K. Christiansen.
The Collembola of Lebanon and Western Syria. Part III: Family Isotomidae.
Psyche 65:59-80, 1958.

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TXE COLLEMBOLA OF LEBANON
AND WESTERN SYRIA
PART I11 FAMILY ISOTOMIDAE1
. .
BY K. CHRISTIANSEN
Grinnell College, Grinnell, Iowa
The work herein described was done under N.S.F. Grant G 4563. I wish to gratefully acknowledge the work of my assistant, Jerry Tecklin, who did all of the preliminary sorting and most of the handling and mounting of the material here studied.
Relatively little has been known about the Isotomidae of the Syrian region. Until the work of Cassagnau and Delamare only a few of the more prominent epigeic forms had been described. With the above mentioned work the recorded species from the area were eight. In the present study four of these forms! were recovered and two others were probably recovered. In addition 16 new records were established.
Anurophorus coiffaiti Cassagnau & Delamare Plate 8, figures 6, 7
Anwophorw coiffaiti P. Cassagnau & Cl. Delamore, 1951 Biospeologica 75 :377,378.
This species was the first of the genus to be described from Western Syria. The specimens at hand agree well with the illustrations and figures given by Delamare and Cassagnau. The P.A.o. (see figure 7) is characteristically oval with a definite indentation or indication of a listel at least ,on the forward margin. The abdominal thickenings chararacteristic of the species vary a great deal but some indication of these can be seen even in the youngest speci- mens. In some forms these fuse into a single large tubercle- like projection. The sense organ of the third antenna1 segment usually has only two short setae between the 1 Published with the aid of a grant from the Museum of Comparative Zoology at Harvard College.
Pnrhe 655tl-W (1958). hup Ylpsfchv rnlclub org/65/65.059 html



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60 Psyche JUNE-SEPTEMBER
guard rods instead of three as indicated in the describers9 illustration.
The species appears in the moist regions, largely in the mountains. Type locality Antelias Lebanon, x 9-51. Other captures: LEBANON : Ain Zhalte Cedars, 177 M., XI '52; B'charra Cedars, 1900 M. v '52; between Dhour Schweir and Bikfaya, VIII '53; Nahr Array between B%a1ssine and Jezzine, 800 M.; Wadi Jahnam (N.W. Tripoli Prov.), VII 31 '52 ; Bikfaya VII '53. SYRIA :
Latakia Oak Grove, just
below Turkish border, VIII '53.
~nurophorus asfouri, n. ST).
Plate '8, figures 1-5
Baody subcylindrical with deep constrictions between segments. Unpigmented prdthorax much wrinkled and about Vy as longas rnesonotum. Clothing of body of rel- atively uniform short smooth acuminate setae, mostly curveld with slightly longer straight setae on last three abdominal segments. Intersegmental membranes clearly demarcated except for fifth and sixth abdominal segments, which are solidly fused. Pigment blue, in scattered scabrous patches over head, antennae, and tergites. Intersegmental mem- branes and legs pale. Eyes eight per side on a medially constricted eye patch. Antenna1 apex with a large bilobed bulb set in a deep pit. Fourth antennal segment clothed Figures 1-5: Anurophorus asfouri n. sp. 1. Habitus, paratype, seen from above and slightly to one side, setae omitted, 90X. 2. Tip of fourth antennal segment, paratype, 1000X. 3. Hind foot, paratype,1000 X. 4. Third antennal segment "sense" organ, paratype, 1000 X. 5. Eyes and P.A.O. right side, paratype, 700 X. Figures 6 and 7: A. coiffaiti. 6. Tip fourth antennal segment, specimen from between Dhour Shweir and Bkfaya Lebanon, 1000 X. 7. Anterior portion eye and P.A.O. Same specimen as above, 1000 X. Figures 8-13: Ballistura levanting n. sp. 8. Habitus, setae and pigment omitted, paratype, 40X. 9. Hind foot, paratype, 1200X. 10. Inner margin of apex of fourth antennal segment, paratype, 1200X. 11.
Inner part base of left dens showing dental projections, paratype, 1200 X. 12. Right mucro seen from above and slightly to one side, para- type, 1200X. 13. Left mucro, side view, holotype, 380X.



