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Survey of Social Insects in the Fossil Record.
Psyche 85(1):85-133, 1978.
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SURVEY OF SOCIAL INSECTS
IN THE FOSSIL RECORD*
BY LAURIE BURNHAM
Museum of Comparative Zoology,
Harvard University,
Cambridge, Massachusetts 021 38, U.S.A.
Biologists have long been intrigued by the complex social systems of various insects. Despite a voluminous literature dealing with the evolution of these systems, immense gaps remain in our understand- ing of insect sociality. Several theories have been proposed to explain the evolution of social behavior in certain groups of insects (e.g., Hamilton, 1964), but none consider this problem with respect to geological time. The present paper does so by examining the fossil record for clues not only on the antiquity of sociality, but also on the nature of these early social insects. Included in this survey are those insects recognized as eusocial: the Isoptera, and three super- families of the Hymenoptera: Vespoidea, Formicoidea, and Apoidea.
ISOPTERA
The termites are remarkable in two regards: 1) as a group, they are fully eusocial, exhibiting a wide range of behavioral modifica- tions and sophistications, and 2) their record in the geological past, although sparse, is highly indicative of an Early Mesozoic origin. This latter point is of particular significance if one considers sociality among insects as a pinnacle of evolutionary success. Wilson (1971, p. 1) states that "[insect societies] best exemplify the full sweep of ascending levels of organization, from molecule to society." The possibility that termites evolved a social organization as far back in geological time as the Jurassic (roughly 190 million years ago) is of great interest, particularly when attempting to draw parallels with the evolution of sociality in the Hymenoptera, a group phylogenetically very remote from the termites. *Manuscript received by the editor July 7, 1978. 85
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86 Psyche [March
Five of the six families' of termites recognized by Emerson (1955) have a fossil record extending at least as far back as the Tertiary. In 1967, Cretatermes carpenter! (Hodotermitidae) was found in an Upper Cretaceous deposit in Labrador (Fig. l), a discovery which immediately placed the origin of the Isoptera no later than the Mesozoic - an extension of 45 million years from previously known specimens. In addition, the advanced phylogenetic position of Cretatermes provides evidence for a much earlier origin of the order than has formerly been recognized (Emerson, 1967). An examination of various fossil localities reveals a widespread termite fauna during the Tertiary Period (Table 1). The Termitidae are found in Miocene deposits of California and Germany; the Rhinotermitidae, Hodotermitidae, and Kalotermitidae are found at various Tertiary deposits throughout the United States and Europe; and the Mastotermitidae have the most widespread Cenozoic distribution of all, having been found at localities in the United States, Europe, South America, and Australia. This latter finding is highly intriguing because the family Mastotermitidae today has but one species, Mastotermes darwiniensis, which is restricted to north- ern Australia.* Emerson (1955) postulates that this widespread 'The sixth family is the Serritermitidae - an aberrant taxon known from only one species.
*A look at past climatic shifts provides additional insight into the redistribution of the termites, particularly with respect to the Mastotermitidae, now solely restricted to Australia. Reconstructions of paleo-climatic patterns may be made fairly accurately on the basis of floral analyses (Reid and Chandler, 1933). The presence of Sequoia stumps in the Florissant Shales of Colorado provides evidence for warmer tempera- tures during the Oligocene (Emerson, 1969). Tiffney (1977) postulates on the basis of fossil angiosperm assemblages that temperatures in New England during the Oligocene were much more equable than at present - the temperatures ranging from 26OC to 9OC in contrast to today's 21å¡ to -lOå¡C Furthermore, extended frosts and hard freezes were unknown. In the more tropical climate of the Oligocene, 'colony activities were presumably carried out year round in a relatively warm, moist environment, explaining the widespread distribution of the Mastotermitidae during the Lower to Middle Tertiary. By the Late Miocene or Early Pliocene, the earth's climate began shifting towards cooler temperatures with the rising level of the continental land masses and increasingly large polar ice caps. My hypothesis is that, unable to adapt to an increasingly colder climate, and possibly to a concomitant change in predator pressures, the Mastotermitidae began to die out during the Tertiary. And, because at this time the Termitidae were undergoing tremendously successful radiation in Africa and South America, the Mastotermitidae became geographically restricted to northern Australia, represented today by only one relict species, Mastotermes darwiniensis.
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19781 Burnham - Social Insects in Fossil Record 87 Figure 1.
Cretafermes carpenters Emerson from lower part of Upper Cretaceous of Labrador. Note humeral suture at wing base. Original photograph of holotype in Prince~on Museum. Length of wing, 7.5 rnm, geographical distribution provides strong evidence to support a Mesozoic origin of the order. He argues (1975) that the breakup of the united land mass Pangaea in the Permian or Lower Triassic must have occurred subsequently to the origin of the Isoptera to explain their distribution in the southern and northern continental land masses and that all five families must have been present in the Late Mesozoic to explain their diversity and distribution by the Tertiary.
In 1971 he looked at a variety of primitive and derived characters of each family and analyzed the geographical distribution of the groups, using plate tectonics to provide the following estimates on the geological origin of the families:
Mastotermitidae - possibly Early Mesozoic. Hodotermitidae - Triassic, or Early Jurassic before the breakup of southern continents.
Kalotermitidae - mid-Jurassic, or Lower Cretaceous, before the separation of Africa and South America.
Rhinotermitidae - Late Jurassic, Early Cretaceous. Termitidae - Cretaceous.
Because termites are such poor fliers and do not mate until the adults have cast their wings, he considers water gaps of more than 50 miles capable of preventing termite dispersal. While I am supportive of the theory that places great importance on the role of a unified land mass in animal dispersal, I do not agree that this can effectively be used to date the origin of the Isoptera.
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TABLE 1 ISOPTERA IN THE FOSSIL RECORD.
Geological Age
CRETACEOUS
Hodotermitidae
* Cretatermes carpenteri Emerson
EOCENE
Mastotermitidae
*Blattotermes wheeleri \Collins
*Zdomastotermes myslicus Ha up t
Kalotermitidae
Neotermes grassei Pi ton
Hodotermitidae
Termopsis mallaszi Pongracz
OLIGOCENE
Mastotermitidae
*Miotermes insignis (Heer)
*Miotermes spectabilis (Heer)
Mastotermes bournemouthensis von Rosen
Mastotermes heeri (Goppert)
Mastotermes balheri von Rosen
Kalotermitidae
* Prokalotermes hageni (Scudder)
*Electrotermes giradi (Giebel)
*Electrotermes affinis (Hagen)
Kalotermes rhenanus Hagen
*Eotermes grandaeva Statz
*Proelectrotermes berendti (Pictet)
Locality
Labrador, Canada
Tennessee, U.S.A.
Geiseltal, Germany
Menat, France
Hungary
Oeningen, Germany
Oeningen, Germany
England
Schlesien, Germany
England
Florissant, Colorado
Baltic Amber
Baltic Amber
Rott, Germany
Rott, Germany
Baltic Amber
References
Emerson, 1967
Emerson, 1965
Emerson, 1965
Emerson, 1969
Snyder, 1949
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, I969
Emerson, 1969
Emerson, 1969
Emerson, I969
Emerson, I969
Emerson, I969
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Hodotermitidae
Archotermopsis tornquisti von Rosen
Termopsis bremii Heer
* Parotermes insignis Scudder
* Parotermes scudderi Cockerel1
* Ulmeriella bauckhorni Meunier
* Ulmeriella cockerelli Martynov
Rhinotermitidae
*Reticulitermes minimus (Snyder)
Reticulitermes fossarum (Scudder)
Reticulitermes antiquus (Germar)
Reticulitermes creedei Snyder
* Parastylotermes robustus (Rosen)
MIOCENE
Mastotermitidae
*Spargotermes costalimai Emerson
Mastotermes vetustus Pongracz
Mastotermes minor Pongracz
Mastotermes haidingeri (Heer)
Mastotermes croaticus von Rosen
*Miotermes procerus (Heer)
* Miotermes randeckenensis von Rosen
* Pliotermes hungaricus Pongracz
Kalotermitidae
Cryptotermes ryshkoffi Pierce
Kalotermes swinhoei (Cockerell)
Kalotermes tristis (Cockerell)
Kalotermes nigrit us Snyder
Baltic Amber
Baltic Amber
Florissant, Colorado
Florissant, Colorado
Rott, Germany
Siberia, U.S.S.R.
Baltic Amber
Florissant, Colorado
Baltic Amber
Creede, Colorado
Baltic Amber
Brazil
Radoboj, Croatia
Radoboj, Croatia
Radoboj, Croatia
Radoboj, Croatia
Radoboj, Croatia
Wiirttemberg, Germany
Radoboj, Croatia
Calico, California
Burma
Burma
Chiapas, Mexico
Snyder, 1949
Snyder, 1949
Snyder, 1949
Cockerell, 19 13
Emerson, 1968
Emerson, 1968
Emerson, 197 1
Emerson, 197 1
Emerson, 197 1
Emerson, 197 1
Emerson, 197 1
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1965
Emerson, 1969
Emerson, 1969
Emerson, 1969
Snyder, 1960
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Geological Age
TABLE 1. (CONCLUDED)
Locality
MIOCENE (continued)
Hodotermitidae
* Ulmeriella latahensis Snyder
* Ulmeriella martynovi Zeuner
Rhinotermitidae
Heterotermes primaevus Snyder
Reticulitermes hartungi (Heer)
Reticulitermes laurae Pierce
* Parastylotermes calico Pierce
*Parastylotermes washingtonensis (Snyder) Termitidae
Gnathamitermes magnoculus rousei Pierce
Macrotermes pristinus (Charpentier)
Latah, Washington
Biebrich, Germany
Chiapas, Mexico
Radoboj, Croatia
Calico, California
Calico, California
Latah, Washington
Calico, California
. Radoboj, Croatia
References
Emerson, 1968
Emerson, 1968
Emerson, 197 1
Emerson, 1971
Emerson, 197 1
Emerson, 1971
Emerson, 197 1
Pierce, 1958
Snyder, 1949
*Extinct genera.
