Ecologica Montenegrina 10: 71-87 (2017) This journal is available online at: www.biotaxa.org/em
The Halacarellus capuzinus group (Halacaridae: Acari), diagnoses, keys and a description of Halacarellus floridearum (Lohmann) ILSE BARTSCH Forschungsinstitut Senckenberg, c/o DESY, Notkestr. 85, 22607 Hamburg, Germany. E-mail:
[email protected] Received: 24 March 2017 │ Accepted by V. Pešić: 14 April 2017 │ Published online: 19 April 2017.
Abstract The genus Halacarellus, with its presently known 50 species, includes several species groups. The numerically most important one is the Halacarellus capuzinus group. The capuzinus group is diagnosed. Of each of the 10 species morphological characters are outlined, together with their geographical and ecological data. The number of setae on the leg segments proved to vary, variants are listed in a table. In addition to a tabular key, with the species' characters, a dichotomous key is given. The poorly known Halacarellus floridearum (Lohmann) is described. Key words: Halacaroidea, Halacarellus capuzinus group, diagnoses, distribution, key.
Introduction The family Halacaridae includes more than 1000 species (Bartsch 2016). The genus Halacarellus, with its presently known 50 species (Bartsch 2009; Bartsch and Gerecke 2011; Tuzovski 2010), holds rank five in respect to the number of species, preceded by the genera Copidognathus, Rhombognathus, Agauopsis, and Halacarus. On the basis of shared character combinations, species of a genus can often be assigned to natural groups. In Halacarellus the group most rich in species is the Halacarellus capuzinus group (Bartsch 1997). It includes H. capuzinus (Lohmann, 1893), H. chersonesus Bartsch, 1998, H. discretus Bartsch, 1998, H. floridearum (Lohmann, 1889), H. kamtchatkaensis Tuzovski, 2010, H. procerus (Viets, 1927), H. psammophilus (Krantz, 1976), H. southerni (Halbert, 1915), H. subcrispus Bartsch, 1978, and H. tropicalis Bartsch, 1984. The capuzinus group is diagnosed and of each species characters and geographical distribution are outlined. To facilitate identification, both a tabular and dichotomous key is given. A table of variants in number of setae on the leg segments telofemur, genu and tibia is added. The original description of Halacarellus floridearum, published more than a century ago, is short and restricted to a few characters, these allow the identification but do not present details nowadays known to be of importance, accordingly a new description and illustrations are presented.
Material and methods The data presented, namely character states, geographical and ecological distribution, have been extracted from publications and re-examined specimens. Very rare character variants, already mentioned in Table 1, Ecologica Montenegrina, 10, 2017, 71-87
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are omitted in the diagnoses of the species and Table 2. A question mark is added in case the state is in need of verification; a preceding question mark means, the following statement may be erroneous. Unless otherwise stated, the length of a segment is that along the dorsal flank. The position of dorsal and ventral setae, given in decimal notations, refers to the dorsal and ventral flank, respectively, from the basal to apical end of that flank. The characters mentioned in the diagnoses and keys are visible at a magnification of about 400 times or less; oil immersion objectives are not necessary. The list of publications preceding the diagnoses of each species is restricted to the ones offering morphological details, descriptions and/or figures of the given species. Abbreviations used in the text: AD, anterior dorsal plate; AE, anterior epimeral plate; ds-1 to ds-4, first to fourth pair of dorsal idiosomatic setae, numbered from anterior to posterior; GA, genitoanal plate; glp, gland pore(s); GO, genital opening; OC, ocular plate(s); P-2 to P-4, second to fourth palpal segment; pas, parambulacral seta(e); PD, posterior dorsal plate; PE, posterior epimeral plate(s); pgs, perigenital setae; sgs, subgenital setae. The legs, their segments and claws are numbered from I to IV from anterior to posterior.
Systematics
Diagnosis of the Halacarellus capuzinus group (Figure 1A–C) Diagnosis. AD with pair of gland pores anterior to the level of ds-1, generally OC with one glp in anterolateral and one in posterior corner of plate and PD with a pair of pores in posterolateral margins; gland pores on OC and PD may be vestigial. Majority of species with three pairs of small dorsal sclerites, circular to ovate and 5–10 µm in diameter or length, rarely sclerites reduced. Pairs of ds-2, ds-3 and ds-4 arising from sclerites (if present) (Fig. 1A and B) or from striated integument. PE with one dorsal and two ventral setae. Female GA with two pairs of pgs; ovipositor extending to or somewhat beyond anterior pair of pgs; each genital sclerite with two to three (to four) sgs; three pairs of internal acetabula arranged in line. Male GA with 40–90 pgs; each genital sclerite with five (rarely four or six) sgs; genital acetabula in posterior part of GO, situated adjacent in a pair of foveae. Trochanters I to III with one seta, trochanters IV without setae (one individual unilaterally with a seta). All basifemora with two setae. Telofemur I with three (rarely four) dorsal setae and one (rarely zero or two) ventral setae, telofemur II with (two to) three dorsal setae and zero or one ventral seta, telofemora III and IV with two dorsal setae and zero or one ventral seta. Genu I with three dorsal and two ventral setae, one of ventral setae bipectinate, genu II with four, rarely three setae, genua III and IV with two (rarely three) dorsal setae and one (rarely two) ventral seta. Tibia I with six to eight ventral setae, basal pair spur-like (Fig. 1C) (except for few unilateral variants with three spurs), the other setae slender, bristle-like. In addition to one smooth ventrolateral seta, tibia II with one or two, tibiae III with two (rarely three) and tibia IV with two bipectinate ventromedial setae, respectively. Tarsus I ventrally with one (or two) spur(s), two delicate eupathidia and a pair of doubled pas (Fig. 1C); tarsi II, III and IV lack ventral setae. Claws on leg I shorter than those on following legs, tines on shaft of claw I smaller than on following tarsi or vestigial; claws of tarsi II to IV with distinct tines (Fig. 2G). Biology. Species of the Halacarellus capuzinus group run through one larval and two nymphal stages (protonymph, deutonymph). The AD of larvae often has small spicules arranged along the frontal margin (Krantz 1976: fig. 21; Bartsch 1972: fig. 25E, F, 1978: fig. 13, 1998: fig. 66). The genital and anal plates of the nymphs are fused to a GA. The protonymphal GA bears a single pair of acetabula, pgs and sgs are lacking, the deutonymphal GA has two pairs of pgs, two pairs of sgs and two pairs of internal acetabula. Remarks. Though the publication with the description of H. floridearum (in the year 1889) predates that of H. capuzinus (in the year 1893), the group was called H. capuzinus group (cf. Bartsch 1997). At the end of the 1990s, H. capuzinus was a well known species, in addition to the first description by Lohmann (1893), its characters had been outlined and illustrated by Schulz (1933), Newell (1947), Bartsch (1972), and Green and MacQuitty (1987), whereas the knowledge of H. floridearum was vague.
