Behaviour and life history of a large carpenter bee (Xylocopa virginica) in the northern extent of its range

Large carpenter bees (Hymenoptera: Apidae: Xylocopa) have traditionally been thought of as exhibiting solitary or occasionally communal colony social organization. However, studies have demonstrated more complex fonns of social behaviour in this genus. In this document, I examine elements ofbehaviour and life history in a North American species at the northern extreme of its range. Xylocopa virginica was found to be socially polymorphic with both solitary and meta-social or semi-social nests in the same population. In social nests, there is no apparent benefit from additional females which do not perfonn significant work or guarding. I found that the timing of life-history events varies between years, yet foraging effort only differed in the coldest and wettest year of2004 the study. Finally, I that male X virginica exhibit female defence polygyny, with resident and satellite males. Resident males maintain their territories through greater aggression relative to satellites.

Nestmate recognition in the large carpenter bee, Xylocopa virginica /

Abstract Many species of social insects have the ability to recognize their nestmates. In bees, sociality is maintained by bees that recognize which individuals should be helped and which should be hanned in order to maximize fitness (either inclusive or individual) (Hamilton 1964; Lin and Michener 1972). Since female bees generally lay eggs in a single nest, it is highly likely that bees found cohabitating in the same nest are siblings. According to the kin selection hypothesis, individuals should cooperate and avoid aggression with same sex nestmates (Hamilton 1964). However, in opposite sex pairs that are likely kin, aggression should increase among nestmates as an expression of inbreeding avoidance (Lihoreau et al. 2007). Female bees often guard nest entrances, recognizing and excluding foreign conspecific females that threaten to steal nest resources (Breed and Page 1991). Conversely, males that aggressively guard territories should avoid aggression towards other males that are likely kin (Shellman-Reeve and Gamboa 1984). In order to test whether Xy/ocopa virginica can distinguish nestmates from non-nestmates, circle tube testing arenas were used. Measures of aggression, cooperation and tolerance were evaluated to detennine the presence of nestmate recognition in this species. The results of this study indicate that male and female X virginica have the ability to distinguish nestmates from non-nestmates. Individuals in same sex pairs demonstrated increased pushing, biting, and C-posturing when faced with non-nestmates. Males in same sex pairs also attempted to pass (unsuccessfully) nOIl-nestmates more often than ncstmates, suggesting that this behaviour may be an cxpression of dominancc in males. Increased cooperation exemplified by successful passes was not observed among nestmates. However, incrcased tolerance in the [onn of head-to-head touching was observed for nestmates in female same sex and opposite sex pairs. These results supported the kin selection hypothesis. Moreover, increased tolerance among opposite sex non-nestmates suggested that X virginica do not demonstrate inbreeding avoidance among nestmates. 3 The second part of this study was conducted to establish the presence and extent of drifting, or travelling to different nests, in a Xylocopa virgillica population. Drifting in flying Hymenoptera is reported to be the result of navigation error and guard bees erroneously admitting novel individuals into the nest (Michener 1966). Since bees in this study were individually marked and captured at nest entrances, the locations where individuals were caught allowed me to determine where and how often bees travelled from nest to nest. Ifbees were captured near their home nests, changing nests may have been deliberate or explained by navigational error. However, ifbees were found in nests further away from their homes, this provides stronger evidence that flying towards a novel nest may have been deliberate. Female bees are often faithful to their own nests (Kasuya 1981) and no drifting was expected in female X virginica because they raise brood and contribute to nest maintenance activities. Contrary to females, males were not expected to remain faithful to a single nest. Results showed that many more females drifted than expected and that they were most often recaptured in a single nest, either their home nest or a novel nest. There were some females that were never caught in the same nest twice. In addition, females drifted to further nests when population density was low (in 2007), suggesting they seek out and claim nesting spaces when they are available. Males, as expected, showed the opposite pattern and most males drifted from nest to nest, never recaptured in the same location. This pattern indicates that males may be nesting wherever space is available, or nesting in benches nearest to their territories. This study reveals that both female and male X virginica are capable of nestmate recognition and use this ability in a dynamic environment, where nest membership is not as stable as once thought.

