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Journal of Comparative Physiology A https://doi.org/10.1007/s00359-021-01492-4

ORIGINAL PAPER

Neuroanatomical differentiation associated with alternative reproductive tactics in male arid land bees, Centris pallida

and Amegilla dawsoni

Meghan Barrett1 · Sophi Schneider2 · Purnima Sachdeva1 · Angelina Gomez1 · Stephen Buchmann3,4 Sean O’Donnell1,5
·

Received: 1 February 2021 / Revised: 19 May 2021 / Accepted: 22 May 2021 © The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature 2021

Abstract

Alternative reproductive tactics (ARTs) occur when there is categorical variation in the reproductive strategies of a sex within a population. These different behavioral phenotypes can expose animals to distinct cognitive challenges, which may be addressed through neuroanatomical differentiation. The dramatic phenotypic plasticity underlying ARTs provides a powerful opportunity to study how intraspecific nervous system variation can support distinct cognitive abilities. We hypothesized that conspecific animals pursuing ARTs would exhibit dissimilar brain architecture. Dimorphic males of the bee species Centris pallida and Amegilla dawsoni use alternative mate location strategies that rely primarily on either olfaction (large-morph) or vision (small-morph) to find females. This variation in behavior led us to predict increased volumes of the brain regions supporting their primarily chemosensory or visual mate location strategies. Large-morph males relying mainly on olfaction had relatively larger antennal lobes and relatively smaller optic lobes than small-morph males relying primarily on visual cues. In both species, as relative volumes of the optic lobe increased, the relative volume of the antennal lobe decreased. In addition, A. dawsoni large males had relatively larger mushroom body lips, which process olfactory inputs. Our results suggest that the divergent behavioral strategies in ART systems can be associated with neuroanatomical differentiation.

Keywords Alternative mating tactics · Sensory differentiation · Solitary bees

Introduction

insects, and include both behavioral and morphological trait variation (Shuster and Wade 2003; Paxton 2005; Oliveira et al. 2008; Shuster 2010). ARTs evolve when fitness gains can be achieved by pursuing divergent mating strategies, leading to selection on phenotypes that maximize the success of two or more specialized morphs (Shuster 2010). Because morphs often develop via phenotypic plasticity, ARTs allow for the study of behavioral and morphological variation within a population that are not dependent upon genotypic differences (Kukuk 1996; reviewed in Oliveira et al. 2008). In this way, ARTs provide a unique and powerful opportunity to explore the evolution of relationships between behavioral and morphological specialization.
Neuroecology theory predicts that relative investment in functionally discrete regions of the brain will be correlated to the cognitive demands of an organism’s environment/ behaviors, due to constraints imposed by the high metabolic cost of producing and maintaining neural tissue (Aiello and Wheeler 1995; Sherry 2006; Liao et al. 2016; Niven 2016; Luo et al. 2017). Energy limitation thus places neural
Alternative reproductive tactics (ARTs) occur when there is categorical variation in the mating-related behaviors or traits of same-sex individuals within a population (Oliveira et al. 2008). ARTs have evolved in diverse animal taxa, including fish, crustaceans, birds, amphibians/reptiles, mammals, and

* Meghan Barrett [email protected]

1

Department of Biology, Drexel University, Philadelphia, PA, USA

2

Upper Dublin, PA, USA

345

Department of Entomology, University of Arizona, Tucson, AZ, USA

Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ, USA

Department of Biodiversity, Earth, and Environmental Science, Drexel University, Philadelphia, PA, USA

Vol.:(0123456789)

