Mary Jane West.
Range Extension and Solitary Nest Founding in Polistes exclamans. (Hymenoptera: Vespidae).
Psyche 75:118-123, 1968.

Full text (searchable PDF, 480K)
Durable link: http://psyche.entclub.org/75/75-118.html

The following unprocessed text is extracted from the PDF file, and is likely to be both incomplete and full of errors. Please consult the PDF file for the complete article.

RANGE EXTENSION AND SOLITARY NEST FOUNDING IN POLISTES EXCLAMANS
( HYMENOPTERA : VESPIDAE) *
BY MARY JANE WEST
Museum of Comparative Zoology, Harvard University It is generally difficult to document temporal changes in the geo- graphic distribution of insect species because of the sporadic nature of collections made primarily for taxonomic study and specimen identification. Therefore some new distribution data providing an unusual amount of evidence for range extension in the social wasp Polistes exclamans Viereck seem of interest. Several states have been added to the known range of P. exclamans in the last twenty years; and P. exclamans is now abundant in east-central Missouri, where it was almost certainly absent two decades ago. Solitary nest founding by P. exclamans queens may enhance the ability of this species to colonize new areas.
Porter (1963) first suggested that P. exclamans may be extending its range. He reported new records from Maryland and New Jersey, noting that these states are considerably north of the northernmost previous records for P. exclamans in the Atlantic Coast states. A collecting trip to St. Louis and Jefferson Counties, Missouri, in July, 1967, provided the first evidence of range extension by Polistes exclamans in the midwestern United States. The purpose of the trip was to collect specimens of Polistes species observed by the Missouri naturalist Phil Rau prior to his death in 1948. Rau re- peatedly mentioned (e.g., in Rau 1929, 1g42a, 1942b) that only four species of Polistes existed in Missouri: P. pallipes (= metricus), P. rubiginosus, P. anndaris, and P. variatus (= fuscatus) (Rau's de- terminations by J. Bequaert). However, I found P. exdamans nests common in barns and abandoned buildings in southern St. Louis County, and collected specimens of P. exclanzans in six of the eight localities where Polistes wasps were found (other species collected were P. metricus, P. fuscatus, and P. mbiginosus). Three of the 1967 P. exclamans localities were frequented by Rau during his studies of Polistes: "Meramac Highlands", an area near the Mera- mac River about eight miles southwest of St. Louis (first mentioned in Rau and Rau, 1918) ; Kirkwood, Rau's home town for 24 years (first mentioned as a study locality in Rau, 1941) ; and a farm near 'Manuscript received by the editor 17 June 1968



================================================================================

19681 West - Polistes exclamans 1 19
the present Beaumont Reservation, 18 miles southwest of St. Louis, the former site of some "woodcutters' cabins" often visited by Rau during the years 1945-1947 (Rau, unpublished manuscript) There is little doubt that Rau would have noticed and reported P. exclamans in the St. Louis region if the species had been present there during his lifetime. He published 39 papers and one book chapter (Rau and Rau, 1918) on the Polistes of 43 named localities in that area during the years 1918-1946, and devoted his last years to writing a book dealing exclusively with the bionomics of Polistes (Rau, unpublished manuscript). Furthermore, Rau would have been unlikely to confuse exclamans with any of the Missouri species he recognized, since it has a distinctive appearance and nest form appreciated and described by Rau when he collected it in Texas (Rau, 1943).
Therefore it seems reasonable to conclude that the present abundance of Polistes exclanzans in southern St. Louis County, Missouri, is due to colonization in that area during the last twenty years.
Examination of museum collections (see Acknowledgments) has revealed specimens of P. exclamans from nine states not considered part of the species range in 195 I (Bohart, 195 I ) (Table I ) . Eight of the new state records are in the northeastern quarter of the species range, suggesting a generally northward expansion in the eastern half of the United States. Since climate-related range fluctuations are quite common in animals, and there are numerous examples of north- ward expansion in North America during the present century (see Mays, 1963), climatic change and/or adaptation to more northern habitats must be considered as possible contributing factors in the expansion of P. exclanzans. However, there are pre-1950 records of P. exclamans from northern Kansas, Nebraska, and Iowa - north of all the new records (Table I ) - suggesting that the expansion may not bear a simple relationship to climate. A possible behavioral basis for range expansion in P. exclamans was suggested by field observations of newly founded Polistes colonies on and near the University of Oklahoma Biological Station (Marshall County, Oklahoma). Nests of P. annularis, P. apachus, P. fuscat,us, P. nzetricus and P. rubiginosus observed between 20 April and I I May, 1966, were commonly attended by more than one female (foundress). However, each except one of more than 100 P. ex- clamans colonies was attended by a single female; the one exception had only two foundresses. Solitary nest founding2 predominated even 'Phil Rau's son, Mr. David Rau, kindly helped in locating these sites. '1 prefer this term to the older "haplometrosis') or "monogyny", and the term "social nest founding" to "pleometrosis" or "polygyny", for reasons given in West, 1967b.




