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appendices
Appendices
Appendix 1
Primer of networks – a guide to the usage of, and terminology
describing networks in this thesis
A network (or interaction network/web) is a graphical representation of a collection of species and the interactions among them. Species can be referred to as nodes and interactions as links. Links refer the presence of an interaction and does not take into account the frequency of interaction.
There are several possible variations and different ways to portray a network of interactions:
• Interaction matrix: an adjacency matrix where plant species represent columns and animal species represent rows. The cell where a plant and animal species intersect can be marked by a ‘1’ to indicate the presence of and interaction and a ‘0’ to indicate no interaction. Alternatively, the cell can represent the number of interactions or a measure of the strength of the interaction. • Bipartite network (two-mode network): a network where only interactions occurring between plants and animals are considered. I.e. if animal b pollinates plant x, then b and x will be connected in a bipartite network. • Unipartite network (one-mode network): a network where only interactions occurring among plants or among animals are considered. A bipartite network can be collapsed into a unipartite network that portrays links among species that share a common interaction partner. I.e. if plant species x and y share pollinator species b, then x and y will be connected in a uni-partite network. • Quantitative/weighted network: a network where links are given a weight. In flower-visitor networks it refers the frequency of interaction. In pollen-transport networks it refers to the amount of pollen transported.
226 Appendices
• Qualitative/presence-absence/binary/unweighted network: a network where only the presence/absence of a link is shown.
Three different types of interactions are considered as networks in this thesis:
• Pollination network: links represent pollination interactions among species • Flower-visitor network: links represent flower-visitor interactions among species • Pollen-transport network: links represent the transport of a pollen species by an animal species.
On the ways to describe the patterns of species interactions in a network:
• Network function: as defined by the interaction type. The function of a pollination network is the movement of gametes effecting pollination and the acquisition of resources by visitors. • Network structure/topology: refers to the patterns (or geometric layout) in a network due to the way species interact with each other. • Network structural property: a specific structural pattern. E.g. nestedness • Network Indices: The variable to describe, characterize or measure some feature of network structural property. Synonymous with network metric and network statistic. See Dormann et al. (2009) for a more comprehensive list. • Forbidden link: two species that cannot interact due to trait incompatibility or spatio-temporal mismatch.
Network indices:
• P: plant species richness • A: animal visitor species richness • Network size (S): combined total of plant and animal species • I: total number of interactions • Iap: number of interactions between animal species a and plant species p. Also referred to as interaction strength. • Species degree: the number of links of a species
227 Appendices
Indices based on unweighted links (qualitative networks)
• Links per species: total number of links in a network divided by network size (L = I/S) • Connectance (C): proportion of realized links (C = AP) (Jordano 1987). • Nestedness (N): Tendency of specialised species to interact with a subset of the interaction partners of more generalised species (Bascompte et al. 2003). Nestedness implies both a high occurrence of degree asymmetry and the existence of a ‘generalised network core’ (relatively small group of highly connected species). A nestedness value of 1 equals perfect nestedness, 0 equals the opposite of nestedness of which there are several distinct patterns (Almeida-Neto et al. 2007), and randomness falls somewhere in between. • Degree asymmetry: the imbalance in the degree between two interacting species (Vazquez and Aizen 2004). • Degree distribution: frequency histogram of the degree of species in a network (Jordano et al. 2003). • Truncated power-law: a degree distribution of pollination networks are usually truncated power law which indicates that most species have few links and few species have many links, more than would be expected by chance. • Modularity/compartmentalisation: The existence groups of species with many intragroup links and few intergroup links (Guimera and Amaral 2005, Olesen et al. 2007). Modularity ranges from 0 (no modularity) and 1 (high modularity).
Indices based on weighted links (quantitative networks)
• Dependence: an estimate of the extent to which a species depends on another species (Jordano 1987, Bascompte et al. 2006). • Interaction strength asymmetry/dependence asymmetry: quantifies the imbalance in the interaction strength/dependence of two interacting species (Bascompte et al. 2006). Positive values indicate a higher specialisation of animals negative values indicate a higher specialisation of plants.
228 Appendices
• H2’ specialisation: a network-level measure of specialisation based on the deviation of a species’ realised number of interactions and that expected from each species’ total number of interactions
(Bluthgen et al. 2006). The H2’ ranges between 0 (no specialisation) and 1 (perfect specialisation). • d’ specialization: a species-level measure of specialisation which expresses how specialised a given species is in relation to what species are on offer (Bluthgen et al. 2006). Species-level equivalent
of H2’ specialisation. The d’ ranges between 0 (no specialisation) and 1 (perfect specialisation). • Generality: mean number of plant species visited per animal species (Brauman and Daily 2008, Dormann et al. 2009). See Dormann et al. (Dormann et al. 2009) for equation. Increasing value indicates a visitor species is visiting an increasingly diverse set of plants. • Vulnerability: mean number of animal species per plant species. Analogous to generality. • Evenness: a measure of the skewness in the distribution of interaction frequencies (Tylianakis et al. 2007). Also know as interaction evenness.
Processes driving network structure:
• Interaction neutrality: Occurrence of species interactions resulting from the random encounter of individuals, so that all individuals have the same probability of interacting regardless of identity (Vázquez et al. 2007, Krishna et al. 2008). • Trait-mediated: the likelihood of an interaction between two species depends on their phenotypic match. There two types of trait- mediated processes: trait complementarity, where the traits of two species correspond (e.g. a visitors nectar requirement matches that offered by a flower); and, exploitation barrier, where traits preclude interactions from happening (e.g. a long corolla length exclude short tongued visitors). Exploitation barrier can be a form of forbidden link.
