knowledge on the tax- The terminology of morphological structures follows onomy and distribution of these mites (Weigmann & van der Hammen (1980) and Weigmann (2006). Murvanidze 2003). In this paper we describe a new species ( sp. n.) and a new sub- Carabodes djaparidzae Austrocarabodes foliaceisetus georgiensis species (Austrocarabodes foliaceisetus georgiensis subsp. subsp. n. n.) and additionally, a regional form of the Nearctic Figs 1–5 Carabodes granulatus Banks, 1895. This paper con- tributes to a revision of the regional fauna of Georgia, Type material�. Holotype /: Georgia: Eastern Geor- a region which links three Palaearctic biogeographi- gia, soil of river Pantishara gorge (at the border to cal regions (Caucasian region, partly alpine – Turkish Azerbaijan), N�� 41°08′30,98′′, E 46°38′24,25′′; and Iranian regions, subtropic to tropic) and, there- altitude 262 m.�� Collected�� by A. Bukhnikashvili, fore, is of high biogeographical interest. This revision 2.vii.2003. Material in permanent slide deposited in includes critical discussion of earlier descriptions of collections of Georgia LEPL Institute of Zoology. Caucasian species and of Georgian records of species described from other regions originally. Diagnosis Notogaster structured with clear tubercles, sculpture of prodorsum formed by irregular ridges, the one of the ventral plate by bright foveolae. Interlamellar

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1 2 3

4 ad3 5

Figs 1–5. Austrocarabodes foliaceisetus georgiensis subsp. n. – 1, Dorsal aspect; 2, genital plates; 3, sensillus; 4, ventral aspect of anal region; 5, Notogastral seta lm and scupture. Scales: at 1: 100 μm; at 5: 100 μm, for 2–5.

setae long phylliform. Sensilli bacilliform, bent up- anal formula: 4g–1ag–2an–3ad. All three pairs of wards, distally granulated. 14 pairs of notogastral se- adanal setae phylliform, widened and dentate. tae phylliform and dentate. All adanal and anal setae Legs. With 1 claw. phylliform. Ecology Description The semi desert vegetation is dominated by the tree General characters. Colour yellowish-brown. Body Pistacia mutica F. et M. length 540 μm, width 295 μm. Length/width- index 1.8. Discussion Prodorsum. Sculpturing with fine pattern of irregu- Three species of Austrocarabodes have been recorded lar ridges. Lamellae rounded. Rostral setae strong, (Shtanchaeva 2001) from the Caucasus area: A. folia- thick bacilliform, dentate, curved inside. Lamel- ceisetus Krivolutski, 1969, A. ensifer (Sellnick, 1931) lar setae situated on the anterior edge of lamellae, and A. arrogans Perez-Iñigo, 1967. thick, strong, barbed. Interlamellar setae the longest The new subspecies resembles most of all A. folia- prodorsal setae, phylliform, with dentate edges. The ceisetus, firstly described from Kirgizia (Krivolutski sensillus bacilliform and pointed, curved upwards, 1971), but there are some evident differences. (1) In distal part slightly granulated, about 90 μm long the original description of A. foliaceisetus, its length is (Fig. 1, 3). indicated as 602 μm; in the identification keys edited Notogaster. Cuticle covered with clear tubercles. by Ghilarov & Krivolutski (1975) the length is given Dorsosejugal line nearly straight. On the shoulders as 610 μm; both records show a slender body size (in strongly chitinized, triangular tubercles present. contrast to all cited European species) with a length/ With 14 pairs of long, phylliform, dentate notogas- width-index of 2–2.2. The new subspecies has a tral setae, length about 60 μm (Fig. 1). body length of 540 μm and a length/width-index of Ventral side. Sculpture of ventral side formed by large 1.8. (2) The sculpture of the body surface of A. folia- bright foveolae. Epimeral formula 3–1–3–3. Genital- ceisetus is formed by well developed rough tubercles