Evolutionary Origin and Maintenance of Sociality in the Small Carpenter Bees

Many arthropods exhibit behaviours precursory to social life, including adult longevity, parental care, nest loyalty and mutual tolerance, yet there are few examples of social behaviour in this phylum. The small carpenter bees, genus Ceratina, provide important insights into the early stages of sociality. I described the biology and social behaviour of five facultatively social species which exhibit all of the preadaptations for successful group living, yet present ecological and behavioural characteristics that seemingly disfavour frequent colony formation. These species are socially polymorphic with both / solitary and social nests collected in sympatry. Social colonies consist of two adult females, one contributing both foraging and reproductive effort and the second which remains at the nest as a passive guard. Cooperative nesting provides no overt reproductive benefits over solitary nesting, although brood survival tends to be greater in social colonies. Three main theories explain cooperation among conspecifics: mutual benefit, kin selection and manipulation. Lifetime reproductive success calculations revealed that mutual benefit does not explain social behaviour in this group as social colonies have lower per capita life time reproductive success than solitary nests. Genetic pedigrees constructed from allozyme data indicate that kin selection might contribute to the maintenance of social nesting -, as social colonies consist of full sisters and thus some indirect fitness benefits are inherently bestowed on subordinate females as a result of remaining to help their dominant sister. These data suggest that the origin of sociality in ceratinines has principal costs and the great ecological success of highly eusociallineages occurred well after social origins. Ecological constraints such as resource limitation, unfavourable weather conditions and parasite pressure have long been considered some of the most important selective pressures for the evolution of sociality. I assessed the fitness consequences of these three ecological factors for reproductive success of solitary and social colonies and found that nest sites were not limiting, and the frequency of social nesting was consistent across brood rearing seasons. Local weather varied between seasons but was not correlated with reproductive success. Severe parasitism resulted in low reproductive success and total nest failure in solitary nests. Social colonies had higher reproductive success and were never extirpated by parasites. I suggest that social nesting represents a form of bet-hedging. The high frequency of solitary nests suggests that this is the optimal strategy when parasite pressure is low. However, social colonies have a selective advantage over solitary nesting females during periods of extreme parasite pressure. Finally, the small carpenter bees are recorded from all continents except Antarctica. I constructed the first molecular phylogeny of ceratinine bees based on four gene regions of selected species covering representatives from all continents and ecological regions. Maximum parsimony and Bayesian Inference tree topology and fossil dating support an African origin followed by an Old World invasion and New World radiation. All known Old World ceratinines form social colonies while New World species are largely solitary; thus geography and phylogenetic inertia are likely predictors of social evolution in this genus. This integrative approach not only describes the behaviour of several previously unknown or little-known Ceratina species, bu~ highlights the fact that this is an important, though previously unrecognized, model for studying evolutionary transitions from solitary to social behaviour.

The Implications of Forager Behaviour for Social Organisation in a Socially Polymorphic Carpenter Bee (Xylocopa virginica)

In social Hymenoptera, the division of labour is a major step in the evolution of sociality. Bees, which express many different kinds of sociality, can be classified according to how individuals share or do not share foraging and reproductive activities (Michener, 1974). The large carpenter bee, Xylocopa virginica, lives in populations with both solitary and social nests. In social nests, reproduction is controlled by the dominant female, who does all of her own foraging and egg-laying, while the subordinates guard the nest only. This study examined foraging behaviour as a way to classify the social hierarchy. Individual females were marked, measured and intensely observed for the foraging season. It was found that a large number of subordinates forage and likely obtain more reproductive fitness than previously thought. The dominance hierarchy is very likely a social queue, in which bees take turns foraging and egg-laying.

Indenture between Barnet Ulman, Benjamin Ulman, Peter Ulman and Philophilus Carpenter

Indenture between Barnet Ulman and Peter Ulman of Niagara Township to Asa Philophilus Carpenter of Niagara Township regarding 300 acres in Lot no. 15 in the 1st Concession and the north half of Lot no. 11 in the 2nd Concession of the Township of Southwood - instrument no. 680. This was listed in folio 287, November 9, 1852.

Indenture between Asa Philophilus Carpenter and Joseph Augustus Woodruff

Indenture (vellum) of deed with dower from Asa Philophilus Carpenter and Margaret of Niagara to Joseph Augustus Woodruff of Niagara for lot no. 15 in the 1st Concession in the County of Elgin. This was registered April 14, 1853 in folio 288 - instrument no. 681, April 2, 1853.