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19781 Burnham - Social Insects in Fossil Record 9 1 Simpson (1952) has made some insightful remarks on the matter. He contests the premise that if a given group of organisms requires a land connection, then disjunctive areas occupied by the group must have been once connected by continuous land. His contention is that there is no group of organisms that cannot be dispersed over water. Given a probability of only one chance in a million that an organism can cross a stretch of water, when geological time is considered the chance that the event will actually take place (over tens of millions of years) becomes significantly greater. It is further argued that successful colonization is dependent on successful invasion and the ability of the intruder to compete with existing species. Chances for survival are much higher when there are numerous, simultaneous arrivals of individuals. In my opinion, the termites support such reasoning, and this can be argued in several ways. Firstly, termites are relatively light- bodied, winged insects. Studies by Simberloff and Wilson (1969) and Glick (1933) on the repopulation of an island by wind trans- ported insects strongly support the possibility that termites are capable of being carried considerable distances in the upper atmos- phere. Furthermore, because termites swarm in such large numbers prior to reproduction, a reasonable possibility exists that they will be dispersed to a new habitat as either a group or at least as a malelfemale pair. A wind current strong enough to blow one individual into the upper atmosphere should be equally capable of carrying multiple individuals, and, according to windflow, of trans- porting them in the same directional pathway. Secondly, termites are ideally suited to dispersal over large bodies of water via floating logs. The more primitive families construct their extensive nesting colonies in wood and logs; as a consequence, it is entirely plausible that a dead tree falling into a body of circulating water could be carried extended distances. Furthermore, this mode of transportation provides the termites with a source of food during their sojourn, and travel en rnasse obviates the prob- lems of reproduction upon arrival. In addition, as Simpson points out, the larger the number of individuals, the more likely it is that they will be successful competitors in the new habitat. I am not presenting this as evidence that the termites did not evolve while the earth's land masses were still contiguous, but am merely pointing out the problems in arguing that land dispersal was essential for termites.
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92 Psyche [March
The Isoptera exhibit strong affinities to the Blattodea; evidence linking the two groups to a common ancestor is well marked between the Mastotermitidae, an archaic termite family, and the Cryptocercidae, a family of generalized cockroaches. This theory of common ancestry is supported by several comparative morphologi- cal and behavioral studies (Emerson, 1965; McKittrick, 1965; Ahmad, 1950; Cleveland, 1934; Hill, 1925). McKittrick (1965) goes so far as to incorporate both groups into the Dictyoptera, an order which also includes the Mantodea. The gut fauna, female genitalic structures, anal expansion of the hind wing, morphology of the proventriculus, and deposition of eggs in ootheca-like masses are much alike in Mastotermes and Cryptocercus. Furthermore, both groups inhabit similar habitats. As a consequence, termites have often been referred to as merely social cockroaches. This degree of relatedness becomes immediately interesting in view of the extensive geological record of the cockroaches.
Fossil cockroaches are first found in deposits from the Upper Carboniferous, which makes them among the oldest insects known. Furthermore, they comprise 80 percent of the fossil insect fauna during that period (Carpenter, 1930) - an indication that they have not only existed, but have flourished, for three hundred million years. If the similarities between termites and cockroaches are indeed the result of monophyletic, rather than convergent or parallel evolution, one might speculate on a much earlier origin for the Isoptera than is shown by the fossil record. McKittrick (1965) admits that the flagellate gut fauna essential for cellulose digestion in both groups may have arisen independ- ently in each; however, she believes that the similarities in two important morphological characters, the female genitalia and the dental belt of the proventriculus, represent primitive characters and are therefore indicative of a common origin for Mastotermes and Cryptocercus. On the other hand, Tillyard (1926, 1936), Cleveland (1934), Imms (1 9 19), Carpenter (personal communication), among others, believe that the termites were derived from more ancient stock and may have evolved during the Late Paleozoic. Hamilton (1978) supports the view that social termites arose from "roach-like ancestors" in the habitat of dead phloem, and suggests that the invasion of Cryptocercus into the same typeof habitat was inde- pendent of the ancestral termite. The possibility of termite bbevolu-
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19781 Burnham - Social Insects in Fossil Record 93 tion under bark" seems immensely feasible; not only is isolation (and, hence, inbreeding) possible, but selective pressures leading to dependence on a cellulose diet would also be high. It seems an excellent explanation for the early separation of the termites and cockroaches from a common protorthopteran (protoblattoid) an- cestor as long ago as the Late Paleozoic. More definite conclusions on the origin of the Isoptera must wait until termites or termite-like insects have been found in pre-Cretaceous strata. HYMENOPTERA
The Hymenoptera belong to the major subdivision of the Insecta known as the Endopterygota. There are no clues elucidating the nature or precise age of the earliest endopterygote insects, but the fossil record does provide insight into the history of the group as a whole. Representatives of two endopterygote orders, Neuroptera and Mecoptera, are found as far back as the Early Permian, some 280 million years ago. This occurrence suggests an origin of the Endopterygota approximately 100 million years after the origin of the true insects.3
The earliest known Hymenoptera have been found in Triassic beds of Central Asia (Rasnitsyn, 1964) and Australia (Riek, 1955). These fossils establish a minimum age for the order of about 220 million years. All the specimens known from this period belong to the suborder Symphyta, and surprisingly enough belong to the existing family Xyelidae.
A major advance in the evolution of the Hymenoptera occurred with the development of a constriction between the first and second abdominal segments; this presumably had the selective advantage of increasing the flexibility of the abdomen, important for both oviposition and defense. Hymenoptera which possess this adapta- tion, a diagnostic character of the suborder Apocrita, are first known from Upper Jurassic deposits of Central Asia (Rasnitsyn, 1975, 1977). These specimens have been assigned to the more primitive division of the Apocrita known as the Terebrantia or The oldest known insects, found in Upper Carboniferous deposits, comprise 11 orders and include the Apterygota (Thysanura), Paleoptera and Exopterygota. It should be noted that here the use of the term insect does not include the Collembola, Protura or Diplura.
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94 Psyche [March
Parasitica; the other division within this suborder is the Aculeata.4 Members of the latter are characterized by modifications of the ovipositor that have enabled its use not only for oviposition, but also as a transport vessel for defensive and prey-paralyzing com- pounds. This structure unquestionably plays an important role in colony defense and might provide an explanation for the restriction of eusociality within the Hymenoptera to the Aculeata. The oldest known aculeate hymenopteron, Cretavus sibericus, was discovered in an Upper Cretaceous (Cenomanian) deposit in Siberia in 1957. Although placed by Sharov (1962) in an extinct superfamily Cretavidea, related to the Scolioidea, it has recently been transferred to the existing family Mutillidae by Rasnitsyn (1977, p. 109). Since 1967, species representing 10 families and 19 genera of aculeate Hymenoptera have been found in Upper Cretaceous deposits in Central Asia (Rasnitsyn, 1977) (Table 2). Evans (1966) believes that such diversity by the Late Cretaceous is indicative of an earlier origin and postulates that the group may have evolved during the Jurassic. However, it must be pointed out that the Cretaceous is one of the longer periods in the earth's history, having a duration of roughly 70 million years, and may have been of sufficient length to account for such diversification.
VESPOIDEA
Included in this group are the three families considered to be "true wasps": The Masaridae and Eumenidae, both of which are solitary, and the Vespidae, where one finds behavioral modifications ranging from subsocial to highly advanced eusocial (P-ichards, 1953, 197 1). It is the Vespidae, by virtue of their sociality, with which I am primarily concerned in this paper.
There are many gaps in our record of the early social wasps and of the Vespoidea in general. Most striking, perhaps, about the fossil record of the wasps is their lack of representation (see Table 3). The "The classification of the Aculeata has recently undergone a major revision by D. J. Brothers (1975), in which the seven previously recognized superfamilies (Bethyloidea, Scolioidea, Pompiloidea, Formicoidea, Vespoidea, Sphecoidea, and Apoidea) are now combined into three: the Bethyloidea, Sphecoidea (subdivided into the Spheci- formes and Apiformes), and Vespoidea (subdivided into the Vespiformes and Formiciformes). However, since this revised classification has not been generally accepted in its entirety, I am employing here the more conventional classification (sensu Riek, 1970; Richards, 1971).
