SOCIETAS PRO FAUNA ET FLORA FENNICA
ACTA ZOOLOGICA FENNICA
152
Cesare Baroni U rh ani and Cedric A. Collingwood The zoogeography of ants (Hymenoptera, Formicidae) in Northern Europe
HELS INKI-HELS INGFORS 1977 ACfA ZOOLOGICA FENNICA
1-4.5 vide Acta Zoologica Fennica 45-.50. 46--.59 vide Acta Zoologica Fennica 6{}.-93. 6{}.-99 vide Acta Zoologica Fennica 100-125.
100. MARIA RBUTER: Untersuchungen iiber Rassenbildung bei Gyratrix hermaphroditus (Turbellaria Neorhabdocoela). 32 S. (1961). 101. MARIA RBUTER: Index Generalis Seriei Acta Zoologica Fennica .51-100 (1948- 1961). 63 s. (1964). 102. W ALTER HAcKMAN: Studies on the dipterous fauna in burrows of voles (Microtus, Oethrionomys) in Finland. 64 pp. (1963). 103. A. M. J. EvERS: Ober die Entstehung der Excitatoren und deren Bedeutung fiir die Evolution der Malachiidae (Col.). 24 S. (1963). 104. JoHAN RBUTER: The international concentration in some hypotrichous Ciliates and its dependence on the external concentration. 94 pp. (1963). 10.5. GbRAN BERGMAN and KAI Orro DoNNER: An analysis of the spring migration of the Co=on Scorer and the Long-tailed Duck in southern Finland . .59 pp. (1964). 106. HENRrK OsrnRHOLM: The significance of distance receptors in the feeding behaviour of the Fox, Vulpes vulpes L. 31 pp. (1964). 107. BJORN KuRTEN: The Carnivora of the Palestine caves. 74 pp. (196.5). 108. BJORN KURrtN: The evolution of the Polar Bear, Ursus mflTitimus Phipps. 30 pp. (1964). 109. fRANK S. ToMPA: Factors determining the numbers of song sparrows, Melospiza melodia (Wilson), on Mandarte Island, B. C., Canada. 73 pp. (1964). 110. PoNTUS PALMGREN: Die Spinnenfauna der Gegend von Kilpisjiirvi in Lappland. 70 s. (196.5). 111. BJORN KURrtN: On the evolution of the European Wild Cat, Felis silvestris Schreber. 29 pp. (196.5). 112. PAUL KRiiGER: Ober die Einwirkung der Temperatur auf das Brutgeschiift und das Eierlegen des Rebhuhnes (Perdix perdix L.) 64 S. (196.5). 113. SAMUEL PANELIUS: A revision of the European gall midges of the subfamily Porricondylinae (Diptera: Itonididae). 1.57 pp. (196.5). 114. Bo-JuNGAR P. WrKGREN: The effect of temperarure on the cell division cycle in Diphyllobothrid plerocercoids. 27 pp. (1966). 11.5. BJORN KuRrtN: Pleistocene bears of North America. 1. Genus Tremarctos, spec tacled bears. 120 pp. (1966). 116. ToR G. KAlu.ING: On nematocysts and similar structures in Turbellarians. 28 pp. (1966). 117. BJC>RN KuRrtN: Pleistocene bears of North America. 2. Genus Arctodus, short faced bears. 60 pp. (1967). 118. FRANK S. ToMPA: Reproductive success in relation to breeding density in pied flycatchers, Ficedula hypoleuca (Pallas). 28 pp. (1967). 119. ANN-MARm FoRSrtN: Revision of the Palearctic HippflTion. 134 pp. (1968). 120. CHARLES HrGHAM: Size trends in prehistoric European domestic fauna, and the problem of local dbmestication. 21 pp. (1968). 121. ANITA RosENGREN: Experiments in colour discrimination in dogs. 19 pp. (1969). 122. ToM RBUTER: Visual pigments and ganglion cell activity in the retinae of tad poles and adult frogs (&na temporarta L.). 64 pp. (1969). 123. GC>RAN GnLENBERG: The energy flow through a Cborthippus pflTallelus (Zett.) (Orthopera) population on a meadow in Tviirminne, Finland. 74 pp. (1969). 124. PAUL KROGER: Zur Rassenfrage der nordeuropiiischen Igel (Erznaceus europaeus L.). 1.5 s. (1969). 12.5. BRIT Goosu ERntSEN: Rotifers from two tarns in southern Finland, with a description of a new species, and a list of rotifers previously found in Finland. 36 pp. (1969). 126. ANN-MARIE FoRSTEN: Variation in and between three populations of Meso hippus bairdii Leidy from the Big Badlands, South Dakota. 16 pp. (1970). 127. BENGT WESTERLING: Rumen ciliate fauna of semi- THE ZOOGEOGRAPHY OF ANTS (HYMENOPTERA, FORMICIDAE) IN NORTHERN EUROPE Cesare Baroni Urbani and Cedric A. Collingwood HE LS I N K 1- H E LS ING FORS N 0 V . 1 9 7 7 ISBN 951-661-023-4 I•SSN 0001-7299 © Societas pro Fauna et Flora Fennica Abstract BARONI URBANI, CESARE & CoLLI NGWOOD, CEDRIC A.: The zoogeography of ants (Hymenoptera, Formicidae) in Northern Europe. 1. - Acta Zoo!. Fen nica 1.5'2:1-'34. 197'7. Results from a numerical ana•lysois of 67 indigenous ant species map ped on 103 area units to include the British Isles, Denmark and Fennoscandia are discussed. 12 sets of species groups clustering at a similarity level :::,. 0.5 represent the major distribution patterns of North European ants. Multiple regressions calculated between species number per area unit and 11 environmental variables showed that main determinants included bright sunshine, July temperatures, relative oceanicity and length of growing season; these showed correlations in decreasing order with the distribution type having the largest number of species ( 40 °/o) representing the numerically dominant component of every area unit. Presence/absence analysis separated 21 biogeographical areas clustering at the 0.75 similarity level. The British Isles, as the largest homogenous area, separates out mainly from the absence of particular species, especially mem bers of the world-wide genus Camponotus. Some characteristic island faunae may be explained by the competitive advantage of post ~glacial first colo nisers. No support for postulating pre-glacial relict species was found. Authors' addresses: C. Baroni Urbani, Natural History Museum, Augus tinergasse 2 CH-4051 Basle, Switzerland; C. A. Collingwood, Agricultural Development and Advisory Service, Leeds, LS16 5PY England. Contents I. Introduction ...... 3 II. Material and methods ...... 4 Ill. The basic distribution types of the ants of North Europe ...... 6 IV. The Biogeographical areas ...... 24 V. Discussion ...... 29 VI. .Summary ...... '1'3 References ...... 33 I. INTRODUCTION The ants of North Europe, both taxo nomena observed together with a state nomically and in their geographical dis ment of conclusions of general interest tribution, are among the better known appear to be ju stified at this time. faunas of the world. Minor changes and additions to present day knowledge are to Acknowledgements. Mauro ·W . Buser of the Institute of Nuclear Phy~ics, University of Basle be expected in the future; in particular kindly prepared a new version of his programme some areas in Central Fennoscandia are CLUSTERTREE allowing the principal compo under recorded. However we consider nent analysis to flow directly from the similarity that the general picture of the population matrix in a single run. The illustrations to this paper were prepared by Miss Eva Weber through as presented here is not likely to change a grant from the Schweizerischer Nationalfonds to an extent that would modify our main zur Foerderung der wissenschaftlichen Forschung conclusions. Therefore an attempt to de (Request No. 3. 581.75). scribe and interpret the distributional phe- 100 100 ~0 400 100 ""'· FIG. 1. The area units used for Fennoscandia, Denmark and the British Isles. For explanation see Table 1. 4 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants II. MATERIALS AND METHODS The limits of the region studied do not corre absence of ants species were to be mapped. We spond exactly with the geographical limits of have already suggested (BARON! URBA 1 & Northern Europe but have been dictated for prac CoLLINGWOOD 1975) that the use of geometrical tical reasons to include those areas best known in grid maps, as recommended by the European In detail from a myrmecological point of view. In vertebrate Survey, could lead to misleading results this paper the British Isles, Channel Islands, Den in a biogeographical analysis because of lack of mark and the whole of Fennoscandia with their detailed information on the one hand and the related islands up to the political boundary be presence of unnatural d1scontinuities due to human tween Finland and Russia are included. The Balt disturbance on the other. Therefore we adopted as ic countries, North Germany, Belgium, Holland, a practical solution the use of area units often North France are excluded for practical reasons stemming from a compromise between geographic because of the lack of detailed information avail al and administrative barriers. The 1{)3 area units able from certain areas while the northernmost are mapped in Figure 1 and listed in table 1 and lands such as the Faroes, Iceland and Spitzbergen