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Figure 1. Halacarellus capuzinus (Lohmann, 1889), A - general dorsal aspect, female; B - AD, right OC and anterior part of PD, male; C - Leg I, male. AD, anterior dorsal plate; ds-2, second dorsal idiosomatic seta; eu, eupathidia; f-pro, frontal process; OC, ocular plate; PD, posterior dorsal plate; sp, spur. Scale bar = 50 µm. (A - personal coll., English Channel, France, Roscoff; B and C - personal coll., Norwegian Sea, Norway, Tromsö)
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Figure 2. Halacarellus floridearum (Lohmann, 1889), A - idiosoma, dorsal, female; B - genitoanal plate, ventral, female; C - genitoanal plate, ventral, male; D - genital opening, ventral, male; E - gnathosomal base, rostrum and basal part of palps, dorsal, female; F - gnathosoma, ventral, female; G - tarsus I, lateral, female (claw and medial parambulacral setae omitted, the other medial setae in broken line). ds-2, second dorsal idiosomatic seta; fov, fovea; glp, gland pore; ov, ovipositor; pgs, perigenital setae; sgs, subgenital setae; so, solenidion; spe, spermatopositor. Scale bar = 50 µm. (A - CeNak, Viets Coll. 5393, Baltic, Poland, Mittelbank); B–G - personal coll., Arctic Ocean, Norway, Gamvik)
Description of Halacarellus floridearum (Lohmann, 1889) (Figures 2A–G, 3A–E) Idiosoma. Length of female (402–)495–555 µm, of male 508–532 µm. Surface of dorsal plates distinctly foveate. AD slightly longer than wide; anterior margin truncate. OC ovate, with distinct gland pore in anterolateral and posterior corner (Fig. 2A). Anterior part of PD ovate. Setae ds-2, ds-3 and ds-4 within striated integument, dorsal sclerites reduced. Posterior margin of AE truncate. PE with one dorsal seta and two ventral setae. GA with foveate ornamentation. Female GA oblong, ovate (Fig. 2B); distance from anterior margin of GA to GO 0.7–0.8 times length of GO; GA with two pairs of pgs; each genital sclerite with three (rarely four) minute sgs; ovipositor extending beyond anterior pair of pgs and almost reaching to anterior margin of GA; internal genital acetabula arranged in line. Male GA ovate (Fig. 2C), anterior-most part may be truncate; distance from GO to anterior margin of GA equalling 1.4–1.7 times length of GO; with 74
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71–89 pgs closely around GO; each genital sclerite with two to three anterior and two to four posterior sgs (Fig. 2D), anterior setae short, spur-like, posterior sgs somewhat longer. Genital acetabula in posterior part of GO, in a pair of foveae (Fig. 2D). Spermatopositor surpassing GO by length of GO. Gnathosoma. Approximately 1.5–1.7 times longer than wide. Tectum truncate (Fig. 2E). Rostrum not markedly slender; its length 0.8 times that of gnathosomal base. Both pairs of maxillary setae on rostrum (Fig. 2F). P-3 with small medial spur. Basis of P-4 with three setae in a whorl, apex with lateral solenidion, a delicate seta and two apical spurs.
Figure 3A–E. Halacarellus floridearum (Lohmann, 1889), A - leg I, medial, female; B - leg II, medial, female; C - leg III, medial, female; D - leg IV, medial, female; E - tibia I, medial, male. F - Halacarellus procerus (Viets, 1927), leg I, ventral (dorsal setae in broken line), female; G - Halacarellus southerni (Halbert, 1915), leg I, medial, female. Scale bar = 50 µm. (A–E - personal coll., Arctic Ocean, Norway, Gamvik; F - personal coll., Baltic, Germany, Schilksee; G CeNak, Viets Coll. 5386, North Sea, Germany, Helgoland) Ecologica Montenegrina, 10, 2017, 71-87
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Legs. Shorter than idiosoma. Tibia of leg I as long as (or slightly longer than) telofemur. Telofemur I 2.3–2.4 times longer than high; ventral seta at 0.7, basal one of dorsal setae in middle of segment (Fig. 3A). Genu I with five setae, ventromedial seta bipectinate. Tibia I with six (rarely five or seven) ventral setae, basal pair spur-like, distal setae slender, bristle-like (Fig. 3E) (rarely three setae spur-like). Tarsus I with three dorsal setae, dorsolateral solenidion, 16 µm long, ventromedial spur, pair of eupathidia and pair of doubled pas (Fig. 2G). Telofemora II, III and IV each with a ventral seta, on telofemur II seta in about middle of segment, on telofemora III and IV ventral and dorsal setae in apical half. Genua II to IV mostly with 2/1, 2/1, 3/1 dorsal/ventral setae, rarely genu IV with 2/1 or 3/2 setae, in the latter case one of ventral setae bipectinate. Tibiae II and IV ventrally with one smooth seta and two bipectinate setae (Fig. 3B and D); tibia III with one smooth seta and three bipectinate setae (Fig. 3C). Tarsus II with solenidion, 16 µm long, on dorsomedial fossa membrane. On tarsi II to IV ventral setae absent; apex of tarsus II with doubled pas, that of tarsi III and IV with pair of pas singlets. All claws with pectines, tines of pectines on tarsus I smaller than on following tarsi. Collecting data. The type specimens had been extracted from red algae collected in the western Baltic (Fehmarn and Langelands Sund) in about 6–23 m depth. Amongst the halacarid collection of K. Viets, stored in the CeNaK (Zoological Museum Hamburg), there is one slide (Nr 5393) labelled Halacarellus floridearum (Lohm.), Lohm. det., Ostsee, Mittelbank, 1889, Lohmann coll. The locality, Middle Bank, is in the South West Baltic Proper, NNE of Ustka (Poland). The female on the slide has a length of 402 µm. The above given description is prepared on the basis of specimens from the Baltic (Poland) and the North Atlantik (Norway, Greenland). Remarks. The original description (Lohmann 1889: 340–341, pl. 6: fig. 41, pl. 8: figs 111, 112, 114– 116) is very short. The figures 111 and 115 show a rather slender idiosoma with reticulated dorsal plates. The idiosomal length is said to be 326–404 µm, the OC to have two gland pores, one in anterolateral and one in posterior corner. The rostrum is slender, the tectum arched. Females are said to have three pairs of pgs, two of them adjacent and levelling about the middle of GO and one near anterior margin of GA, males to have numerous setae around the GO, telofemur I three dorsal setae, tibia I three pairs of ventral setae, with a short, spur-like basal pair, tibia III four long ventral setae. The given length of 326–404 µm is unusual small, compared with the data given above (about 500 µm), and is expected to be too low.