Indenture between Robert Miller of the County of La Porte, Indiana and Asat Carpenter of the Town of Niagara regarding 50 acres in the Township of Southwold, Middlesex

Indenture between Robert Miller of the County of La Porte, Indiana and Asat Carpenter of the Town of Niagara regarding 50 acres in the Township of Southwold, Middlesex. There are 2 copies of this document tin which the wording seems slightly different. The 1st copy is slightly burned on the side and extremely faded, making it completely illegible. The 2nd copy is legible. Feb. 15, 1853

Women's Volleyball Team (1981-82)

Back Row: Maura Purdon (Coach), Kelly Grantham, Liz Jansen, Diane Thiesen, Louise Argenta, Rhonda, Oatman, Kelly Fahlenbock (Asst. Coach) Middle Row: Kaaren Quartermain, Sherri Crossman, Sue Crowley, Kin Zamecnik Front Row: June LeDrew, Darlene Danis Absent: Margo Schijns

Effect of pH on non-photochemical quenching in spinach photsystem 2 particles as measured by picosecond fluorescence decay kinetics

Single photon timing was used to study picosecond chlorophyll a fluorescence decay kinetics of pH induced non-photochemical quenching in spinach photosystem 2 particles. The characteristics of this quenching are a decrease in chlorophyll a fluorescence yield as well as a decrease in photochemistry at low pH. Picosecond kinetics of room temperature fluorescence temporally resolve the individual components of the steady state fluorescence yield into components that are related to primary energy conversion processes in photosystem 2. Four components were resolved for dark adapted (Fo), light saturated (Fm), and chemically reduced (Nadithionite) photosystem 2 reaction centres. The fastest and slowest components, indicative of energy transfer to and energy capture by the photosystem 2 reaction centre and uncoupled ("dead") chlorophyll, respectively, were not affected by changing pH from 6.5 to 4.0. The two intermediate components, indicative of electron transfer processes within the reaction centre of photosystem 2, were affected by the pH change. Results indicate that the decrease in the steady state fluorescence yield at low pH was primarily due to the decrease in lifetime and amplitude of the slower of the intermediate components. These results imply that the decrease in steady state fluorescence yield at low pH is not due to changes in energy transfer to and energy capture by the photosystem 2 reaction centre, but is related to changes in charge stabilization and charge recombination in the photosystem 2 reaction centre.

John Smith, M.P fonds, 1930, 1949-1950, 1954, 1957-1962, n.d.

John Smith (1894-1977), son of Daniel Smith and Annie Douglas was a native of Scotland, immigrating to Canada in 1913. He first worked as a coach builder, then as a carpenter, finally developing his own contracting business. During WWI he served overseas with the 10th Battery, RCA as a sergeant. In 1924 Smith married Jean Wood, and together they had a daughter Irene (Hugh Langley). Smith first entered politics in 1940 serving as an alderman for the next 11 years. In 1954 he was elected mayor of the city of St. Catharines, and was twice returned to office by acclamation, serving until 1957 when he successfully ran as the Progressive Conservative candidate representing Lincoln County in the federal election. He won the election by a 10 000 vote majority. He served his constituents in Ottawa until he was defeated in the 1962 election. After leaving politics Mr. Smith was active in his community. He spearheaded the establishment of the St. Catharines Museum, and then was appointed its first director in 1966, serving in that capacity until 1972. He was an active member of the board of governors of the St. Catharines General Hospital and a life member and former president of the Lincoln County Humane Society. In 1971 he was voted Citizen of the Year for the city of St. Catharines. John Smith died on February 8, 1977 and was buried at Victoria Lawn Cemetery. Source: The St. Catharines Standard, February 9, 1977, page 1

Photo album, St. Catharines area, ca. 1913-1919.

One hardcover photo album containing black and white photos. Many of the photos were taken in the St. Catharines area. Included are photos of Port Dalhousie, Port Weller, Niagara Falls, Niagara-on-the-Lake and St. Catharines. There are also photos of Braeside, Ont. and the Ottawa valley. Various local landmarks are included, such as the armoury in St. Catharines, Montebello Park, and Martindale pond. Some of the events captured include a train wreck that occurred in St. Catharines in 1914, the visit of the Governor General to St. Catharines in 1914 (featuring the Carnegie library and Post Office and federal building decorated with flags), and an airplane that crashed into a body of water, possibly a plane from an air training camp in Beamsville during World War I. There are also two photos of champion Niagara district basketball teams, possibly taken in the gymnasium building located behind the former St. Catharines Collegiate building (later Robertson School) on Church Street. One photo includes Norman Byrne, Gladys Ansell, Miriam Marshall, Irene Stoter (?), Mildrerd Houston, A. Gardner, and Madeline Jenner. The other photo includes George Moase, W. Bennett, Norman Byrne, Jack Bain, Mr. Brackenbury, Cyril Merriman, Jim Galway, Harry Erskine, and Roy Carpenter.