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Journal of Comparative Physiology A

systems under selective pressure for optimal investment (Niven and Laughlin 2008). Because successful mate location behaviors are expected to be strongly linked to reproductive success, different sensory mate location strategies could be associated with neuroanatomical differentiation in the regions of the brain that support those sensory systems. We hypothesized that conspecific animals utilizing ARTs would have different brain architecture patterns, related to the cognitive demands of their morph-specific mating behaviors. chase after females ‘upwind’ of them, as males are oriented 360° around the vegetation—not just on the downwind side— and always orient facing the largest open area visually, away from the vegetation and not necessarily into the wind (Barrett, personal observation).
Similarly, the large ‘majors’ of A. dawsoni exhibit a fixed strategy, patrolling female emergence sites and likely using cuticular hydrocarbon cues to locate females waiting within emergence tunnels, before fighting other males and/or guarding potential mates (Alcock 1997; Simmons et al. 2003). Smaller A. dawsoni ‘minor’ males rarely patrol emergence sites, instead typically hovering or patrolling near blooming host plant vegetation and use visual cues to locate females (Houston 1991; Alcock 1997).

The male morphs of Centris pallida Fox, found in the

Sonoran Desert of the USA and Northern Mexico, and Amegilla dawsoni Rayment (Dawson’s burrowing bee), found in the deserts of Western Australia, use alternative sensory mate location strategies. In C. pallida, large-morph males are morphologically distinct based on coloration and hind leg morphology (called ‘metanders’ with ‘swollen’ legs in Snelling 1984 or ‘largest males’ in Alcock et al. 1977). These males have a fixed mate location strategy, using chemosensory cues to patrol close to the soil surface in search for females buried underground. Males land near buried females and repetitively touch the soil with their antennae (typical insect odor-tracking behavior on a surface; Wenner 1974), before digging up the buried female (Alcock et al. 1977). Chemosensory cues are both sufficient and necessary for locating females–males will dig up dead, buried females that are not moving or visible but will not dig up vibrating objects (Alcock et al. 1976). Other males will fight with the digging male for the opportunity to mate with the emerging females, with the largest male typically winning and copulating with the female (Alcock 1976, 2013; Alcock et al. 1976, 1977).
In summary, the large-morph males (e.g. metanders or majors) of both species are behaviorally fixed on using chemosensory cues to locate females (Alcock et al. 1977; Alcock 1997; Simmons et al. 2003), while the small-morph males are more behaviorally flexible but rely more heavily on visual cues to locate mates when hovering. We predicted brain structure would differ between the male morphs in both species, with relative increases in tissue investment in brain regions that support mating-tactic relevant cognitive abilities.
We analyzed whether relative brain investment patterns differed between the morphs of each species in their antennal lobes (AL), which receive chemosensory information from the antennae, and their optic lobes (OL), which receive visual information from the eyes (Kenyon 1896). We tested for a negative correlation in investment between visual and olfactory brain regions, which are seen in comparative studies of other insects (often described as trade-offs: Niven and Laughlin 2008, Stöckl et al. 2016; Kessey et al. 2019; Özer and Carle 2020). In addition to analyzing investment in peripheral sensory brain regions, we asked whether the relative volumes of the mushroom body (MB) calyces (we analyzed the lip and collar separately) differed between the morphs. The mushroom bodies are involved in learning, memory, and sensory integration, and receive olfactory input to the MB calyx lip and visual input to the MB calyx collar (Fahrbach 2006; Paulk and Gronenberg 2008). We thus predicted that the large-morph males that are fixed on chemosensory cues for mate location would have relatively larger MB lips and ALs, and relatively smaller MB collars and OLs, as compared with the small-morph males that rely more heavily on vision but are behaviorally flexible.
In contrast to the fixed mate location strategy the large-male morph uses, small-morph males can be behaviorally flexible. They may patrol the ground like large-morph males or alternatively hover near vegetation, chasing after and mating with any females or mating pairs they locate visually (Alcock 1976, 1979, 1984; Alcock et al. 1977). Visual cues are likely the only or primary sensory strategy used while hovering. First, hovering males chase any insect passing through their visual field (even those several meters away, C. pallida males, and non-C. pallida insects). Second, males only chase nearby hovering males when they enter their visual field, even though they are often hovering less than a meter away for several hours. If olfaction were in use, males would detect, orient towards, and chase after nearby males even when not in their visual field (e.g. drones of Apis mellifera, Brandstaetter et al. 2014). Third, male C. pallida do not discriminate between the odors of male or female C. pallida (when digging Alcock et al. 1976; and even in close contact, Alcock and Buchmann 1985), making it unlikely that they are tracking the odor of a single, fast-moving female bee in the midst of a turbulent aggregation of thousands of fast-moving C. pallida males. Fourth, males frequently