Species role: the characterisation of a species depending on the pattern of their intra-module and inter-module links (Guimera and Amaral 2005). First defined by Guimera and Amaral (2005) using species ‘within module degree’ (z)
229 Appendices and ‘participation coefficient’ (P). Species with a similar pattern of links in the network are allocated the same role. Species role communicates the importance of a species in a network: how many other species rely on them, and what sort of effects to the network can be expected if they are lost. Four roles are defined, in this thesis:
• Hub: species with many links especially within their own module, but can have many outside of their module as well (z ≥ 2.5, 0 ≤ P ≤ 1). • Connector: network ‘glues’, linking many different modules together (z ≤2.5, P ≥ 0.62). • Peripheral: species with several links mainly within their own module, but can also link to another module (z ≤ 2.5, 0.05 < P < 0.62). • Ultra-peripheral: species often with only one or two links but only within their own module (z ≤0 2.5, P ≤0.05).
230 Appendices
231 Appendices
Appendix 2
A2-1. Abundance of invertebrates caught visiting flowers of each plant species using net sampling at each site and during each sampling period. Plant names follow the Australian Plant Name Index.
Jun-10 Nov-10 Jul-11 Total Total species abundanc Family Plant species FR KS MC FR KS MC FR KS MC richness e Aizoaceae Trianthema pilosa 26 59 3 1 63 152 35
Amaranthaceae Alternanthera sp. A 2 2 1
Amaranthaceae Ptilotus atriplicifolius 3 12 15 5
Amaranthaceae Ptilotus latifolius 1 15 2 34 15 2 69 27
Amaranthaceae Ptilotus polystachyus 7 11 18 12 48 11
Araliaceae Trachymene glaucifolia 103 91 120 13 19 47 393 70
Asteraceae Asteraceae sp. A 1 1 1
Asteraceae Calotis erinacea 9 1 129 1 37 6 183 35
Asteraceae Calotis plumothera 2 2 2
Asteraceae Helichrysum sp. A 2 1 3 3
Asteraceae Helipterum molle 6 25 31 6
Asteraceae Myriocephalus sp. A 1 1 1
Asteraceae Pterocaulon sphacelatum 31 52 83 21
Boraginaceae Halgania cyanea 7 7 6
Boraginaceae Trichodesma zeylanicum 19 1 11 7 27 55 21 18 159 28
Brunoniaceae Brunonia australis 6 13 19 12
Chenopodiaceae Chenopodiaceae sp A 6 6 1
Chenopodiaceae Salsola kali 1 1 1
Euphorbiaceae Adriana tormentosa 8 8 2
Euphorbiaceae Euphorbia drummondii 74 57 132 70 88 220 130 35 69 875 132
Fabaceae Acacia bivenosa 19 22 24 65 17
Fabaceae Acacia coriacea 3 3 3
Fabaceae Acacia dictyophleba 50 39 19 34 32 174 25
Fabaceae Acacia ligulata 60 2 138 27 93 320 43
Fabaceae Acacia stenophylla 1 1 1
Fabaceae Crotalaria cunninghamii 2 4 34 38 84 162 23
Fabaceae Crotalaria eremaea 1 10 8 10 84 22 28 30 193 61
232 Appendices Fabaceae Crotalaria smithiana 4 12 16 1
Fabaceae Fabaceae sp. A 3 3 2
Fabaceae Psoralea eriantha 2 2 2
Fabaceae Senna artemisioides 2 2 1
Fabaceae Senna pleurocarpa 9 2 11 8
Fabaceae Swainsona phacoides 1 2 3 2
Fabaceae Tephrosia rosea 5 4 65 44 42 1 9 170 30
Frankeniaceae Frankenia gracilis 7 7 4
Goodeniaceae Goodenia cycloptera 24 14 12 28 29 26 17 14 7 171 30
Goodeniaceae Lechenaultia divaricata 10 10 4
Goodeniceae Scaevola depauperata 3 1 21 82 8 47 1 163 29
Goodeniceae Scaevola parvibarbata 4 25 3 32 10
Goodeniceae Scaevola parviflora 19 5 1 25 11
Haloragaceae Haloragis gossei 4 4 1
Lamiaceae Dicrastylis costelloi 92 6 15 113 41
Lamiaceae Dicrastylis llewellynii 11 11 8
Lamiaceae Newcastelia cephalantha 16 122 138 26
Lamiaceae Newcastelia spodiotricha 146 172 318 36
Malvaceae Abutilon otocarpum 43 43 1
Malvaceae Rulingia loxophylla 1 1 1
Malvaceae Sida fibulifera 2 16 18 29 65 26
Malvaceae Sida trichopoda 5 2 7 6
Malvaceae Triumfetta winneckeana 3 68 71 10
Myrtaceae Eucalyptus pachyphylla 316 316 18
Portulacaceae Calandrinia balonensis 11 26 14 81 132 23
Proteaceae Grevillea juncifolia 8 1 6 15 6
Proteaceae Grevillea stenobotrya 221 61 3 285 37
Rubiaceae Oldenlandia pterospora 8 8 6
Sapindaceae Atalaya hemiglauca 1 1 1
Scrophulariaceae Eremophila macdonnellii 12 68 1 81 13
Scrophulariaceae Eremophila sp. A 2 2 2
Solanaceae Solanum sp A 10 3 13 4
Violaceae Hybanthus aurantiacus 2 2 2
Unknown 1 3 3 2
Total 407 248 411 636 1085 1067 451 479 436 5220 327 The five shades of cell colour indicate flowering intensity (darker shade equates to greater flowering intensity) of that species at that specific time and place.
