E. S. McCluskey.
Generic Diversity in Phase Rhythm in Formicine Ants.
Psyche 80:295-304, 1973.

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GENERIC DIVERSITY IN PHASE OF RHYTHM
IN FORMICINE ANTS
BY E. S. MCCLUSKEY
Departments of Physiology and of Biology Loma Linda University, Lorna Linda, California 92354 Ants are abroad through most of the day and night. But the species composition of this 24-hour patrol changes from one part of the day to the next (Talbot 1953 ; Wilson 1971 ). For example, in Michigan the maximum foraging activity of Lasius neoniger is at night, of Myrmica americana in the early morning and late after- noon, and of Formica pallidefulva nitidwentris in the middle of the day (Talbot 1946, 1953).
Likewise the mating flights of ants occur at different hours for different species (Kannowski 1963 ; Talbot 1945). The flight times may be similar for closely-related species (Kannowski 1959a). If one were to look at many species of one genus, would he find them to be similar as to time of day of foraging or of mating flight? Or does each genus span the 24 hours in terms of its various species? The aim of this report is to quantitatively compare species diversity with generic diversity of such phase relationships in one tribe of ants, the Formicini. The biosystematics of much of this group, particu- larly of the genera Lasius, Acanthomyops, and Formica, is relatively well known on morphological and zoogeographic grounds (Wilson and Regnier 1971 ).
The comparisons are based on a compilation of literature records for as many species and genera as possible in this tribe (Figs. I and 2). A genus was included if there were records for three or more species. About a third of the species of Acanthomyops, of Cataglyphis, of Lasius, and of Myrmecocystus are represented in the records cited here, but a smaller fraction of the large genus Formica, These genera are all from North Temperate latitudes. (For a preliminary report see McCluskey, 1972.)
The workers could be classified as nocturnal, diurnal, etc. But in the absence of single or definite beginning points or midpoints of activity in most of the literature records, another method was used to reduce each rhythm to one point for comparison with other species: If the ants do not normally come above ground at all (e.g., Acan- th~omyops species), the species is plotted as an X at the extreme left (Fig. I) ; if nocturnal only, one position farther to the right; if out Pu&e 80:295.304 (1973). http:llpsyche cnlclub ore/SWS&295 html



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as late as sunrise, another position to the right; etc. As far as pos- sible, only summer records were used so as to make directly com- parable.
It can be seen that the species in Acanthomyops, in Formica, and in Cataglyphis are closely grouped within each genus. The species in Lasius appear less so, but they barely overlap those of Formica or Cataglyp his.
x (XI x
FORMICA
X X X
(X) x
Figure I




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19731 McCluskey - Generic Diversity 397
Scoring an X in the leftmost column of Fig. I as "I", next to the left ("night") as "2" and finally the rightmost column as "5", permits an analysis-of-variance comparison of the variation within a genus with the variation between the genera: SS d f ms F p<
among genera 66.1 4 16.5 16.5 ,001
within genera 36.9 37 1.0
Thus the likeness within genera is greater than that between genera. A different type of analysis confirms this conclusion. The "none" aboveground activity was now omitted, since that is not really a time character (thus eliminating A c a n t ~ p s and some Lhs species) ; the "night" species were arbitrarily considered as (out until) 5 AM, "sunrise" 7 AM, "morning" 10 AM and "midday" I PM. These hours were treated as angles of a circular distribution, and the mean angles of the different samples (genera) were compared by Watson - .
Fig. 1. Worker surface activity (limited mainly to summer records, for most direct comparison of species). Each X represent0 one species and shows its nearest approach to midday; those based on the moat limited cited records are enclosed in
0. The leftmost column "NONE" indicates
that the species does not usually come above ground at all. Following are species and literature sources represented: FORMICA: iirdeyi (Wheeler
and Wheeler 1963), dakotemis (Talbot 1971)) emctoides (Andrews 1927, 1929; McCook IW), fu~a (Morisita 19391, finca iemmi (Brian 1955), lades (Wheeler and Wheeler 1970)) neogagates (TaIbot 1953), ob- swipes (Weber 193S), pal!idefufva didiventris (Talbot 1946, 1953, 1965), polyctena (Btuna 1954; Chauvin 1965a,b), $ratensis (Stebaev 1971; Stebasv and Reznikova 1972), sibylla (Wheeler 1917), submUgra (Talbot and Kennedy IWO), suhmlens (Ayre 1958, i959), subfiiissa (Stebaev 1971 ; Stebaev and Reznikova 1972), svbpolila (Mallis 1941), dkei (Holmqaist EGS). LASIVS: emarginutus (Tohmi 19691, flaws (Bernard 1968; Odum and Pontin 1961 ; Tatbot 1965 ; Wilson l9SS), fdiginosus (Wilson 19551, m h s (Kannowski 19S9b; Talhot 1965), seamffer (Talbot 1946, 1953, l965), niger (Eidmann 1926 ; Morisita 19391, diem (Wilson l9S5), sitkamsh (now $a!Uiarsfi) (Tahot 1965; Wilson 1955), ipec-al;wntris (Talbot IMS), umhrat~ll (Starcke 1937 ; Talbot 1965 ; Wihoa 1955). ACAATTHOMYOPS: claviger, interjectus, ldipes, and murfhyi (Talbot 1963 ; Wheeler and Wheeler 1963). MYRSfECOCYSTVS: lugubris (Cole 1966), metKger orbiceps (now placodops} (Wheeler 190Sb), w e d s (Cole 1966; LaRivers 1968), msxicaitw horlidear-urn (McCook 1SE2), mimicus (Cazier and Statham 1962; Leonard 1911), mojave (Cole 1966; LaRivcrs 196s; Leonard 1911), pyramicux (Smith 1951), deeleri (Snelling 1971 ) . CATAGLYPHIS: albicans (Mye 1968), alliicuns viaiiroidei (Tohm6 l969), a!ftsguamls (TohmS l969), bicoiur (Delye 1968 ; Pickles 1944; Tohm4 1969 ; Wehner and Duelli 1971), bicohr J&M (Gupta 1970), bombytina (Wye 19631, frig'tda (Tohmh 1969), IUCMI (Baroni Vrbani 1969; DUye 1964). An annotated table giving the details of support for Figs. 1 and 2 is available from the author.