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62 Psyche
[ JUNE-SEPTEMBER
with a mixture of curved and straight acuminate setae, and a number of slightly curved blunt "sensory" setae. The latter type folrm about one third of the clothing of the apical half of the segment. Third segment clothe'd with a number of curved acuminate setae, and in addition six blunt setae as f~ollows: on the dorsal surface two long rods enclose between them two short oval knobs (about one third as long as the rods) ; on the ventral surface there are two additional curved rods about one half as long as the dorsal ones. Second and first antennals segments each with a circlet of acuminate setae. P.A.O. broadly oval and more than twice a.s large as the largest eye. Eyes eight per side with eyes D, H, and E, being smaller than the remainder. Integument granulate save for the last two abdominal tergites which are finely reticulate. Legs with tibiotarsi subsegmented. Unguis without lateral teeth but with an extremely small inner tooth. Empodial appendage lanceolate, slightly less than half as long as unguis, and lacking any apical bristle. Unguis with three weakly clavate tenent hairs, the median being strikingly longer than the remainder.
The de~cripti~on of this form brings to four the number of species of Anurophorus having the fifth and sixth abdominal segments fused into a single mass. Although these species are generally similar they may be readily separated on the basis of the characteristics listed below: SEPARATION OF SPECIES OF Anwophorus HAVING THE FIFTH AND SIXTH ABDOMINAL SEGMENTS FUSED Organ
Species P.A.O. eyes/side ungual tooth
A. konseli
broad1 y oval 4 -
A. asfouri
broadly oval 8 +
A. oredonensis narrow, indented
5 -
A. coiffaiti
narrow, indented 8 (7) -
In addition to these, special characteristics such as the abdominal thickenings of A. coiffati and the antenna] chaetotaxy can be used for additional separation. Type locality : Kammouha LEBANON (25 specimens), 1900 M., VII 19 '52; also taken from Latakia, SYRIA, oak grove just below Turkish border, VIII '53.




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19581 Christiansen - Collembola 63
Tetracanthella pilosa Schott .
Tetracanthella pilosa Schott, 1891. Ent: Tidskr. 12 : 191.
The specimens at hand agree with thedescriptions and figures in Stach quite well. The only small difference noted was that the furcula was relatively longer. The .species is quite rare in the region under consideration. It has been taken from : LEBANON : Ain Z'Halte Cedars, above forest, XI' 52 ; SYRIA : Pine Forest, Northern Latakia, VIII '53. Folsomides americanus Denis
Folsomides americanus Denis 1931. Bul. Lab. Zool. Portici 26:69-170. I have nothing to add to the comments of Cassagnau and Delamare (op. cit.) concerning thi8s species except that comparison with North American material shows it to be identical with the specimens at hand. This material had previously been identified as Folsorruides parvus. The species is widespread and abundant through the lowland and lower mountain regions' of the area under consideration. Taken from LEBANON : Beirut, campus American Uni- versity, many dates from 1951 through 1953; Souk el Gharb, VIII '53, between Kfardebiene & Bskinta, v '53: Belt ed Dine, XI '53 ; Saida, XI '51 ; 14 Km. South of Saida, XI '53 ; 2 km. East of Rayak, VII '52 ; Farayha, v '53. SYRIA Latakia, oak grove just below Turkish border, VIII '53. Folsomia
This genus is the best represented Isotomid genus in the area. There are probably five species occuring in the area, and the nature of the population isolation and vari- ation within several of the species concerned merits a great deal more study. Cassagnau and Delamare (op. cit.) de- scribe one new species and mention the capture of three single specimens they determined as Folsomia henemmi, I feel certain that these were in fact single-eyed specimens of the same species I have identified as F. penicula Bagnall. For this reason I have not included ksened in the list below. It is quite probable that larger series in each case would have shown the more typical four-eyed forms in each population.
Folsomia quadrioculata Tullberg
Isotoma quadriomlata Tullberg, 187 1. Ofv. Kongl. Vet. Akadm. Forhandl. 27 (I ) : 143-155.