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6 7 8

ad3

Figs 6–9. Austrocarabodes ensifer. – 6, Dorsal aspect; 7, sensillus; 8, notogastral seta la; 9, ventral aspect of anal region. Scales: at 6: 100 μm; at 8: 100 μm, for 7–9. and ridges; the Georgian subspecies has a sculpture proportions to A. f. georgiensis subsp. n. The sculp- with fine clear tubercles on the notogaster and ir- ture of the prodorsum is formed by roundish foveo- regular ridges on the prodorsum. (3) The sculpture lae. The sensilli are with 50–55 μm length shorter of the ventral side is quite different: A. foliaceisetus and have a distincly thickened head (��Fig��. 7). Accord- has well developed net-pattern on the whole ventral ing to Ghilarov & Krivolutski (1975), A. ensifer has region (similar to that of A. ensifer); on the contrary, rounded shoulders and no tubercles, but Perez-Iñigo A. f. georgiensis subsp. n. has weakly developed sculp- (1971, 1997) illustrated chitinized triangular tu- ture formed by big light foveolae. (4) According to bercles. This character is also shown by recent find- the original description, the shoulder curvature of ings of A. ensifer in Portugal by one of the authors A. foliaceisetus is weakly developed and lacks any (G. W.), which are in concordance with the rede- triangular tubercle, which is present in A. f. geor- scription of Mahunka (1986) from Greece, where giensis subsp. n., as in A. ensifer and other species. the type locality is situated. Strong differential char- Future findings will allow to assess the variability of acters of A. ensifer are the thin anal setae and adanal the differentiating characters. For the present, we setae ad3, whilst the same are phylliform in both sub- avoid to declare it a new species. We think it more species of A. foliaceisetus. The ventral plate has a net- probable that the citations from the Caucasus region like structure. For reasons of comparison we present (Shtanchaeva 2001) concern the new subspecies than figures of A. ensifer from South Portugal (Figs 6–9). the Kirgizian nominal subspecies. A. ensifer is distributed in Europe from the Iberian With a body length of 495–565 μm and a length/ region through the Mediterranean area to the Cau- width-index of 1.8, A. ensifer is similar in size and casus region; the references in Shtanchaeva (2001)