================================================================================

Psyche
Table 1
New state records of Polistes exclamans Viereck. State
Illinois1
Indiana
Kentucky
Maryland
New Jersey
New Mexico
Tennessee
Virginia
Locality Date
Williamson Co.
Posey Co.
Perry Co.
Floyd Co.
Nelson Co.
Bardstown
Prince George Co.
// I/ //
Public Landing
Plummers Island
Salisbury
Cambridge
Columbia
Boone Co.
St. Louis Co., Jackson Co.
Penn State Forest
Lebanon State Forest
Metuchen
Eddy Co.
Roswell
Benton Co,
Gloucester
Eagle Rock
York Co.
Collection
Southern Illinois Univ.'
Purdue Univ.
I/ //
I/ I1
Purdue Univ.
I/ //
Univ. of Maryland
Los Angeles County Mus.
Cornell Univ.
// /I
see Porter, 1963
I/ // //
Univ. of Calif. Davis
Univ. of Missouri
Mus. of Compar. Zool.
see Porter, 1963
// // /I
// I/ //
Ccrnell Univ.
Purdue Univ.
Los Angeles County Mus.
Univ. of Maryland
Los Angeles County Mus.
// /I // /I
at "popular" nesting sites: eighteen single-foundress exclamans colo- nies located in close proximity beneath the porch roof of a small abandoned cottage were dispersed, rather than clustered, beneath the shelter. Furthermore, P. exclamans was the only Polistes species among the six common in the area to occupy artificial nesting places erected at the Biological Station on I April - the species was ex- ceptional in colonizing these newly available sites. Eickwort (in *The collection of the Illinois Natural History Survey contains IS speci- mens of P. cxclamans from Macoupin Co. Illinois for the years 1890- 1910, and the University of Missouri collection contains one specimen re- corded from Boone Co., Missouri,
1940. However, these states are listed
as "new records" here because they were not included in Bohart, 1951, and because of the paucity of P. exclamans in early collections from these areas, 2collection examined by R. M. Bohart.




================================================================================

19681 West - Polktes exclanzans 12 1
press) reports that solitary nest founding also redom mi nates in Kan- sas populations of P. exclanzans.
These associated tendencies of P. exclamans foundresses to disperse and to move into newly available nesting sites would promote range expansion. In contrast, social foundresses tend to remain and re- produce near their place of origin.
In Polistes fuscatus and P. can-
adensis, foundress associations are composed of siblings which begin new colonies near the parental nest site (West, 1967a), and there is indirect evidence that this occurs in other species having social nest founding (see Hamilton, 1 964 ; West, I 967a). The mode of nest founding of P. exclamans within the newly oc- cupied portions of its range is unknown. While most foundresses were solitary in the populations of P. exclamans observed in Okla- homa and Kansas (above), the same species comn~only exhibits social nest founding in Texas (Rau, 1943 ; Caskey, 1955). Similarly, Rau ( 1942b)
found only solitary foundresses in P. fuscatus (= "vari- atus") in Missouri, whereas in Michigan social nest founding was most common in that species (West, I 967a b) . Geographic variation in mode o'f nest founding has also been noted in the European wasp P. gallicus (see Hamilton, 1964). Thus, while it has been traditional in the literature on social wasps (e.g., Wheeler, 1922) to regard mode of nest founding as a constant feature of a species, it is prob- ably better to consider most Polistcs "polyethic" in this respect, with mean size of foundress group differing from locality to locality and from year to year. Various factors, including parasitization (Eick- wort, in press), nest site availability, number of overwintering sib- lings, and dominance relations among foundresses, may affect the frequency of solitary nest founding and the size of foundress groups (see West, 1967a). As the observations of this study suggest, soli- tary nest founding might be expected to increase in frequency in areas where there is opportunity for (and advantage in) expansion; and the ability of some Polistes species to colonize new sites might be limited or delayed by selection for social nest founding. Distribution data from labels of determined specimens of P. excla?mns were sent to me by entomologists at the following in- stitutions: United States National Museum (A. Alenke) ; Univer- sity of California, Davis (R. M. Bohart) ; University of Kansas (G. W. Byers) ; Kansas State University (H. D. Blocker) ; Illinois