233 A2-2. Rarefaction curves for pan trap (broken line) and net sampling (solid line). Transects within each dune zone at each location at each site were pooled and used to generate average estimates (n = 12, ± SE).
Appendices
A2-3. List of bee species and abundance sampled by pan traps and nets.
Family Species Net sampling Pan traps Apidae Amegilla chlorocyanea 19 55 Apidae Exoneurella eremophila 58 53 Apidae Thyreus warooensis 3 Colletidae Brachyhesma barrowensis 44 Colletidae Chrysocolletes strangomeles 2 Colletidae Euhesma loorea 1 Colletidae Euhesma sp. D 31 14 Colletidae Euhesma sp. J 9 Colletidae Euhesma sp. T 2 3 Colletidae Euhesma sybilae 18 1 Colletidae Hylaeus albimandibulatus 2 Colletidae Hylaeus albozebratus 65 13 Colletidae Hylaeus calandriniae 12 33 Colletidae Hylaeus circinatus 3 Colletidae Hylaeus sp. AB 1 Colletidae Hylaeus sp. AG 1 Colletidae Hylaeus sp. AH 3 Colletidae Hylaeus sp. E 3 1 Colletidae Hylaeus sp. F 75 1 Colletidae Hylaeus sp. H 3 11 Colletidae Hylaeus sp. K 9 Colletidae Hylaeus sp. L 1 Colletidae Hylaeus sp. Z 12 3 Colletidae Hylaeus trimerops 13 Colletidae Hylaeus vittatifrons 4 Colletidae Hylaeus wyndhamensis 26 Colletidae Hylaeus zebrinus 4 48 Colletidae Leioproctus alloeopus 7 Colletidae Leioproctus argentifrons 2 1 Colletidae Leioproctus finkei 13 39 Colletidae Leioproctus impatellatus 8 2 Colletidae Leioproctus lucidicinctus 63 8 Colletidae Leioproctus sequax 98 174 Colletidae Leioproctus sp. A 25 61 Colletidae Leioproctus sp. AD 2 Colletidae Leioproctus sp. AE 29 3 Colletidae Leioproctus sp. B 4 Colletidae Leioproctus sp. S 17 Colletidae Leioproctus sp. V 1 Colletidae Leioproctus sp. W 4 4 Colletidae Leioproctus tarsalis 3 Colletidae Xanthesma flava 16 Colletidae Xanthesma lucida 42
235 Appendices
Colletidae Xanthesma nigrior 3 Halictidae Homalictus sphecodopsis 33 Halictidae Homalictus urbanus 340 346 Halictidae Lasioglossum alacarinatum 105 97 Halictidae Lasioglossum albopilosum 2 24 Halictidae Lasioglossum ebeneum 5 Halictidae Lasioglossum immaculatum 217 41 Halictidae Lasioglossum platychilum 120 82 Halictidae Lasioglossum cognatum 27 Halictidae Lasioglossum eremaean 18 40 Halictidae Lipotriches australica 41 3 Halictidae Lipotriches flavoviridis 73 15 Halictidae Lipotriches semipallida A 4 Halictidae Lipotriches sp 89 3 Halictidae Lipotriches sp AC 3 Halictidae Lipotriches sp G 1 Megachilidae Coelioxys reginae 5 1 Megachilidae Megachile apicata 2 Megachilidae Megachile aurifrons 1 Megachilidae Megachile barvonensis 3 Megachilidae Megachile bidentis 2 Megachilidae Megachile boharti 2 Megachilidae Megachile captionis 2 3 Megachilidae Megachile giddioorla 11 Megachilidae Megachile maculariformis 3 Megachilidae Megachile sequior 17 1 Megachilidae Megachile serricauda 6 1 Megachilidae Megachile tarltoni 1 Megachilidae Megachile warrambuccis 1
236 Appendices
A2-4. Partially nested ANOVA results comparing species richness and total abundance between methods (log transformed).
