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Redescription of the bee-associateEllingsenius fulleri (Hewitt andGodfrey) (Arachnida, Chelonethi,Cheliferidae) with new records fromAfrica, Europe and the Middle EastMark L.I. Judson aa Department of Pure and Applied Biology, Baines Wing ,University of Leeds , Leeds, LS2 9JTPublished online: 17 Feb 2007.

To cite this article: Mark L.I. Judson (1990) Redescription of the bee-associate Ellingseniusfulleri (Hewitt and Godfrey) (Arachnida, Chelonethi, Cheliferidae) with new records fromAfrica, Europe and the Middle East, Journal of Natural History, 24:5, 1303-1310, DOI:10.1080/00222939000770771

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JOURNAL OF NATURAL HISTORY, 1990, 24, 1303-1310

Redescription of the bee-associate Ellingsenius fulleri (Hewitt and Godfrey) (Arachnida, Chelonethi, Cheliferidae) with new records from Africa, Europe and the Middle East

MARK. L. I. JUDSON

Department of Pure and Applied Biology, Baines Wing, University of Leeds, Leeds, LS2 9JT

(Accepted4May1990)

Ellingseniusfulleri (Hewitt and Godfrey), a bee-associated pseudoscorpion, is re- described. New records are given for Mozambique, Oman, Iran, Cyprus and Spain. Ellingsenius somalicus Beier, from Somalia, is regarded as a junior subjective synonym. The presence of E.fulleri in Europe is probably due to human introduction. It is suggested that Hansenius Chamberlin is the sister genus of Ellingsenius Chamberlin and that E.fulleri is a relatively plesiomorphic species of Ellingsenius.

KEYWORDS: Chelonethi, pseudoscorpions, Cheliferidae, Ellingsenius fulleri, rede- scription, distribution, Apidae.

Ellingsenius Chamberlin is an unusual genus of pseudoscorpions restricted to the nests of bees. Eight species have been described from southern, central and eastern Africa, and India. New material of Ellingsenius fulleri, previously known only from South Africa, extends this range to Spain, Cyprus, Iran and Oman.

The role of Ellingsenius species within bee hives is still a matter of some doubt. Beier (1948) reviewed the earlier literature and concluded that they were probably commensals ('phagophiles' in his terminology), preying on other commensals and parasites. However, Vachon (1954) found that E. hendrickxi Vachon showed a distinct preference for feeding on the host bees, at least in captivity. He also noted their reputation among local apiarists as bee predators. Recently, Murthy and Venkatara- manan (1987) have reported that E. indicus Chamberlin never eats bees, preferring small, slow-moving microarthropods instead (they also mention oophagy in their summary, but do not refer to this in the text--presumably this does not refer to bees' eggs).

Although Hewitt and Godfrey's description (1929) of E.fulleri is adequate for identification of the species, many characters currently employed are omitted, hence a full re-description is given here. The measurements used follow Chamberlin (1931), except that the length of the telofemur of leg I is measured as the intercondylar distance of the posterior face; the lengths of the hand and chela of the palp include the pedicel.

Ellingsenius fulleri ( Hewitt and Godfrey) (Figs 1-22)

Chel([er exiguus (nec Tullgren) Ellingsen, 1912:92 [misidentification, see Hewitt and Godfrey, 1929: 333-334]. Chel(/erfulleri Hewitt and Godfrey, 1929:331 334, Figs la, 7a-h; pl. 22, Fig. 11. Types from

0022 2933/90 $5-00 © 1990 Taylor & Francis Ltd.

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1304 M.L . I . Judson

Dunbrody, Cape Province (Albany Museum, Grahamstown, South Africa) [not examined]. Ellingseniusfulleri, Chamberlin, 1932: 36; Beier, 1932: 275. Ellingsenius somalieus Beier, 1932: 275, Fig. 298; Beier, 1932: 66-67. Holotype female from Somalia, 'Afmadu Oltregiula [ = Oltreguiba], (Coll. Mennozzi [ = Menozzi])' [not examined: apparenty lost]. New synonymy.