Exploring Perceptions of Accessibility, Necessity and Use of Social Support for Wilderness Therapy Field Instructors

Organizations offering therapeutic wilderness programming have a responsibility to ensure the well-being of their front line employees. A system of social support that is formed through communication with others, either personally or professionally, can assist field instructors in effectively managing the demands arising from their work. Phenomenological analysis of semi-structured interview transcripts from seven participants provided insight on perceptions of necessity, accessibility and use of social support. Fourteen main themes and thirteen subthemes emerged from the data. Findings are presented using the six components of Parsons’ (1980) staff development model and strongly suggest program managers consider and apply specific measures aimed at increasing the social support for front line field instructors in a wilderness therapy work context.

Schram Family of Wainfleet Township documents, 1839-1901 (not inclusive)

George William Schram (1809-1885), son of Garrett Schram and Leah Van Etten, married Orpha Pearson on Nov.13, 1835. His son Marsena John Schram (farmer) was born in May of 1840, in Canada. He died on Nov. 17, 1926 in Wexford County, Michigan. He was married in 1867 to Sarah (1825-1887).Marsena married again on April 18, 1910 to Ann Clarinda Warner (1861-1924). He was working as a carpenter at this time. They had another son, William who was born about 1838 and he married Sabina Chambers on Jan. 21, 1862. The 1861 census for Wainfleet lists siblings of Marsena John Schram as Sarah J. (age 14), Georgiana (age 5), and William (age 21). The Schrams lived on Concession 5 and owned approximately 144 acres of land. David Thompson was born Feb. 4, 1873 and died Feb. 19, 1951. He married Sally Ann Wilson on Sept. 7, 1825 in Pelham. She died about 1840 in Indiana Ontario (near Cayuga). Lemuel Victor Hogue was born Dec.1, 1854 and died Jan. 12, 1929. He was married to Elizabeth Wills who was born Aug. 2, 1861 and died Mar. 8, 1926. Sources: http://www.findagrave.com/cgi-bin/fg.cgi?page=gr&GRid=99294842 http://wc.rootsweb.ancestry.com/cgi-bin/igm.cgi?op=GET&db=seadragon5&id=I91708

Proposal for Construction of Brock’s Monument, Queenston, newspaper fragment, 1824

Brock’s Monument is owned by Parks Canada and maintained by the Niagara Parks Commission in collaboration with the Friends of Fort George and Niagara National Historic Sites. It is located in Queenston Heights Park atop the Niagara Escarpment. On March 14, 1815, Parliament passed an act to erect a monument to the memory of General Isaac Brock. A design by engineer Francis Hall was selected. He envisioned a 135 ft. tall Tuscan column, made out of stone with a winding staircase inside. By the spring of 1824, work had begun on the monument. In June of that year, the cornerstone was laid and William Lyon Mackenzie was in attendance at the ceremony. It was on October 13th, 1824 (the anniversary of Brock’s death) that 6000 people traveled to Queenston to inter the remains of Brock and Lieutenant-Colonel Macdonell. This was the second burial for both. After 3 years the tower had reached 135 feet, but there was no inscription at the base, the fence around the observation deck had not been installed and there was no statue of Brock. Hall submitted a plan to finish the statue, but he was turned down and a simple ornament was placed where the Brock statue should have been. A massive blast of gunpowder destroyed the monument in 1840. It is alleged that an American sympathizer with the Upper Canada Rebellion set off the blast. Brock and Macdonell’s bodies were reburied in the Hamilton Family Cemetery in Queenston. The present monument was rebuilt in 1853. William Thomas (designer of St. Michael’s Cathedral in Toronto) was the architect. Brock and Macdonell were once again laid to rest in separate vaults at the statue. In 1968, Brock’s Monument was declared a national historical site. In 2005, it was closed to the public due to safety concerns, but it reopened in 2010. Source: http://www.thecanadianencyclopedia.com/articles/brocks-monument-queenston-heights

Alloparental care in a solitary bee

Research into the evolutionary origins of sociality in insect colonies is changing emphasis from understanding how eusociality is maintained to how insects transition from solitary to social lifestyles. The pygmy carpenter bees (Ceratina spp.) offer an excellent model for investigating such factors as they have been historically thought of as solitary but have recently been shown to be socially polymorphic, which may indicate that they are currently in a transitive phase. By utilizing behavioural observation and experimental removal protocols, I show that extended parental care, as well as sibling care in Ceratina calcarata plays an important role in offspring development. I found, upon removal of the mother, that specifically produced ‘dwarf’ female offspring take over parental care roles in the nest. The existence of alloparental care and generational overlap suggests that although they are classified as solitary bees, C. calcarata possess the prerequisite behavioural repertoire for sociality.