although they have probably never been used to are known to have no indigenous ants at all. gether in the same paper, and in the same way The next decision to be made was the size and as in the present study, they have been widely shape of the area units on which the presence/ employed in the literature and are well known TABLE 1. List of Area units British Isles Norway 69 Viirmland 70 Dalarna 35 0stfold & Akershus 1 Peninsula 71 Giistrikland 36 Hedmark 2 Channel 72 Hiilsingland 3 Thames 37 Opland 38 Buskerud 73 Medelpad 4 Anglia 74 Hiirjedalen 39 Vestfold 5 Severn 75 .Tiimtland 6 S. Wales 40 Telemark 41 Aust-Agder 76 Angermanland 7 N. Wales 77 Viisterbotten 8 Trent 42 Vest-Agder 43 Rogaland 78 orrbotten 9 Mersey 79 Asele Lappmark 10 Humber 44 Hordaland 80 Lycksele Lappmark 11 Tyne 45 Sogn og Fjordane 81 Pile Lappmark 12 Lakes 46 M0re og Romsdal 82 Lule Lappmark 13 Munster 47 S0r-Tr0ndelag 83 Torne Lappmark 14 Leinster 48 Nord-Tr0ndelag 15 Connaught 49 S. Nordland 16 Ulster 50 N. Nordland Finland 17 W . Lowlands 51 Troms 18 E. Lowlands 52 Finnmark wi 84 Alandia 19 E.Highlands 53 Finnmark ne 85 Regio aboensis 20 W. Highlands 86 Nylandia 21 N. Highlands Sweden 87 Karelia australis 88 Satakunta 22 North Isles 54 Sk!ne 23 Channel Islands 89 Tavastia australis 55 Blekinge 90 Savonia australis 56 Halland Denmark 91 Ostrobottnia australis 57 Smll.land 92 Tavastia borealis 24 South Jutland 58 Oland 93 Savonia borealis 25 East Jutland 59 Gotland 94 Karelia borealis 26 West Jutland 60 Gots ka Sandon 95 Os trobottnia media '27 North-west Jutland 61 tJstergotland 96 Ostrobottnia kajanensis 28 North-east Jutland 62 Viistergotland 97 Ostrobottnia borealis S '29 Funen 63 Bohusliin 98 Ostrobottnia borealis N 3'0 Lolland, Falster, Mon 64 Dalsland 99 Kuusamo 31 South Zealand 65 Niirke 100 Lapponia kemensis W 32 North-west Zealand 66 Sodermanland 101 Lapponia kemensis E 33 North-east Zealand 67 Uppland 102 Lapponia enontekiensis 34 Bornholm 68 Viistmanland Hl3 Lapponia inarensis ACTA ZOOLOGICA FENNICA 152 5 TABLE 2. List of species. Hypoponera punctatissima (Roger) Lasius niger (Linnaeus) Ponera coarctata (Latreille) Lasius emarginatus (Olivier) Stenamma westwoodii (Westwood) Lasius alienus (Forster) Myrmica rubra (Linnaeus) Lasius brunneus (Latreille) Myrmica ruginodis (Nylander) Lasius flavus (Fabricius) Myrmica sulcinodis (Nylander) Lasius mixtus (Nylander) Myrmica rugulosa Nylander Lasius umbratus (Nylander) Myrmica gallieni Bondroit Lasius rabaudi (Bondroit) Myrmica specioides Bondroit Lasius bicornis (Forster) Myrmica sabuleti Meinert Lasius carniolicus Mayr Myrmica scabrinodis Nylander Lasius fuliginosus (Latreille) Myrmica schencki Emery Formica exsecta Nylander Myrmica lobicornis Nylander Formica pressilabris Nylander Sifolinia karavajevi (Arnol'di) Formica foreli Emery Diplorhoptrum fugax (Latreille) Formica forsslundi Lohmander Formicoxenus nitidulus (Nylander) Formica suecica Adlerz Harpagoxenus sublaevis (Nylander) Formica truncorum Fabricius Leptothorax acervorum (Fabricius) Formica rufa Linnaeus Leptothorax muscorum (Nylander) Formica polyctena Forster Leptothorax nylanderi (Forster) Formica aquilonia Yarrow Leptothorax corticalis (Schenck) Formica lugubris Zetterstedt Leptothorax tuberum (Fabricius) Formica pratensis Retzius Leptothorax interruptus (Schenck) Formica nigricans Emery Leptothorax unifasciatus (Latreille) Formica sanguinea Latreille Myrmecina graminicola (Latreille) Formica uralensis Ruzsky Anergates atratulus (Schenck) Formica fusca Linnaeus Tetramorium caespitum (Linnaeus) Formica lemani Bondroit Strongylognathus testaceus (Schenck) Formica gagatoides Ruzsky Tapinoma erraticum (Latreille) Formica transkaucasica Nassonow Camponotus herculeanus (Linnaeus) Formica cunicularia Latreille Camponotus ligniperda (Latreille) Formica rufibarbis Fabricius Camponotus vagus (Scopoli) Formica cinerea Mayr Camponotus fallax Nylander Polyergus rufescens (Latreille) Plagiolepis vindobonensis Lomnicki to most North European naturalists. Indeed the information on the distribution of the indigenous area units for Denmark and Fennoscandia are species known to occur in Northern Europe. Pa precisely those used in the current series of volu pers by GosswALD et al. (1965) an·d by GASPAR mes of the Fauna Entomologica Scandinavica. (1971) also show general distribution maps of A large part of the information on the ant some of the species but in a few cases give a very distribution used here is already published and a different picture from that presented here, mainly few papers dealing with individual countries through the non critical reproduction of old lite (FoRSSLUND 1957; CoLLI NGwooo & BARRETr rature references. Some records however in these 1964; COLLINGWOOD 1974) but in addition many papers appear biogeographically acceptable but records for the other territories have been assemb where their sources are unknown they have been led through the help of local naturalists including excluded from our data input. A justification for in particular C. Bisgaard and C. Sk0tt for Den the classification adopted here and of previously mark and H. Wuorenrinne for Finland and from unpublished but verified locality records is given various scattered publications. All the species and in COLLINGWOOD (in press). their records have been carefully checked from Data consist of a small mxn matrix in which museum and private collections during recent the species are entered as rows and the area units years. as columns. The presence of a species in the given A list of all the species included in the analysis area unit is coded as "1" and its absence as "0". is given in table 2 and their distributions are de The comparison between species (rows) is termed tailed in a sequence of maps. These maps represent the R analysis and the comparison between bio the data input for our analyses and provide an geographical units (columns), Q analysis. Accord easily understandable presentation of our updated ing to the type of analysis performed, each row 6 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants or each column represents an Operational Taxo The R analysis of the data matrix and the nomic Unit (OTU). successive clustering were used to identify the The Q and R analyses of the original data basic distribution types in the selected area. From matrix were carried out by the programme the reasonable symmetry properties of s .... and of CLUSTERTREE available at the computing cent the low distortion of the original similarity matrix re at the University of Basle. The similarity mat during clustering by single link, distributions clus rix was calculated by the Sn coefficient of tering at a level of ~ 0.5 were interpreted as being BARONI URBANI & BusER (1976): similarity be similar and representing the major generalised tween two OTU's given by distribution patterns. Clustering with s .... <0.5 should connect true differences and are not con Vad + a s.... = sidered here. "\lad+a+b+c In addition we attempted a stepwise multiple regression between the number of species per area where, a = number of attributes (presences) in unit of each distributional type and a set of en common, b = number of attributes present in the vironmental variates, translating the original data first OTU and absent in the second, c = number into meristic values. These were abstracted from of attributes absent in the first OTU and not source works on climatology by LANDSBERG et al. in the second, d = number of attributes absent in (1965) and WALLEN (197 0) . The following para both OTU's compared but present in others. The meters were used: Average annual rainfall (6 calculation of the similarity coefficient gives a categories), July mean temperature (4 categories), symmetrical m X m (or n X n) similarity matrix on Average bright sunshine hours per year (5 cate which the similarity between every possible pair gories), Average annual atmospheric pressure (6 of OTU's can be found. From the numerous tech categories), Average length of growing season (6 niques that would allow a graphic representation categories), Inland. versus island climate (3 cate of the results with a minimum distortion of the gories) , Latitude (6 categories), Longitude ( 4 ca similarity matrix, we used the clustering by single tegories), January mean temperature (5 catego linkage in which