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64 Psyche [ JUNE-SEPTEMBER
A great deal has been written about the group of species related to the form named above. Out of this there has de- veloped a we1 ter of species : brevicauda, qitadrioculata, manolachei, nana, microchaeta, penicula and mu1 tiseta. Further than this when we considered the excellent work of Agrell upon variation in eye number we can add to this group at least the one-eyed forms such as regularis, dip- lopthalmia, agrelli, ksened, pseudodiplopthdima, sirnilis, and monopthalma. Mlost of these so-called species differ by very few characteristtics, indeed often only by one. These are commonly chaetotaxy differences and there is not the slightest bit of evidence to deny the possibility that they represent single gene variations. Thus the differences seen could well represent in some cases balanced polymorphism, and in others' ecotypical 'selection for a given gene. Further than this most of the species here mentioned have been studied at one time from a few relativley homogenous habitats and there is 'thus little opportunity to study the variation through a wide range of conditions. I wish to suggest that there is at present insufficient evidence to sort out accurately from the forms mentioned ablove more than a few forms and be able to say with any degree of assurance that they actually represent different species. The speci- mens studied below appear to be classifiable as two species, F. quadrioculata and F. penicula. If we accept thils[ thesis, then it is possible to sort out the specimens at hand quite readily on the basis of the ventral manubrial chaetotaxy. If we sort them out thus, it appears that with few exceptions a given collection or sample is of one form or the other. If an attempt is made to sort out the populations on any other basis, this is not so and most samples are mixed. Further, in a statistical sense it is possible to sort out these two species upon about five additional characteristics ; how- ever, none of these shows very good correlation with the manubrial chaetotaxy. For purposes of illustrating the dis- tribution of some of these characteristics through the pop- ulation, a large number of specimens from twelve localities was sorted out to quadrioculata or penicula on a basis of .
the-ventral manubrial setae. Fifteen specimens were chosen from each species-and checked for a number of character- i7ties. The results are indicated below.



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19581 Christiunsen - Collembola 65
% quudrioculata % penicula
Description of character having having
Eyes unequal in size 47 86
Largest setae posterior
margin head twice smallest 33 100
Brush-like mass long setae
(4-5 rows) at end abdomen 73 100
Gap of P.A.O. wider than lip 93 47
Dentes with median dorsal seta 47 93
Other differences examined (ex., angling P.A.O. dorsal chaetotaxy, occurence one-eyed forms, shape of m,ucro) showed even less differentiation, or differences so fine as to be impractical. In many populations of both species an occasional one-eyed form is to be found, anld a few pop- ulations consist of such forms exclusively, without differing in any other visible respect from the four-eyed populations. The net result of this is to indicate that in the region studied there are two forms separable upon a basis of their manubrial chaetotaxy. It is convenient to consider them as two species but it must be understood that the real nature of their genetic relationship has yet to be established. In the Western Syrian region penicula is widely dis- tributed throughout the Lebanese mountains and the coastal plain, whereas quadrioculata is largely limited to the higher elevations of the Lebanon Mountains. Distribution of qmdrioculata: LEBANON: Ain Z'Halte Cedars, 1700 M. VII '52 & XI '52; between K'Fardebiene & Beskinta, 1900 M., VI '53; Hadeth Cedars, 1700 M., VII '53 B'Charra Cedars, 1900 M., v '53; Barouk Cedars, 1700 M.. VI '53.
Folsomia pewicvla Bagnall
Fokomia penicula Bagnall 1939. Ent. Mo. Mag. 75 :56-59. Fokomia tetropthalma Gisin 1946. Mitt. Scweie. Ent. Ges. 20:219. (NEW SYNONYMY).
Folsomia multiseta Stach 1947. Acta Monographica Polska Akademia umiejetnosci 138 : 172-177. (NEW SYNONYMY). PenicuZa was described for forms having two pairs of eyes with the posteriormost pair smaller than the fore pair and having a brush-like mass of setae at the end of the abdomen. In addition to this the members of the species are