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19781 Burnham - Social Insects in Fossil Record 9 5 Table 2. Genera of aculeate Hymenoptera known from Cretaceous deposits (based on Rasnitsyn, 1977, and Evans, 1973). All genera are extinct. SCOLIOIDEA
Mutillidae
7SCOLIOIDEA
Scolioidae
Angarosphecidae
Falsiformicidae
Cretavus Sharov, 1962;
Rasnitsyn, 1977
Oryctopterus Rasnitsyn, 1977
Angarosphex Rasnitsyn, 1977
Falsiformica Rasnitsyn, 1977
7SCOLIOIDEA-BETHYLOIDEA
7Scolebythidae Cretabythus Evans, 1973
BETHYLOIDEA
Bethylidae
Cleptidae
Archaepyris Evans, 1973
Celonophamia Evans, 1973
Procleptes Evans, 1969
Hypocleptes Evans, 1973
Protamisega Evans, 1973
Dryinidae Cretodryinus Rasnitsyn, 1977
POMPILOIDEA
Pompilidae
FORMICOIDEA
Formicidae
SPHECOIDEA
Sphecidae
7SPHECOIDEA
?S phecidae
VESPOIDEA
Masaridae
Pompilopterus Rasnitsyn, 1977
Sphecomyrma Wilson and Brown, 1967
Cretomyrma Rasnitsyn, 1977
Paleomyrmex Rasnitsyn, 1977
Lisponema
Evans, 1969
Pittoecus Evans, 1973
Archisphex Evans, 1969
Taimyrisphex Evans, 1973
Curiovespa Rasnitsyn, 1975
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96 Psyche [March
absence of Vespidae from Baltic Amber (Lower Oligocene) and other fossil resins, in which ants are abundant, is probably due to their relatively large size, which reduces the likelihood of their entrapment in the sticky tree resin. Spradbery (1973, p. 316), attributes their scarcity in sedimentary deposits to "the behavioral characteristics and paper nest structures which do not lend them- selves to fossilization." As with any other fossil, the absence of an insect in the paleontological record provides no proof as to its actual occurrence in the past; one can only reconstruct and evaluate paleofaunas on the basis of those organisms that are represented. Therefore, it is conceivable that wasps were present earlier than the record indicates, but that conditions conducive to their preservation were lacking. The following does, however, provide information on the diversity of the group as we know it. Cretaceous
The earliest record of the Vespoidea extends back to the Upper Cretaceous (Turonian). Two species of vespoid wasp have been found in a deposit of this age in the USSR - both assigned to the genus Curiovespa (Rasnitsyn, 1975). Unfortunately, nothing is known about the body structure of these insects but on the basis of their wing venation they are placed in the family Masaridae. The presence of two distinct species in the same deposit suggests that some diversification of the Vespoidea had taken place as early as the Upper Cretaceous, although nothing is known about the morphological character of these early wasps.
Paleocene
No Vespoidea from this period are known. Eocene
The Eocene beds of Green River have yielded a surprisingly diverse assemblage of aculeates, but most of these belong to the Terebrantia or Sphecoidea; the only vespoid recovered from this deposit, Didineis solidescens, is of uncertain systematic position (Evans, 1966, p. 393). Scudder (1890) described this specimen as a sphecid of the subfamily Nyssoninae. However, Evans (1966) examined the type and concluded that it did not belong to the family Sphecidae, but was probably a eumenid, and tentatively assigned it to the genus Alastor.
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19781 Bwnham - Social Insects in Fossil Record 97 Figure 2.
Vespoid wasp from Eocene of British Coiumbia. Original photograph of specimen in Royal Ontario Museum, Toronto. Length offorewing, 12 mm. Piton (19401, in a thesis on the Eocene fossil beds of Menat, France, described an assemblage of Vespoidea found in this sedi- mentary deposit. However, because the six specimens he described are all assigned to extant genera, and do not show the characters essential for such generic designation, Piton's taxonomic determina- tions are perforce questionable. Particularly dubious is his place- ment of one specimen in the family Vespidae, genus Poliszes. Because the morphological features necessary for accurate taxo- nomic placement are obscured in this fossil, I prefer to place it in Vespoidea incertae sedis. The remaining five specimens are assigned to the Eumenidae incertae sedis.
Another vespoid species was recently recovered from a Middle Eocene deposit in British Columbia (M. V. H. Wilson, 1977). Al- though not formally described, the fossil clearly shows the charac- teristic venation of the vespoid complex (see Fig. 21, but could be either a vespid or a eumenid. Of course, one has no way of stating
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9 8 Psyche [March
with certainty that these early vespids were social. Within the Vespidae, divisions into subfamily and tribe are based primarily on behavioral rather than morphological characters. Furthermore, the morphological differences between the castes in any given species are often not obvious in the preserved fossils. Oligocene
True vespids are first found in the Upper Oligocene shales of Florissant, Colorado and Rott, Germany, two highly productive fossiliferous deposits. These beds and other various localities listed in Table 3 have turned up an assemblage comprised of four genera and 14 species. It is quite remarkable that three of the four genera represented are extant and this supports the possibility that the Vespidae were essentially modern by the Oligocene. Furthermore, the diversification of taxa suggests a much earlier origin for the family than is evidenced by the fossil record. Miocene
Scarcely any Vespidae are known from the Miocene, although this is most likely due to the overall dearth of deposits from this epoch. One vespid has been described from a deposit in Germany. This is Polistes kirbyanus and clearly belongs to the subfamily Polistinae. Other wasps from Miocene deposits have yet to be discovered, but one can assume that the wasp fauna of this age would be barely distinguishable from the wasp fauna of today. The following review of the fossil history of the Formicidae provides important information on their dominance, distribution, and supposed habits during the Mesozoic and Cenozoic eras. In contrast to the Vespoidea, ants are the most abundant insects in Tertiary formations. This may be attributed to their foraging behavior on and around trees, which enhances their chances of preservation in amber. A rough total of 20,000 specimens represent- ing some 200 species of ants has been studied (Table 4); this massive amount of work far exceeds the paleontological investigations carried out on any other family of insects. Several comprehensive monographs on the subject have been written, including The Ants of the Baltic Amber (Wheeler, 1914), and The Fossil Ants of North America (Carpenter, 1930), which are drawn on extensively in the following pages.
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19781 Burnham - Social Insects in Fossil Record 99 Cretaceous
The Cretaceous Period has, without question, provided more information on the early evolution of the ants than any other period, primarily because of the discovery in 1967 of two perfectly preserved worker ants in a New Jersey amber deposit. No doubt exists as to the primitive nature of these Cretaceous ants - both are members of the same species, Sphecomyrma freyi Wilson and Brown, and possess a mixture of wasp and ant characters. The petiole is distinctly ant-like, although the mandibles, which are short and bidentate, are very wasp-like (see Fig. 3A). A new subfamily, Sphecomyrminae, was named to accommodate S. freyi (Wilson, Carpenter, and Brown, 1967), and is considered ancestral to all known formicid subfamilies (see Taylor, 1978). Since the discovery of Sphecomyrma, several other Cretaceous ants have been found, and these provide strong evidence that the family was widespread during this period. Dlussky (1 975) described two new genera and three species, Cretomyrma arnoldii, C. uni- cornis, and Paleomyrmex zherichini (from a Late Cretaceous amber deposit in Yantardak, USSR) which he assigned to the Sphecomyr- minae. It is of interest that the type of P. zherichini is the first winged male ant to be found in a Cretaceous deposit and provides the only indication of wing venation in the Sphecomyrminae (Fig. 3B). The figured specimen of Cretomyrma unicornis raises doubts as to its position in the Formicidae for it is a badly mangled, poorly preserved specimen and might be better assigned to Hymenoptera incertae sedis.5 A fifth specimen, apparently a worker, has recently been discovered in the Cretaceous amber of Manitoba, Canada. Although not yet described, it undoubtedly belongs to the subfamily Sphecomyrminae (Wilson, personal communication). Paleocene
No ants from the Paleocene are known, undoubtedly because so few fossiliferous beds containing insect remains from this epoch SDlussky (1975) also described several other "ants" which were found in Upper Cretaceous deposits in the Kzyl-Zhar of Russia. Three genera (3 species) were placed in the subfamily Ponerinae: Petropone petiolata, Cretopone magna, and Archaeo- pone kzylzharica. These are all fragmentary specimens, and, as figured by Diussky, present no characters which would place them unequivocally in the Formicidae. They much more obviously belong in Hymenoptera incertae sedis, as does Dolichomyrma longiceps from the Upper Cretaceous of Kzyl-Zhar, which Dlussky put into Formicidae incertae sedis.
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TABLE 3. VESPOIDEA IN THE FOSSIL RECORD. Geological Age
CRETACEOUS
Masaridae
*Curiovespa curiosa Rasnitsyn
*Curiovespa magna Rasnitsyn
EOCENE
Eumenidae
?Alastor solidescens (Scudder)
?Rhygchium andrei Piton
?Odynerus manevali Piton
?Ancistrocerus eocenicus Piton
?Ancistrocerus berlandi Piton
?Eumenes projaponica Piton
PVespidae
?Polistes vergnei Piton
OLIGOCENE
Eumenidae
Rhynchium sp. Theobald
Odynerus terryi Cockerel1
Odynerus wilmattae Cockerel1
Odynerus oligopunctatus Theobald
?Odynerus praesulptus Cockerel1
Odynerus percantusus Cockerel1
?Alastor rottensis Statz
"Pseudonortania"'\ sepulta Timon-David
Locality
Kazakh, U.S.S.R.
Kazakh, U.S.S.R.