each OTU joins the most similar ries), Boreal versus oceanic climate (5 categories), one in the matrix at a distance corresponding to Mean altitude (2 categories). A constant pro its similarity; once one OTU is ascribed to a given portion of the sum of squares > 1 °/o was used set, it remains there until the end of clustering. to limit the number .of variables in the regression. m. THE BASIC DISTRIDUTION TYPES OF THE ANTS OF NORTH EUROPE The dendrogram based on the similar East Jutland in Denmark. No correlations ity between species distributions is pre with environmental variables were shown sented in Figure 2. By cutting this dend by the analysis. rogran at the 50 Ofo similarity level, we obtained 12 basic distribution types as described below. Distribution Type B This is presented in Figure 3 and in Distribution Type A cludes the species Hypoponera punctatis sima (Roger), Sifolinia karavajevi (Ar The sole species in this distribution nol'di), Myrmica specioides Bondroit and type is Formica foreli Emery. This is a Leptothorax interruptus (Schenck) (Figu local species with a scattered distribution res 4-7). They have in common their in Central and Eastern Europe (DLuSSKY regular presence on Thames and scatter 1967). The only 2 records for North Eu ed records in the southern part of the rope are from North Zealand and North studied area. The regression analysis for ACTA ~OOLOGICA FENNICA 152 7 % SIMILARITY lo I~ I~ I~ I~ 18 Formica forell Emery ! Hypoponera punctatissima lRogerJ Myrmlu spedoides Bondroit i Sifolinla karavajevi '-'rnol'!l'il i LeptothOrax Interruptus tSchenct} ! lasius brunneus G..atreille) I Anergates atratulus lSthenck} i Strongylognathus testaceus (Schenck1 Ponera c.oarctata n..atreille) i Tapinoma erraticum a..atreillel '--- Diplorhoptrum lugn O..iltreillel Stenamma westwoodii tWestwood) leptothorax nylanderl fllrsterl I Myrmecina gra minkDi a (Utreille) lasius rabaudi (8ondroitJ 0 ~ formiu cunicularla latreille ! Camponolus vagus I'Scopolij Lasius u:rntoncus Mrtr Myrmica gallienl Bondroit Myrmica schencki Emery ~ Formica rufibarbis Fabricius l ltpfothorax tuberum fllbrkius) '-----1 Camponotus ligniperda tl..Jtreilld Fo rmica pratensis Rrtzius leptothorax muscorum 1Ny1aoder) Myrmiu sabuleU Meinert Formica fusu Unnatus - Myrmica scabrinodis Nyfander Myrm+ca rubra tlinn;,eusl lasius njger G.inn~tus) r- ~ lasius flavus hbrk:iusl Tetramorium c.espitum Linn.eusl Fo rmica rufalinnaeus l Lasius fulic} inosus flatreiUel ! Usius alienus F6rstuJ I Usius mirtus fNylande rJ I i L.asius umbratus (Nytander) ! Formlcoxenus niHdulus INyf andert ! Formica truncorum Flbrklus ! I Formica WlgUinea Latreille I L_ LL.r Camponotus herculeanus Linn.eus} - ! FormiuexsectaNytancler ! Myrmia ruginodis CHyl ander) ~ leptolhorax urvorum f"lbriciusl I Myrmiu loblcornis Nylander i Myrmic.a sulclnodis Otylander} - ! formia cinerea Mayr Formica pol)dena Ft:irster I Formica nigriuns Emery i y Myrmlu rugulosa N)'lander 1..---- ! Formica pressillbris Nylander I Formica suedca Adltrz Formica forulundi lohmander i Harpagoxenus sublaevis ~ andtr ) i Formic.l uralensb Ruzsly I ~ '--- Formica transbuasicJ Nassonow ! Fo;mici qagatDides Ruzs);y I formica aqu Uonia Yar row ! Formica lugubris Zrttersteclt. I ~ I :Formla lem~ni Sonclroit Camponotus fallu Nyfander I ltf)tothoru cortblls tSthtnc::k) lasius bk.ornis FOrsterl i Polytnjus rufescens LJtreilleJ i leptothonx unHasclatus · i..JlreJUe) Laslus tmarginatus fCMiYierl i Pl~g iolepls vlndotlontnsJs t.omnlcki 1~8~--~----~~~~----~----~~g~--~-----ei~~---L----~~~~----J---~Io 30N3!13::1=110 % FrG. 2. Dendrogram rJf species distribution similarity. this distribution type (Table 3) suggests 2 factors contribute only 13 Ofo of the a double and antithetical influence of 2 observed variation although coupled with factors: a positive influence of the July a high R value and additional environ mean temperature which should account mental data are needed for a full inter- for the southern pattern within a wide . pretation. H. punctatissima is a cosmo longitudinal range and a negative in politan species long established in Europe fluence of inland climate. However these where it colonises outside rubbish tips, 8 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants ' presence 1\1 > 0.0 •11 >33.3 100 1 00 JOG •oo toO •m FIG. !1. Distribution Type B. TABLE 3. Step-wise multiple regression between distribution type B and 11 environmental variables. Variable entered Cumulative proportion Regression of sum of squares reduced coefficient July mean temperature 0.107 rQ .2314 3.199 Inland vs. island clima 0.13{) - 0.1453 -1.627 Intercept --{).098. Multiple correlation coefficient 0.348 (p<0.01). FIG. 4. Hypoponera punctatissima (Roger), FIG. 5. Sifolinia luzravajevi (Arnol'di), FIG. 6. Myrmica specioides Bondroit, FIG. 7. Leptothorax interruptus (Schenclc). ACTA ZOOLOGICA FENNICA 15'2 9 sawdust heaps, mines in addition to heat remnants of this type of woodland occur ed premises. For the purposes of this in the south up to a broad band extending paper its distribution relates only to re across the English Midlands to the East corded locations away from buildings. S. ern Counties and in several localities karavajevi is parasitic on Myrmica. M. there is a characteristic association of specioides and L. interruptus are widely Lasius brunneus and Leptothorax nylan but sparsely distributed through Central deri in old oaks throughout this area, with Europe and in the north are found on S. westwoodii also usually present on the coastal sand or dry lowland heath. woodland floor. Such associations occur throughout Belgium and the oakwoods of Distribution Type C Northern and Central France. However, This distribution type (Figure 8) inclu in Norway L. brunneus occurs in similar des a group of species with a wide range remnants of old oak woodland in res in Central and Southern Europe: Formica tricted areas round Oslofjord without L. cunicularia Latr., Lasius rabaudi Bond nylanderi while in Central Sweden L. roit, L. brunneus (Latr.), T apinoma erra brunneus is also not uncommon in old ticum (Latr.), Myrmecina graminicola oaks but not apparently associated with (Latr.), Stenamma westwoodii W estwoo·a, L. nylanderi which is found more frequ Diplorhoptrum fugax (Latr.), Strongylo ently in the southwest of the country. gnathus testaceus (Schenck), Anergates Similarly in Zealand L. brunneus occurs atratulus (Schenck), Leptothorax nylan locally in oak without L. nylanderi. deri (Foerster) and Ponera coarctata (Latr.) (Figures 9-19). Lasius brunneus, Leptothorax nylanderi Stenamma west woodii and Myrmecina graminicola are species usually associated with deciduous woodland mainly Quercus robur. This reaches its northern distribution in Cent ral Sweden; there are no records for these species from Finland where pedunc ulate oak is restricted to the southwest extremity of the country. In England % presence lliill > 0. 0 !11 > 33.3 • > 66.6 o 100 2 0 0 aoo •oo .oo kno FIG. 8. Distribution -r'ypc: C. C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants FIG. 9. Formica cunicularia Latreille, FIG. 10. Lasius rabaudi (Bondroit), FIG. 11. Lasius brunneus (Latreille), FIG. 12. Tapinoma erraticum (Latreillle), FIG. 13. Myrmecina graminicola (Latreille), FIG. 14. Stenamma westwoodii (Westwood), FIG. 15. Diplorhoptrum fugax (Latreille), FIG. 16. Strongy lognathm testaceus (Schenck), FIG. 17. Anergates atratulus (Schenck), FIG. 1 . Leptothorax nylanderi (Forster), FIG. 19. Ponera coarctata (-Latreille). The most influential environmental in common their total absence of records variable according to the regression from Finland where apparently suitable (Table 4) for this distribution type is lati climatic conditions are present at least in tude. The other species in this group are the South. This pattern may be explained not at all related to the distribution of by the regression where length of grow oak but are largely thermophilous nor ing season and July mean temperature mally associated with lowland sandy are negatively correlated with these spec heath. Except for L. rabaudi they have ies within the set of variables considered, ACTA ZOOLOGICA FENNICA 152 11 so that although they are of generally southern distribution, notably Formica cunicularia, Diplorhoptrum fugax, T api noma erraticum and Ponera