Materialexamined. Mozambique: Machava, near Maputo, 1 June 1980, A. Quadros leg., in wild bees' nests, 1 5', 1 9 (BM NH; bee specimens deposited in entomology collection), 28 November 1982, 3 ~', 2 9, 1 9 partial carcass (author's collection). Spain: Barcelona, La Llagosta, 19 June 1984, T. Ye'Lamos leg., 1 ~ (Gardini collection); Murcia, associated with beehives, sp. 207-2, 1 ~ (BM NH, CIE A19860). Cyprus: (no locality) 7 September 1983, Y. Antoniou leg., 2275, on honey bees, 2 ~, 49 (BM NH, CIE A15340, 1 ~, 1 ~ in author's collection). Iran: Shiraz, A. Ahmadi leg., on Apis mellifera, 1 ~, 1 9 (BM NH, CIE A15870).

Description. Carapace: eyes with weak lenses; two deep furrows present, anterior procurved, 0"31~0-43, posterior recurved, 0' 13-0.25 length of carapace from posterior margin; anterior disc with 109-127, median with 45-64 and posterior with 15-24 (total 172-206) dentate setae, those of anterior and median discs set on large tubercles; granulation even, but coalesced anteriomedially on posterior disc; male with posteriolateral keels.

Tergites: I X I divided (I-III weakly so in male); keels present on first 6-9 tergites of male, the last keel weaker than the rest; setae dentate, weakly biseriate, formula of half tergites, 8-12: 9-13: 9-14: 12-15: 12-16:13 17: 11-17: 11-16:12 15: 12-16: 10-14: 2; granulation dense, anteriorly fused to form longitudinal ridges, XI with denticulate granulation; lateral lyrifissures of tergites IV-X situated on pleural membrane in female (IV with a single fissure, V-X with two), lateral lyrifissures of male on lateroventral extensions of tergites, with the exception of one on the pleurum below tergite X.

Coxae: all evenly granulate; palpal coxa with about 6 setae on manducatory process--apical short, subapical long and acuminate, 'sensory seta' short--total number of setae about 30~40, anterior foramenal seta long; setae of pedal coxae, I 10-15: II 11-18: III 16-26: IV 50-70; coxal sacs of male vestigial, atrium sometimes weakly developed, one or more fragments of sac usually present in coxa, but not attached to opening (e.g. Fig. 10).

Sternites: anterior genital sternite of male with 31 59 setae of which 6-7 along the posterior border are bifid, posterior with 8-20 setae, including 2 longer acuminate setae, and a group of 4~8 internal setae on each side (Fig. 7); female with about 30 setae on the anterior, and 6-9 on the posterior genital sternite (Fig. 11); remaining sternites divided, setae ofhalfsternites (male and famale) 4~7: 6-9: 6-8: 7-10: 7-12: 9-12: 8-12: 7-11: 2, sternites VI-XI biseriate, XI with a long, acuminate, submedian seta on each half; anterior spiracle without setae, posterior with 1 seta; IV XI with numerous acini glands ('microlyrifissures'), particularly abundant on XI, along posterior half; lateral lyrifissures present on pleural membrane ofsternites VI XI (both sexes).

Genitalia of dactylocheliferine form. Lateral rods of male not fused anteriorly; dorsal apodeme large; eversible sacs (ram's horns organs) small; posterior faces of median and ventral diverticula covered by spicules ('pseudopoils' Vachon, 1938: 27) (Figs 1-3). Female with oval median cribriform plate and 2 large membraneous sacs (Fig. 12).

Cheliceral hand bearing 5 setae, b and sb dentate; gs of moveable finger tripled

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Redescription o f Ellingseniusfulleri 1305

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FIGS 1 7. Ellingseniusfulleri, males from Mozambique (unless otherwise noted). 1, Genitalia, dorsal view (aes apodeme of eversible sac, da dorsal apodeme, ddv dorsal diverticulum, ejca ejaculatory canal atrium, /a lateral apodeme, lr lateral rod, m muscle, td terminal dilation of vas deferens), detail shows spinules of median diverticulum; 2, ventral view showing eversible sac (es) and atrium of posteriodorsal gland (apdg); 3, dorsal view of dorsal apodeme and lateral rods (Cyprus); 4, right chelicera; 5, moveable finger of left chelicera of small female; 6, flagellum and base of serrula exterior; 7, genital sternites, right setae of anterior genital sternite omitted (crossed areoles indicate dentate setae). Scale lines: a, Figs 5 6; b, Figs 1-4, 7; divisions = 0.1 ram.