Green River, Wyoming
Menat, France
Menat, France
Menat, France
Menat, France
Menat, France
Menat, France
Cereste, France
Florissant, Colorado
Florissant, Colorado
Cereste, France
Florissant, Colorado
Florissant, Colorado
Rott, Germany
Camoins, France
References
Rasnitsyn, 1975
Rasnitsyn, 1975
Evans, 1966
Piton, 1940
Piton, 1940
Piton, 1940
Piton, 1940
Piton, 1940
Piton, 1940
Theobald, 1937
Cockerell, 1909a
Cockerell, 19 14
Theobald, 1937
Cockerell, 1906
Cockerell, 19 14
Statz, 1936
Timon-David, 1944
================================================================================
Vespidae
?* Paleovespa gillettei Cockerel1
?* Paleovespa florissantia Cockerell
?*Paleovespa scudderi Cockerel1
?*Paleovespa relecta Cockerel1
* Paleovespa baltica Cockerel1
*Paleovespa wilsoni Cockerell
Polistes industrius Theobald
Polistes signata Statz
?Polybia anglica Cockerel1
Polybia oblita Cockerell
Vespa bilineata Statz
Vespa cordifera Statz
Vespa nigra Statz
MIOCENE
Vespidae
Polistes kirbyanus Cockerel1
?Vespa attavina Heer
? Vespa crabroniformis Heer
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Florissant, Colorado
Cereste, France
Rott, Germany
Isle of Wight, England
Isle of Wight, England
Rott, Germany
Rott, Germany
Rott, Germany
Oeningen, Germany
Parschlug, Germany
Radoboj, Croatia
Bequaert, 1930
Bequaert, 1930
Bequaert, 1930
Bequaert, 1930
Cockerell, l909b
Cockerell, 19 14
Theobald, 1937
Statz, 1936
Cockerell, 192 la
Cockerell, 192 1 b
Statz, 1936
Statz, 1936
Statz, 1936
?Of uncertain position within the Vespoidea - clearly Diploptera, but further determination impossible. tPseudonortania Timon-David is a junior homonym of Pseudonortania Soika, 1936. *Extinct genera.
Cockerell, 1914
Heer, 1849
Heer, 1867
================================================================================
Psyche [March
Figure 3A.
Sphecomyrma freyi Wilson and Brown from the lower part of Upper Cretaceous of New Jersey. Drawing of holotype worker in Museum of Comparative Zoology, modified from'wilson, Carpenter, and Brown (1967). Length of body, 3.5 mm.
Figure 3B. Paleomyrmex zherichini Rasnitsyn from the lower part of the Upper Cretaceous of U.S.S.R. Drawing of holotype male in Paleontological Institute, Moscow, from Rasnitsyn, 1977. Length of body, 5.4 mm. have been discovered. Mention is made by Brues (1936) of a piece of petrified wood containing what he considers ant borings, highly resemblant of borings made by Camponotus today. Although there is no clear-cut evidence that these borings represent Camponotus activity, or insect activity of any kind, it is conceivable that Camponotus was present in New Mexico during the Paleocene; several species have been dscribed from the Florissant Shales, Colorado (Upper Oligocene), and one from the Baltic Amber
================================================================================
19781 Burnham - Social Insects in Fossil Record 103 (Lower Oligocene). In addition, it must be remembered that the Paleocene did not begin for at least 40 million years after the appearance of Sphecomyrma freyi.
Eocene
Very few fossil ants have been found in deposits of this age, and the determinations of many of these ant species are in doubt. Scudder (1877, 1878) described four supposed ants from the Green River formation, and five ants (1877) from the Quesnel Beds in British Columbia. Generic identifications on all of these fossils are to be considered dubious at best, and more likely incorrect (Car- penter, 1930).
In 1920, two species, Oecophylla bartoniana and Formica heter- optera, were described by Cockerel1 from an Eocene deposit in Bournemouth, England. Wheeler (1928) considered these ants for- micines, but because the descriptions were based on wing fragments, he questioned their generic determinations. Similarly, Cockerell's Formica eoptera (1923a) from the Eocene of Texas is of uncertain position at both the generic and subfamily levels. Archimyrmex rostratus (Cockerell, 1923b) from the Eocene shales of Colorado is probably a myrmicine (Carpenter, 1930), and is the only Green River ant that can be placed with any certainty in a subfamily. Carpenter (1929) described Eoponera berryi from the Wilcox formation of Tennessee, and placed this ant in the subfamily Ponerinae. He suggests that it may be closely allied to the Neotrop- ical genus Dinoponera. This is of interest because Eoponera berryi is the oldest known ant (Lower Eocene) to be assigned to a living subfamily of Formicidae.
Wilson (personal communication) mentions the recent discovery of three ants in a Middle Eocene amber deposit near Malvern, Arkansas, each belonging to a different subfamily. One belongs to the Dolichoderinae, genus Iridomyrmex; one is a formicine closely allied to the genus Paratrechina, and considered a relatively primi- tive, or "typical euformicine"; the last is a new genus of myrmicine, unique by virtue of its inflated postpetiole. These ants have yet to be formally described but they are nevertheless of paramount interest. The presence of these subfamilies in North America in the Eocene is strongly suggestive of their rapid evolution and dispersal during the Paleocene and perhaps during the Cretaceous.
================================================================================
104 Psyche [March
Oligocene
The Baltic Amber is, most certainly, the best studied of all Tertiary insect deposits, and has revealed a great deal about the nature and diversity of Oligocene ants.6 As of 1928, 11,7 11 ants (93 species) were examined from this deposit. Of this number, 1461 were studied by Mayr (1 868); 690 by Andre (1895); and 9,560 by Wheeler (1914, 1928).
An examination of the ant fauna reveals wide representation at the subfamily and generic levels. All extant subfamilies of Formici- dae are found in the amber with the exception of the Dorylinae and Leptanillinae. The absence of the Dorylinae is probably not due to selective exclusion on the part of the amber, but more likely indicates their absence from that part of the European continent during the Oligocene. Wheeler (1914) speculates that the foraging behavior of doryline ants should readily lead to entrapment in tree resin, but, in all probability, this group was then, as it is now, confined to the tropics. It is not surprising that the Leptanillinae are absent from the Baltic Amber; this is a small subfamily once considered a tribe of the Dorylinae, consisting of one genus and a few species; and although pantropical is hypogaeic and rarely encountered.
The Dolichoderinae and Formicinae together constitute 97 per- cent of all specimens and evidence indicates that these amber ants were already extraordinarily specialized. Workers of Iridomyrmex goepperti were found in a piece of amber (originally in the Konigs- berg collection) with several aphids. On the basis of this discovery, Wheeler (1914) concludes that Homoptera were attended by ants then much as they are today. The finding of several genera of paussid beetles (e.g., Arthropterus, Cerapterites and Eopaussus) in the Baltic Amber (Wasmann, 1929) suggests that myrmecophiles were established at this time. Perhaps most remarkable of all was the discovery of two Lasius schiefferdeckeri workers - each found with a mite attached to the base of the hind tibia, in precisely the ^'Because the
Baltic Amber was secondarily deposited in a clay bed of Lower
Oligocene age, it is necessarily older than the glauconitic sand ("blue-earth" clay) in which it lies. How much older is uncertain. In some published accounts it is referred to as Eocene. However, since the composition of the Baltic Amber ant fauna is very similar to that of the Florissant Shales and other born fide Oligocene deposits, I am following Zeuner (1939, p. 26) in referring to the amber as Lower Oligocene.
================================================================================
19781 Burnham - Social Insects in Fossil Record 105 same position on each. This demonstrates almost certainly that by the Lower Oligocene mites had acquired distinct preferences for attachment on specific regions of their host's integument. Almost as valuable as the Baltic Amber in providing a large and diverse assemblage of fossil ants is the Upper Oligocene deposit in Florissant, Colorado, studied by Carpenter (1930). The ant fauna of this deposit is strikingly similar to that of the Baltic Amber in many respects. It is interesting to note that roughly the same percentage of extant genera is found in both places; in the Florissant Shales this figure is given as 60 percent (Carpenter, 19301, in the Baltic Amber 56 percent (Wheeler, 1914). Iridomyrmex is clearly a dominant genus in the Baltic Amber, and although not so common in the Florissant Shales, a closely allied genus, Protazteca, comprises more than 25 percent of all specimens (Brown, 1973). Another similarity between the two deposits is the relative percentages of the various subfamilies. As in the amber, the Dolichoderinae are predominant, comprising 60 percent of the total number of ants. The Formicinae comprise another 25 to 30 percent, and the Myrmicinae in each deposit are represented by five percent or less of the total specimens. This suggests that the ant fauna in the northern hemisphere was essentially homogenous during the Oli- gocene.
The remaining deposits of Oligocene age from which ants have been described are of relatively minor importance. Most of the specimens are fragmentary and the determinations dubious; never- theless, a mention of them is certainly necessary. Specimens from Gurnet Bay, Isle of Wight, England, have been studied by Cockerel1 (1915) and Donisthorpe (1920). Cockerel1 described eight species of ants from this deposit but, because his generic determinations are based chiefly on highly variable measurements of wing fragments, they are of dubious significance. Donisthorpe examined a total of eight genera and fourteen species belonging to the subfamilies Ponerinae, Dolichoderinae, and Formicinae. Surprising is the large number of Oecophylla workers recovered (245); this genus is now restricted to Africa, India, and Australia, and is much more numerous in the Gurnet Bay deposit than in the Baltic Amber or Florissant Shales. This might be due to the difference in latitude between the deposits which would account for a warmer climate at Gurnet Bay later into the Tertiary than at the more northern deposits.
================================================================================
106 Psyche [March
Another Lower Oligocene deposit which has provided beautifully preserved fossil ants is Aix-en-Provence, France. Several species have been described by Theobald (19371, who recognized four subfamilies: Myrmicinae (1 species); Ponerinae (1 species); Doli- choderinae (1 genus, 2 species); and Formicinae (3 genera, 9 species). Also described by Theobald (1937) is an Oligocene collec- tion from Haut-Rhin, France, in which he recognizes the same four subfamilies (16 genera, 34 species). This fauna is very similar to that found in the Baltic Amber; in fact, Theobald has found five species which he considers identical to species in the Baltic Amber. In a deposit in Gard, France, Theobald (1937) describes two species, one a myrmicine, the other a dolichoderine.