Materials and methods

Specimen collection

Centris pallida males (n=27) were collected in late April and early May of 2018 at Tonto National Forest in Arizona

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Journal of Comparative Physiology A

Fig. 1 Characteristic coloration difference in large and small C. pallida male morphs. Largemorph (metandric; Snelling

1984) C. pallida males are

a light gray coloration with ‘swollen’ hind femurs (bee on the right) while small-morph C. pallida males are a dark brown on the thorax, with thin hind femurs (bee on the left). Bees were collected in 2018, from the same population used in this study

(33.552, − 111.566). A. dawsoni males (n = 19) were collected on July 29, 2019 at the Carnarvon Pistol Range, Western Australia (− 24.917, 114.725). Dense emergence and nesting aggregations at both sites have persisted for several decades. C. pallida males were transported in a cooler on ice to a lab where they were weighed on an analytical balance to the nearest 0.1 mg; males of A. dawsoni were transported on ice from the field to a lab for dissection within six hours. Heads were cut from the thorax and placed immediately into Prefer fixative (Anatech, Ltd.) following transport (A. daw-

soni) or weighing (C. pallida).

ethanol concentrations, acetone, and then increasing concentrations of plastic resin. Resin was composed of 5.5 g EMbed 812, 5.7 g DDSA (dodecenyl succinic anhydride), 0.65 g DBP (dibutyl phthalate), and 0.31 g of DMP (2,4,6-tri(dimethylaminoethyl)phenol; all Electron Microscopy Sciences products). Heads were incubated at 60 °C for 72 h inside cylindrical molds, until the resin hardened.
Heads were cut along the frontal plane into 14 um thick sections using a rotary microtome and disposable steel histology blades. Sections were mounted on gelatin-coated microscope slides and stained with Toluidine blue stain, then cleared in a series of distilled water, increasing ethanol concentrations, and Histochoice clearing medium (SigmaAldrich) and kept in the oven at 60 °C for 4 h. Slides were coverslipped under DEPEX transparent mounting medium (Electron Microscopy Sciences).
For C. pallida, males were classified as the ‘large’ male morph if they were found patrolling, digging, or fighting and had the gray/white coloration and leg morphology distinctive to this ‘large’ behaviorally inflexible morph (Fig. 1; Alcock 1976; Alcock et al. 1977; Snelling 1984). They were classified as ‘small’ male morphs if they were collected hovering near vegetation, or if they were found patrolling or digging but did not have the distinct morphology/coloration of the large morph (Alcock et al. 1977; Snelling 1984).
For A. dawsoni, males were classified as small-morph
(minor) males if they were collected near vegetation away from emergence areas. Alcock (1997) classified emergencesite patrolling males as large-morph (major) males if their head width was greater than 6.3 mm; Houston (1991) and Alcock (1997) found a distinct separation in head width around 6.0–6.2 mm between majors and minors. This criterion was used to classify large-morph males at patrolling sites in our study.
A compound light microscope-mounted digital camera was used to photograph every section containing brain tissue at 2560×1920 pixel resolution at 2.5× microscope objective and a 1× camera mount. Digital photographs were taken using LAS V4.9 software, with sharpness set to robust.