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1306 M. L. I. Judson

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FIGS 8 12. Ellingseniusfulleri. 8,9, Abnormal anterior tergites of male (8) and female (9) from Mozambique; 10, median part of coxae IV of male (Barcelona), showing rudimentary coxal sacs (cs) and openings (ocs); 11, female genital sternites (Mozambique); 12, female genitalia, dorsal view (Mozambique) (lcp lateral cribriform plate, mcp median cribriform plate, ms membraneous sac). Scale lines: a, Figs 8,9; b, Figs 11,12; c, Fig. 10; divisions = 0.1 ram.

(Fig. 4); spinneret with 5 or 6 rami, serrula exterior of 18-22 blades; serrula interior lamellar, with 4 apical, dentate blades; flagellum with 3 blades, anterior bearing 3-7 spinules, other blades usually simple (Fig. 6); dentition of fingers variable; hand reticulate, moveable finger with moderate, scaly granulation.

Palps (Figs 13-16) evenly granulate; setae denticulate and set on small tubercles on mesal surface of the trochanter, femur and tibia; hand without the long, dense setation found in E. sculpturatus (Lewis); femur and tibia each with a pronounced collar distally over articulation with following segment; hand of both sexes with a large number of pores (about 100) mesally at base of fixed finger; fingers slightly gaping, but without

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Redescription of Ellingseniusfulleri 1307

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FIGS 13-16. Ellingseniusfulleri (Mozambique), palps, 13, Right chela, male, with details ( x 4) of lamina defensor and diploid sensillum; 14, right palp of small female; 15, right patp of normal female; 16, right palp of male (tubercles and setae drawn only on trochanter, femur and tibia of male). Scale lines: a, Fig. 13; b, Figs 14-16; divisions = 0.5 ram.

strong modifications; fixed finger with 29-39 teeth, the first (distal) tooth slightly displaced ectally; numerous short, chemosensory hairs on ectal surface near tip of fixed finger; 1-8 acini pores between esb and est and 1-5 at level of ist; two dorsal furrows present, one running from ib to ist, the other from it to near the end of the finger; moveable finger with 32-45 teeth; chemosensory hairs apparently absent; 0-3 pores distad of sb; diploid sensillum slightly closer to st than to sb; granulation on mesal surface of moveable finger extending along most of its length; each finger with a single

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1308 M . L . I . Judson

haploid sensillum near the end; nodus ramosus of both fingers almost reaching level of st; trichobothriotaxy as illustrated (Fig. 13).

Legs normal, telofemur and tibia of posterior legs with a distal collar; setae dentate dorsally and simple ventrally; tarsi without TS; subterminal setae dentate; claws simple and retractile; tarsus of male first leg modified, dorsum slightly produced distally, often with a slight indentation (presumably to act as a stop for the anterior claw), anterior claw elongate, basally curved and lying over posterior claw (Figs 17-22).

Measurements (in mm; ratios in parentheses) of males and females combined (males tend to be slightly larger and more robust than the females, though the ranges overlap), except for leg I:

Body 2.f~3.1; carapace 0.98-1.17 x 0.92-1-09 (0-92-1.2). Palp: femur 0"91-1.13 x 0.31-0.42 (2.7-3-1); tibia 0-88-1-14 × 0-37-0.52 (2-1-2-5);

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FIGS 17-22. EllingseniusJulleri, males. 17,18 Distal part ofright tarsus and claws ofleg l, dorsal (17) and lateral (18) (Mozambique); 19-21 lateral view of right tarsus I of males (most setae only indicated by areoles) from Iran (19), Mozambique (20) and Cyprus (21); 22, right leg I (Mozambique). Divisions of scale lines = 0.1 mm.