Meunier (1917) has described four ant species from an Upper Oligocene deposit in Rott, Germany. These have been assigned to three genera: Formica, Ponera, and Myrmica. The specimens are well-preserved, as may be seen in Meunier's photographs, but his generic determinations are questionable. In 1957, two female reproductives of the same species were discovered in an Upper Oligocene deposit in Argentina. The authors described the species as Ameghinoia piatnitskyi and placed it in the subfamily Ponerinae (Viana and Haedo-Rossi, 1957). E. 0. Wilson (personal communication) is highly sceptical of the placement of A. pia~nitskyi in the Ponerinae, and thinks that it is very clearly a myrmeciine, This is quite extraordinary because no other fossil ants have been recovered from South America, and more importantly, if Wilson is correct, this is the first indication that the Myrmiciinae were so widespread by the Oligocene.
Miocene
The deposits of Miocene age which have provided the greatest number of ant specimens have been the Oeningen beds in Germany, and the Radoboj formation in Croatia. Approximately 60 species of ants from these places were described by the Swiss myrmecologist Heer (1 849, 1856, 1867), but his generic assignments are necessarily questionable in terms of present-day concepts of a formicid genus. Regrettably, the type specimens which are essential to a revision of this fossil fauna are believed to be lost.
================================================================================
19781 Burnham - Social Insects in Fossil Record 107 A few species were described by Emery (1 891) in Sicilian am be^, presumed to be Miocene, but these, like the specimens studied by Heer, are of questionable generic position.7 Another Miocene amber deposit has been found in Chiapas, Mexico, from which some one hundred ants have been recovered. Unfortunately, the majority of these are fragmentary, or otherwise too poor for determination. The assemblage does, however, suggest that the ant fauna in Mexico during the Miocene was essentially the same as might be found in that region today (Brown, 1973). Fujiyama (1970) described a single ant from the chhjabaru formation in Japan (middle Miocene) which he named Aphaeno- gaster axila, thought to be closely allied to the subgenus Dero- myrmu. This is not particularly unusual inasmuch as Aphaenogasler is a world-wide genus, and several species are found in Japan today. Perhaps the most interesting of all Miocene material is an ant colony of Oecophylla leakeyi found in Kenya (Wilson and Taylor, 1964). This is the first record of an actual, although fragmented, ant colony and contains a total of 366 specimens: 197 larvae, 105 worker pupae, and at least 64 workers. No Nearctic fossils of Oecophylla are known, but the species is well represented in European Tertiary deposits. Wilson and Taylor suggest on the basis of these fossil specimens that Oecophylla is a morphologically stable paleotropical genus which has persisted through most of the Tertiary with very little specialization.
The Apoidea form an interesting complex of social insects. Unlike the other social insect groups that are consistent in their degree of social achievement at the ordinal level (Isoptera), family level (Formicidae), and virtually the subfamily level (Vespinae), the Apoidea present a wide spectrum of social behavior at the generic level. Evidence suggests that eusociality has arisen in the bees at least eight times (Michener, 1962; Wilson, 197 I), which may explain this variance. Nevertheless, it is noteworthy that of roughly 20,000 existing species of bees only a small minority are thought to be presocial and eusocial (Wilson, 1971). Why sociality in the Apoidea 'These generic determinations are currently being reviewed by Dr. W. L. Brown, Jr.
================================================================================
TABLE 4. FORMICOIDEA IN THE FOSSIL RECORD. Geological Age
CRETACEOUS
S phecomyrminae
*Sphecomyrma freyi Wilson and Brown
?*Sphecomyrma sp.
*Cretomyrma arnoldii Dlussky
*Cretomyrma unicornis Dlussky
* Paleomyrmex zherichini Dlussky
EOCENE
Myrmicinae
*Archimyrmex rostratus Cockerel1
Formicinae
Oecophylla bartoniana Cockerel1
Formica eoptera Cockerel1
Formica heteroptera Cockerel1
?Paratrechina sp.
Ponefinae
*Eoponera berryi Carpenter
Dolichoderinae
Iridomyrmex sp.
OLIGOCENE
Myrmiciinae
*Ameghinoia piatnitskyi Viana and Haedo-Rossi Ponerinae
Brachyponera dubia Theobald
Locality
New Jersey, U.S.A.
Manitoba, Canada
Taymyr, U.S.S.R.
Taymyr, U.S.S.R.
Taymyr, U.S.S.R.
Florissant, Colorado
Bournemouth, England
Texas, U.S.A.
Bournemouth, England
Arkansas, U.S.A.
Tennessee, U.S.A.
Arkansas, U.S.A.
Argentina
Haut-Rhin, Germany
References
Wilson, Carpenter
and Brown, 1967
Wilson, pers. comm.
Rasnitsyn, 1975
Rasnitsyn, 1975
Rasnitsyn, 1975
Carpenter, 1930
Cockerell, 1920
Carpenter, 1930
Cockerell, 1920
Wilson, pers. comm.
Carpenter, 1929
Wilson, pers. comm.
Viana and Haedo-Rossi, 1957
z
e
Theobald, 1937a ~r
================================================================================
*Arch@onera wheeleri Carpenter
* Prionomyrmex longiceps Mayr
* Procerapachys annosus Wheeler
*Procerapachys favosus Wheeler
* Bradoponera meieri Mayr
Ectatomma europaeum Mayr
* Eleclopunera dubia Wheeler
Platyihyrea primaeva Wheeler
Euponera calcarea Theobald
Euponera succinea (Mayr)
Euponera crawZeyi Donisthorpe
Euponera globiventris Theobald
Ponera atavia Mayr
Ponera minuta Donisthorpe
Ponera elegantissima Meunier
Ponera hypolitha Cockerel1
Ponera rhenana Meunier
?Ponera graciZicornis Mayr
*Emplastus emeryi Donisthorpe
*Syntaphus wheeleri Donisthorpe
Pseudomyrmicinae
Pseudomyrma extincta Carpenter
Myrmicinae
Aphaenogaster mayri Carpenter
Aphaenogaster donisthorpei Carpenter
Aphaenogaster maculipes Theobald
Aphaenogaster maculata Theobald
Aphaenogaster sommer$eldti Mayr
Aphaenogaster oligocenica Wheeler
Florissant, Colorado
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Haut-Rhin, Germany
Baltic Amber
Isle of Wight, England
Haut-Rhin, Germany
Baltic Amber
Isle of Wight, England
Rott, Germany
Isle of Wight, England
Rott, Germany
Baltic Amber
Isle of Wight, England
Isle of Wight, England
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Haut-Rhin, Germany
Aix-en-Provence, France
Baltic Amber
Baltic Amber
Carpenter, 1930
Wheeler, 191 4
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 1914
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Theobald, 1937
Wheeler, 1914
Donisthorpe, 1920
Theobald, 1937
Wheeler, 19 14
Donisthorpe, 1920
Meunier, 1923
Cockerell, 19 15
Meunier, 19 17
Wheeler, 19 14
Donisthorpe, 1920
Donisthorpe, 1920
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Theobald, 1937
Theobald, 1937
Wheeler, 19 14
Wheeler, 19 14
================================================================================
TABLE 4. (CONTINUED)
Geological Age
OLIGOCENE Myrmicinae (continued)
Aphaenogaster mersa Wheeler
Sima klebsi Wheeler
Sima ocellata Mayr
Sima simplex Mayr
Sima angustata Mayr
Sima lacrimarum Mayr
Sima klebsi Theobald
Sima oligocenica Theobald
Monomorium pilipes Mayr
Monomorium mayrianum Wheeler
Erebomyrma antiqua (Mayr)
Erebomyrma thorali Theobald
Vollenhovia beyrichi (Mayr)
Vollenhovia prisca (Andre)
Stenamma berendti (Mayr)
*Electromyrmex klebsi Wheeler
*Agroecomyrmex duisburgi (Mayr)
Myrmica longispinosa Mayr
Myrmica archaica Meunier
*Nothomyrmica rudis (Mayr)
* Nothomyrmica intermedia Wheeler
*Nothomyrmica rugosostriata (Mayr)
* Nothomyrmica petiolata (Mayr)
Leptothorax gracilis Mayr
Locality
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Haut-Rhin, Germany
Gard, France
Baltic Amber
Baltic Amber
Baltic Amber
Haut-R hin, Germany
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Rott, Germany
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
References
Wheeler, 19 14
Wheeler, 1914
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 1914
Theobald, 1937
Theobald, 1937
Wheeler, 1914
Wheeler, 1914
Wheeler, 1914
Theobald, 1937
Wheeler, 1914
Wheeler, 19 14
Wheeler, 1914
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Meunier, 19 15
Wheeler, 19 14
Wheeler, 1914
Wheeler, 19 14
Wheeler, 1914
Wheeler, 19 14
================================================================================
Leptothorax glaesarius Wheeler
Leptothorax longaevus Wheeler
Leptothorax hystriculus Wheeler
Leptothorax placivus Wheeler
Leptothorax gurnetensis Cockerel1
Leucotaphus cockerelli Donisthorpe
*Stiphromyrmex robustus (Mayr)
* Parameranoplus primaevus Wheeler
Stigmomyrmex venustus Mayr
* Enneamerus reticulatus Mayr
Solenopsis maxima (Forster)