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regarding A. ensifer might be based on material of Ventral side. Sculpture with rosette-like foveolae. A. f. georgiensis subsp. n., indeed. But the material Epimeral formula 3–1–2–3, all setae are smooth, was not available for us. pointed. Genital-anal�� formula 4g��–��1ag–��2a–��3ad. The species A. arrogans, originally described from Genital setae short, about 5 μm, smooth, thin; ag Spain, is 500–550 μ��m in length and broader than setae 7 μm, smooth, thin. ad1, ad2 phylliform, barbed A. f. georgiensis subsp. n. (length/width-index of 1.6, about 40 μm. ad3 thin and smooth (Fig. 14). measured behind shoulders), it has distinctly pro- Legs. With 1 claw. truding shoulder curvations in contrast to A. ensifer and A. foliaceisetus; all adanal setae are leaf-like, too, Material examined. Georgia: 1/, Western Georgia: but the anal setae an are fine. The sensillus of A. arro- Surami Range, N 42°07′21.88′′; E 43°33′18.74′′. ��Col- gans is setiform and about 90–95 ��μm�� long (cf. Perez- lected by M. Murvanidze, 6.vii.2005, ��in litter and soil of Iñigo 1971, 1997), as in A. f. georgiensis subsp. n. Carpinus-Fagus forest with individual Alnus and Castanea. Understorey with Struthiopteris filicastrum ��All. ��and Hedera From Spain also A. intermedius Ruiz et al., 1989, is colchica (K. Koch) K. Koch��. ��Material in permanent slide known, which is very large (540–660 μm length) deposited in collections of Georgia LEPL Institute of and slightly broader than A. f. georgiensis�� subsp. n. Zoology. (length/width-index 1.7), it has fine ad3 and an setae, as A. ensifer; the sensillus is unique: setiform with fine setulae in the distal part (cf. Perez-Iñigo 1997). Discussion The species is widely distributed in Northern Amer- ica, but has not yet been found in the Palaearctic re- Carabodes cf. granulatus Banks gion. Its typical habitat is forest litter, rotten wood Figs 10–14 and soil, preferably with lichens and moss (Reeves Carabodes granulatus Banks, 1895: 129. 1988). The Georgian species shows evident similari- Carabodes omo Jacot, 1937: 241. Type data (synonymised ties with C. granulatus in shape of sensilli, interla- by Norton 1978) mellar and notogastral setae, sculpture of notogaster Carabodes granulatus; Norton 1978: 614 (redescription) and shape of adanal setae, as far as the indications of the American authors are given. The identification Diagnosis key of Reeves & Behan-Pelletier (1998) of the North Anterior part of prodorsum widely foveolate, poste- American species leads to C. granulatus; a similar Eu- rior part with larger foveolae and chitinized longitu- ropean species is not known to us. dinal ridges; dorsosejugal groove narrow. Interlamel- Nevertheless, there are some differences: (1) the la- lar and notogastral setae are phylliform and barbed; mellar setae of C. granulatus in the SEM picture in rostral and lamellar setae setiform to spiniform; head Reeves & Behan-Pelletier (1998) looks as thin as the of sensilli finger-like divided. Sculpture of notogaster rostral setae; yet, Jacot (1937) indicates for C. omo: and ventral plate with rosette-like foveolae. Adanal a body length of 430 μm and “rostral and lamellar setae ad1, ad2 phylliform, ad3 fine and short. bristles stout, spinelike, faintly burred”, which seems to fit to the setae in our ��Fig��. 10. (2) The lengths of Description the interlamellar and notogastral setae of C. granula- General characters. ��Colour reddish-brown. Body tus (��Fig��. 41 in Reeves & Behan-Pelletier 1998) seem length 415 μm, width 213 μm. to be longer than in the Georgian specimen; still, the Prodorsum. Sculpture in the anterior part of Fig��. 11 in Norton (1978) gives the same size range prodorsum formed by wide-set arranged foveolae (20–30μm) for the interlamellar and notogastral and in posterior part by larger foveolae and irregular setae. (3) Reeves & Behan-Pelletier (1998) indicate longitudinal ridges. Lamellae broad. Rostral setae unequal adanal setae for C. granulatus (without fur- nearly smooth, curved inside; lamellar setae thick ther details); Jacot (1937) declares all ventral setae setiform curved and barbed. Interlamellar setae as “short, except ad2 and ad3, the latter being half situated near to the lamellae, phylliform, barbed, length of notogastral bristles, and weakly clavate- about 30 μm long. Sensilli about 40 μm long, fimbriate”. It seems to us that the numbering of an- with a short stalk and a distally finger-shaped setae by Jacot is the opposite of the one given in the head. Prodorsum with narrow dorsosejugal groove contemporary literature, and he may have described (Fig. 10). the setation as in our ��Fig��. 14. �Y��et, the Georgian spec- Notogaster. Sculpture formed by bright rosette-like imen has longer adanal setae than notogastral setae; foveolae surrounded by dark cuticula surface, with Jacot describes the opposite characteristic. There is fine granula (Fig. 13). With 10 pairs of phylliform, no certainty of the specific identity with C. granula- barbed notogastral setae, length 20–30 μm (Fig. 12). tus, but the likelyhood is high.

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10 11 Figs 10–14. Carabodes cf. granulatus. – 10, Dorsal aspect; 11, Sensillus; 12, Noto- gastral seta lm; 13, Noto- gastral pattern; 14, Ventral aspect of anal region. Scales: at 10: 100 μm, for 10, 14; at 11: 50 μm, for 11–13. 12 13