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Redescription of Ellingseniusfulleri 1309

hand 0.82-0.97x0.45 0.63 (1.5-1.9); chela 1-43-1-71 (2.9-3.2); moveable finger

Leg I d': basifemur 0"32~).35x0.194).21 (1-6-1.8); telofemur 0.43-0.48x 0.18~).19 (2.4-2.5); tibia 0.44-0.47x0.15 (3.0-3.2); tarsus 0-35~.41 x0.144).16 (2.3-2-9) (~ Iran 0-47 x 0.14 (3.4)).

Leg I ~: basifemur 0.28~).33x0.20 (1.4-1.6); telofemur 0-42~).49x0.19-0.20 (2.2-2.4); tibia 0.43-0.49 x 0-13 (3-3 3.6); tarsus 0.38~3-44 x 0.104).11 (4-0-4.1).

Leg IV: basifemur 0.254).30 x 0.20-0.24 (1.2 1.5); telofemur 0.634).73 x 0-24-0.31 (2-4-2.7); total femur 0.97 1.11 (3.0-3.4); tibia 0.62-0-71 x 0.164)-17 (3.9-4.3); tarsus 0.47-0.55 x 0.114). 13 (3.9-4.5).

Abnormalities. A number of abnormalities were noted in specimens from Mozam- bique. One male and one female have tergal abnormalities: in the male left tergite II is fused to III laterally and lacks a keel, left IV is represented by an enlarged keel, right IV is fused to V medially and has a reduced keel (this may represent partial helicomery, with fusion, between tergites IV and V) (Fig. 8); in the female right tergite II is reduced and tergites III IV are fragmented (Fig. 9). The carcass (coxae, legs and left chelicera only) appears to represent an abnormally small female; the chela bears the usual 12 trichobothria, but there are only two galeal setae on the moveable finger of the chelicera (Fig. 5). It is certainly not an exuvium, as there are still muscles in the appendages. The measurements of this specimen are:

Palp: femur 0.75 x 0.27 (2.8); tibia 0.76 x 0-32 (2.4); hand 0.73 x 0.40 (1.8); chela 1.27 x 0.40 (3-2); moveable finger 0-62 (h/mf 1.2).

Leg I: basifemur 0-24 x 0' 17 (1-4); telofemur 0-35 x 0.15 (2.3); tibia 0-36 x 0-12 (3.0); tarsus 0.40 x 0.09 (4-4).

Leg IV: basifemur 0.24 x 0.19 (1.3); telofemur 0"58 x 0.24 (2.5); total femur 0"74 (3.14); tibia 0.54 x 0.16 (3-5); tarsus 0.42 x 0.11 (3.8).

Remarks. Hewitt and Godfrey (1929) described E.fulleri from material collected at Dunbrody, Bergvliet, Douglas and Willowmore in Cape Province. There have been no subsequent records of this species.

Ellingsenius somalicus Beier (1932) was based on a single female from Somalia which Beier separated fromfulleri on the basis of its less robust palps (femur 3-0 times, tibia 2.6 times as long as broad). However, the dimensions and proportions of the material studied here are rather variable and the values given by Beier fall within the ranges observed. Beier also stated in his description that the subterminal tarsal setae were simple--this is probably an error, as indicated by Chamberlin (1949, p. 51). Although the holotype of E. somalicus appears to be lost (it is not present in the collections of the Museo Civico di Storia Naturale Verona (Dr Ossella), Museo Civico di Storia Naturale 'Giacomo Doria' Genoa (Dr Capoccacio), Instituto di Entomologia Universita di Bologna (via Dr Gardini), Museo Zoologico 'La Specola' Florence (Ms Mascherini), or Naturalhistoriches Museum Wien (Dr Gruber)), I have little hesitation in reducing this species to a junior subjective synonym offidleri.