Species richness Total abundance Source df MS F P MS F P Method 1 817.0 27.32 <0.05 1.634 38.00 <0.01 Trip 2 1662.9 53.64 <0.001 2.375 91.35 <0.001 Site 2 117.3 4.60 ns 0.314 9.81 <0.05 Dune 1 1094.5 243.22 <0.01 0.381 11.55 <0.05 Location(site) 3 25.5 1.42 ns 0.032 0.74 ns Method x Trip 2 366.8 33.35 <0.01 1.058 27.84 <0.01 Method x Site 2 44.6 1.49 ns 0.175 4.07 ns Method x Dune 1 5.1 0.11 ns 0.165 0.75 ns Method x Location(site) 3 29.9 1.67 ns 0.043 1.00 ns Trip x Site 4 179.8 5.80 <0.05 0.079 3.04 ns Trip x Dune 2 154. 8.24 <0.05 0.121 6.72 <0.05 Trip x Location(site) 6 31.0 1.73 ns 0.026 0.60 <0.05 Site x Dune 2 66.1 14.69 <0.05 0.124 3.76 ns Dune x Location(site) 3 4.5 0.25 ns 0.033 0.77 ns Method x Trip x Site 4 22.4 2.04 ns 0.215 5.66 <0.05 Method x Trip x Dune 2 42.3 1.51 ns 0.081 0.90 ns Method x Trip x Location(Site) 6 11.0 0.61 ns 0.038 0.88 ns Method x Site x Dune 2 42.3 0.95 ns 0.022 0.10 ns Method x Dune x Location(site) 3 44.5 2.49 ns 0.220 5.12 <0.01 Trip x Site x Dune 4 30.9 1.65 ns 0.033 1.83 ns Trip x Dune x Location(Site) 6 18.7 1.04 ns 0.018 0.42 ns Method x Trip x Site x Dune 4 3.2 0.11 ns 0.028 0.31 ns Method x Trip x Dune x Location(Site) 6 28.1 1.57 ns 0.090 2.09 ns Error 72 17.9 0.043
237 Appendices
A2-5. Species richness and abundance per transect (n = 24) of invertebrate orders (average ± SE). Hymenoptera split in bees, ants and wasps. Bees are grouped by Method x Trip x Site so mean values are from n = 8 transects.
Coleoptera
ptera Hemi
238 Appendices
A2-6. Pairwise comparisons for the significant ANOSIM method (a) x trip (b) x site (c) interaction comparing invertebrate assemblages collected using net sampling and pan traps.
Pairwise Test t P t P t P Method x Trip x Site a) Net, Pan FR KS MC Jun 10 3.344 <0.01 3.140 <0.01 2.483 <0.05 Nov 10 3.366 <0.01 2.913 <0.01 2.142 <0.05 Jul 11 2.979 <0.05 2.800 <0.01 2.160 <0.05 b) Net Jun 10, Nov 10 2.392 <0.05 2.777 <0.01 2.365 <0.05 Jun 10, Jul 11 2.594 <0.05 1.476 ns 1.997 ns Nov 10, Jul 11 2.758 <0.05 2.777 <0.05 2.055 <0.05 Pan Jun 10, Nov 10 2.192 <0.05 2.738 <0.01 3.961 <0.01 Jun 10, Jul 11 2.627 <0.05 2.597 <0.05 2.862 <0.05 Nov 10, Jul 11 2.649 <0.05 1.978 <0.05 2.382 <0.05 c) Net Jun 10 Nov 10 Jul 11 FR, KS 1.969 <0.05 2.403 <0.01 1.593 ns FR, MC 2.023 <0.05 1.800 <0.05 1.310 ns KS, MC 2.071 <0.05 1.779 <0.05 1.165 ns Pan FR, KS 3.525 <0.001 1.746 <0.05 1.784 <0.05 FR, MC 3.117 <0.01 2.628 <0.01 1.577 ns KS, MC 3.212 <0.01 1.961 <0.05 1.875 <0.05
239 Appendices
A2-7. Nonmetric multidimensional scaling (nMDS) ordination of assemblages of common invertebrate species. Solid symbols represent nets, hollow symbols represent pans, and shapes represent the sampling periods.
240 Appendices
A2-8. Comparison of assemblages of common invertebrate species sampled using PERMANOVA. Common invertebrates were arbitrarily defined as any species with a total abundance of 20 or greater of which there were 61 invertebrate species.
Source df MS Pseudo-F P(perm) Method 1 40968 24.90 <0.001 Trip 2 29644 12.16 <0.001 Site 2 14785 5.06 <0.0011 Dune 1 5127 2.29 ns Location(Site) 3 2919 1.77 <0.01 Method x Trip 2 23330 14.18 <0.001 Method x Site 2 11886 7.23 <0.001 Method x Dune 1 5444 2.46 <0.05 Trip x Dune 2 3314 1.60 ns Trip x Site 4 9534 3.91 <0.001 Site x Dune 2 3540 1.58 ns Trip x Location(Site) 6 2439 1.48 <0.01 Dune x Location(Site) 3 2238 1.36 ns Method x Site x Dune 2 4228 1.91 ns Method x Trip x Site 4 7351 4.47 <0.001 Method x Trip x Dune 2 2851 1.47 ns Method x Dune x Loc(site) 3 2213 1.35 ns Trip x Site x Dune 4 2508 1.21 ns Method x Dune x Loc(site) 6 2067 1.26 ns Method x Trip x Site x Dune 4 2948 1.52 ns Method x Trip x Dune x Lo(site) 6 1933 1.18 ns Residual 81 1645 1 = P-value determined using Monte Carlo sampling.
241 Appendices
Appendix 3
A3-1. Characteristics of the visitation, pollen-transport, and null pollen-transport networks when singletons were removed from the analysis.