Neuroanatomical measurements

ImageJ (Schneider et al. 2012) was used to quantify the area of brain regions by outlining the neuropil of the target region on every other section using the freehand selections tool, quantifying the number of pixels in the structure, and converting the pixel count to area using a photograph of a stage micrometer taken at the same resolution and magnifi- cation. We then multiplied the area by section thickness and finally added up all the volumes across all slides to obtain total brain region volumes in cubic millimeters. We quantified the following regions: mushroom bodies (calyx lip,

Histological sectioning

Brain tissue was dissected completely out of the head capsule and dehydrated through a series of increasing

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Journal of Comparative Physiology A

calyx collar, and basal ring + peduncles together), central complex, central brain mass (protocerebrum, protocerebral bridge, subesophageal ganglion), antennal lobes, optic lobes (medulla and lobula only). We did not measure Kenyon body cell volumes.

df = 25, p = 0.0013; A. dawsoni: n = 19, t = 2.60, df = 17,

p = 0.0188). In contrast, the large-morph males, which rely exclusively on a chemosensory mate location strategy, had relatively larger antennal lobes (Fig. 2b; unpaired

t test; C. pallida: t = 3.26, df = 25, p = 0.0032; unpaired t test with Welch’s correction; A. dawsoni: t=3.38, d f=7.62, p=0.0104).

Statistical analysis

Optic and antennal lobe volumes both increased as

ROB volume increased (C. pallida, OL: y=0.98 x+0.13,

R2 = 0.71, F = 61.04, p < 0.0001, AL: y = 0.04 x − 0.003,

R2 =0.68, F=54.27, p<0.0001; A. dawsoni, OL: y=0.54

x + 0.27, R2 = 0.40, F = 11.30, p = 0.0037, AL: y = 0.07 x − 0.02, R2 = 0.57, F = 22.51, p = 0.0002). However, in

both C. pallida and A. dawsoni as the relative volume of the

optic increased, the relative volume of the antennal lobes
GraphPad Prism v. 8.3.0 (GraphPad Prism for Windows 2018) was used for all statistical analyses and data were confirmed to meet the assumptions of parametric tests before those analyses were performed using Anderson–Darling and Shapiro–Wilk normality tests and an F test was used to test for equal variance. Relative brain region volumes were calculated by dividing the brain region of interest by the total brain volume minus the brain region of interest (as in O’Donnell et al. 2018, termed Rest of Brain [ROB]). The differences in relative brain region volumes between morphs of each species were assessed using standard unpaired t tests, except when variances were unequal in which case a Welch’s correction to the unpaired t test was used. Linear regressions were used to assess the relationship between brain region volumes, and the relationship between relative optic and antennal lobe volumes.

Results

Body size variation

Small- and large-morph C. pallida males differed in mean head width and wet body mass (unpaired t tests; head width: n = 27, t = 7.63, df = 25, p < 0.0001; body mass: t = 10.89, df = 25, p < 0.0001). Large-morph males had a minimum head width of 5.35 mm and minimum body mass of 0.25 g (ranges, head width: 5.35–6.02 mm; body mass: 0.25–0.35 g), while small-morph males had a maximum head width of 5.27 mm and maximum body mass of 0.21 g (ranges, head width: 4.53–5.27 mm; body mass: 0.11–0.21 g).
Small- and large-morph A. dawsoni males differed in

mean head width (unpaired t test; n=19, t=13.21, d f=17,

p < 0.0001). Minimum head width of large-morph males was 6.51 mm (range 6.51–7.20 mm) and the maximum head width of small-morph males was 5.86 mm (range 5.28–5.86 mm).

Fig. 2 Smaller optic lobes (OL) and larger antennal lobes (AL) in

large-morph males of C. pallida and A. dawsoni. a C. pallida and

A. dawsoni large-morph males have relatively smaller optic lobes

(unpaired t tests; C. pallida: n=27, t=3.63, d f=25, p=0.0013; A. dawsoni: n=19, t=2.60, d f=17, p=0.0188) and b relatively larger

antennal lobes than the small-morph males of their species (unpaired

t tests; C. pallida: t=3.26, d f=25, p=0.0032; A. dawsoni: t=3.38,

d f=7.62, p=0.0104). Means with error bars representative of standard deviations. *p<0.05, **p<0.01