Ellingseniusfulleri is evidently the most widely distributed species of the genus, but its presence in Europe is rather surprising. Other species of Ell#lgsenius are known from southern and eastern Africa and from southern India, usually with fairly restricted ranges. It may be that this reflects the distribution of commerical apiaries, rather than the true distribution of Ellingsenius. The recent records from Europe are hard to account for: had the species always been present one would expect it to have

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1310 Redescription of Ellingsenius fulleri

been collected by apiarists before. I assume that the records from Spain, at least, represent recent dispersal, probably by man. An interesting parallel is the record of E. sculpturatus, a southern African species, from California (Chamberlin 1932). However, Mr Macfarlane (personal communication) has suggested an alternative explanation--that the records may be the result of recent collecting activity due to interest in the mite Varroa, a bee pest.

Mahnert (personal communication) has examined a single male offulleri from Oman (Dhofar, Salal, 17 '~ 00'N 54 ° 10'E, M. D. Gallagher and R. B. Whitcombe leg., 8 October 1985).

Ellingsenius is probably most closely related to the African genus Hansenius with which it shares many features, including the form of the male and female genitalia and male tarsus of leg I. Indeed, the characters separating Ellingsenius from Hansenius (galeat seta tripled, increased sclerotization, strong collars on legs, and association with bees) are probably all autapomorphies of Ellingsenius; the monophyly of Hansenius remains to be demonstrated. Comparison with Hansenius suggest that E.fulleri is probably a relatively plesiomorphic member of Ellingsenius, based on the position of trichobothrium ist and the weak development of the tubercles of the palp.

Acknowledgements I am indebted to Mr D. Macfarlane (Commonwealth Institute of Entomology, BM

NH) for providing me with material; to Prof. Dr J. A. Travassos Santos Dias (Centro de Zoologia, Lisbon; formerly at Museu de Histdria Natural, Maputo) for obtaining the specimens from Mozambique; to Dr G. Gardini for allowing me to look at his unidentified collections during a visit to the University of Genoa and for information regarding the holotype of Ellingsenius somalicus; and to Dr V. Mahnert for the record of E.fidleri from Oman.

References

BEIER, M, 1932a, Pseudoscorpionidea II. subord. Cheliferinea, Das Tierreich, 58, 1-294. BEIEn~,, M, 1932b, Zur Kenntnis der Cheliferidae (Pseudoscorpionidea), Zoologische Anzeiger,

100, 53-67. BEIER, M., 1948, Phoresie und Phagophilie bei Pseudoscorpionen, Osterreichische Zoologische

Zeitsehrift, l, 441-497. CHAMBERLXN, J. C., 1931, The arachnid Order Chelonethida, Stanford University Publications,

Biological Sciences, 7 (1), 1-284. CHAUB~RL~N, J. C., 1932, A synoptic revision of the generic classification of the chelonethid

family Cheliferidae Simon (Arachnida), Canadian Entomologist, 64, 35-39. CHAr~B~RLIN, J. C., 1949, New and little-known false scorpions from various parts of the world

(Arachnida, Chelonethida), with notes on structural abnormalities in two species, American Museum Novitates, No. 1430, 1-57.

ELLINGSEN, E., 1912, The pseudoscorpions of South Africa based on the collections of the South African Museum, Cape Town, Annals of the South African Museum, 10, 75-128.

HEWITT, J. and GODfrEY, R., 1929, South African pseudoscorpions of the genus CheliJer Geoffroy, Annals of the Natal Museum, 6, 305-336~

MtJRTHY, V. A. and VEN~ATARAMANAN, R., 1987 Contribution to the biology of the pseudoscorpion Ellingsenius indicus Chamberlain, Indian Bee Journal (1985), 47, 34-35.

VACHON, M., 1938, R6cherches anatomiques et biologiques sur le reproduction et te developpment des Pseudoscorpions, Annales des Sciences naturelles, Paris' (Zool.), (11) 1, 1-207.

VACHON, M., 1954, Remarques sur un Pseudoscorpion vivant dans les ruches d'Abeiltes au Congo Belge, Ellingsenius hendriekxi n. sp., Annales du Musbe royal du Congo Beige, N. S.. Zool, 1,284-287.

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