Solenopsis valida (Forster)
Solenopsis major Theobald
Solenopsis superba Forster
Solenopsis forsteri Theobald
Solenopsis blanda Theobald
Pheidole tertiaria Carpenter
Messor sculpteratus Carpenter
Pogonomyrmex fossilis Carpenter
Lithomyrmex rugosus Carpenter
Lithomyrmex striatus Carpenter
*Cephalomyrmex rotundatus Carpenter
Dolichoderinae
* Protaneuretus succineus Wheeler
* Paraneuretus tornquisti Wheeler
* Paraneuretus longipennis Wheeler
*Mianeuretus mirabilis Carpenter
Dolichoderus oviformis Theobald
Dolichoderus coquandi Theobald
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Isle of Wight, England
Isle of Wight, England
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Haut-Rhin, Germany
Haut-Rhin, Germany
Haut-Rhin, Germany
Haut-Rhin, Germany
Haut-Rhin, Germany
Haut-Rhin, Germany
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Baltic Amber
Baltic Amber
Florissant, Colorado
Haut-Rhin, Germany
Haut-Rhin, Germany
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Cockerell, 191 5
Donisthorpe, 1920
Wheeler, 1914
Wheeler, 19 14
Wheeler, 1914
Wheeler, 1914
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Wheeler, 1914
Wheeler, 1914
Wheeler, 1914
Carpenter, 1930
Theobald, 1937
Theobald, 1937
================================================================================
TABLE 4. (CONTINUED)
Geological Age
OLIGOCENE Dolichoderinae (continued)
Dolichoderus bruneti Theobald
Dolichoderus explicans Theobald
Dolichoderus affectus Theobald
Dolichoderus balticus Theobald
Dolichoderus balticus (Mayr)
Dolichoderus oviformis Theobald
Dolichoderus antiquus Carpenter
Dolichoderus rohweri Carpenter
Dolichoderus cornutus (Mayr)
Dolichoderus passalomma Wheeler
Dolichoderus elegans Wheeler
Dolichoderus mesosternalis Wheeler
Dolichoderus vexillarius Wheeler
Dolichoderus sculpteratus (Mayr)
Dolichoderus tertiarius (Mayr)
Dolichoderus longipennis Mayr
Dolichoderus britannicus Cockerel1
Dolichoderus gurnetensis Donisthorpe
Dolichoderus ovigerus Cockerel1
Dolichoderus vectensis Donisthorpe
Iridomyrmex goepperti Theobald
Iridomyrmex goepperti Mayr
Iridomyrmex geinitzi Theobald
Iridomyrmex geinitzi (Mayr)
Locality
Haut-Rhin, Germany
Haut-Rhin, Germany
Haut-Rhin, Germany
Aix-en-Provence, France
Baltic Amber
Gard, France
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Isle of Wight, England
Isle of Wight, England
Isle of Wight, England
Isle of Wight, England
Haut-Rhin, Germany
Baltic Amber
Haut-Rhin, Germany
Baltic Amber
References
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Wheeler, 19 14
Theobald, 1937
Carpenter, 1930
Carpenter, 1930
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 1914
Wheeler, 1914
Wheeler, 19 14
Wheeler, 19 14
Cockerell, 191 5
Donisthorpe, 1920
Cockerell, 19 15
Donisthorpe, 1920
Theobald, 1937
Wheeler, 1914
Theobald, 1937
Wheeler, 1914
================================================================================
Iridomyrmex breviantennis Theobald
Iridomyrmexflorissantius Carpenter
Iridomyrmex obscurans Carpenter
Iridomyrmex constrictus (Mayr)
Iridomyrmex samlandicus Wheeler
Iridomyrmex oblongiceps Wheeler
Protazteca elongata Carpenter
Protazteca quadrata Carpenter
Protazteca capitata Carpenter
Liometopum miocenicum Carpenter
Liometopum oligocenicum Wheeler
Liometopum scudderi Carpenter
Elaeomyrmex gracilis Carpenter
Elaeomyrmex coloradensis Carpenter
Asymphylomyrmex balticus Wheeler
Pityomyrmex tornquisti Wheeler
Miomyrmex impactus (Cockerell)
Miomyrmex striatus Carpenter
Petraeomyrmex minimus Carpenter
Formicinae
Plagiolepis succini Andre
Plagiolepis klinsmanni Mayr
Plagiolepis kueno wi Mayr
Plagiolepis squamifera Mayr
Plagiolepis singularis Mayr
Plagiolepis solitaria Mayr
*Rhopalomyrmex pygmaeus Mayr
Dimorphomyrmex theryi Emery
Dimorphomyrmex mayri Wheeler
Haut-Rhin, Germany
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Baltic Amber
Baltic Amber
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Baltic Amber
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Theobald, 1937
Carpenter, 1930
Carpenter, 1930
Wheeler, 1914
Wheeler, 1914
Wheeler, 19 14
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Wheeler, 19 14
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Wheeler, 1914
Wheeler, 1914
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Wheeler, 1914
Wheeler, 1914
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 1914
Wheeler, 1914
Wheeler, 1914, 1929
Wheeler, 19 14
================================================================================
TABLE 4. (CONTINUED)
Geological Age
OLIGOCENE Formicinae (continued)
Gesomyrmex annectens Wheeler
Gesomyrmex expectans Theobald
Gesomyrmex miegi Theobald
Gesomyrmex hoernesi Theobald
Gesomyrmex hoernesi Mayr
* Prodimorphomyrmex primigenius Wheeler
Oecophylla superba Theobald
Oecophylla brischkei Mayr
Oecophylla brevinodis Wheeler
Oecophylla megarche Cockerel1
Oecophylla atavina Cockerel1
Oecophylla perdita Cockerel1
Prenolepis henschei Mayr
Prenolepis pygmaea Mayr
Lasius schiefferdeckeri Mayr
Lasius pumilus Mayr
Lasius epicentrus Theobald
Lasius chambonensis Piton and Theobald
Lasius tertiarius Zalessky
Lasius punctulatus Mayr
Lasius nemorivagus Wheeler
Lasius edentatus Mayr
Tetramorium peritulus (Cockerell)
Eoformica eocenica Cockerel1
Locality
Baltic Amber
Haut-Rhin, Germany
Haut-Rhin, Germany
Haut-Rhin, Germany
Baltic Amber
Baltic Amber
Haut-Rhin, Germany
Baltic Amber
Baltic Amber
Isle of Wight, England
Isle of Wight, England
Isle of Wight, England
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Aix-en-Provence, France
Lac Chambon, France
Ukraine, U.S.S.R.
Baltic Amber
Baltic Amber
Baltic Amber
Florissant, Colorado
Florissant, Colorado
References
Wheeler, 1914
Theobald, 1937
Theobald, 1937
Theobald, 1937
Wheeier, 1929
Wheeler, 1914
Theobald, 1937
Wheeler, 19 14
Wheeler, 1914
Donisthorpe, 1920
Cockerell, 19 15
Cockerell, 191 5
Wheeler, 1914
Wheeler, 1914
Wheeler, 19 14
Wheeler, 19 14
Theobald, 1937
Piton and Theobald, 1935
Zalessky, 1949
Wheeler, 1914
Wheeler, 1914
Wheeler, 19 14
Wilson, 1955
Cockerell, 1921c
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Formica flori Mayr
Formica flori Theobald
Formica horrida Wheeler
Formica phaethusa Wheeler
Formica clymene Wheeler
Formica constricts (Mayr)
Formica strangulata Wheeler
Formica tripartita Theobald
Formica alsatica Theobald
Formica serresi Theobald
Formica latinodosa Theobald
Formica oculata Theobald
Formica minutula Theobald
Formica sepulta Theobald
Formica robusta Carpenter
Formica cockerelli Carpenter
Formica grandis Carpenter
Formica masculipennis Piton and Theobald Formica pitoni Theobald
Formica bauckhorni Meunier
Formica auxillacensis Piton and Theobald Glaphyomyrmex oligocenicus Wheeler
Pseudolasius boreus Wheeler
Dryomyrmex fuscipennis Wheeler
Dryomyrmex fuscipennis Theobald
Dryomyrmex claripennis Wheeler
Glaphyromyrmex oligocenicus Theobald
Camponotus mengei Mayr
Ckzmponotus mengei Theobald
Baltic Amber
Haut-Rhin, Germany
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Haut-Rhin, Germany
Haut-Rhin, Germany
Aix-en-Provence, France
Aix-en-Provence, France
Aix-en-Provence, France
Aix-en-Provence, France
Aix-en-Provence, France
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Auxillac, France
Lac Chambon, France
Rott, Germany
Auxillac, France
Baltic Amber
Baltic Amber
Baltic Amber
Haut-Rhin, Germany
Baltic Amber
Haut-Rhin, Germany
Baltic Amber
Haut-Rhin, Germany
Wheeler, 1914
Theobald, 1937
Wheeler, 19 14
Wheeler, 1914
Wheeler, 1914
Wheeler, 1914
Wheeler, 19 14
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Piton and Theobald, 1935
Piton and Theobald, 1935
Meunier, 19 17
Piton and Theobald, 1935
Wheeler, 19 14
Wheeler, 19 14
Wheeler, 1914
Theobald, 1937
Wheeler, 19 14
Theobald, 1937
Wheeler, 19 14
Theobald, 1937
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TABLE 4. (CONCLUDED)
Geological Age
OLIGOCENE Formicinae (continued)
Camponotus vehemens Forster
Camponotus longiventris Theobald
Camponotus saussurei Theobald
Camponotus penninervis Theobald
Camponotus fuscipennis Carpenter
Camponotus microcephalus Carpenter
Camponotus petrifactus Carpenter
Camponotus brodiei Donisthorpe
MIOCENE
Ponerinae
Ponera umbra Popov
M yrmicinae
Aphaenogaster axila Fujiyama
Formicinae
Camponotus obesus Piton
Camponotus tokunagai Naora
* Pseudocamponotus elkoanus Carpenter
Solenopsis longaevus Heer
Formica cantalica Piton
Lasius crispus Piton
Lasius martynovi Popov
Oecophylla leakeyi Wilson and Taylor
Locality
Haut-Rhin, Germany
Aix-en-Provence, France
Aix-en-Provence, France
Aix-en-Provence, France
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Isle of Wight, England
Caucasus, U.S.S.R.