Relative optic and antennal lobe volumes

The small-morph C. pallida and A. dawsoni males, which

rely primarily on a visual mate location strategy, had relatively larger optic lobes compared with the large-morph

males (Fig. 2a; unpaired t tests; C. pallida: n=27, t=3.63,

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Journal of Comparative Physiology A

decreased (Fig. 3: C. pallida, y=−0.01 x+0.05, R2 =0.17, F=5.22, p=0.03; A. dawsoni, y=−0.03 x+0.08, R2 =0.30, F=7.24, p=0.016).

t test with Welch’s correction; MB lip: t = 1.63, df = 10.3,

p=0.13; MB collar: t=0.05, d f=24.37, p=0.96; unpaired t

test; MB basal ring and peduncle: t=0.10, d f=25, p=0.92; central complex: t = 1.74, df = 25, p = 0.09). In addition, there was no relationship between the relative volumes of the optic and antennal lobes and the corresponding MB region (C. pallida, OL-collar: R2 =0.05, F=1.44, p=0.24;

AL-lip: R2 =0.08, F=2.47, p=0.12; A. dawsoni, OL-collar:

R2 = 0.07, F = 1.20, p = 0.29; AL-lip: R2 = 0.08, F = 1.50,

Relative volumes of other brain regions

There were no significant differences between the C. pallida morphs in the relative volumes of the MB lip, collar, or basal ring and peduncles, or the central complex (Fig. 4a; unpaired

Fig. 3 Negative correlation in relative OL and AL volumes in C. pal- lida and A. dawsoni males. As relative OL volume increases, rela-

tive AL volume decreases in a C. pallida (y=−0.01 x+0.05; n=27,
R2 =0.17, F=5.22, p=0.031) and b A. dawsoni (y=−0.03 x+0.08; n=19, R2 =0.30, F=7.24, p=0.016) males

Fig. 4 Larger relative MB lip in A. dawsoni large-morph males; no differences in relative volumes of other MB regions or the cen-

tral complex between morphs in C. pallida or A. dawsoni. a There

are no significant differences in the relative volumes of the MB lips, collars, basal ring+peduncles, or the central complexes of C. pal- lida male morphs (unpaired t tests with Welch’s correction; MB lip:

n=27, t=1.63, d f=10.3, p=0.13; MB collar: t=0.05, d f=24.37,

p=0.96; unpaired t test; MB basal ring and peduncle: t=0.10,

d f=25, p=0.92; central complex: t=1.74, d f=25, p=0.09). b A.

dawsoni large-morph males have relatively larger MB lips than the small-morph males (unpaired t test; n=19, t=2.55, d f=17, p=0.0209). There are no differences in the relative volumes of the MB collars, basal ring+peduncles, or the central complex of A.

dawsoni male morphs (unpaired t test; MB collar: t=0.40, d f=17,

p=0.70; MB basal ring and peduncle: t=1.19, d f=17, p=0.25; unpaired t test with Welch’s correction; central complex: t=1.32, d f=9.57, p=0.22). Means with error bars representative of standard deviations. *p<0.05

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Journal of Comparative Physiology A

p = 0.24). There was also no relationship between the relative volumes of the two MB regions (C. pallida, collarlip: R2 = 0.007, F = 0.17, p = 0.69; A. dawsoni, collar-lip: R2 =0.11, F=2.15, p=0.16), demonstrating that the negative correlation between sensory systems occurs only in the peripheral processing lobes. manipulation of the visual environment of “Dark flies” (Drosophila melanogaster that had been reared in darkness since 1954, followed by 65 generations in the light) showed a negative correlation between OL and AL volumes evolved simultaneously in this new environment, as OL volume increased and AL volume decreased (Özer and Carle 2020). Closely related hawkmoth (Sphingidae) species that rely on different senses for foraging behavior (vision- vs. olfactionbased) show differentiation in both lower- and higher-order brain regions, with larger neuropil volumes to support the preferred foraging method (Stöckl et al. 2016).
Amegilla dawsoni large-morph males had a relatively larger MB lip region compared with the small-morph males (Fig. 4b; unpaired t test; t = 2.55, df = 17, p = 0.0209). No other regions of the brain differed in their relative volumes