coarctata, they are not positively correlated with temperature. A possible hypothesis would be the influence of a historical factor in Southern England where an older fauna ~ presence directly derived from continental Europe rl1 > 0.0 would be well established. In Southern 11 > 33.3 Scandinavia this might have given way • > 66.6 to later invading species from Eastern Europe except in such island refugia as Oland and Gotland which were not o 100 200 )00 4M .... km reached by the later invaders. FIG. 21. Distribution Type E: Lasius carniolicus One apparent anomaly is Stenamma Mayr and Myrmica gallienei Bondroit. westwoodii in the Bergen district of Nor way but the record (CoLLINGwooo 1974) is based on specimens collected in a bo poli (Figure 20). This is a South European tanic garden which may well have been species abundant in the Mediterranean introduced on imported plant material. area but sparse and discontinuous north However, areas of coastal Norway in the of latitude 4 7°. It has been recorded very southwest are climatically not dissimilar locally in Central and Southern Poland from parts of Southern Ireland where (PISARSKI 1975). Old records from Got this species is indigenous. land and Oland and South Finland were Anergates atratulus and Strongylo doubted by F ORSSLUND, 194 7 but are sup gnatus testaceus are satellite or parasite ported by specimens in the Lund Uni species dependant on their host Tetra versity Museum and the species has re morium caespitum (L.) and have an evi cently been rediscovered in the extreme dently restricted range within the larger southwest of Finland by MERISUO and KA range of that species. PYLA, 1975. This is a distinctly thermo philous species even in South Europe and evidently able to persist still in North Distribution Type D Europe on sun exposed, climatically fa voured sites. Correlations with the broad The sole species segregated in this dis regional and climatic environmental var tribution type is Camponotus vagus Sco- iables analysed in this paper including sunshine hours and latitude are low. This and the very scattered distribution of the species well to the north of its present day abundance would support the sugg estion that C. vagus may be a relict from the warmer sub-boreal climatic optimum. Distribution Type E Two arenicolous species only are m eluded in this distribution type: Lasius camiolicus Mayr and Myrmica gallieni . ,...... Bondroit (Figure 21). M. gallieni is a sand FIG. 20. Distribution Type D: Camponotus vagus dune and salt marsh species with a con (Scopoli) . centration of records in the coastal Baltic 12 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants TABLE 4. Step-wise multiple regression between distribution type C and 11 environmental variables. Cumulative proportion Regression Variable entered of sum of squares reduced coefficient Latitude 0.300 - 1.2385 - 2.688 Annual rainfall 0.362 - '0 .416 -2.120 Length of the growing season 0.400 ---'() 0 8'0 1 7 - 3.226 Boreal vs. oceani·c clima 0.424 0.0334 1.264 Bright sunshine hours 0.443 0.7&27 2.884 Mean annual atmosph. pressure 0.494 1.2085 3.738 Longitude 0.511 l.Ol04 2.236 July mean temperature 0.5_6 -0.7061 - 1.738 Intercept 2.774. Multiple correlation coefficient 0.725 (p< <0.01). recorded under its synonym M . jacobsoni South Switzerland, Austria, the Balcans, Kutter (KUTTER 1972) but occurring more South Italy and the Pyrenees Orientales. locaJ.ly in other parts of Europe. L. car According to the regression analysis the niolicus by contrast is probably a true re only significant variable for this distribu lict recorded only from Gotland with a tion is a negative correlation with inland known distribution in Europe restricted to climate. % presence !IJ > 0.0 Ill > 33.3 • > 66.6 o too 200 ~ •oo soo km FIG. 22. Distribution Type F. ACTA ZOOLOGICA FE NICA 152 13 Distribution Type F This includes most of the commoner species found in the region (Figure 22). The 27 species partitioned off in this group are Myrmica schencki Emery, M. sabuleti Meinert, M. rubra (L.), M. scab rinodis Nylander, M. sulcinodis Nylan der, M. lobicornis Nylander, M. rugino dis Nylander, Leptothorax tuberum (Fab ricius), L. muscorum Nylander, L. acer vorum Fabricius, Formicoxenus nitidulus Nylander, T etramorium caespitum (L.), Camponotus ligniperda Latreille, C. her culeanus (L.), Lasius alienus (Foerster), L. umbratus (Nylander), L. mixtus (Nylan- FIG. 26. M yrmica scabrinodis Nylander. FIG. 2 3. Myrmica schencki Emery. FIG. 27. Myrmica sulcinodis Nylander. FIG. 24. Myrmica sabuleti Meinert der), L. fuliginosus (Latreille), L. niger (L.), L. flavus (Fabricius), Formica rufi barbis Fabricius, F. pratensis Retzius, F. rufa L., F. truncorum Fabricius, F. san guinea Latreille, F. fusca L. and F. ex secta Nylander (Figures 23-49). It is noteworthy that although all are common European species F. truncorum, Camponotus ligniperda, C. herculeanus and L. muscorum are not represented in the British Isles. This is evidently of less importance than the general association with bright sunshine which the regression shows (Table 5) accounts for over 50 Ofo FIG. 25. Myrmica rubra (Linnaeus) of the environmental variables and is 1'4 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants TABLE 5. Step-wise multiple regression between distribution type F and 11 environmental variables. Cumulative proportion Regression Variable entered of sum of squares reduced coefficient Bright sunshine hours 0.504 3.2372 5.185 July mean temperature 0.543 1.5531 1.692 Inland versus island clima 0.568 2.5808 3.235 Length of the growing season ().591 0 .7 837 2.305 Intercept -6.788. Multiple correlation coefficient 0.760 (p< FIG. 30. Leptothorax tuberum (Fabricius) FIG. 28. Myrmica lobicornis Nylander. t ------FIG. 31. Leptothorax muscorum (Nylander). FIG. 29. Myrmica ruginodis Nylander. highly significant for this distribution group. The implication of this particular segregation from the analysis is that the distribution type selects out the species that could be present according to the climatic possibility as well as the species FIG. 32. Leptothorax acervorum (Fabricius} ACTA ZOOLOGICA FENNICA 152 15 actually present. Finland in this analysis is a discriminant area where the British Is les and even Ireland with its reduced fau na are not. The British Isles in fact are the only area in the world within the climatic ranges of the genus Camponotus where species of this genus have never been re corded. The Scandinavian species have differing ecological valencies but C. her culeanus at least should be able to estab lish in plantations in Northeast Britain FIG. 33 . Formicoxenus nitidulus (Nylander) FIG. 36. Camponotus herculeanus (Linnaeus). FIG. 35 . Tetramorium caespitum (Linnaeus) Lasius alienus (Forster) FIG. 35. Camponotus ligniperda (Latreille). FIG. 38. Lasius umbratus (Nylander). 16 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants FIG . .39. Lasius mixtus (Nylander). FIG. 42 . Lasius flavus (Fabricius). through to the more local L. muscorum, M. schencki and C. ligniperda. The in clusion of Northeast Finnmark is probab- 9' FIG. 40. Lasius fuliginosus (Latreille). FIG. 4.3. Formica rufibarbis Fabricius ly related to the relatively higher number of species recorded there than from im mediately adjacent area units through its geographical situation, sheltered from ex posure to western gales and the more in tense summer linking it with more south ern areas of Scandinavia. For example, Lasius flavus, M. rubra and Formicoxenus nitidulus are recorded from Northeast Finnmark but not from adjacent areas FIG. 41. Lasius niger (Linnaeus) and is sometimes imported into the count ry in timber from Eastern Europe. The lower density of species to the north in this group is clearly related to the positive association with summer cli mate and the species included range from the universally distributed Myrmica ru ginodis and Leptothorax acervorum FIG. 44. Formica pratensis Retzius. ACTA ZOOLOGICA FENNICA 152 17 M. scabrinodis, C. herculeanus, L. niger, L. flavus and F. sanguinea have been re cognised from sub-fossil deposits ranging in age between 600 and 4.850 years in South Finland (KoPONEN & NuoRTEVA, 1973). Distribution Type G This is rather similar to the previous FIG. 45. Formica ru/a Linnaeus. distribution but excludes the British Isles entirely (Figure 50). The five species in clude Formica nigricans Emery, F. polyc tena Foerster, F. cinerea Mayr, F. pressi labris Nylander and Myrmica rugulosa FIG. 46. Formica truncorum Fabricius. FrG. 48. Formica fusca Linnaeus. FIG. •1. Formica sanguinea Latreitle. while F. sanguinea occurs throughout Fin land but is rare or unrecorded from Northwest Norway. Several species in this distribution group including M. rubra, M. ruginodis, FrG. 49. Formica exsecta ylander. 