Pollen-transport network Visitation Null model network Observed Mean (95% CI) Plant species 51 27 27.22 (22.00–32.00) Bee species 28 13 17.01 (13.00–21.00) Links 146 55 72.02 (64.00–80.00) Visits 654 95 95
Connectance 0.102 0.157 0.159 (0.123–0.210) Nestedness 0.885 0.807 0.857 (0.775–0.912)
H2' 0.501 0.783 0.469 (0.400–0.541) Vulnerability 3.323 1.577 4.111 (3.373–4.956) Evenness 0.864 0.736 0.975 (0.961–0.985)
Mean links per bee species 5.21 ± 2.28 4.23 ± 2.44 4.29 (3.45-5.43) Mean links per plant species 2.81 ± 0.55 2.43 ± 0.92 2.66 (2.24-3.17) Mean bee specialisation index d' 0.55 ± 0.06 0.67 ± 0.13 0.54 (0.46-0.62) Mean plant specialisation index d' 0.36 ± 0.07 0.44 ± 0.08 0.42 (0.35-0.49)
242 Appendices
A3-2. Examples of bees visiting various flowers. Clockwise from top left: Hylaeus zebrinus (Colletidae) visiting Calotis erinaceae (Asteraceae), bee = 5 mm; Hylaeus trimerops visiting Euphorbia dummondii (Euphorbiaceae), bee = 4 mm; Lasioglossum immaculatum (Halictidae) visiting Grevillea stenobotrya (Proteaceae), bee = 4 mm; Lipotriches flavoviridis (Halictidae) visiting Calandrinia balonensis (Portulacaceae) and Scaevola depauperata (Goodeniaceae), bee = 5 mm; Lasioglossum cognatum (Halictidae) visiting Wahlenburgia gracilis
(Campanulaceae), bee = 5 mm. Photos by Tony Popic.
243 Appendices
Appendix 4
A4-1: PERMANOVA results of main test assessing species composition differences among networks. Networks were sampled at the Location(Site) level during three periods (June 2010, November 2010, June 2011), and at three sites (FR, KS, MC – two Locations were nested within each site). Within each location there were also two dune zones (crest and swale).
Source df MS Pseudo-F P(perm) Time 2 25443 13.18 <0.01 Site 2 9567 3.96 <0.0011 Dune 1 29417 12.01 <0.05 Location(Site) 3 2416 1.33 <0.05 Time x Site 4 6002 3.11 <0.01 Time x Dune 2 8024 3.64 <0.01 Site x Dune 2 6131 2.50 ns Time x Location(Site) 6 1931 1.07 ns Location(Site) x Dune 3 2450 1.35 <0.05 Time x Site x Dune 4 3950 1.79 <0.05 Time x Location(Site) x Dune 6 2202 1.21 <0.05 Residual 36 1813 1 = P-value determined using Monte Carlo sampling due to insufficient permutations
A4-2: Pairwise tests of the significant Time x Site interaction.
Pairwise Test t P Time Jun 10, Nov 10 3.55 <0.05 Jun 10, Jun 11 3.38 <0.05 Nov 10, Jun 11 3.93 <0.05 Site FR, KS 2.23 <0.011 FR, MC 1.84 <0.011 KS, MC 1.90 <0.011 1 = P-value determined using Monte Carlo sampling due to insufficient permutations
244 Appendices
Appendix 5
A5-1: The realised roles of plant species and visitor species in all 18 flower-visitor networks grouped by roles. Adapted from Guimera and Amaral (2005).
hub Plants A 5 connector 4 peripheral ultraperipheral 3
2
1
0 z -within moduledegree -1
-2 0 B0.2 0.4 0.6 0.8 1 5 Visitors 4
3
2
1
0 z -within moduledegree -1
-2 0 0.2 0.4 0.6 0.8 1 P -par cipa on coefficient
245 Appendices
A5-2. The participation coefficient and within module degree were combined to produce one variable that described the importance of species in a network.
Toward the red colour (top right, values of up to 2) are hub and connector species, and toward the blue colour (bottom left, values of down to 0) are ultra-peripheral species.
C
246 A5-3. Rainfall, observed network dimensions, qualitative indices and quantitative indices for the 18 networks from spinifex grassland. Rainfall is the cumulative rainfall from the nine months before the network was sampled. See A1 for index details.
Observed network dimensions Qualitative Indices Quantitative Indices Interaction Visitor Plant strength Rainfall species species Network Links per Conn. Nested. Number Modularity asymmetry H2’ Time Site (mm) richness richness size Links Interact. species (0 – 1) (0 – 1) modules (0 – 1) (-1 – 1) (0 – 1) Generality Vulnerability Jun-10 FR1 447.2 33 12 45 64 176 1.42 0.16 16.0 6 0.48 0.18 0.48 2.71 6.09 Jun-10 FR2 447.2 51 15 66 98 231 1.48 0.13 12.2 6 0.51 0.07 0.44 3.26 6.88 Jun-10 KS1 360.4 38 9 47 55 130 1.17 0.16 15.2 6 0.53 0.40 0.67 1.74 10.59 Jun-10 KS2 360.4 35 12 47 54 118 1.15 0.13 11.4 8 0.55 0.10 0.65 1.90 6.52 Jun-10 MC1 311.4 34 13 47 58 217 1.23 0.13 9.2 8 0.50 0.16 0.43 2.66 6.67 Jun-10 MC2 311.4 38 13 51 60 194 1.18 0.12 9.3 6 0.54 0.13 0.45 2.28 6.92 Nov-10 FR1 654.2 67 20 87 110 251 1.26 0.08 9.5 9 0.61 0.07 0.65 2.00 6.63 Nov-10 FR2 654.2 70 19 89 119 385 1.34 0.09 9.6 10 0.56 0.21 0.56 2.55 7.34 Nov-10 KS1 589.2 63 20 83 116 599 1.40 0.09 15.5 10 0.56 0.23 0.60 2.79 6.00 Nov-10 KS2 589.2 67 26 93 119 486 1.28 0.07 8.6 12 0.61 0.14 0.65 2.37 6.23 Nov-10 MC1 526.2 94 24 118 176 634 1.49 0.08 8.8 9 0.55 0.20 0.54 3.13 8.86 Nov-10 MC2 526.2 73 15 88 121 433 1.38 0.11 12.3 7 0.60 0.27 0.67 2.31 8.30 Jun-11 FR1 519.2 39 11 50 60 261 1.20 0.14 12.2 6 0.55 0.28 0.40 2.91 6.05 Jun-11 FR2 519.2 47 10 57 69 190 1.21 0.15 16.6 6 0.56 0.35 0.62 2.21 7.71 Jun-11 KS1 577.8 46 13 59 70 199 1.19 0.12 19.3 8 0.60 0.09 0.56 2.96 4.77 Jun-11 KS2 577.8 62 14 76 88 280 1.16 0.10 11.5 10 0.63 0.30 0.57 2.49 7.90 Jun-11 MC1 393.0 34 12 46 52 256 1.13 0.13 11.4 6 0.58 0.11 0.61 2.58 4.47 Jun-11 MC2 393.0 38 14 52 58 180 1.12 0.11 13.0 8 0.61 0.13 0.58 3.14 4.26 A5-4: The observed number of plant and visitor species in each role across all 18 networks. The expected values are based on the null that the number of plant and visitor species in each role would be the same proportion as the total number of observed plants and visitors. Observed and expected numbers were compared using a chi-square test.