Chojabaru, Japan
Joursac, France
China
Elko, Nevada
Radoboj, Croatia
Joursac, France
Joursac, France
Caucasus, U.S.S.R.
Kenya
References
Theobald, 1937
Theobald, 1937
Theobald, 1937
Theobald, 1937
Carpenter, 1930
Carpenter, 1930
Carpenter, 1930
Donisthorpe, 1920
Popov, 1933
Fujiyama, 1970
Piton and Theobald, 1935
Naora, 1933
Carpenter, 1930
Poncracz, 1928
Piton and Theobald, 1935
Piton and Theobald, 1935
Popov, 1933
Wilson and Taylor, 1964
*Extinct genera.
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19781 Burnham - Social Insects in Fossil Record 117 is so highly polyphyletic remains unanswered, and is a problem unlikely to be resolved by the geological past. However, the fossil record does provide intriguing information on the evolution of the bees and indicates that their sociality may well have been established prior to the Oligocene. The following survey of the fossil Apoidea is indicative of the diversity of bees which have been found (Table 5). Those species which were described by early 19th century entomologists (Latreille, Heer, Heyden, etc.) are excluded from this coverage because these were uniformly assigned to modern genera.8 Cockerel1 (1909) claims that most of these species actually belonged to quite different and extinct genera. Oligocene
The earliest bees in the fossil record are found in the Baltic Amber, of Lower Oligocene age. The bees in this deposit are well- diversified (Zeuner and Manning, 1976), and the most prevalent apoid genus in the amber, Electrapis, is thought to have been social. Cockerel1 (1909) based this conclusion on the occurrence of many specimens of Electrapis meliponoides crowded together in a small piece of amber, a suggestive but certainly not conclusive deduction. Zeuner (1944, 1951), however, believed Electrapis to be social based on its pollen collecting apparatus. The extent to which social behavior was developed in this genus nevertheless remains a matter of conjecture. Electrapis is considered by some to be directly ancestral to the highly eusocial Apis, although Kelner-Pillault (1974) disagrees with this relationship. She suggests that Electrapis is actually a long extinct genus which possessed many primitive characters and represents an evolutionary side-line of the Apoidea. Both hypotheses are highly conjectural.
The presence of long-tongued bees such as Electrapis suggests that the Baltic Amber bees were rather specialized. Tongue structure is assumed to have evolved in response to various morphological changes (i.e., longer corollas) which took place during the evolution of the angiosperms (Michener, 1974). Short-tongued bees such as the colletids are considered the more primitive members of the Apoidea and are representative of bee radiation that occurred at a time when most of the angiosperms had shallow flowers (Michener, 1974).
8For a listing of these specimens, see Zeuner and Manning (1976).
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118 Psyche [March
In Late Oligocene deposits, the Apoidea are extremely well represented. Six major families of bees are known from this epoch: Haiictidae, Andrenidae, Melittidae, Megachilidae, Anthophoridae, and Apidae. A total of 29 genera arc represented, many of which are extant. Several specimens belonging to Chalcobombus and Bumbus are described from deposits in both Europe and North America suggesting widespread radiation of this specialized group of bees by the Early Oligocene. In the Late Oligocene, bees very similar to Apis melttfera are found. Manning (1952) feels that some species from the Rott Shales possess almost all the necessary characters for place- ment in the genus Apis (Fig. 4). Moreover, in the Dominican Amber of Oligocene-Miocene age, several Trigona workers are found, providing convincing proof that social behavior was well established at this time (Michener, 1974).
Figure 4. Apis henshawi Cockerel1 from Upper Oligocene of Rott, Germany. Original photograph of holotype in Museum of Comparative Zoology. Length of body, 15 mm.
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19781 Burnham - Social Insects in Fossil Record 119 Miocene
By the Miocene, the bee fauna is essentially modern. In Chiapas Amber from Mexico, bees have been discovered that are so similar to an existing Neotropical species that they have been assigned to the same subgenus, Trigona (Nogueirapis), and are scarcely different at the specific level (Wille, 1959). Fujiyama (1970) mentions the discovery of a fossil bee in a Japanese Miocene deposit and states that, "There is no room for doubt that this is a species of honeybee." A review of the fossil record reveals the following about the evolution of the bees. 1) We know that the Early Oligocene fauna is a mixture of primitive and advanced genera, although it appears to be dominated by fairly advanced species. By the end of this epoch, the fauna is modern in overall character. 2) We know that sociality had clearly arisen by the end of the Oligocene, and possibly much earlier. And 3) by the Miocene, the bees were virtually indistinguish- able from the bees of today. Six families of bees are represented in the Oligocene: including the phylogenetically advanced Apidae with six genera and 22 species. Such diversity of relatively advanced bees is indicative of either a much longer history of the group than is evidenced by the fossil record, or a fairly short history characterized by the rapid speciation and explosive radiation of the group. The bees are clearly derived from the spheciform wasps, al- though nothing is known about the nature of this sphecid ancestor (Wilson, 1971; Michener, 1974). In 1964, just prior to his death, F. J. Manning was investigating a sphecid from the Jurassic beds of Lerida Province, Spain, which "he thought might be (or be closely related to) the ancestor of the bees" (Zeuner and Manning, 1976, p. 155). This would be an astounding find if true, and it is unfortunate that nothing more is known - either about the specimen or about Manning's reasons for thinking it ancestral to the bees. The distinction between the Sphecoidea and the Apoidea is sufficiently subtle as to make determinations of fossil compressions extremely difficult. The presence of plumose hairs and enlarged basitarsi, characters which are important apoid features, rarely survive preservation unless the insect is preserved in amber. The origin of the bees remains a subject of much speculation. It is believed that "insect-plant interactions played a key role in the origin of the angiosperm flower and component structures" (Hickey
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120 Psyche [March
and Doyle, 1977, p. 92). Conversely, angiosperms have been instrumental to the evolutionary success of the Apoidea. On the basis of the evolutionary dependence of the two groups, can anything be said about their relationship in geological time? Two possibilities present themselves: 1) the angiosperms evolved first and were initially wind pollinated9 or pollinated by arthropods other than Hymenoptera (e.g., Coleoptera, Diptera, Thysanoptera, pos- sibly spiders); and 2) the first bees evolved from sphecid wasps prior to the origin of the angiosperms by adapting themselves to feeding on pteridosperm pollen or reproductive organs. A closer look at these possibilities is warranted. Coleoptera and Diptera are found in the fossil record at least by the Triassic. This supports the argument that they could have served as vectors for dispersal of angiosperm pollen. The question arises, if these insects were capable of performing essential roles as pollinators, why didn't angiosperms arise earlier in the Mesozoic than the Cretaceous? Regal (1977) suggests that the limiting factor to angiosperm dis- persal was the presence of seed-carrying birds and mammals. He argues that this method of seed dispersal, acting in conjunction with insect pollination, provided the selective advantages behind the subsequent primary radiation of the angiosperms. This is a sound argument, but says little about the insects which may have been pollinating these early plants. It would seem that successful dispersal of flowering plants is dependent on efficiency at two levels - pollination and seed dispersal. The explosive radiation of the angiosperms during the Cretaceous indicates that the more special- ized insect pollinators, the bees, may have been present in order to explain this success.
This might support the possibility that pollen collecting bees had already evolved by the time the first angiosperms appeared. Accord- ing to Wilson (1971, p. 79, the "Apoidea can be loosely character- ized as sphecoid wasps that have specialized in collecting pollen instead of insect prey as larval food." The possibility, however speculative, exists that bees evolved in response to the food source presented by the pteridosperms but subsequently abandoned this resource when the angiosperms appeared. Certainly one way of accounting for the explosive radiation of the angiosperms would be å´^Stebbin (1970, p. 323) suggests that the earliest angiosperms were not wind pollinated.
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1 9781 Burnham - Social Insects in Fossil Record 121 TABLE 5. APOIDEA IN THE FOSSIL RECORD.10 Geological Age
EOCENE
PApidae
Probombus hirsutus Piton
OLIGOCENE
Halictidae
*Cyrtapis anomalis Cockerel1
Halictus ruissatelensis Timon-David
Halictusflorissantellus Cockerel1
Halictus miocenicus Cockerel1
Halictus scudderiellus Cockerel1
Andrenidae
Andrena wrisleyi Salt
Andrena clavula Cockerel1
Andrena grandipes Cockerel1
Andrena hypolitha Cockerel1
Andrena lagopus Latreille
Andrenapercontusa Cockerel1
Andrena sepulta Cockerel1
*Lithandrenu saxorum Cockerel1
*Pelandrenu reducta Cockerel1
*Libellulapis antiquorum Cockrell
*Libellulapis wilmattae Cockerel1
Melittidae
*Ctenoplectrella dentata Salt
*Ctenoplectrella viridiceps Cockerel1
*Ctenoplectrella splendens Kelner-Pillault *Glyptapis fuscula Cockerel1
*Glyptapis mirabilis Cockerel1
*Glyptapis neglecta Salt
*Glyptapis reducta Cockerel1
*Glyptapis reticulata Cockerel1
Melitta willardi Cockerel1
Megachilidae
Anthidium mortuum (Meunier)
Anthidium exhumatum Cockerel1
Anthidium scudderi Cockerel1
* Dianthidium tertiarium Cockerel1
*Lithanthidium pertriste Cockerel1
Heriades bowditchi Cockerel1
Heriades halictinus Cockerel1
Heriades laminarum Cockerel1
Heriades mersatus Cockerel1
Locality
Menat, France
Florissant, Colorado
Marseille, France
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Rott, Germany
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
'åÁSe Zeuner and Manning (1976) for reference citations,
================================================================================
Psyche
TABLE 5. (CONTINUED)
Geological Age
[March
Locality
OLIGOCENE Megachilidae (continued)
Heriades mildredae Cockerel1
Heriades priscus Cockerel!