18 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants TABLE 6. Step-wise multiple regression between distribution type G and 11 environmental variables. Variable entered Cumulative proportion Regression of sum of squares reduced coefficient Bright sunshine hours 0.!!57 0.4592 3.208 Mean annual atmosph. pressure 0.!!68 --().701!! -!!.957 Latitude 0.422 -0.5605 --2.450 Boreal vs. oceanic cliina 0.442 0.0397 2.!!56 Inland vs. island clima 0.458 tQ.!!.W5 1.651 Intercept 1.620. Multiple correlation coefficient 0.660 (p< <0.01). ...... - ...... FIG. 52. Formica polyctena Forster. ~ presence > 0.0 > 33.3 > 66.6 FIG. 50. Distribution Type G. . ,...... ,.. .,...... FIG. 5·1. Formica nigricans Emery. FIG. 5!!. Formica cinerea Mayr. ACTA ZOOLOGICA FENNICA 152 19 .... IN Me- ..... FIG. 55. Myrmica mgulosa Nylander. Nylander (Figures 51-55). These all have a high concentration in Denmark but are rather more scattered in Southern ~ FrG. 56. Distribution Type H . • - t .... - FIG. 54. Formica pressilabris Nylander. % presence ;g > o.o ~ > 33.3 • >66.6 ,o '00 200 )00 aoo lOO q 20 C. Baroni Urbani &: C. A . Collingwood: The zoogeography of ants TABLE 7. Step-wise multiple regression between distribution type H and 11 environmental variables. Variable entered Cumulative proportion Regression of sum of squares reduced coefficient Longitude 0.271 0.1533 0.465 Inland vs. island clima 0.3'39 1.1010 4.300 Bright sunshine hours ().394 0.1536 0.793 Mean annual atmosph. pressure 0.416 - 0.6649 - 2.823 January mean temperature 0.434 - 0.4983 - 1.746 July mean temperature 0.445 0.3749 1.375 Intercept 0.686. Multiple correlation coefficient 0.645 (p< <0.01). and Central Fennoscandia. F. cinerea in pressure, latitude and boreal versus ocea this region is largely a sand dune species. nic climate are significant variables to According to the regression analysis gether accounting for 44 Ofo of the factors (Table 6) bright sunshine, atmospheric influencing this distribution type. FIG. 57. Formica uralensis Ruzsky, FIG. 58 . Formica forsslundi Lohmander, Fig. 59. Formica tran.s kaucasica Nassonow, FIG. 00. Formica suecica Adlerz, FIG. 61. Harpagoxenus sublaevis (Nylander). ACTA ZQOLOGICA FENNICA 152 21 % presence • > 0.0 ~ > 33.3 • >66.6 o too 2oo :NIO . oo 100 ""- FIG. 62. Distribution Type I. Distribution Type H aggressive species which may account for its restriction to bogland habitats in North This distribution type (Figure 56) inclu Europe contrasting with its recorded ha des Formica uralensis Ruzsky, F. forsslun bits in the Urals, South Siberia and Mon di Lohmander, F. transkaucasica Nasso golia where it is a dryland steppe species. nov (= F. picea Nyl.) F. suecica Adlerz and Harpagoxenus sublaevis (Nylander) (Figures 57-61). The main variables ac cording to the regression analysis are lon gitude, implying a mainly eastern conti nental distribution, inland versus island climate and hours of bright sunshine which together account for over 39 °/o of the factors influencing the distribution of this group (Table 7). Only F. transkaucasica occurs locally in the British Isles and this species together with F. forsslundi and F. uralensis are characteristic of lowland for est bog in Northern Europe. RosENGREN (1969) demonstrated that F. uralensis is a weak competitor, easily displaced by more FIG. 63. Formica lugubris Zetterstedt. 22 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants Scandinavia but found in the higher mountain areas of Central Europe. Distribution Type I This includes the northern boreal spe cies Formica lugubris Zetterstedt, F. aqui lonia Yarrow, F. lemani Bondroit and F. gagatoides Ruzsky (Figures 62-66). F. gagatoides is an arctic species restricted FIG. 64. Formica aquilonia Y arrow F. suecica is endemic to Fennoscandia on ly recorded outside this area from Estonia in the Baltic States. H. sublaevis is a wide ly distributed but rather uncommon spe cies associated with Leptothorax acervo rum and L. muscorum not restricted to FIG. 66. Formica gagatoides Ruzsky . ...... FIG. 67. Distribution Type J: Leptothorax corti calis (Schenck) and Camponotus fallax (Nylan FIG. 65. Formica lemani Bondroit. der). TABLE 8. Step-wise multiple regression between distribution type I and 11 environmental variables. Cumulative proportion Regression Variable entered of sum of squares reduced co effi ci cnt Mean altitude 0.396 1() .7332 3.314 Latitude 0.541 0.6719 4.<>21 Bright sunshine hours 0.617 -0.1707 -U98 Inland versus island clima TABLE 9. Step-wise multiple regression between distribution type K and lol environmental variables. Cumulative proportion Regression Variable entered of sum of squares reduced coefficient Bright sunshine hours 0.071 0.0631 . 2.776 Intercept -0.146. Correlation coefficient 0.266 (p<0.01) . Distribution Type J This includes two species only, Lepto thorax corticalis (Schenck) and Campono tus fallax (Nylander) (Figure 67). None of the variables examined are important in influencing their distribution. Both spe cies are very local in Central Europe, mainly associated with old deciduous woodland and are probably relict species with a discontinuous or mosaic distribu tion north of latitude 50°. C. fallax be ... ,...... comes much more common in South Eu . rope and the Mediterranean area includ FIG. 68. Distribution Type K: Lasius bicornis ing North Africa but L. corticalis is spar (Fiirster) and Polyergus rufescens (Latreille). sely distributed in Europe and is one of the least known of the European Lepto thorax species. to Northeastern Europe and Siberia north of latitude 60° (DLuSSKY, 1967) and not found in the British Isles or Central Eu Distribution Type K rope. The other species are characteristic of woodland and upland areas of North This includes Polyergus rufescens (Lat Britain, the mountains of Central Europe reille) and Lasius bicornis (Foerster) (Fi and are also widely and abundantly dis gure 68). The only significant variable for tributed throughout the northern conife this group is hours of bright sunshine rous forest belt from Norway to Kam which however only accounts for 7.1 °/o chatka. According to the regression ana of the observed variation according to the lysis, the distribution of these species is regression analysis (Table 9) suggesting positively correlated with altitude and la that these also are relict species in North titude and also with bright sunshine with Europe with little relationship to the pre increasing latitude (Table 8). sent environment. P. rufescens is not un- TABLE 10. Step-wise multiple regression between distribution type L and 11 environmental variables. Regression Variat!le entered Cumulative proportion t of sum of squares reduced coefficient Latitude 0.063 -0.1392 -1.978 Length of the growing season 0.109 -0.1039 -2.443 Inland versus island clima 0.156 -0.1486 -2.765 Boreal versus oceanic clima 0.192 -0.0104 -2.344 Mean annual atmosph. pressure 0.206 0.0810 1.769 Bright sunshine hours 0.218 0.0444 1.196 Intercept 1.319. Multiple correlation coefficient 0.421 (p<0.01}. 