ad3

One of the diagnostic characters of the species are collected by ��T. Arabuli, 22.iv.2002. Paratypes: 2/ the phylliform interlamellar and notogastral setae. 1?, same data as Holotype; 2�?, Georgia, Great C. bellus Aoki, 1959, C. chirstlus Mahunka, 1987, Caucasus Range, Omalo Plateau;�� N 42°15′45.19′′, and C. foliatus Morell, 1990, have similar setae. Dif- E 45°13′01.11′′, altitude 1765 m; collected by ferences are as follows. E. Kvavadze, 7.viii.2004. Type material stored in With 550–660 μm, C. bellus is much longer than glycerine, deposited in collections of Georgia LEPL C. granulatus. Posterior part of prodorsum strongly Institute of Zoology. chitinized, notogastral sculpture is made with tubercles (Aoki 1959). Diagnosis With 500–566 μm, C. chirstlus is also longer; with Notogaster with “flower-shaped” pattern formed by long, fusiform, bent sensillus, large dorsosejugal bright foveolae surrounded by chitinized tubercles. groove at prodorsum, and notogastral sculpture with Interlamellar setae long, curved inside. Sensillus polygonal pattern and tubercles (Mahunka 1987). head distally finger-shaped. 10 pairs of notogastral Carabodes foliatus differs in non phylliform setae. Setae c2 long, strong, erect; other notogastral notogastral setae of the p-row, a different form of setae and adanal setae ad1 and ad2 phylliform, dis- sensillus and special apophyses in the dorsosejugal tally widened and barbed. region�� (cf. Perez-Iñigo 1997). Description General characters. ��Colour reddish-brown. Body�� Carabodes djaparidzae sp. n. length 510 μm, width 260 μm. Fig. 15-18 Prodorsum. Sculpturing of prodorsum is formed with Type material. Holotype /, Georgia: Gombori bright foveolae and irregular longitudinal ridges. The Range, Shuamta; N ��42°04′42,17′′, E 45°32′38,58′'; dorsosejugal groove is moderately broad. Lamellae

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15 16

19 17

ad3

Figs 15–19. Carabodes djaparidzae sp. n. – 15, Dorsal aspect; 16, notogastral pattern and seta lm; 17, Notogas- tral seta p1; 18, ventral aspect of anal region; 19, Carabodes dubius: notogastral pattern and seta lm. Scales: at 15: 100 μm; at 16: 50 μm, for 16, 17, 19; at 18: 100 μm.

rounded. Rostral and lamellar setae are curved in- Ecology side, slightly granulated. Interlamellar setae are 90 Shuamta: in soil of a Carpinus forest; Omalo Plateau: µm long, strong setiform, smooth, slightly curved moss in Populus tremula-Pinus-forest; understorey inside and positioned on the lamellae. The sensilli with different grasses. are directed laterally, head distally finger-shaped (Fig��. 15). Etymology Notogaster. With “flower-shaped” pattern formed by The name C. djaparidzae sp. n. is dedicated to the first bright foveolae surrounded by chitinized pavement- oribatidologist of Georgia, Dr. N. I. Djaparidze. like tubercles between which the bright foveolae seem interconnected by bright lines (Fig��. 16). With Discussion 10 pair of notogastral setae; c2 setae are about 85 Based on the descriptions of two new species, Kuli- μm long, strong, directed forward. Other setae are jev (1977) created the new genus Flexa and desig- 38–50 μm long, phylliform, distally widened and nated Carabodes dubius Kuliev, 1968 as type species. barbed, especially at the distal edge (Fig��. 15, 17). Mahunka (1987) considered the differentiating Ventral side. Epimeral formula 3–1–3–3. Genital- characters of minor taxonomic value with high vari- anal formula 4g–1ag–2an–3ad; ad1 and ad2 are ability (shape of shoulder and of femora III and IV; distally widened and barbed, ad3 and an setae are orientation of notogastral setae c2), and he sank Flexa smooth setiform (Fig 18). again as a junior synonym of the genus Carabodes. Legs. With 1 claw. We agree with the statement of Mahunka and regard Flexa as a junior synonym of Carabodes.