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66 Psyche [ JUNE-SEPTEMBER
characterized by having six or more ventral manubrial setae in adult animals. In 1946 Gisin pointed out that this species was probably identical with Kseneman9s F. dip- lopthalma var. Tetropthalma. Since Kseneman had indicated that in his opinion this was a matter of population vari- ation and thus not subspecific in, rank, this name would fall under the category of "infr1asubspecif ic form" accolrd- ing to the rules "A name given to any Qfrasubspecific form if elevated to subspecific or specific ran@ by a subsequent reviser shall rank in its new status for purposes of priority as from the date on which it was slo elevated, and shall be attributed to the author by whom it was so elevated." Thus Folsomia tetropthalma becomes a junior synonym of F. penicula.
The species is extremely common in Lebanon and Syria, and in a variety of habitats. The specimens at hand display characteristics which destroy the separation shown between this species and multiseta by Stach (1947). The table below indicates how this species violates both sets of limitatimons: penicula ace. multiseta ace.
Stach Stach Leb. penicula
Ventral maubrial
setae 12-13 14-17 11-16
~.~.o./ommatidium 3.5-4 4-5 3.5-6
ventral setae on dens 12 16-22 12-18
In view of this I feel best to consider mzdtiseta a synonym of the locally variable penicula.
EXPLANATION OF PLATE 9
Figures 14-21 : Isotomina salaymehi n. sp. 14. Habitus, specimen from Beirut, 90 X. 15. Tip of fourth antennal segment, paratype, 1200 X. 16. Chaetotaxy last two abdominal segments, side view, paratype, 600 X. 17. Eyes and P.A.O. Unusual specimen with complete division eye patch, paratype, 1200 X. 18. Eyes and edge P.A.O. showing normal con- dition, paratypc, 600 X. 19. Right mucro, paratype.1100 X. 20. Abdomin- al "sensory" seta with neighboring normal seta, ptaarype, 1100 X. 21. Hind unguis, paraytpe, 1200 X. Figure 22-25: Vertagopus ciliatus n. sp. 22. Habitus (slightly distoried by mounting) paratype, 90 X. 23. Apex fourth antennal segment, holotype, 1200 X. 24. Long ciliate setae of fifth and sixth abdominal segments, holotype, 1200 X. 25. Eyes and P.A.O. left side paratype, 1200 X.




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68 Psyche [ JUNE-SEPTEMBER
One last point which must be discussed is the variation in the number of eyes. Agrell, Kseneman, Stach and a num- ber of other investigators have accepted the fact that the number of eyes in at least some species
of Fokomia is
varible. Gisin in 1946 has stated that this condition has not been proven for any species of the genus. Among the specimens I have identified as F. peniczda are occasional specimens with a single pair of eyes, and in one case a speci- men lacking eyes entirely. Beyond this, an occasional (2) small sample may consist of all one-eye paired forms. In all cases, and particularly where the four-eyed and two- eyed forms occurred in the same population, a slow, care- ful comparison was made between the two-eyed and four- eyed forms and not the slightest difference other than the eyes could be found. To assume that these represent dif- ferdnt species, particularily where the one-eyed paired forms represent two specimens in a sample of thousands, is to my view sheer folly. I feel therefore that I can state with authority that in the species I call penicula there are both two-eyed and four-eyed forms. The two-eyed forms are undoubtedly synonymous with those specimens iden- tified by Cassagnau as Ksenmani. It is entirely possible that these two species are synonyms but before this can be ascertained the whole group must be thoroughly reviewed. Distribution : LEBANON : Ain Z'Halte Cedars, many dates ; B'Charra Cedars, v '53; Kammouha, VIII '52; Barouk Cedars, VI '53 ; Chamlane, XII '53 ; Vic. Beit ed Dine, XI '53 ; Beirut, American University campus, many dates. SYRIA: North Latakia pine forest, VII '53 ; Latakia, oak grove just below Turkish border, v '53.
Folsomia candida Willem
Folsomia candida Willem 1902. Ann. Soc. Ent. Belgique, 46 : 275-283. The specimens at hand agree very well with the figures and descriptions given in Stach. The only notable dif- ference was the absence of an inner tooth on the unguis. In this respect the present specimens are similar to Bag- nail's distincta, but I agree with Stach that there is insufficent grounds for separating this species at the present time. The evidence of Delamare (1950 )and this material point to a locally variable widespread form. The