24 G. Baroni Urbani & G. A. Gollingwood: The zoogeography of ants common in South Europe but L. bicornis, thorax unifasciatus (Latreille), Plagiole as with Leptothorax corticalis above, is pis vindobonensis Lomnicki and Lasius sparsely distributed and one of the least emarginatus (Olivier). These all occur in known Lasius species in Central and South the Channel Islands but L. unifasciatus Europe. Its Scandinavian distribution rests has recently been found also in Oland in with one locality record only in Smaland Sweden. The main variables selected by whereas P. rufescens has a wider distri the analysis include negative correlations bution with several records in South Swe with latitude, length of growing season, den. inland versus island climate and boreal versus oceanic climate (Table 10). All three species are abundantly distributed Distribution Type L in South Europe with main ranges ex tending into Northern France and Central This distribution type includes Lepto- Poland. IV. THE BIOGEOGRAPlflCAL AREAS The interrelations between the area shown by the dendrogram illustrated in units on the basis of their ant distributions Figure 70. Two levels of similarity have were studied by using principal compo arbitrarily been selected for our purpose: nent analysis. An arrangement of the area the 50 Ofo and 75 Ofo similarity levels. units according to the three major axes At the 50 Ofo similarity level, the stud is represented in a three dimensional ied area divides straightforwardly into model, Figure 69. This allows the identifi north and south on the basis of numbers cation of grouping corresponding with the of species present. At this partition, the major geographical regions such as the southern zone includes the whole of the British Isles, Denmark and Fennoscandia. British Isles exclusive of the Northern It is to be noted that all the Fennoscan Islands, Denmark, South Norway up to dian localities show little variation ac and including Hedmark, Opland, Buske cording to Factor Ill (14.9 °/o contribu rud and Hordaland, South Sweden up to tion) while all the British area units are and including Medelpad, Halsingland strongly positively correlated with it. All and Dalarna, and South Finland includ the units at the top of the model corre ing Karelia borealis, Savonia borealis and spond to areas within the British Isles Ostrobottnia media, Figure 71. with the sole exception of the faunistical In this southern zone most of the cha ly poor North Isles. The Danish areas racteristic species of distribution type F appear relatively close one with another are present, notably F. rufa, F. fusca, L. and are characterised by their common fuliginosus, L. mix.tus, L. umbratus, L. positive correlation with Factor I (44.1 Ofo niger, L. alienus, M. sabuleti, L. musco contribution) and a negative correlation rum, L. tuberum and T. caespitum. With with Factor II (27.2 Ofo contribution). a few exceptions these are all absent from However no general criteria for isolat the northern zone and all are widely ing these groupings in a standard way distributed in Denmark and Southern was found and despite the implied Fennoscandia and with the exception of loss of information we preferred the sing L. muscorum are also present in the Bri le link cluster analysis for a clearer re tish Isles. This division corresponds ap presentation of the most characteristic proximately with that between middle groupings of the area units. These are boreal and south boreal climates as defin- ACTA ZOOLOGICA FENNICA 152 25 -1 FIG. 69. Arrangement of J{).'l area units along .'l axes from similarity matrix analysis of ant fauna. Bri tish areas marked by dot, Danish by triangle, Fennoscandian areas unmal'ked. ed by KALLIOLA (1973) on the basis of mani and F. gagatoides are characteristic vegetation types. In Finland this is near and present with the more generally dis the northern limit of Formica rufa and tributed M. ruginodis, M. sulcinodis and F. pratensis and of the southern pine zone L. acervorum. No species however is en (WUORE RINNE 1975). tirely restricted to this zone and even the In the northern zone, the north boreal arctic F. gagatoides overlaps the two zones species represented in distribution type I in the mountains of Opland and Horda including F. aquilonia, F. lugubris, F. le- land in Norway, although the distribution 26 c. Baroni Urbani & c. A. Collingwood: The zoogeography of ants "SIMILARITY I~ lg lg Jel I~ 18 lg : CNN'wiiWndS ll'lniMUh i '"-' i c...... '"""S.WIIt1 !: ..... ! ...... lll... i NorthWnlJutl.lnd WHt JutiMd !: S«rlt!JutiM'Id : : ...... '"""" i (.st.Jutllnd..... ! , i ...... i -~ ! ...... """ --IM:Shlaustnfit I ...... ,...... : - s.... liiUStnfis -JIDrttllat JuiMit -ltnlil &l.strlfb ...... --- ....., ~~h tld'StW!Iol'alh- ICM'Itirlllllmfh ·-,,_.. Q. Osl.fd ...uustnn - 0 ·-...... "' 1:0 --.. ·a~ -lilllti\Wtstl~ ,...., «<"' - u...... «< ...... ·- 0 E ...... _ «< ·--- ... -..... 1:0 0... '"0 L._- .,c:l -..._.W.lMintJ; A -,_...... 0 L-..__'"" ci ·· ~" ~ ...... t.llnd,.F ,_ Ylt , Mln -·- OJinllttaliiOII'tllfis.M ,.rt -...~blnllls,.S _ pwt .,_ '-- lJd.Sitl~ A..oo- ·-,--.a...,..,.... ._ ,...._ ...... --·~"-"* .• ...... , --...,...... ftna.lft i i I I~ lg I~ l::l I ~ la lo 3:)11311~10" ACTA ZOOLOGICA FENNICA 15~ 27 lOO 200 300 400 lOO km FIG. 71. Map of area un-its. Partitioned at 5{) 0/o similarity level. of this species is mainly within the boun species which are more common in South England daries of the north boreal climatic area as than in South Scandinavia and are not found in illustrated by KALLIOLA (1973) . the British Isles to the north of the demarcated At the 75 Ofo similarity level a total of area. twenty-one biogeographical areas are se 3. Rest of the British Isles (excluding the parated as shown in Figure 72. These in orth Isles) - this area corresponds exactly with clude the following areas each of which is the first partition of BARONI URBANI & COLLING more or less characterised by the presence wooo ( 1975) from their analysis of the British or absence of certain key species. vice-county distribution. The area includes a rela tively impoverished fauna and lacks some species l. Channel Isles - these are clearly separat that are widely distributed in Northern Fenno ed by the common presence of P. vindobonensis scandia including F. truncorum, F. gagatoides and and L. emarginatus which are entirely absent C. herculeanus. from the rest of the area studied. This partition 4. The orth Iseles and North Tr0ndelag - corresponds with distribution type L except for these two area units are very poor in number of L. unifasciatus which also occurs on Oland. species with only L. acervomm and F. lemani re 2. South England - this area includes Penin corded from both areas and C. herculeanus and sula, Channel, Thames and Anglia corresponding F. aquilonia present in North Trondelag but ab in part with distribution type C. The area includes sent from the North Isles. Ponera coarctata which is not present in Denmark 5. Northwest, West and South Jutland - or Fennoscandia and also many other southern these have a number of species in common but 28 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants lack several species found in East Jutland includ 8. Northwest Zealand - Formica and Lasius ing M. rugulosa, F. nitidulus, L. mixtus, C. ligni species are well represented but only M. rugulosa perda, F. cunicularia, F. polyctena and F. aquilo of the genus Myrmica has been recorded. nia. 9. Funen and ortheast Zealand - these 6. South Fennoscandia and E. Jutland - this have in common the presence of H. punctatissima area includes a large part of South Sweden includ from outdoor locations and F. nitidulus, a rather ing Blekinge, SHine, Siidermanland, Upland, Os local species in Denmark. tergiitland, Narke, Vastergiitland, Smaland, Bo IQ. Lolland, Falster, Miin - these islands huslan and South Finland including Alandia, have no representatives of the F. rufa group apart Regio aboensis, Nylandia, Tavastia australis, Sa from F. truncowm but are well represented by vonia australis and Karelia australis. This area Lasius and Myrmica species including the local has a rich fauna with most of the less co=on M. gallieni and M. specioides. These two species species found in Scandinavia including F. cinerea. have also occurred at Taktom in ylandia but are 7. Bornholm - this island group has a poor not normally found together in their main areas fauna which has probably not been fully investi of distribution. gated. There are no records for Myrmica species 11 . South Zealand - this has a rather poor other than M. schencki but the uncommon Aner fauna but includes the uncommon Stenamma gates atratulus is present with its host T. caespi westwoodii (SK0TI 1973). tum (BISGAARD 1942). Other species recorded in 12. Oiand and Gotland - these have several clude L. tuberum, F. trunco rum, F. rufibarbis, L. rare species including C. vagus, D. fugax and T. (uliginosus, L. rabaudi, L. fla vus and L. niger. erraticum. Each island has in addition a number FIG. 72. Map of area units. Partitioned at 75 °/o Similarity Level. ACTA WOLOGICA FENNICA 152 29 of uncommon species separately including M. gal F. lemani as well as the more southern F. rufa, lieni, L. interruptus, L. unifasciatus, Strongylogna F. pratensis and F. fusca. Lasius species are poor thus testaceus and L. carniolicus. ly represented with only L. flavus and L. niger 13. Gotska Sandon - this also has a peculiar recorded from all area units, L. mixtus from Hor fauna with few Lasiw or Myrmica but including daland and Rogaland, and L. fuliginosus from the the very local L. interruptus and L. nylanderi in Finnish areas. common with Gotland. 