Halacarellus capuzinus group, list of species, diagnostic characters and distribution Halacarellus capuzinus (Lohmann, 1893) (Figure 1A–C) Halacarus capuzinus Lohmann, 1893: 199–201, figs 1–5. Halacarus (Halacarellus) capuzinus, Viets 1927b: 22. Halacarellus capuzinus, Schulz 1933: 98, 99, figs 4, 5. Halacarus (Halacarellus) capuzinus, Viets 1936: 548. Halacarus (Halacarellus) capuzinus, Sokolov 1952: 86, 87, fig. 34,1–2. Halacarellus capuzinus, Bartsch 1972: 205, 207, fig. 25A–L. Halacarellus capuzinus, Bartsch 1976: 103. Halacarus (Thalassarachna) capuzinus, Newell 1947: 118–123, figs 179–191. Thalassarachna capuzina, Green and MacQuitty 1987: 122–123, fig. 49A–D. non Halacarellus capuzinus, Dahl and Wieser 1955: 69–71, fig. 2a–c. non Halacarus (Halacarellus) capuzinus, Petrova 1972: 583–585, figs 4a, b, 5a–d. Characters. Idiosomal length of female 250–367 µm, of male 285–328 µm; specimens from the eastern and western Atlantic and the adjacent North Sea and Baltic, studied by the author, have a length of 273–353 µm (females) and 285–332 µm (males). AD with short, frontal process (Fig. 1A and B). Anterior margin of PD truncate, following lateral margins evenly diverging. Gland pores on OC vestigial. Setae ds-2, ds-3 and ds-4 situated on small sclerites. Anterior margin of female and male GA slightly rounded or almost truncate. In females distance between anterior margin of GO and GA equalling 1.3 times length of GO, in males 1.5–1.7 times. Ovipositor far from reaching to anterior margin of GA. Females with two pairs of sgs, males with five pairs. Male GO surrounded by 60–77 pgs. Gnathosoma short, about (1.1–)1.3–1.5(–1.6) times longer than wide. Short rostrum hardly extending to the level of seta on P-2. Telofemora I to IV with 1, 1, 0, 0 ventral 76
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setae, respectively. Length:height ratio of telofemur I 2.2–2.6: 1. Ventral seta of telofemur I in apical half. On telofemora I and III basal one of dorsal setae in middle of segment, on telofemur IV that seta in apical half. Tibia I about as long as telofemur I. Most common combination of dorsal/ventral setae on genua I to IV: 3/2, 2/2, 2/1, 2/1 and on tibiae I to IV: 5/6, 4/3, 3/3, and 3/3 (variants in Table 1). Two ventromedial setae of tibiae II, III and IV coarsely bipectinate, ventrolateral seta smooth. Tarsus I with single short ventromedial spur. Claws I with accessory process but no tines along shaft of claw (Fig. 1C). Table 1. Number of dorsal/ventral setae on the telofemora, genua and tibiae of the species Halacarellus capuzinus, H. chersonesus, H. discretus, H. floridearum, H. procerus, H. southerni, and H. subcrispus. Number of legs with the given character state in parentheses. leg H. capuzinus I II III IV
telofemur
genu
tibia
3/1(39),4/1(1) 3/1(20) 2/0(20) 2/0(20)
3/2(25) 2/1(5),2/2(13),3/1(2) 2/1(20) 2/1(20)
5/4(5),5/6(18),6/5(1) 4/2(1),4/3(18),5/3(1) 3/3(20) 3/3(20)
2/2(1),3/2(19) 2/2(20) 2/1(20) 2/1(20)
5/6(4),6/6(41),7/6(1),5/7(1) 4/2(51),5/2(3),4/3(2) 3/3(20) 3/3(20)
3/2(25) 2/2(28) 2/1(27) 2/1(25)
5/5(1),5/6(2), 5/7(30),6/7(3),5/8(1) 3/3(2),3/4(1),4/2(1),4/3(32),4/4(2),5/3(1),6/4(1) 3/3(26) 3/3(24)
3/2(20) 2/1(20) 2/1(20) 2/1(1),3/1(18),3/2(1)
5/5(1),5/6(2),6/6(14),7/6(2),6/7(1) 6/3(16),6/4(2),6/5(1),7/4(1) 3/4(20) 3/3(20)
H. chersonesus I 3/0(27),4/0(8),3/1(17),4/1(4) II 3/0(55),3/1(3) III 2/0(20) IV 2/0(20) H. discretus I 3/1(39),4/1(1) II 2/1(3),3/0(1),3/1(37) III 2/0(27) IV 2/0(25) H. floridearum I 3/1(20) II 3/1(16),4/1(4) III 2/1(20) IV 1/1(1),2/1(19) H. procerus I II III IV
3/1(20) 3/1(20) 2/0(20) 2/0(20)
1/2(1),3/2(19) 2/2(20) 2/1(20) 2/1(1),3/1(19)
6/7(2),6/8(18) 4/2(2),4/3(16),3/4(1),5/3(1) 3/3(20) 3/3(20)
H. southerni I II III IV
3/1(13),3/2(1) 3/1(15) 2/1(16) 2/1(14)
3/2(15) 2/1(13),2/2(1) 2/1(16) 2/1(16)
5/5(1),5/6(14) 4/2(15),3/3(1) 3/3(14) 3/3(14)
H. subcrispus I II III IV
3/1(20) 3/1(19) 2/0(19) 2/0(19)
3/2(20) 2/1(1),2/2(18) 2/1(19) 2/1(19)
6/6(1),6/7(19) 4/2(1),4/3(18) 3/3(19) 3/3(19)