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C. djaparidzae sp. n. belongs to the species group alpine meadows of Switzerland (Schweizer 1956), of “Flexa”type Carabodes with long forwarded c2 and in the Italian and Austrian Alps. The notogastral setae. Five additional species of “Flexa”type Cara- ornamentation is similar to that of C. djaparidzae sp. bodes are recorded from the Caucasian countries: n. and C. tarbae, as redescribed by Weigmann (2006). Carabodes dubius, C. scopulae Kuliev, 1968, C. bidens The notogastral setae are about 50 μm in length and Djaparidze, 1990, C. horreo Djaparidze, 1990 and are setiform to bacilliform, at most slightly widened, C. tarbae Shtanchaeva, 2004 (Djaparidze 1990; sparsely barbed. The adanal setae ad1 and ad2 are Shtanchaeva 2001, 2004). strong setiform, ad3 and the anal setae are fine (cf. The notogastral�� setae of C. dubius are fusiform and redescriptions of Bernini & Baratti 1991; Weigmann barbed all around (cf. Shtanchaeva 2004) with mi- 2006). The redescription of “C. intermedius” from nor length about 25–30 μm. This diagnostic char- the Caucasus region of Georgia in Murvanidze & acter is evaluated together with recent findings Arabuli (2003) refers to C. djaparidzae sp. n. in Georgia (Surami Range, mixed forest soil; leg. M. Murvanidze) and is illustrated in ��Fig��. 19, which is in conformity with the original description and Acknowledgements the redescription of Shtanchaeva (2004). The noto- M. Murvanidze wishes to express her gratitude to gastral setae in C. djaparidzae sp. n. are quite differ- the German Academic Exchange Service (DAAD) ent: flattened leaf-like, distally widened and sparsely for giving the opportunity to conduct research in barbed (Fig��. 17). The redescription of “C. dubius” Germany. She also wishes to thank Dr. E. Kvavadze, by Mahunka (1987) shows notogastral setae as being T. Arabuli and A. Bukhnikashvili for sampling of the typical for C. djaparidzae sp. n.; thus, we suppose material in different regions of Georgia. that his redescription does not regard C. dubius. C. scopulae has distally widened notogastral setae, similar to those of C. djaparidzae sp. n., but the References sculpturing of the prodorsum and the notogaster are Aoki, J.-I., 1959. Zur Kenntnis der Oribatiden im Pilz. I. quite different. The posterior part of prodorsum of Berichte über einige Arten aus Nikko. – Annotationes C. scopulae is sculptured with fine longitudinal ridg- zoologicae Japonenses 32: 156–161. es. The anterior border of the notogaster is rounded Bernini, F. & M. Baratti, 1991. Notulae Oribatologicae forwards and makes the dorsosejugal groove very LIII. Carabodes intermedius Willmann, 1951 (Acarida, narrow. The notogastral pattern is characterized by Oribatida) new findings in Italy. – Redia 74: 211–223. bowed tubercles, which form rosette-like structures Djaparidze, N. I., 1990. Two new species of mites (Orib- atei) of genus Flexa Kuliev, 1977 in Georgian fauna. isolated from each other (cf. Shtanchaeva 2004). – Bulletin of the Academy of Sciences of the Georgian C. bidens differs in bacilliform notogastral setae, SSR 137: 405–408. [in Russian] slightly widened distally, if at all. The sculpture of Ghilarov M. S. & D. A. Krivolutski, 1975: Opredelitel the posterior part of the prodorsum is formed by Obitaiushchikh v Pochve Kleshchei. Sarcoptiformes. several longitudinal ridges. The notogastral setae c2 (The identification keys of soil inhabiting mites. Sar- of C. bidens are distinctly longer (120 μm) than the coptiformes). – Nauka, Moscow, 491 pp. [in Russian] ones of C. djaparidzae sp. n. (85 μm) (cf. Djaparidze Hammen, L. van der, 1980. Glossary of acarological termi- 1990). nology. Vol. I: General terminology. – W. Junk., The Sculpture of prodorsum of C. horreo is formed by Hague, 244 p. strongly chitinized longitudinal ridges at the poste- Jacot, A. P., 1937. New moss-mites, chiefly Midwestern. II. rior part and transversal ridges on the anterior part. – American Midland Naturalist 18: 237–250. The sculpture of the notogaster is formed by bright Krivolutsky, D. A., 1971. Some new oribatoid mites from foveolae, diffuse and indistinct rosette-like pattern. Eastern Kirgisia. – Zoologicheskij Zhurnal 50: 939– The notogastral setae are thick, bacilliform and short 942. [in Russian] Kuliev, K. A., 1977. Flexa Kulijev nov. gen. typovoi Cara- (28–35 μm) (Djaparidze 1990). bodes dubius Kulijev, 1968 semeistvo Carabodidae The sculpturing of prodorsum and notogaster in C. L. Koch, 1837. – Doklady Akademii Nauk Azerbai- C. tarbae is similar to that in C. djaparidzae sp. n. jan SSR 33: 64–67. [in Russian] The notogastral setae of C. tarbae are variable in Mahunka, S., 1986. A survey of the family Carabodidae shape: from smooth and pointed in the centrodorsal C.L. Koch, 1836 (Acari: Oribatida). – Acta zoologica region to slightly widened bacilliform and barbed in Academiae Scientiarum Hungaricae 32: 73–135. the posterio-marginal region (Shtanchaeva 2004). Mahunka, S., 1987. A survey of the family Carabodidae All listed species were found in the mountains of the C.L. Koch, 1836 (Acari: Oribatida), II. – Acta zoolog- Caucasus. An additional species of the “Flexa-type” ica Academiae Scientiarum Hungaricae 33: 399–434. is C. intermedius Willmann, 1951, which occurs in Murvanidze, M. & T. Arabuli, 2003. Carabodes intermedius Willmann, 1951 (Acari, Oribatida, Carabodidae), Downloaded from Brill.com09/25/2021 03:19:43PM via free access 200 Tijdschrift voor Entomologie, volume 150, 2007