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present specimens are similar to the forms described and figured from Ardeche by Delamare in the absence of an inner tooth and the proportions of the dens, as well as in the existence of an occasional triangular P.A.O. However, in the chaetotaxy of the manubrial venter and the structure of the unguis, the present specimens are much more like those figured by. Stach for candida.
Distribution : LEBANON : Beirut, campus American Uni- versity, v '52; between Jezzine and B'Kassine, IX '52; Hermel, VII '52 ; Kammouha, VIII '52.
FoZsomia cavicola Cassagnau & Delamare
Folsomia cavicola P. Cassagnau & Cl. Delamare Deboutteville op. cit. :381,382.
I have not recovered this species but it appears to be distinguished from candida in Lebanon by the absence of an apical filament on the unguis, presence of a large internal lingual tooth, and the blunt mucronal teeth. Also the apical tooth is strongly upcurved in contrast to the straight tooth of candida. Known only from Ghita cave.
Folsomm sp.
A single specimen taken (damaged) from the Nahr Array in Lebanon merits mention. The P.A.o. is divided by a prominent listel, and the last three abdominal segments are weakly fused. The furcula is short and the mucro three-toothed. The eyes appear to be eight/side but are hard to distinguish. While this is probably a new species of Folsomia the single damaged
specimen is insufficient
material for its description.
Proisotorna minuta Tullberg
fsotoma minuta Tullberg 1871. op. cit.
~143-155.
The specimens at hand appear quite similar to those described and figured by Linnianemi, and Stach. There appears to be some variation in the setae on the tenaculum (most of the present forms have two setae) and in the eyes. In a number of specimens it appears that eyes G. and H are definitely smaller than the remainder. The size of the basalmost mucronal tooth is variable. Distribution: LEBANON:H~~~~~, VII '52; 2 Km. N. of Saida, XI '53 ; Antilias. XI '52 Nahr Array, v '52 ; Beirut, American University campus. SYRIA: Latakia, below Turk- 1'41 how'l~r. Opk Grove, VTI '53; North nine forest



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70 Psyche
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Proisotom minima, (Absolon)
Isotoma minima Absolon 1901. 2001. Anz.24 (634) : 32,33. Only three specimens were recovered from one locality: LEBANON : Aaranita, VII '52.
Ballisrum schotti Dalla Torre
Proisotoma schotti (Dalla Torre) 1885, 46 Progr. St. Gymn. Innsbruck. Handschin recorded this species from Palestine. Ballistura levantina, n. sp.
Plate 8, Figures 8-13
Facies Hypogastruru-like. Body and appendages thick. Uniformly blue-black in color except for slihgtly paler legs. Clothing of trunk and head of smooth, short curved setae of relatively uniform length. Sixth abdominal segment with a dozen longer setae about twice to three times as long as the shortest setae of the same segment. Antennae slightly but definitely longer than head. Fourth antennal segment without apical bulb, clothed with a variety of short, smooth setae. These are of three main types, straight to curved acuminate setae, straight truncate setae, and long mostly curved blunt "sensory" setae. In addition the dorsal surface bears a short triangular spine-like seta in a clear pit.
Sense organ of third antennal segment of two oval central pegs, one of which is strongly curved. Remainder of antennae clothed with normal acuminate setae. P.A.O. elliptical, eyepatch. Tibiotarsus without subsegment. Ten- ent hair not clavate and unguis untoothed. Empolial ap- pendage with large inner basal lamella and clear apical filament. Manubrion without ventral setae and with num- erous torsal setae. Dentes thick but tapering. Mucro boat shaped with thick granulate central axis, and with toothed lateral lamellae. Mucro from site view very deep, more than y2 as deep as long. Tenaculum with four teeth and a single
bristle. Cmorpus with an anterior projection much longer than the rami.
The species is similar to B. crassicauda in most respects. The differences between the two species are all small but in accumulation they indicate a real difference. Since previous abundant records of the group have shown little variation I feel the differences listed below merit specific separation.




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19581 Christhsen - Collembola 71