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Distribution. Western North Atlantic: United States from Connecticut to New Brunswick. Eastern North Atlantic: France (English Channel), Eire (South East Galway, County Mayo?), Northern Ireland (Down, Strangford Narrows), Wales (Gower), Scotland (Outer Hebrides), Norway (Tromsö); North Sea: Germany; Baltic: southern and northern Sweden, Finland, Germany, Poland (Lohmann 1893; Newell 1947; Bilio 1966; Jansson 1968; Bartsch 1972, 1976, 1978; Jarvekjul 1979; Kautsky et al. 1981; Waterston 1981; Pugh and King 1985; Somerfield 1988, 1991). Records from the North Pacific (Kamchatka) and Black Sea (Ukraine) (Vorobyeva and Yaroshenko 1979; Tolstikov et al. 2005) are in need of confirmation. Habitat. Sandy beaches and subtidal sediments, abundant in turf of Baltic salt marshes; also found amongst subtidal algae (e.g. Fucus sp., Phaeophyta). Present in marine and brackish coastal water, even at salinities of no more than 1‰. Remarks. The length of the idiosoma given by Lohmann (1893), namely 210–220 µm, is expected to be too small. Dahl (1948) mentioned H. capuzinus from the Öresund (Sweden), extracted from algae in 5–15 m depth. Dahl and Wieser (1955) presented a short description, according to this, individuals from the Öresund have a length of 410–460 µm, 6/7 dorsal/ventral setae on tibia I and the anterior margin of the GA is slightly concave (notched), both in females and males. These characters do not fit to H. capuzinus but to H. subcrispus. The species reported from the Black Sea (Bulgaria) (Petrova 1972: p. 583–585, figs 3a, b, 4a,b, 55a–d) has an anteriorly arched AD (no frontal process), its palps and chelicerae are longer and more slender than in H. capuzinus. The remarkable long palps and chelicerae refer to H. chersonesus. A species called Halacarus areolatus was described by Halbert (1915: 123, pl. VIII, fig. 39a, b). The species, from Blacksod Bay, County Mayo, Eire, is said to be allied to H. floridearum, to be small, its legs long and slender, telofemur I to bear two dorsal and three ventral setae, tibia I four dorsal and six ventral setae (though according to fig. 39b there are five dorsal setae), and the basal pair of ventral setae to be spurlike. According to Bartsch (1985a), it is possible that H. areolatus is a synonym of H. capuzinus. Somerfield and O'Connor (1992) re-studied a mite on an unlabelled slide in the Halbert collection (National Museum of Ireland), that mite is large (550 µm), but as it is badly damaged this is not the accurate length, else the characters agree with those of H. capuzinus (Somerfield and O'Connor 1992). Halacarellus capuzinus is the smallest species in the capuzinus group. Considering that Halbert (1915) in the description of Halacarus areolatus referred to H. floridearum, which according to Lohmann (1889) has a length of 326–404 µm, and presented length data of H. southerni, namely 385 µm, it is surprising that the specimen from Blacksod Bay, called 'a small species', surpasses the others in length. A re-evaluation of the slide in the Halbert collection is recommended.
Halacarellus chersonesus Bartsch, 1998 Halacarellus chersonesus Bartsch, 1998: 152–155, figs 26–45. Halacarellus chersonesus, Pešić 2004: 40, figs 6, 7. ? Halacarus (Halacarellus) capuzinus, Petrova 1972: 583–585, figs 4a, b, 5a–d. Characters. Idiosomal length of female 360–400 µm, of male 369–371 µm. Anterior margin of AD slightly arched. PD with truncate anterior margin, then lateral margins evenly diverging. Gland pores on OC vestigial. Setae ds-2, ds-3 and ds-4 on small sclerites. Anterior margin of GA in female and male truncate. In female each genital sclerite with two sgs, in male with five sgs. Ovipositor far from reaching to anterior margin of GA. Male with 80–82 pgs. Distance from anterior margin of GA to that of GO in females 1.3–1.5 times length of GO, in males about twice that length. Gnathosoma very slender, 1.8–2.2 times longer than wide. Rostrum extending to or even beyond end of P-2. Telofemur I 2.8–3.0 times longer than high. Telofemur I with 3(–4) dorsal and 0(–1) ventral setae; ventral seta, if present, in apical half, basal-most dorsal seta in middle, rarely closer to basis of segment. Telofemora II generally, telofemora III and IV always without ventral setae. Basal one of two dorsal setae on telofemora III in middle and on telofemora IV in apical half of segment. Genu I generally with 3/2 dorsal/ventral setae, genua II with 2/2, III and IV with 2/1 setae (variants in Table 1). Tibia I and telofemur I almost equal in length. Most common combination of setae on tibiae I and II: 6/6 and 4/2 dorsal/ventral setae, respectively. Tarsus I with single short ventromedial spur. Claws on tarsus I with accessory process but no tines along shaft.
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Distribution. Mediterranean and Black Sea: Monte Negro, Crimea (Sevastopol) (Bartsch 1998; Pešić 2004). As mentioned above (H. capuzinus: Remarks), specimens from Bulgaria Petrova (1972), called H. capuzinus, most likely belong to H. chersonesus. Habitat. Sandy beaches. All present records are from low saline waters.