Observed plant Observed visitor chi- Network Role species (expected) species (expected) square d.f. Significance Hub 62 (14) 0 (48) 211.76 1 P < 0.001 Connector 14 (22) 81 (73) 3.39 1 P > 0.05 Peripheral 67 (51) 156 (172) 6.97 1 P < 0.01 Ultra-peripheral 129 (185) 692 (635) 22.54 1 P < 0.001 Total 272 929
Appendices
A5-5: The average number of bee species that visited zygomorphic plant species (black
2 symbols) increased with rain (F(1,17) = 32.53, P < 0.001, r = 0.65) but not with
2 actinomorphic species (F(1,17) = 0.73, P > 0.05, r = 0.00; red symbols; a). The average role of zygomorphic plant species increased with network bee species richness (F(1,17) =
2 38.77, P < 0.001, r = 0.69), but not for actinomorphic plant species (F(1,17) = 0.67, P > 0.05, r 2 = 0.00; a). Each symbol represents a different network.
A 6 a 5
4 Zygomorphic 2 3 r = 0.65 species 2
1 Average # bee species visi ng plant 0 200 300 400 500 600 700 9 month cumula ve rainfall
1.4 B b 1.2
1
0.8 Zygomorphic r2 = 0.57 0.6 Average role 0.4
0.2
0 0 5 10 15 20 25 30 Bee species richness
249 Appendices
A5-6: Relationship between flower size and the variation in visitor size. Lower flower sizes may restrict the size of visitors that can visit, but no difference in the variation in visitor size was observed as flower size changed.
6 Jun-10 Nov-10 5 Jun-11
4
3
2
1 Standard devia on of visitor size 0 0 0.5 1 1.5 2 2.5 Log(flower size)
A5-7: The distribution of visitor sizes varied among sampling periods. There was a peak in visitors between 3-5 mm for all three sampling periods. June and November 2010 also had a peak in visitors around 8-9 mm but June 2011 did not.
0.50
0.45 Jun-10 0.40 Nov-10 0.35 Jun-11 0.30 0.25 0.20
Propor on of total 0.15 0.10 0.05 0.00 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
Visitor body size (mm)
250 Appendices
A5-8: Average role of plant species according to their nectar available pooled across networks. There was no significant difference in average role of plants with different nectar volumes (ANOVA, F(2,233) = 0.254, P = 0.78).
0.9 0.8 0.7 0.6 0.5 0.4 n=41 n=133 n=62
Average Role 0.3 0.2 0.1 0 Absent Glisten More than glisten
Available nectar
251 Appendix
Appendix
Appendices
Appendix 7
This appendix contains a manuscript currently being reviewed by the journal Records of the Australian Museum. It describes five new species of
Leioproctus (Protomorpha) Rayment (Hymenoptera: Colletidae). My contribution to this study was collecting bees in the Simpson Desert.
254 Appendices
Five New Species of Leioproctus (Protomorpha) Rayment
(Hymenoptera: Colletidae)
MICHAEL BATLEY1* AND TONY J. POPIC2
1Australian Museum, 6 College Street, Sydney NSW 2010, Australia
2Desert Ecology Research Group, School of Biological Sciences, The
University of Sydney, Sydney NSW 2006, Australia
Abstract
Five new species of Leioproctus (Protomorpha) Rayment from western
Queensland are described: L. crispus, L. gibber, L. gurneyi, L. latifrons and L. nix. A revised key to species is provided.
Introduction
In the most recent revision of the subgenus Leioproctus (Protomorpha)
(Maynard 1991) nine species were recognised, though only five were named. The remaining four were known from females only. In a recent study of plant- pollinator networks in the arid-zone grassland of the north-eastern Simpson
Desert (Popic et al. 2013) seven L. (Protomorpha) species were found, five of them undescribed. Female of four species had specialised hairs on the face or vertex of the head, a feature not previously seen in this subgenus and one that has been
255 Appendices suggested as indicative of particular pollen collection methods (Müller 1996,
Thorp 2000, Gonzalez & Chavez 2004, Alqarni et al. 2012, Rightmyer et al. 2011).