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  • Male Mating Behaviour and Mating Systems of Bees: an Overview Robert John Paxton

    Male Mating Behaviour and Mating Systems of Bees: an Overview Robert John Paxton

    Male mating behaviour and mating systems of bees: an overview Robert John Paxton To cite this version: Robert John Paxton. Male mating behaviour and mating systems of bees: an overview. Apidologie, Springer Verlag, 2005, 36 (2), pp.145-156. hal-00892131 HAL Id: hal-00892131 https://hal.archives-ouvertes.fr/hal-00892131 Submitted on 1 Jan 2005 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie 36 (2005) 145–156 © INRA/DIB-AGIB/ EDP Sciences, 2005 145 DOI: 10.1051/apido:2005007 Review article Male mating behaviour and mating systems of bees: an overview1 Robert John PAXTON* School of Biology and Biochemistry, Queen’s University Belfast, 97 Lisburn Road, Belfast BT9 7BL, UK Received 9 November 2004 – Accepted 9 December 2004 Published online 1 June 2005 Abstract – Considerable interspecific diversity exists among bees in the rendezvous sites where males search for females and in the behaviours employed by males in their efforts to secure matings. I present an evolutionary framework in which to interpret this variation, and highlight the importance for the framework of (i) the distribution of receptive (typically immediate post-emergence) females, which ordinarily translates into the distribution of nests, and (ii) the density of competing males.
  • Rangelands, Western Australia

    Rangelands, Western Australia

    Biodiversity Summary for NRM Regions Species List What is the summary for and where does it come from? This list has been produced by the Department of Sustainability, Environment, Water, Population and Communities (SEWPC) for the Natural Resource Management Spatial Information System. The list was produced using the AustralianAustralian Natural Natural Heritage Heritage Assessment Assessment Tool Tool (ANHAT), which analyses data from a range of plant and animal surveys and collections from across Australia to automatically generate a report for each NRM region. Data sources (Appendix 2) include national and state herbaria, museums, state governments, CSIRO, Birds Australia and a range of surveys conducted by or for DEWHA. For each family of plant and animal covered by ANHAT (Appendix 1), this document gives the number of species in the country and how many of them are found in the region. It also identifies species listed as Vulnerable, Critically Endangered, Endangered or Conservation Dependent under the EPBC Act. A biodiversity summary for this region is also available. For more information please see: www.environment.gov.au/heritage/anhat/index.html Limitations • ANHAT currently contains information on the distribution of over 30,000 Australian taxa. This includes all mammals, birds, reptiles, frogs and fish, 137 families of vascular plants (over 15,000 species) and a range of invertebrate groups. Groups notnot yet yet covered covered in inANHAT ANHAT are notnot included included in in the the list. list. • The data used come from authoritative sources, but they are not perfect. All species names have been confirmed as valid species names, but it is not possible to confirm all species locations.
  • Biodiversity Summary: Rangelands, Western Australia