Halacarellus discretus Bartsch, 1998 Halacarellus discretus Bartsch, 1998: 155–159, figs 46–67. ? Halacarus (Halacarellus) procerus, Petrova 1972: fig. 5e. Characters. Idiosomal length of female 350–421 µm, of male 378–422 µm. Anterior margin of AD slightly arched, anterior margin of PD truncate, then PD evenly diverging. Gland pores on OC vestigial. Pairs of ds2, ds-3 and ds-4 on small sclerites. In both female and male anterior margin of GA broadly rounded to truncate. Female genital sclerites with two, those of male with five (to six) pairs of sgs. Ovipositor far from reaching to anterior margin of GA. Male GO surrounded by 52–66 pgs. Interval between anterior margin of GA and that of GO in females 1.4–1.6 times, in males 1.6–1.8 times length of GO. Length:width ratio of gnathosoma 1.5–1.6:1. Rostrum extending to end of P-2. Telofemur I cylindrical, 2.8–3.1 times longer than high. Telofemora I and II with a ventral seta, telofemora III and IV without such seta. Ventral seta on telofemur I in apical half of segment. Basal one of dorsal setae on telofemora I and III in about middle of segment, on telofemur IV seta in apical half. Length of tibia I same or slightly less than that of telofemur I (0.9–1.0:1). Genua I to IV with 3/2, 2/2, 2/1, and 2/1 dorsal/ventral setae. Tibia I generally with 5/7 dorsal/ventral setae, tibia II with 4/3 setae (variants in Table 1); and tibiae III and IV both with 3/3 setae. Tarsus I with single short ventromedial spur. Shaft of claws I without tines. Distribution. Black Sea: Ukraine (Odessa) (Bartsch 1998). A specimen collected in Bulgaria and called H. procerus (Petrova 1972: fig. 5e) bears a single ventral spine on tarsus I, accordingly it is no H. procerus, it may belong to H. discretus. Habitat. Sandy beaches.
Halacarellus floridearum (Lohmann, 1889) (Figures 2A–G, 3A–E) Halacarus floridearum Lohmann, 1889: 340–341, pl. 6: fig. 41, pl. 8: figs 111, 112, 114–116. Halacarus (Halacarellus) bergensis Viets, 1927a: 129–131, figs 61–66. Halacarus (Halacarellus) floridearum, Viets 1927b: 22, 23, fig. 56. Halacarus (Halacarellus) bergensis, Viets, 1927b: 23, fig. 57. Halacarus (Halacarellus) floridearum, Viets 1936: 549, fig. 643. Halacarus (Halacarellus) bergensis, Viets 1936: 549, fig. 644. Halacarellus floridearum, Bartsch 1998: 159–160. Halacarus (Halacarellus) bergensis, Sokolov 1952: 89, 90, fig. 37,1–5. non Halacarellus floridearum, Schulz 1935: 85, fig. 1–3 (cf. H. southerni). Characters. Idiosomal length of female 402–555 µm, of male 508–532 µm. Anterior margin of AD truncate, that of PD broadly rounded. OC with two distinct gland pores. Setae ds-2, ds-3 and ds-4 within striated integument (Fig. 2A), no sclerites present. Each female genital sclerite with three (to four) minute sgs, two (to three) setae in anterior and one in posterior position, each sclerite of male with two anterior and two to four posterior sgs. Anterior margin of both female and male GA widely ovate. Ovipositor almost reaching to anterior margin of GA (Fig. 2B). Males with 71–87 pgs. In females distance from GO to anterior margin of GA equalling 0.7–0.8 times length of GO, in males 1.4–1.7 times. Gnathosoma 1.5–1.7 times longer than wide. Rostrum extending to about end of P-2 (Fig. 2E and F). Telofemur I 2.3–2.4 times longer than high. All telofemora with ventral seta; this seta in apical half. Dorsal setae on telofemora III and IV in apical half, basal one of dorsal setae on telofemur I in middle of segment. Length of tibia I same or slightly more than that of telofemur I. Most common combination of setae on genua I to IV: 3/2, 2/1, 2/1, 3/1, on tibiae I to IV: 6/6, 6/3, 3/4, 3/3 dorsal/ventral setae (Fig 3A–D), respectively (variants in Table 1). Tibia III with one
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slender ventral and three bipectinate ventromedial setae (Fig. 3G). Tarsus I with single short ventromedial spur. Claws on tarsus I with distinct tines, though tines smaller than those of claws on following tarsi. Distribution: Western and eastern North Atlantic and White Sea: Greenland (Diskö, Godhavn); Norway (Bergen, Tromsö, Gamvik), Russia (White Sea); North Sea: The Netherlands (Den Helder); Baltic: Germany (Langelands Sund, Fehmarn, Greifswalder Bodden), Poland (a mite on a slide in CeNaK labelled Mittelbank, 1889, coll. Lohmann, Nr. 5393), Estonia (Lohmann 1889; Viets 1927a, b; Seifert 1938; Bartsch 1998; Jarvekjul 1979; Bartsch and Schmidt 1979; Bartsch and Smit 2006; Krapivin 2012; artsdatabanken 2017). Records from eastern Canada (northwestern Atlantic) and Ukraine (Black Sea) (Anderson 1933; Vorobyeva and Yaroshenko 1979) are in need of verification. Habitat. The present records are from lower tidal and subtidal zones, deepest record, namely 160 m, is from Godhavn, Greenland. Halacarellus floridearum has been extracted from algae and muddy sediments, also from the mantle cavity of the mussel Mytilus edulis Linnaeus, 1758. According to its presence in the eastern part of the Baltic (Poland, Estonia, 6–7 S‰), the species is adapted to life at reduced salinities.