19. Vestfold, Opland, Dalsland, Viistmanland, 14. Giistrikland, Aust-Agder, Vest-Agder - Hedmark, Buskerud, 0stfjold and Akershus, these have F. rufibarbis in common but lack M. Viirmland, Dalarna, Medelpad - this area has schencki and L. muscorum found in adjacent records for most of the more local Formica species areas. including F. suecica, F. pressilabris, F. forsslundi 15. Telemark - this has no records for seve and F. uralensis. ral common species of Myrmica and Formica 20. Hiirjedalen, Jiimtland, Angermanland, known from adjacent areas. Viisterbotten, Lycksele Lappmark, Asele Lapp 16. M0re og Romsdal, Sogn og Fjordane - mark, Ostrobottnia kajanensis, Os trobottnia bo these have both C. herculeanus and C. ligniperda realis, Kuusamo, E and N Finnmark - most of as well as the north boreal species included in the commoner Myrmica species are recorded in Distribution Type I but are poor in Myrmica this area, also F. nitidulus, either or both L. flavus species. and L. niger, F. uralensis, F. aquilonia, F. sangui 17. S0r Tr0ndelag - the common speci-es nea and other common northern Formica species. M. ruginodis and C. herculeanus are not recorded. 21. Lule Lappmark, Tome Lappmark, Pite This is a poorly represented area with records Lappmark, South Nordland, North Nordland, only for F. aquilonia, F. gagatoides, F. lemani, Troms, East and West Lapponia kemensis, Lap L. flavus, M. rubra, M. sulcinodis and L. acer ponia inarensis, Lapponia enontekiensis - this vorum. northern area is characterised by the complete 18. Hordaland, Rogaland, Hiilsingland, Savo absence of Lasius species and a restricted fauna nia borealis, Ostrobottnia australis, Tavastia bo including M. ruginodis, M. sulcinodis, M. lobi realis, Karelia borealis, Satakunta - this is evi comis, L. acervorum, C. herculeanus, F. lugubris, dently a transitional zone with several northern F. gagatoides, F. lemani, F. truncorum and F. boreal species such as F. aquilonia, F. lugubris and exsecta. V. DISCUSSION It is of interest to demonstrate briefly phenomena relating to the North Euro how the distribution types recognised con pean ant fauna should be mentioned. The tribute to the fauna of each of the biogeo British Isles within the regions inhabited graphical areas separated out by our ana by ants are unique as a relatively large lysis (Figure 73). Since both distribution area entirely without indigenous species types and biogeographical areas are de of the worldwide genus Camponotus. rived through two different approaches However, there is at least one, probably from the same data matrix, conclusions credible, record of a fossil species describ cannot be rigidly drawn because of the ed from the Oligocene of the Isle of implied circular reasoning. However, the Wight, C. brodei, referred to the tropical species groupings obtained show some sig temperate subgenus Colobobsis (DoNI - nificant correlations with the environ THORPE 1920). The genus Camponotus is mental variables used and it seems rea assumed to have been wiped out of the sonable to regard them as natural sets of British Isles and Northern Europe during some biological significance. the Pleistocene glaciations and to have First the more peculiar and well known repopulated Northern Europe after the . . ~ ~ 0 0 ...... 0 0 ~ ~ 0 0 "' "' 0 0 1:1" 1:1" ~ ~ "" "" ...... c::: c::: ;;· ;;· n n ~ ~ 0 0 0 0 n n ~ ~ :s :s [ [ "' "' SI:> SI:> .. .. 0 0 ""' ""' 0 0 ...... "" "" ...... 0 0 b:l b:l .. 0 0 00 00 :. :. ~ ~ 0"1 0"1 "d "d '< '< 0"1 0"1 • • 52, 52, , 2 2 51 , , - 1,10 ,SO 10 , 49 49 34 34 I I 100 , 83 unitl unitl unit unit 4 . 21 21 14 14 81,82, 81,82, 103 103 Area Area Area I 17 17 1 1 , 9'/ 75, 75, 96 96 , 71f, 71f, , , 80 , 53 2S,28,Sif,SS, 79 I I 1 78 t t unitl unitl units units 17 17 t t 6 6 13 13 20 20 98,99 98,99 76 76 Area Area Area Area t I I 27 27 1 1 H H 24,26, 24,26, FG FG I I E E its its un CD CD 8 8 5 5 19 19 12 12 Area Area I I LA LA , , K K J J types types HI HI ,48 ,48 43,44,72 22 31 31 FG FG I I E E unit unit unit• unit• units units CD CD 91,92,93,91t,95 91,92,93,91t,95 , 8 8 18 18 11 11 Area Area 88 Area Area Are• Are• Distribution Distribution 14 14 LA LA ' ' K K J J I I 9,10,11, 9,10,11, '17,18 ,16 ,16 7 , 8 , 47 47 I I • • E E F G H 21 21 unit unit , units units CD CD 20 , 17 17 8 8 3 3 10 10 Area Area 19 12,13,14,15 12,13,14,15 Area Area LA LA K K J J '11 '11 , HI HI 45 1,2,3,1.l,S,6 1,2,3,1.l,S,6 I I E E F G nits nits unite unite u CD CD 2 2 9 9 8 8 16 16 Area Area Area Area LA LA K K J J HI HI 23 23 32 32 ltO ltO FQ FQ I I E E nit nit u unit unit unit unit CD CD ea ea ea 1 1 8 8 8 8 15 15 Ar Ar Area Area A A % % 10 10 110 110 <40 <40 10 10 :zo :zo 10 10 :zo :zo :zo :zo 110 110 110 110 <40 <40 <40 <40 . . · · · · · · c; _, _, t::l t::l 0 0 c:: c:: ~ ~ ...... n n I:S I:S c;· c;· I:S I:S 0 0 a: a: ...... s s ~ ~ a; a; s. s. I:S I:S .. .. ...... I"> I"> ...... ;;; e: e: 0 0 I"> I"> o. o. r:r r:r 0 0 n n 0 0 1:1.. 1:1.. ::; "' "' n n ~ ...... n n "' "' !'l !'l ~ ~ ...... Er Er n n ...... " " 0 0 11> 11> 1:1" 1:1" .. .. "' "' ...... n n ;;· ;;· e. e. "' "' s· s· !" !" 0"1 0"1 "0 "0 '< '< 0"1 0"1 0"1 0"1 "0 "0 0"1 0"1 "d "d ACTA ZOOLOGICA FENNICA 152 31 separation of the British Isles from the tically significant E-W pattern implying European continent (EMERY 1920). The more or less important correlations with last land connection between England and longitude but most of the species of these the continent corresponds to the Dogger groups are present also in South England Bank formation dating back to at least at least. However, distribution type G 5 000 years ago (LrNDROTH 1935). This (Figure 50) shows a purely Scandinavian fits with the analysis of subfossil insects pattern and includes 7.5 Ofo of the studied recovered from deposits at Piilonsuo in species· These species show a typical South Finland by KoPO NEN & NuoRTEVA north-south distribution with factors such (1973) who found Camponotus herculea as sunshine and latitude contributing to nus in deposits up to 4 000 years in age the regression. Yet the absence of the spe but not older, while ant remains older cies of this group presents the main pecu than 5 000 years were not detected. Since liarity of the British fauna which is thus at least one very common Camponotus characterised more by its absences than species (C . mengei Mayr) with over 100 by the presence of significant species. specimens has been described from the Baltic amber of the lower Oligocene There are at least three well known and (WHEELER 1914) it seems reasonable to documented attempts to explain the pre conclude that the genus was widespread sence of some organisms in North Europe in North Europe during tertiary times by a pre-glacial or inter-glacial origin and disappeared during the Pleistocene (LrNDROTH 1935, 1953; HoLDHAus 1954). glaciations, recolonising the area only Probably the best known of these is the during the fairly warm sub-Boreal period. work of HoLDHAUS (op. cit.) on the boreo However the most interesting result alpine distribution type. This term is ap from our analysis is that the four Campo plied to many different organisms regu notus species now living in North Europe larly present in North Europe and on the pertain to three different distribution typ mountains of Southern Europe, in parti es. Thus there is not a single "Campo cular the Alps, and absent from or more notus pattern" which excluded the British local in Central Europe. Such a distribu Isles from its colonisation range but the tion pattern is shown also by some ant genus repopulated North Europe through species as Formica uralensis, F. aquilonia at least three different paths and there and F. lugubris. This disjunction is sup seems no reason to postulate a similar posed to have originated during the Riss reaction to past