Heriades saxosus Cockerel1
Megachile praedicta Cockerel!
Osmia carbonum Heyden
Anthop horidae
Ceratina disrupta Cockerell
Xdocopa friesei Statz
Tetralonia berlandi Theobald
Anthophora melfordi Cockerel1
*Anthophorites gaudryi Oustalet
*Protomelecta brevipennis Cockerell
Apidae
*Chalcobombus hirsutus Cockerell
*Chalcobombus humilis Cockerel1
*Chalcobombus martialis Cockerel!
Bombus florissantensis (Cockerell)
*Sophrobombus fatalis Cockerel!
Trigona dominicana Wille and Chandler
Trigona eocenica Kelner-Pillault
* Electrapis apoides Manning
*Electrapis meliponoides (Buttel-Reepen) *Electrapis indecisus (Cockerell)
*Electrapis tristellus (Cockerell)
*Electrapis palmnickenensis (Roussy)
*Electrapis minuta Kelner-PiIlault
*Electrapis bombusoides
Electrapis proava (Menge)
Electrapis tornquisti Cockerel1
Apis cuenoti Theobald
Apis henshawi Cockerel1
Apis henshawi dormitans (Cockerell)
Apis henshawi kaschkei (Statz)
Apis aquitanensis de Rilly
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Florissant, Colorado
Rott, Germany
Florissant, Colorado
Rott, Germany
Gard, France
Florissant, Colorado
Corent, France
Florissant, Colorado
Baltic Amber
Baltic Amber
Baltic Amber
Florissant, Colorado
Baltic Amber
Dominican Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Baltic Amber
Cereste, France
Rott, Germany
Rott, Germany
Rott, Germany
Aix-en-Provence, France
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Burnham - Social Insects in Fossil Record 123
Geological Age
TABLE 5. (CONCLUDED)
Locality
MIOCENE
Halictidae
Halictus schemppi (Armbruster)
Andrenidae
Andrena primaeva Cockerel1
Megachilidae
Lithurge adamitica (Heer)
Megachile amaguensis Cockerel1
Osmia antiqua Heer
Osmia nigra Zeuner and Manning
Anthophoridae
Xylocarpa jurinei (Heer)
Xylocopa hydrobiae Zeuner
Xylocopa senilis Heer
*Anthophorites thoracica Heer
*Anthophorites longaeva Heer
*Anthophorites mellona Heer
*Anthophorites titania Heer
*Anthophorites tonsa Heer
*Anthophorites veterana Heer
Apidae
Bombus abavus Heer
Bombus proavus Cockerel1
Trigona succini (Tosi)
Trigona sicula (Tosi)
Trigona silacea Wille
Trigona devicta Kerr and Maule
Apis armbrusteri armbrusteri Zeuner
Apis armbrusteri scharmanni (Armbruster) Apis armbrusteri scheeri (Armbruster)
Apis armbrusteri scheuthlei (Armbruster) Apis catanensis avolii Roussi
Apis melisuga (Handlirsch)
Randeck, Germany
Oeningen, Germany
Oeningen, Germany
Siberia, U.S.S.R.
Oeningen, Germany
Oeningen, Germany
Oeningen, Germany
Biebrich, Germany
Oeningen, Germany
Radoboj, Croatia
Radoboj, Croatia
Oeningen, Germany
Oeningen, Germany
Oeningen, Germany
Oeningen, Germany
Oeningen, Germany
Latah, Washington
Sicilian Amber
Sicilian Amber
Chiapas, Mexico
Burma Amber
Wurttemburg, Germany
Wurttemburg, Germany
Wurttemburg, Germany
Wurttemburg, Germany
Sicilian Amber
Italy
*Extinct genera.
================================================================================
124 Psyche [March
the explanation that the insect pollinators so important to their success were pre-adapted as pollination vectors. It is interesting to note at this point that bees have been observed foraging on conifer pollen in areas where other food resources are scarce. Ray Angelo (personal communication, May, 1978) reports observing Colletes sp. foraging in high numbers on Juniperus virginiana pollen cones. This is noteworthy in two respects: 1) this conifer is the only readily available pollen source in the particular habitat where observations took place (Concord, Mass.), and 2) the bees foraging on the tree are members of the primitive bee family Colletidae. This suggests that they are generalized enough to have retained the ability to forage on gymnosperm pollen. Nevertheless, the hypothesis that bees evolved before the advent of the angiosperms is highly speculative, and remains a difficult theory to prove. The possibility of a pre-angiosperm origin for the bees implies that the Apoidea, and possibly sociality in the Apoidea, may be older than indicated by the fossil record. An inherent problem, of course, is whether or not these early bees would be recognizable as such, or would be mistaken for sphecid wasps. The discovery of additional Cretaceous amber might well provide valuable insight into this problem. SUMMARY
Wheeler writes in his 1928 book, "from the lowest to the highest forms in the series, all animals are at some time in their lives immersed in some society." It is the elaboration or evolution of these habits that leads to the eusocial behavior found in the Isoptera and certain groups of the Hymenoptera. The preceding account has examined insect sociality from a paleontological perspective in the hope that it will provide insight into the antiquity of this behavioral phenomenon. In addition, it has provided information on certain aspects of the evolution of the four major groups of social insects. The Isoptera are highly eusocial at the ordinal level and evidence suggests an ancient origin for the group. The oldest fossil termite known is from a Late Cretaceous deposit in Canada. The presence of a distinct humeral suture at the wing base indicates that social behavior was developed in the Isoptera at this time. It is further- more presumed that the termites arose in the early Mesozoic or possibly earlier, and from "protoblattoid" or blattoid stock. The hypogaeic lifestyle of most termites is not conducive to their
================================================================================
19781 Burnham - Social Insects in Fossil Record 125 preservation as fossils and this may explain their absence in pre- Cretaceous deposits.
The first Hymenoptera appear in the Triassic and belong to the primitive family Xyelidae (Symphyta). Social Hymenoptera are not, however, found in the fossil record until the Upper Cretaceous. The ant species discovered in deposits of this age are more primitive than any now existing and have been of paramount importance in our understanding of ant phylogeny. By the mid-Tertiary, the ant fauna was extremely diverse; by the Miocene, the genera were essentially modern, and geographic distribution of the ants was apparently similar to that of today.
The Vespoidea although not very numerous in fossil deposits, have been found as far back as the Late Cretaceous, represented by one specimen assignable to the Masaridae. The presence of several vespoids in Eocene deposits strongly supports the possibility that social wasps evolved during the Late Cretaceous or Early Paleocene. Apoidea extend into the fossil record only as far as the Oligocene, although it is speculated that they may have evolved much earlier. This is suggested by the fact that the bee fauna was essentially modern by the end of the Oligocene and also because the inter- dependence of angiosperms and bees suggests a co-evolutionary relationship beginning sometime in the Cretaceous. Any discussion of sociality in the geological past must necessarily involve a certain amount of speculation. Morphological characters play an essential role in the analysis of an insect's social status, an example of this being the presence of the humeral suture in Cretatermes. In those social insect groups possessing very little morphological variation between castes, recognition of such social distinctions in the fossils is virtually impossible. It is generally assumed that extinct species belonging to extant genera possessed a similar type of social behavior in the past as is exhibited by the group today. To speculate further about the social habits of fossil insects is simply not possible. The mechanisms behind the evolution of eusociality in the insects remain unknown, yet the success of this form of social behavior is unquestioned. Only the recovery of additional material will provide evidence to further elucidate our understanding of the paleontological record of these insects. As the record now stands, it is possible to state with a fair degree of certainty that insect sociality had evolved by the middle of the Cretaceous and perhaps much earlier.
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Psyche
[March
rn
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19781
Burnham - Social Insects in Fossil Record ACKNOWLEDGEMENTS
This study was originally intended as a brief survey of the social insects in the fossil record but underwent rapid expansion shortly after its initiation. This is partly due to my burgeoning interest in the subject matter, partly due to the vast amount of material requiring my attention, and partly due to the stimulation and encouragement received from friends in the academic community at Harvard. It is to the following friends that I extend my thanks and appreciation: K. M. Horton, Paul Strother, Robert E. Silberglied, Kenneth Miyata, and N. E. Woodley. Special thanks are given to F. M. Carpenter for his continuous guidance and advice, for his patience as my photographic assistant; and most of all, deep appreciation is extended to him for providing the inspiration integral to the success of this study. The Royal Ontario Museum, Toronto, is gratefully acknowledged for the loan of the Eocene vespoid. In addition, partial financial support is acknowledged to National Science Foundation Grant DEB 78-09947 - F. M. Carpenter, principal investigator.
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128 Psyche [March
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19781 Burnham - Social Insects in Fossil Record 129 1969.
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Three new Cretaceous aculeate wasps (Hymenoptera). Psyche 76(3): 251-261.
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