Terminology, methods and measurements
The morphological terminology follows that used by Michener (Michener and Fraser 1978, Michener 2007) including use of the word hair and description of legs in their normal positions. Relative dimensions quoted in the descriptions were measured using an eye-piece graticule on a stereomicroscope with the zoom objective set to give a reading of 50 divisions for the head width. Abbreviations used for the measurements are as in Houston (1990) and are as follows: AOD antennocular distance; ASD antennal socket diameter; FL flagellum length; HL head length; HVO height of vertex above lateral ocelli; HW head width; IAD interantennal distance; LID; lower interorbital distance; OOD ocellocular distance; SL scape length; SW scape width; UFW upper width of face; UID upper interorbital distance;
WOC width of ocellar cluster. Measurement of the length of the hind tarsus excludes the claws. Metasomal terga are referred to as T1, T2 etc. and sterna as S1,
S2 etc. S7 and S8, the “hidden sterna” of males, exhibit useful diagnostic characteristics and were extracted for examination. Geospatial coordinates are
GPS readings. The specimens examined are lodged in the Australian Museum (AM) collection.
Pollen from bees and flowers was stained with Calberla's fluid and examined with a compound microscope.
Descriptions are arranged alphabetically.
Key to species of Leioproctus (Protomorpha) (modified from Maynard,
1991)
256 Appendices
1. Male (hind tibia without scopa) ...... 2 Female (hind tibia with scopa) ...... 10
2. Hind coxa covered in long, dense hair ...... 3 Hind coxa lacking long, dense hair ...... 4
3. Ventral margin of hind basitarsus strongly curved; frons obscured by dense, white hair ...... tarsalis Ventral margin of hind basitarsus almost straight; frons not obscured by
hair ...... plautus
4. Hind tibia with row of stout setae on anterior margin ...... 5 Hind tibia unmodified ...... minutus
5. Hind basitarsus with narrow basal projection ...... alloeopus Hind basitarsus without basal projection ...... 6
6. Anterior margin of hind basitarsus with large apical projection ...... crispus Hind basitarsus with at most a slight apical extension ...... 7
7. Hind tibial spurs flattened and strongly recurved ...... 8 Hind tibial spurs not flattened, almost straight ...... 9
8. Hind femur with broad projection on outer face; outer hind tibial spur ca 0.3× as long as inner spur ...... nix Hind femur without projection; outer hind tibial spur ca 0.7× as long as
inner spur ...... latifrons
9. Hind tibial setae stout; mandible simple ...... gibber Hind tibial setae weak; mandible with preapical tooth ...... gurneyi
10. Inner hind tibial spur strongly curved ...... 11 Inner hind tibial spur almost straight ...... 12
11. Apical margins of T1-4 transparent ...... alloeopus Apical margins of T1-4 opaque ...... sp.A*
12. Frons entirely covered by dense, white, branched hair ...... 13
257 Appendices
Frons medially without dense, branched hair ...... 14
13. Facial fovea obscured by hair, inner margins ill-defined ...... tarsalis Facial fovea not obscured by hair, inner margins sharply defined ...... sp.B*
14. Frons and clypeus with long, weakly-branched, curly hair (Fig 9) ...... crispus Frons and clypeus without curly hair ...... 15
15. Frons and/or vertex with stiff, pale orange setae ...... 16 Frons and vertex without stiff, pale orange setae ...... 18
16. Vertex strongly protuberant medially ...... gibber Vertex not protuberant ...... 17
17. Vertex with single row of stiff setae; clypeus convex ...... gurneyi Vertex and frons laterally with many stiff setae; clypeus flat ...... latifrons
18. Medially, facial fovea narrower than, base of scape ...... 19 Medially, facial fovea as wide as, or wider than, base of scape ...... 20
19. Hair of median clypeus branched ...... minutus Hair of median clypeus branched ...... sp. C*
20. Pygidial plate not striate; body length ca 9 mm ...... fallax Pygidial plate striate ...... 21
21. Paraocular area with dense, white plumose hair; body length ca 7 mm ...... nix Paraocular area with sparse hair; body length ca 5 mm ...... sp. D*
* Species recognised, but not named by Maynard, 1991.
Some morphological features of diagnostic importance.
Labrum. The labrum of four species has an elongated apical area so that the labrum is divided horizontally by a strong carina (Fig 1). The apical part is strongly concave while the basal part is concave, has a medial depression or, rarely, is
258 Appendices convex. The labrum fifth species has a very short apical area and appears smoothly convex.
Male hind tibia. Males of all the newly described species, and most of those previously described, have a modified hind tibia, with the apical half compressed on the anterior side (Fig 2). The narrow ridge thus formed bears 6 to 8 stiff setae that are frequently hooked. The maximum width of the hind tibia relative to the tibial length is a useful external character for separating species.
Male hind tibial spurs. All the males have two hind tibial spurs with minutely ciliate or finely pectinate margins. The outer spur is shorter than the inner, but difference in length varies between species. The shape of the inner spur varies from almost straight to flattened and strongly recurved (Fig 3).