Halacarellus kamchatkaensis Tuzovski, 2010 Halacarellus kamchatkaensis Tuzovski, 2010: 67–70. figs 1–11. Characters. Idiosomal length of female 435 µm, of male 420–425 µm. Anterior margin of AD truncate; anterior 1/6 part of PD almost rectangular. Setae ds-2, ds-3 and ds-4 on small sclerites. Anterior margin of female GA ovate, that of male widely rounded. Male with about 46–54 pgs. In both females and males GO large, in females GO 2.8 times, in males 1.3 times longer than distance between anterior margin of GO and that of GA; length ratio of distance GO–GA to length of GO in females and males equalling 0.3:1 and 0.7:1, respectively. Gnathosoma 1.5 times longer than wide. Rostrum not reaching to end of P-2. Telofemur I 3.1 times longer than high. Telofemora I and II with one, telofemora III and IV without ventral setae. Ventral seta on telofemur I and dorsal setae on telofemora III and IV in apical half of segment; basal-most of four dorsal setae on telofemur I near middle of segment. Genua I to IV with 3/2, 2/2, 2/1, 2/1 dorsal/ventral setae, respectively. Tibia I somewhat longer (1.1 times) than telofemur I. Tibiae I and II with 11 and 5 setae respectively; setae on tibia II in 3/2 (?) dorsal/ventral combination. Both tibia III and IV with 3/3 dorsal/ventral setae. Claws on tarsus I with accessory process but no distinct tines. Distribution. Southern edge of Kamchatka Peninsula, Kuril'skoe Lake. Habitat. Extracted from sand and pebbles of a freshwater lake (Tuzovski 2010).
Halacarellus procerus (Viets, 1927) (Figure 3F) Halacarus (Halacarellus) procerus Viets, 1927a: 127–129, figs 57–60. Halacarus (Halacarellus) procerus, Viets 1927b: 22, figs 54, 55. Halacarus (Halacarellus) procerus, Sokolov 1952: 91, 92, fig. 39,1–4. Halacarellus procerus, Bartsch 1972: 207, fig. 27A, B. Halacarellus procerus, Bartsch 1976: 103, figs 10–23. Halacarus (Halacarellus) procerus, Konnerth-Ionescu 1979: 121–123, fig. 1A–D. Thalassarachna procera, Green and MacQuitty 1987: 126–127, fig. 51A–D. Halacarellus procerus, Bartsch 1998: 166–167, figs 104–106. non Halacarus (Halacarellus) procerus, Petrova 1972: 586, fig. 5e. non Halacarus (Halacarellus) procerus, Makarova 1978: 131–134, figs 1, 2. Characters. Idiosomal length of female (350?–)391–496 µm, of male 372–496 µm. Anterior margin of AD and PD truncate; lateral margins of PD evenly diverging. OC with two very small gland pores. Setae ds-2, ds-3 and ds-4 situated on small sclerites. Anterior margin of GA truncate in both female and male. Ovipositor far from reaching to anterior margin of GA. Male with 56–80 pgs. Distance from anterior margin of GA to that of GO in females 1.2–1.6 times length of GO, in males about 1.7–1.9 times. Length of gnathosoma 1.4–1.6 times the width. Rostrum reaching to about the level of seta on P-2. Telofemur I 80
BARTSCH
cylindrical, (2.8–)3.0–3.2 times longer than high. Telofemur and tibia I almost equal in length. Telofemora I and II with one ventral seta each, telofemora III and IV with two dorsal but no ventral setae. On telofemur I ventral seta in apical half, basal-most of dorsal setae in middle of segment, on telofemur III basal one of dorsal setae near middle or moved slightly towards basis of segment, on telofemur IV both setae in apical half. Most common combination of dorsal/ventral setae on genua I to IV: 3/2, 2/2, 2/1, 3/1, on tibiae I to IV: 6/8 (Fig. 3F), 4/3, 3/3, and 3/3 (variants in Table 1), one of ventral setae on tibia III and IV slender, two setae bipectinate. Tarsus I with two short ventral spurs (Fig. 3F). Claws on tarsus I with accessory process, tines inconspicuous. Distribution. Northwestern Atlantic: United States, Massachusetts; Northeastern Atlantic and Baltic: Azores (coll. H. Kunz, unpublished), Spain (La Coruňa), France (English Channel), Eire (Dublin, West Cork), Northern Ireland (Strangford Narrows), Germany (North Sea and Baltic), Denmark, Sweden (Öresund ?), Norway (Tromsö); Black Sea: Romania (Viets 1927a; Dahl 1948; Bartsch 1976, 1978, 1985a, b; Bartsch and Schmidt 1979; Giere 1979; Somerfield 1988, 1991; Konnerth-Ionescu 1979). The specimen from Romania (Black Sea) is unusual small (length of idiosoma 350 µm) but has the two ventral spurs on tarsus I (Konnerth-Ionescu 1979: fig. 1C), whereas all other species of the capuzinus group have a single spur. Further records from the Black Sea, from Ukraine (Vorobyeva 1977; Vorobyeva and Yaroshenko 1979) are in need of confirmation. Habitat. Tidal and subtidal sand. Remarks. Dahl (1948) and Dahl and Wieser (1955) presented records from the Öresund and Kattegat, the mites were taken amongst algae in 5–15 m depth. According to a short description (Dahl and Wieser 1955), these specimens had a length of 410–460 µm, three pairs of sclerites in the dorsal integument, six dorsal and seven setae on tibia I (ventrally two spurs and five bristles) and a GA with slightly notched anterior margin. The species described is no H. procerus, most likely a H. subcrispus. The specimens taken by Dahl (1948) may be correctly identified. A species from the Gulf of Anadyr (Bering Sea), recorded by Makarova (1978), is no H. procerus. The description (p. 131–134, figs 1, 2) is contradicting. According to the scale, the female illustrated (fig. 1, idiosoma, dorsal aspect) has a length of about 420 µm, on page 134 the length is said to be 447–506 µm. Characters of the species are: telofemur I with three dorsal setae and one ventral seta, genu I with two dorsal and two ventral setae, tibia I with 11 setae (five dorsal and three pairs of ventral setae) (plus a seta represented by a porus?), tibia II with three dorsal and three ventral setae, tarsus I with a single ventromedial spur. The given combination of characters fits to several species of the capuzinus group but not to H. procerus. The specimens from the Gulf of Anadyr are most similar to H. psammophilus, a species known from the coast of Oregon, US.