a new species for the Caucasus fauna. – Bulletin of the Schweizer, J., 1956. Die Landmilben des Schweizerischen Georgian Academy of Sciences 168: 112–113. Nationalparkes. 3. Teil: Sarcoptiformes Reuter 1909. Norton, R. A., 1978. Notes on Nathan Banks’ species of – Ergebnisse der wissenschaftlichen Untersuchun- the mite genus Carabodes (Acari: Oribatei). – Proceed- gen des schweizerischen Nationalparks (N. F.) 5: ings of the Entomological Society of Washington 80: 215–377. 611–615. Shtanchaeva, U. Y., 2001. Catalog of oribatid mites (Acari- Perez-Iñigo, C., 1971. Acaros Oribatidos de Suelos de formes, Oribatida) of the Caucasus. – Acarina 9: Espana peninsular e Islas Baleares (Acari, Oribatei). 177–221. Parte 3. – Eos, Revista Espaniola de Entomologia 46: Shtanchaeva, U. Y., 2004. Flexa is a mountain genus of 263–350. oribatid mites (Oribatida, Carabodidae). – Zoolog- Perez- Iñigo, C., 1997. Acari, Oribatei, Gymnonota I. icheskij Zhurnal 83: 679–692. – Fauna Iberica 9: 1–374. Weigmann G. & M. Murvanidze, 2003. Contribution to Reeves, R. M., 1988. Distribution and habitat compari- the fauna of oribatid mites of Georgia. 2. Carabodes sons for Carabodes collected from Conifer branches and Lamellocepheus. – Spixiana 26: 221–226. with descriptions of brevis Banks and higginsi n. sp. Weigmann, G., 2006. Hornmilben (Oribatida). – Die (Acari: Oribatida: Carabodidae). – Proceedings of the Tierwelt Deutschlands 76: 1–520. Entomological Society of Washington 90: 373–392. Reeves, R.M. & V. Behan-Pelletier, 1998. The genus Cara- bodes (Acari: Oribatida: Carabodidae) of North Amer- ica, with descriptions of new western species. – Cana- Received: 20 December 2006 dian Journal of Zoology 76: 1898–1921. Accepted: 7 February 2007

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