    Biodiversity Summary: Rangelands, Western Australia

    Biodiversity Summary for NRM Regions Guide to Users Background What is the summary for and where does it come from? This summary has been produced by the Department of Sustainability, Environment, Water, Population and Communities (SEWPC) for the Natural Resource Management Spatial Information System. It highlights important elements of the biodiversity of the region in two ways: • Listing species which may be significant for management because they are found only in the region, mainly in the region, or they have a conservation status such as endangered or vulnerable. • Comparing the region to other parts of Australia in terms of the composition and distribution of its species, to suggest components of its biodiversity which may be nationally significant. The summary was produced using the Australian Natural Natural Heritage Heritage Assessment Assessment Tool Tool (ANHAT), which analyses data from a range of plant and animal surveys and collections from across Australia to automatically generate a report for each NRM region. Data sources (Appendix 2) include national and state herbaria, museums, state governments, CSIRO, Birds Australia and a range of surveys conducted by or for DEWHA. Limitations • ANHAT currently contains information on the distribution of over 30,000 Australian taxa. This includes all mammals, birds, reptiles, frogs and fish, 137 families of vascular plants (over 15,000 species) and a range of invertebrate groups. The list of families covered in ANHAT is shown in Appendix 1. Groups notnot yet yet covered covered in inANHAT ANHAT are are not not included included in the in the summary. • The data used for this summary come from authoritative sources, but they are not perfect.
  • The Long and the Short of Mate Attraction in a Psylloid: Do Semiochemicals Mediate Mating in Aacanthocnema Dobsoni Froggatt? AQ1

    The Long and the Short of Mate Attraction in a Psylloid: Do Semiochemicals Mediate Mating in Aacanthocnema Dobsoni Froggatt? AQ1

    3/1/2016 e.Proofing The Long and the Short of Mate Attraction in a Psylloid: do Semiochemicals Mediate Mating in Aacanthocnema dobsoni Froggatt? AQ1 Umar K. Lubanga 1,* Email [email protected] AQ2 Falko P. Drijfhout 2 Kevin Farnier 1 Martin J. Steinbauer 1 1 Department of Ecology, Environment & Evolution, La Trobe University, Melbourne, Victoria, 3086 Australia 2 Chemical Ecology Group, Keele University, Keele, UK Abstract Mating is preceded by a series of interdependent events that can be broadly categorized into searching and courtship. Long­range signals convey species­ and sex­specific information during searching, while short­range signals provide information specific to individuals during courtship. Studies have shown that cuticular hydrocarbons (CHCs) can be used for mate recognition in addition to protecting insects from desiccation. In Psylloidea, four species rely on semiochemicals for long­ range mate attraction. Psyllid mating research has focused on long­range mate attraction and has largely ignored the potential use of cuticular hydrocarbons (CHCs) as mate recognition cues. This study investigated whether CHCs of Aacanthocnema dobsoni have semiochemical activity for long­ and short­range communication prior to mating. Using a solid sampler for solvent­less injection of whole psyllids into coupled gas Keele Research Repository http://eproofing.springer.com/journals/printpage.php?token=fTLCbmYf5q­dL­hwXUtKOtGR7lmEl13uLyeeDHpseRR1bx_QL_zXOQ­v1yprovidedftezDr by 1/26 View metadata, citation and similar papers at core.ac.uk CORE brought to you by 3/1/2016 e.Proofing chromatography/mass spectrometry, we found quantitative, sex­ and age­related differences in CHC profiles. Males had higher proportions of 2­MeC28, 11,15­diMeC29, and n­C33 alkanes, while females had higher proportions of 5­MeC27, 3­MeC27, 5,15­diMeC27, n­C29 and n­C30 alkanes.
  • Evolutionary Studies of Dawson's Burrowing

    Evolutionary Studies of Dawson's Burrowing

    MOLECULAR ECOLOGY OF DAWSON’S BURROWING BEE Amegilla dawsoni (HYMENOPTERA: ANTHOPHORINI) Maxine Beveridge (BSc Hons. PG Dip.) School of Animal Biology The University of Western Australia This thesis is presented for the degree of Master of Science Of The University of Western Australia 2006. SUMMARY In the last two decades, the use of microsatellites has revolutionized the study of ecology and evolution. Microsatellites, or short tandem repeats (STRs), are stretches of DNA repeats, 1 to 5 nucleotides long, where the number of repeats varies between individuals. They are co-dominant, highly variable, neutral markers, and are inherited in a Mendelian fashion. Microsatellite loci were isolated from Dawson's burrowing bee, Amegilla dawsoni, a large, fast-flying solitary nesting bee endemic to the arid zone of Western Australia. Twelve polymorphic loci were found with an observed number of alleles ranging from two to 24 and observed heterozygosities between 0.17 and 0.85. These loci were used to examine two aspects of this bee’s molecular ecology; its population structure and mating system. The population structure of Dawson’s burrowing bee was examined using the 8 most variable microsatellite loci. Adult female bees were collected from 13 populations across the species range. The mean number of alleles per locus ranged from 4 to 38 and expected heterozygosity was uniformly high with a mean of 0.602. Pairwise comparisons of FST among all 13 populations ranged from 0.0071 to 0.0122 with only one significant estimate and an overall FST of 0.001. The entire sample collection was in Hardy-Weinberg equilibrium and there was no evidence of inbreeding with a mean FIS of 0.010.
  • Burdens of Otters in England and Wales: with Case Studies of Populations in South West England