Halacarellus psammophilus (Krantz, 1976) Thalassarachna psammophila Krantz, 1976: 249–252, figs 16–19. Characters. Idiosomal length of female about 440 µm (according to Krantz 1976: fig. 16), males slightly shorter, total length of female 561–583 µm, of male 495–523 µm. Anterior margin of AD arched; PD with narrow truncate anterior margin and evenly diverging lateral margins. Setae ds-2, ds-3 and ds-4 on small dorsal sclerites. Anterior margin of male and female GA arched. Females with two pairs of sgs, males with five pairs of sgs and about 64 pgs. Interval between anterior margin of GA and that of GO in female about 1.2, in male 2.1 times length of GO. Gnathosoma about 1.5(?) times longer than wide. Telofemur I cylindrical, length assumedly somewhat more than 3.0 times the height. Telofemora I and II with a ventral seta, such seta absent on telofemora III and IV. Ventral seta on telofemur I in apical half, basal-most dorsal seta on telofemora I and III in middle of segment, that on telofemur IV in apical half of segment. Genua I to IV with 5, 4, 3, 3 setae. Tibia I longer than telofemur I (1.2 times ?). Tibiae I to IV with 12, 7, 6, 6 setae. Tarsus I with single ventromedial spur. Claws on tarsus I with accessory process but no distinct tines. Distribution. Northeastern Pacific: United States, Oregon (Lincoln County). Habitat. Gravelly tidal sand in brackish water area (Krantz 1976). Remarks. Makarova (1978) described a species which was called H. procerus, it was collected in the Bay of Anadyr, Russia. According to the description (cf. above) these specimens are rather similar to H. psammophilus, perhaps even conspecific. Ecologica Montenegrina, 10, 2017, 71-87
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Halacarellus southerni (Halbert, 1915) (Figure 3G) Halacarus (Halacarus) southerni Halbert, 1915: 122, 123, pl. VIII, fig. 3a, b. Halacarellus southerni, Bartsch 1985a: 31–32, figs 2–14. Thalassarachna southerni, Green and MacQuitty 1987: 128–129, fig. 52A–D. ?Halacarellus floridearum, Schulz 1935: 85, figs 1–3. Halacarus (Halacarellus) floridearum nov. var. Schulz, Viets 1936: 549, fig. 643. Characters. Idiosomal length of female 360–427 µm, of male 360–400 µm. Anterior margin of AD with arched lamella, this about 10 µm long. Anterior margin of PD truncate to arched, then margins evenly diverging. OC with distinct gland pores, one anteriorly, one posteriorly. Dorsal sclerites reduced, less than 2 µm in diameter, ds-2, ds-3 and ds-4 arising from striated integument. Female GA ovate in outline, in male anterior margin of GA truncate, each genital sclerite with three minute sgs. Ovipositor distinctly extending beyond anterior pair of pgs, halfway between anterior margins of GO and GA. Male with 68 pgs. Distance from anterior margin of GA to that of GO in females 1.8–2.1 times length of GO, in males about 2.6–2.9 times. Ratio length:width of gnathosoma 1.5–1.6:1. Rostrum almost reaching to the level of seta on P-2. Telofemur I 2.3–2.5 times longer than high, with dorsal hump in basal half at about 0.3–0.4 (Fig. 3G). Tibia I shorter than telofemur I (0.9:1). Telofemur I with one, rarely two ventral setae, seta(e) at about 0.4 (with reference to length of ventral margin). Telofemora II to IV each with one ventral seta. All telofemora with basal one of dorsal setae in basal half of segment. Number of dorsal/ventral setae on genua I to IV: 3/2, 2/1, 2/1, 2/1, and on tibiae I to IV: 5/6, 4/2, 3/3, 3/3 (variants in Table 1). Tarsus I with single ventromedial spur. Claws on tarsus I with accessory process but no tines along shaft of claws. Distribution. Eastern North Atlantic: Eire (Mayo, West Cork); Irish Sea: Northern Ireland (Strangford Narrows); North Sea: England (Yorkshire), Germany (Helgoland), Sweden (Kristineberg) (Halbert 1915; Schulz 1935; Bartsch 1985a, b, 1991; Green and MacQuitty 1987; Somerfield 1991). A record of unproven identity is from the Adriatic Sea: Italy (Isola Tremiti) (Carriglio 2010). Habitat. Present records are from subtidal sediments, from about 11–56 m depth.
Halacarellus subcrispus Bartsch, 1978 Halacarellus subcrispus Bartsch, 1978: 10–14, figs 10–24. Characters. Idiosomal length of female 407–478 µm, of male 414–457 µm. Anterior margin of AD and PD truncate, anterior margin of PD narrow, then lateral margins evenly diverging. OC with two delicate gland pores. Setae ds-2, ds-3 and ds-4 on small sclerites. In both female and male anterior margin of GA wide, generally slightly concave in the median, rarely truncate. In female each genital sclerite with two sgs, in male with five sgs. Ovipositor far from reaching to anterior margin of GA. GO of male surrounded by 55–65 pgs. In female interval between GO and anterior margin of GA equalling 1.2–1.6 times length of GO, in male 1.8–1.9 times. Gnathosoma slender, 1.5–1.6 times longer than wide. Rostrum extending just beyond the level of seta on P-2. Telofemur I cylindrical, 2.8–3.4 times longer than high. Telofemora I and II with one, telofemora III and IV without ventral setae. Ventral seta on telofemur I and dorsal setae on telofemora III and IV in apical half of segment, basal-most dorsal seta on telofemur I in middle of segment. Genua I to IV with 3/2, 2/2, 2/1, 2/1 dorsal/ventral setae, respectively (variants in Table 1). Tibia and telofemur I almost equal in length. Tibiae I to IV generally with 6/7, 4/3, 3/3, and 3/3 dorsal/ventral setae. Tarsus I with single short ventromedial spur. Claws on tarsus I with accessory process, tines along claw lacking. Distribution. Northern Atlantic: Norway (Tromsö), Spitsbergen, western coast of Greenland (Diskö or Qeqertarsuaq) (Bartsch 1978). Halacarids from the Öresund, by Dahl and Wieser (1955) listed as H. procerus (as a junior synonym of H. capuzinus), may be representatives of H. subcrispus (cf. above, H. procerus: Remarks). In that case, the record would extend the known area of distribution to southern Sweden. Habitat. Tidal and shallow water sediment; also in low saline brackish water.