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298 Psyche
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AM HOURS PM
6 8 10 12 14 16 18 20
F
FORMICA
F
F F F F F F
Fig. 2. Flight hours. Each I? represents for one species the halfway po,int between the earliest and latest literature records of flight; () indicate the most fragmentary records. The following are representeid: FORMICA: dakotensis (Talbot 1971), fusca (Kannowski 1959a; Talbot 1965), montana (Kannowski 1963 ; Kannowski and Johnson l969), neogagates (Talbot 1966), obscuripes (Clark and Comanor 1972; Talbot 1971), obscuriventris (Talbot 1964), opacbentris (Scherba 1961), pallidefulva nitidiventris (Tal- bot l948), pergandei (Kannowsld and Johnson 1969), pratensis (Eidmann 1928 ; Wheeler l908a), rufa (Don,isthorpe 1927 ; Standen 1909), ruftbarbis (Forel 1874a), sanguined (Forel 1874b), subintegra (Kannowski 1963; Talbot and Kennedy 1940), subnitens (Ayre 1957), ulkei (Scherba 1958 ; Talbot 1959). LASIUS: alienus (Gosswald 1932 ; Hall l887), brunneus (Forel 1874b ; Schenck l852), carniolicus (Kutter l946), emarginatus (Fore1 1874b, 1928), fiavus (Donisthorpe 1927 ; Forel 1874b ; Talbot 1965 ; Wilson 1955), fuliginosus (Wilson l955), minutus (Kannowski 1959a ; Talbot 1965)) nearcticus (Wilson 1955), neonigrr (Kannowski 1963; Talbot 1945, 1965; Wilson and Hunt 1966), nevadmis
(Cole 1956), niger (Donisthorpe 1927;
Forel 1874b)) pallitarsis
(Medler 1958 ; Talbot 1965), speculiventris (Kan- nowski 1959a; Talbot 1965), subumbratus (Kannowski 1971), umbratus (Crawley 1913 ; Forel 1874a, 1875 ; Kannowski 1963 ; Rau 1934)*. ACANTHOMYOPS: claviger (Talbot 1963, 1973), interjectus (Talbot 1963), latipes (Gregg 1963 ; Talbot 1963, 1973), murphyi (Talbot 1963), subglaber (Talbot 1973).
*The morning record (Rau 1934) is most unusual for this species (Kan- nowski 1963) and I omitted it in plotting the midpoint in the graph. Craw- ley (1913) and Fore1 (l874a, 1875) may possibly refer to a sibling species of umbratus (cf. Wilson 1955).




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19731 McCluskey - Generic Diversity 299
and Williams' ( 1956; cf. Batscheltet 1965, but only for a two- sample case) test : if the samples are considered random, Fl, y_q - [ (N-q) (XR-R) 1 / [ (q-1) (N-XR,) 1 = 3.58 and P < .05 [N = 33 (species), q = 4 (genera), Ri refers to the combination vector for all the species in each genus, and R refers to the com- bination vector of all genera].
Figure 2 shows that for the three genera of this tribe with enough species records, the flight hours of the species within a genus are strikingly similar, exceptions being the one morning species of Lasius and the two afternoon species of Formica. Comparison of the flight hours by the Watson and Williams test again shows the likleness within genera to ble greater than that between genera (P < .OOI). It seems noteworthy that a single rhythm character would char- acterim a genus this well. A preliminary test for generality of the within-genus likeness of flight hour was made on eleven genera from five subfamilies of ants (all of those from which literature records of at least three species per genus were at hand). The Watson and Williams test gives a value of P < .OOI ; this is true also if the three formicine genera are omitted and only the other eight considered. By including all species records no matter what tribe, it is pos- sible in a few cases to rank workers from early to late within each given locality ( McCluskey, unpublished) . Again species appear grouped generically, giving further suggestive evidence of a taxo- nomic or historical explanation of the phase differences (as opposed to a strictly ecological or geographical explanation). An example of a physiological character earlier shown to correlate well with a previously-established classification is that by Priesner ( I 968). Male moths throughout the family Saturniidae were shown to react to pheromone from the females in a manner roughly cor- responding to taxonomic grouping.
It should be added that some of the phase relationships considered here might in fact be explained by response to temperature, since these are records from the field rather than a constant temperature laboratory. But this consideration does not change the fundamental nature of the conclusion drawn. Further, certain ants which have been studied in constant temperature exhibit a phase relationship to the light cycle similar to that in the field ( McCluskey 1963, 1965, 1969 ; McCluske~ and Carter I 969).
Literature records of field mating flight and worker rhythms in- dicate the phase to be much more alike from species to species within



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300 Psyche
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a genus than between genera. The analysis is based on about a third of the world species in each of several genera of the tribe Forrnicini. Comparison in more genera, from several subfamilies, again indicates this prominent within-genus likeness of behavior. ACKNOWLEDGEMENTS
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