    Burdens of Otters in England and Wales: with Case Studies of Populations in South West England

    Analysis of the Population Genetics and Polybrominated Diphenyl Ether (PBDE) Burdens of Otters in England and Wales: With Case Studies of Populations in South West England Submitted by Angela Pountney, to The University of Exeter as a thesis for the degree of Doctor of Philosophy in Biological Sciences, September 2008. This thesis is available for library use on the understanding that it is copyright material and that no quotation from the thesis may be submitted and approved without proper acknowledgement. I certify that all material in this thesis which is not my own work has been identified and that no material has previously been submitted and approved for the award of a degree by this or any other University. ……………………. 0 Abstract Otter populations declined drastically across many areas of England and Wales during the 1960s to 1980s. The main cause of this decline is thought to have been high concentrations of organic pollutants, in particular PCBs and dieldrin. Here we look at the health of the present day otter population, focussing on the numbers of otters, the genetic diversity of populations and investigating a possible new organic pollutant threat, polybrominated diphenyl ethers (PBDEs). A non-invasive spraint genotyping study of the otter population inhabiting the River Camel in Cornwall not only revealed that the river was capable of supporting a minimum number of 12 otters over a 9 month period, but gave insight into the ranges and genetic relationships of the individuals using the river system. A further population genetic study was carried out focussing on the River Itchen in Hampshire, a population which declined drastically to just a few isolated individuals before receiving otters through a captive breeding programme.
  • An Extraordinary Natural Legacy

    An Extraordinary Natural Legacy

    An Extraordinary Natural Legacy An assessment and recommendations for the proposed expansion of Western Australia’s conservation reserve system March 2019 Centre for Conservation Geography Citation: Lucinda Douglass1, 2, Carol Booth1, Simon Kennedy1 and Joel Turner1 (2019) An extraordinary natural legacy: An assessment and recommendations for the proposed expansion of Western Australia’s conservation reserve system. Commissioned by The Pew Charitable Trusts. Centre for Conservation Geography. 1Centre for Conservation Geography 2University of Queensland Copyright: © Centre for Conservation Geography Designer: Iannello Design Printer: UniPrint The Centre for Conservation Geography (conservationgeography.org) is a research group established in 2011 to provide expert technical support and advice to government and non- government decision-makers and stakeholders. The centre’s primary focus is providing world’s-best-practice decision support to planning for nature conservation. Based in Australia, we are a multidisciplinary team with expertise in marine and terrestrial protected area planning. Our skills include scientific research, evidence-based policy development, biogeography including GIS analyses and mapping, and science communications and advocacy. We can support conservation decision-making across the world’s ecoregions, and currently have projects in Australia, Canada and Antarctica. Contact: [email protected] Acknowledgement of country: The authors acknowledge the Traditional Owners of the lands that are the focus of this report and their continuing connection to these lands. We pay respect to them and their cultures, and to their elders past and present. We acknowledge the inextricable link between natural values and Aboriginal heritage values and that the knowledge of Traditional Owners will be vital for maintaining both. Other acknowledgements: The Centre for Conservation Geography and the report’s authors thank the many people who helped make this report possible.