================================================================================

122 Psyche [June
Natural History Survey (W. E. LaBerge) ; University of Arkansas (E. P. Rouse) ; University of Missouri (W. R. Ems) ; a,nd Los Angeles County Museum of Natural History (E. M. Fisher). R.
M. Bohart kindly sent data from other collections examined by him, and H. E. Evans examined specimens of P. exclamans at the Amer- ican Museum of Natural History. I have personally obtained data from collections at Cornell University and the Museum of Compara- tive Zoology.
This research was supported by NSF grant GB-336 to the Uni- versity of Michigan Museum of Zoology and a grant from the Com- mittee on Evolutionary Biology at Harvard University. Howard E. Evans, William G. Eberhard, Janice Matthews and Robert Matthews made helpful criticisms of the manuscript. Kath- leen Eickwort kindly made available the pre-publication manuscript of her pa,per on Polistes exclanz~ns.
BOHART, R. M.
1951. Family Vespidae, in C. F. W. Muesebeck, K. V. Krombein, H. K. Townes, editors, Hymenoptera of America North of Mexico, U.S.D.A. Agriculture Monograph, 2: 875-907. CASKEY, D.
1955. The behavior of Polistes exclamans. Unpublished Master's Thesis, Baylor University.
EICKWORT, K.
Separation of the castes of Polistes exclamans and notes on its biology (Hym. Vespidae) .
Insectes Sociaux. In press.
HAMILTON, W. D.
1964. The genetical evolution of social behavior. 11. J. Theoret. Biol. 7: 17-52.
MAYR, E.
1963. Animal Species and Evolution. Belknap Press, Cambridge, Mass. xiv + 797 pp.
PORTER, C. C.
1963. Some new locality records for the social wasp Polistes exclamans Viereck (Hymenoptera, Vespidae). Ent. News 74(7) : 176. RAU, PHIL.
1929. The habitat and dissemination of four species of Polistes wasps. Ecology lO(2) : 191-200.
1941. Observations on certain lepidopterous and hymenopterous para- sites of Polistes wasps. Ann. Ent. Soc. Amer. 34: 355-366. 1942a. Habitat preferences of the Polistes wasp. Ent. News 53: 293-95. 1942b.The nesting habits of Polistes wasps as a factor in taxonomy. Ann. Ent. Soc. Amm. 3 5 (3) : 335-338.
1943. The nesting habits of Mexican social and solitary wasps of the family Vespidae. Ann. Ent. Soc. Amer. 36: 515-536. Unpublished manuscript. The bionomics of Polistes wasps.



================================================================================

19681 west - .Pol&es exclamam 123
RAU, PHIL AND NELLIE RAU.
1918. Wasp Studies Afield. Princeton Univ, Press, Princeton, N. J. xv -l- 372 pp.
WEST, M. J.
1967a. The social biology of polistine wasps. Doctoral Dissertation, University of Michigan, Ann Arbor (University Microfilms, Ann Arbor, Michigan).
1967b. Foundress associations in polistine wasps: dominance hierarchies and the evolution of social behavior. Science 157(3796): 1584- 1585.
WHEELER, W, M.
1922.
Social Life Among the Insects.
Constable & Co., Ltd,, London.
375 pp*




================================================================================

THE MATING BEHAVIOR OF
GROAfPHADORHINA PORTENTOSA (SCHAUbI)
(BLATTARIA, BLABEROIDEA) BLABERIDAE!
OXYHALOINAE) : AN ANOfi!IALOUS PATTERN
FOR A COCKROACH1
Department of Zoology
The Uni\~ersity of Texas at Austin
This con~n~unication is the fourth in a series of largely descriptive papers dealing with the mating behavior of cockroaches. (see Barth) 1961 ; Barth, 1964 ; Roth and Barth) I 967 ; and Earth, m,s in prep- aration). The aim of this series is twofold: first to provide back- ground information for experimental studies, and second to provide the detailed comparative information necessary for a study of the evolution of mating behavior within the Blattai-ia. A more gmeral introduction to the series may be found in Barth (1964). The anomalous mating behavior of the Madagascar cockroach, Grompha- dorhina portentosa, (Schaunl) forms the subject of this conmuni- cation.
AIATERIALS AND METHODS
Stock cuIt~11-es of G. portentosa were maintained as described by Barth (1964) for Byrsoiria funzigata. The observations on mating behavior weTe made in the evening (the normal active period for these animals) under red illumination is specially designed obser- vation chambers (for details) see Barth, 1964). The ethological terms employed in the description of the behavior patterns have been previously defined by Bai-th ( I 964) .
RESCLTS AND CONCLUSIONS
Gromphadorhina portentosa is a large, heavy-bodied wingless spe- cies found under debris on forest floors in Madagascar (see Plate 6 'No. 4 in a series of papers entitled "The Mating Behavior of Cock- roaches".
'Much of this work was carried out at the Biological Laboratiories) Har- vard University. Financial suppost from National Science Foundation Predoctoral Fellowships and N.S.F. Grant G 19962 is gratefully acknowl- edged.
Manuscrifit received by the editor 5 April, 1968



================================================================================