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Table 2. Characters states of species of the Halacarellus capuzinus group (rare character states are in parentheses; in respect to number of setae on telofemora, genua and tibiae only the most abundant variants are mentioned). abs, absent; AD, anterior dorsal plate; ap-h, apical half; arch, arched; bs-h, basal half; cyl, cylindrical; di, evenly diverging; ds, dorsal seta(e); ds-2, ds-3, ds-4, pairs of second to fourth dorsal idiosomatic setae; f, female; f-pro, frontal process; GA, genitoanal plate; GO, genital opening; GO–GA, distance between anterior margins of GO and GA; l-arch, arched lamella; L:H, ratio length to height, L:W, ratio length to width; m, male; md, middle of segment; me, striated integument; nr, number; PD, posterior dorsal plate; pect s, bipectinate seta(e); pgs, perigenital setae; pos, position; pre, present; rect, rectangular; roun, rounded; s, seta(e); scl, sclerite; sgs, subgenital setae; sp, spur; tru, truncate; v, ventral; vm, ventromedial seta; vs, ventral seta(e); I-3, II-4, III-5, telofemur I, genu II, tibia III, respectively.
Halacarellus tropicalis Bartsch, 1984 Halacarellus tropicalis Bartsch, 1984: 185–187, figs 1–8. Characters. Idiosomal length of female 524 µm. Anterior margin of AD arched. OC with two gland pores. Anterior narrowed part of PD truncate, following lateral margins evenly diverging. Pairs of ds-2, ds-3 and ds-4 on small sclerites. Anterior margin of female GA rounded to truncate. Each genital sclerite with two sgs. Ovipositor far from reaching to anterior margin of GA. Interval between anterior margin of GA and that Ecologica Montenegrina, 10, 2017, 71-87
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of GO equalling 1.4 times length of GO. Gnathosoma slender, about 1.6 times longer than wide. Rostrum extending to the level of seta on P-2. Telofemur I cylindrical, 3.2 times longer than high. Telofemora I and II with a ventral seta, telofemur III without such seta. Ventral seta on telofemur I in apical half, basal one of dorsal setae on telofemora I and III in middle of segment. Genua I to III with 3/2, 2/2 and 2/1 dorsal/ventral setae. Tibia I slightly longer than telofemur I (1.1:1). Tibia I to III with 7/6, 4/3 and 3/3 dorsal/ventral setae. Tarsus I with single short ventromedial spur. Claws on tarsus I with accessory process, tines lacking. Distribution. Caribbean Sea: Venezuela. Habitat. Coarse sediment with sand and coral shells (Bartsch 1984).
Key to species of the Halacarellus capuzinus group In addition to the tabular key (Table 2), a dichotomous key is presented on the following lines. Both keys are prepared on the basis of adults. For identification, several specimens should be examined. As listed in Table 1, the number of setae on the telofemora, genua and tibiae may differ, but unusual numbers are restricted to one side. Though mentioned in the keys, one should keep in mind that length:width or length:height data are highly influenced by the mounting; evaporation of the mounting medium may cause shrinkage and hence deformation of the mite enclosed, pressure on the coverslip or a slightly oblique orientation of the mite will have an influence on the measurement . 1 -
Tarsus I with two ventromedial spurs. Tibia I with (13–)14 setae. ............................................... procerus Tarsus I with single ventromedial spur. Tibia I with (9–)11–13 setae. ..................................................... 2
2
Anterior 1/6 of PD almost rectangular. In both females and males distance between anterior margin of GO and that of GA smaller than length of GO. .................................................................. kamchatkaensis PD with truncate or ovate anterior margin, lateral margins evenly diverging to middle of plate. Distance between anterior margins of GO and GA larger than length of GO. ......................................................... 3
-
3 4 -
5
-
6 -
Anterior margin of AD with frontal process. Small sized, length of idiosoma mostly less than 350 µm. ..................................................................................................................................................... capuzinus Anterior margin of AD truncate or slightly arched. Idiosoma generally more than 350 µm in length ...... 4 Pairs of ds-2, ds-3 and ds-4 arising from striated dorsal integument, dorsal sclerites reduced. Length:height ratio of telofemur I 2.5 or less. ........................................................................................... 5 Pairs of ds-2, ds-3 and ds-4 arising from rounded or ovate dorsal sclerites, 5–10 µm in diameter. Telofemur I cylindrical, length 2.8 times or more the height .................................................................... 6 Tibia III with three bipectinate setae. Genu IV generally with four setae. Telofemur I highest in about middle of segment, its ventral seta in apical half of segment. Tarsus I with tines along claw shaft. .................................................................................................................................................. floridearum Tibia III with two bipectinate setae. Genu IV generally with three setae. Telofemur I highest at end of basal third, ventral seta in basal half of segment. No tines along shaft of claws I........................ southerni Tibia II with two ventral setae, one seta bipectinate, one smooth, bristle-like. Telofemur I often without ventral seta. Gnathosoma slender, 1.8–2.2 times longer than wide. .........................................chersonesus Tibia II with three ventral setae, two setae bipectinate, one smooth, bristle-like. Telofemur I with one ventral seta. Length:width ratio of gnathosoma less than 1.7:1. ................................................................ 7
7 -
Tibia I with 12 setae. ................................................................................................................................. 8 Tibia I with 13 setae. ................................................................................................................................. 9
8
Anterior margin of female and male GA almost truncate. Length:width ratio of female GA 1.4:1, in males 1.3:1. Telofemur and tibia I almost equal in length............................................................. discretus Anterior margin of female and male GA arched. Length:width ratio of female GA 1.9:1, of male GA 1.5:1. Telofemur I shorter than tibia I (according to fig. 16 in Krantz 1976). ...................... psammophilus
-
9 -
Tibia I with seven ventral setae. Anterior margin of female and male GA concave. ................. subcrispus Tibia I with six ventral setae. Anterior margin of female GA arched .......................................... tropicalis
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Acknowledgements Thanks to Dr D. Harms and N. Dupérré (Arachnology, CeNak, Hamburg) for loan of slides of the K. Viets' halacarid collection, namely the specimens of Halacarellus floridearum (det. Lohmann), H. floridearum var. Schulz and 'H. bergensis'.
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