or present environmental glaciation although for some organisms situations for such ecologically disparate the northern part of the area will also species as C. herculeanus, C. vagus and have been recolonised in post glacial tim C. fallax or for their nearest hypothetical es. (DE LATTIN 1966). However to postula ancestors. te non glaciated refugia in North Europe Another result from our analysis is during Pleistocene glaciations for animals worth comment. Both from the principal with a high potential dispersal capacity component analysis and from the cluster such as ants seems unnecessary. Many analysis, the British Isles are separated as such organisms indeed supposed to have the largest and most characteristic area had a Pleistocene origin fit at least equal within the boundary of our study with the ly well the contemporary distribution of exception of the myrmecologically poor the alpine climate as mapped by LANDS est North Isles. Despite this there is no BERG et al. (1965). distribution type peculiar to this area and The second distribution type of sup entirely missing from Scandinavia and posed Pleistocene origin in North Europe only one indigenous species, Ponera co has been termed by LrNDROTH (1935) as arctata, but no endemic species known on Boreo-British. Such species, of Wiirm ori ly from the British Isles. Some of the dis gin, should be restricted now essentially tribution types separated out such as dis to the British Isles and to Scandinavia but tribution types C, H and I have a statis- none of the North European ant species 32 C. Baroni Urbani & C. A. Collingwood: The zoogeography of ants shows clearly enough a pattern of this period of activity and food resources are type. relatively limited. We have already suggested that the There is evidence that substantial areas peculiarities of the ant fauna of Oland of the Norwegian coast constituted ice and Gotland where climate or other fea free refuges during the Pleistocene gla tures do not contrast sharply with their ciations on which several species of plants nearest continental lands, can be explain and animals should have survived (PE ed by interspecific competition favouring TERSEN 194 7; LINDROTH 1953). However the first chance arrivals on a limited in these same regions are still today under sular surface; a similar explanation may the beneficial influence of the Gulf apply to the presence of F. rufibarbis on stream, yet none of the studied ants shows the Scilly Isles contrasting with its total a preferential distribution there. Although absence from the nearest British main a relatively high species concentration in land where the similar F. cunicularia is Northeast Norway comes out clearly in abundant. The pre-existence of any one both the Q and R analysis this is more number of a species group of a dominant likely to be an expansion of the richer genus such as Formica would be prejudi eastern continental fauna into a relatively cial for the settlement of another member favourable area than a relict fauna of of the same species group. Pleistocene origin and no similar pattern Moreover the presence of a given spe is present on the more beneficially expos cies in a given locality clearly implies the ed west coast. Although it is possible to existence of contemporary appropriate suppose that the most typically north environmental conditions independently boreal elements included in our analyses of the origin of that species. There is evi (Distribution type I, Figure 62) could have dence that the ant geographical picture survived in ice free refugia, there is not for the more observable species of the enough evidence to prefer this hypothesis F. mfa group is continuously changing in to the simpler explanation of a post Pleis response to man made environmental tocene colonisation from the south. changes (HuGHE 1975· BREE 1976). It should be mentioned here that ant At the same time in the British Isles zoogeography has some peculiarities in the more local species appear to have that the biological unit is not the individ been rather stable over the past 100 years ual but the colony which is relatively non in relatively unaltered biotopes (CaLLING motile in space and usually restricted to a wooD 1971 ). The results of our regressions well defined foraging territory (BRIA demonstrate the important role played by 1965). On the other hand ant dispersal environmental factors such as sunshine capacity though differing for different and summer temperature and despite the species may be relatively great at the time limitations of the large generalisations on of the nuptial flight and over periods of homogeneity within our area units we hundreds of years most species would have have been forced to make, this study can a potential to colonise all available free be regarded as a first rough approach to spaces within a region. However, compe evidentiate and explain phenomena of tition, including defence of a territory for this type. Considering the known conti food exploitation (DwssKY 1965) and nuous variation of environmental factors predation of mated females, would favour in time, there seems to be little need to the first colonisers of a given area. Such search for explanations for the present species would play a dominant role day ant population in North Europe bey through North Europe where the annual ond a few thousand years. ACTA ZOOLOGICA FENNICA 152 33 VI. SUMMARY A numerical analysis of the 67 indige The same numerical analysis carried nous ant species mapped on 103 area units out on the presence-absence of species per was carried out to describe and to attempt area unit permits the recognition of 21 a general interpretation of the population biogeographical areas clustering at the of Northern Europe. Northern Europe is 0. 75 similarity level. These results, sup regarded here, for purely practical rea ported also by principal component ana sons, as including the British Isles, Den lysis of the same similarity matrix, indi mark and the whole of Fennoscandia up cate the whole British Isles (with the to the political boundary with Russia. single exception of the myrmecologically Twelve sets of species distributions have poorest North Isles) as the most peculiar been recognized for clustering at a simi and largest homogeneous biogeographical larity level ~0 . 5 and are regarded as re area within the studied territory. The presenting the major distribution patterns reasons for this peculiarity are mainly due of the North European ants. Multiple re to absences than to the presence of ende gression analyses were calculated between mic or of particularly significant species. the number of species per area unit of The best known absence, i.e. that of the each distribution type and 11 environ world distributed genus Camponotus, is mental varia'bles coded into major inter explained with different reasons for at vals assumed to be uniform within a unit. least 3 of the 4 known North European Some results showed high correlation va species and the hypothesis of a homoge lues with environmental variables such as neous Camponotus colonization in North annual number of bright sunshine hours, ern Europe excluding the British Isles July temperature, relative oceanicity and through some common dispersal pattern length of the growing season which are is rejected. regarded as the dominant factors in de Some highly characteristic faunae, like termining the general picture of ant dis those of the isles of Oland and Gotland tribution in Northern Europe· One distri are explained as probably due to historic bution type (type F) includes 27 species al factors such as the competitive advan and shows a decreasing correlation with tages of the first randomly arrived colo the previously mentioned 4 environmental nizers on a limited insular surface in post variables in that same order. 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CoLLINGwooo: T he z oogeography of an ts (Hymenop tera, Formicidae) in orthern Eu rope. 34 pp. (1 977) . Helslngtn Yllop ..... t -n etsfikt~a t Exrhange - Attslarurh - Erhange SOCIETAS PRO FAUNA ET FLORA FENNICA Snellmaninkatu 9-11 Snellmansgatan 00170 Helsinki 17 Helsingfors Fo,. sale - Ve,.kartf - En vent Akateeminen Kirjakauppa - Akademiska Bokhandeln 00100 Helsinki 10 Helsingfors ISBN 951-661-019-6 ISSN 0001--7299 PRINTACO