Male sterna. Males of all the newly described species have apical fringes of hair on S3 and S4 and a raised longitudinal ridge basally on S6 (Fig 4). The only noticeable variation between the species was in the density of the apical hair tuft on S6.
Figs 1–4. (1) Leioproctus latifrons, female, lower head showing labrum.
(2),(3) Leioproctus nix, male: (2) part of hind leg showing tibia (3) ventral view of apex of hind tibia showing the spurs (4) Leioproctus gibber ventral view of metasoma showing shape and hair pattern of S3–S6.
259 Appendices
Species Descriptions
Authorship: in accordance with Article 50.1 and Recommendation 50A of
International Code of Zoological Nomenclature, MB is designated as the author of the new names.
Leioproctus (Protomorpha) crispus Batley n. sp.
Type. Holotype 24 km SW Charleville, Queensland (26.532°S 146.036°E),
28 Aug 2009, M. Batley, ex Eremophila gilesii in AM (K. 273317).
Specimens examined. The holotype and the following paratypes. Queensland: 8 km E Charleville
(26.415°S 146.306°E), 27 Aug 2009, M. Batley, ex Petalostylis labicheoides, in AM (K. 273301); , 68 km S
Charleville (26.988°S 146.032°E), 29 Aug 2009, M. Batley, ex Stemodia glabella, in AM (K. 273393); 2 , 6
Kunnamuka Swamp (23.353°S 138.261°E), 25-28 Apr 2010, pan trap, in AM (K. 362301-8).
Diagnosis
Males with flagellum longer than head width, large ventroapical projection on hind basitarsus and weak setae on hind tibia. Female frons and clypeus closely covered with erect, weakly-branched, curly hair. The labrum of both sexes smoothly convex.
Figs 5–8. Leioproctus crispus male: (5) hind tibia and basitarsus (6) S7, ventral view on left (7) S8, ventral view on left (8) genitalia, ventral view.
260 Appendices
Fig 9. Leioproctus crispus female: front view of head showing clypeal hair and labrum.
Descriptions
Male.— Head width 2.1 mm, body length 6.8 mm. Relative dimensions: HW
50, HL 46, UID 39, UFW 36, LID 30, HVO 3, WOC 21, OOD 10, IAD 8, ASD 5, AOD 7,
SL 18, SW 5, FL 71.—Inner orbits strongly convergent ventrally; clypeus transversely convex, longitudinally almost flat; labrum smoothly convex; flagellum longer than head width; metanotum with fine contiguous punctures and weak nodule; basal half area of propodeum subhorizontal rounding smoothly onto vertical half. Hind tibia slender, maximum width 0.22× length, bearing 7 stiff setae, declivous and weaker than those of other species; hind tibial spurs large, weakly recurved, outer 0.83× as long as inner; hind basitarsus flat, narrow, width 0.25× length, with large distal projection on anterior margin (Fig 5). Metasomal T2 with postgradular area weakly depressed; S6 with polished medial ridge; S7,8 and genitalia as in Figs 6-8.—Integument black except flagellum brown becoming orange-brown towards apex, mandible orange-brown with dark red tip, anterior faces of fore and mid tibia and inner faces fore and mid tarsi yellow-brown, outer faces all tarsi brown, metasomal terga with posterior margins translucent brown, sterna dark brown.—Clypeus densely punctate, supraclypeal area impunctate,
261 Appendices polished; frons with strong, dense, medium punctures, fovea not impressed; scutum, scutellum and mesepisternum strongly and densely punctate, polished; metanotum with fine contiguous punctures and weak tubercle; area of propodeum coarsely rugose, propodeum laterally with large, contiguous punctures; metasoma terga strongly punctate except wide apical margins.—Face below mid frons densely covered with adpressed, silver-white pubescence, vertex with close, pale brown, erect, plumose hair, gena with similar white hair; scutal hair open, white, long, finely plumose, scutellum and metanotum with sparse, long, finely plumose, pale brown hair, somewhat denser around margins; metasomal T1 with open, long, white hair on basal half, T1,2 with white hair apicolaterally, T2-4 with broad basal bands of close white hair; S3,4 with long apical fringes, slightly shorter medially,
S6 with weak fringe around apex but no apical tuft.
Female.— Head width 2.3 mm, body length 7.5 mm. Relative measurements: HW 50, HL 46, UID 38, UFW 40, LID 33, DMA 32, HVO 2, WOC 19,
OOD 10, IAD 9, ASD 4, AOD 9, SL 16, SW 4, FL 27.—Inner orbits convergent ventrally; clypeus transversely convex, base elevated in front of eyes; labrum uniformly convex, smooth and polished (Fig 9); mandible narrow with weak, rounded outer ridge; flagellum 0.54× as long as head width; metanotum with fine contiguous punctures and weak nodule; basal half area of propodeum subhorizontal rounding smoothly onto vertical half. Hind basitibial plate carinate, apically rounded; inner hind tibial spur with ca 5 well-spaced teeth; pygidial plate emarginate.—Integument black except flagellum brown, mandible amber with dark red tip, interior faces of tibiae and tarsi brown, metasomal terga with posterior margins translucent brown.—Clypeus polished, densely punctured except apically and along medial line; supraclypeal area impunctate, polished;
262 Appendices frons with smaller contiguous punctures; fovea impunctate, dull, reaching lateral ocellus, weakly impressed with indistinct margins, as wide as basal width scape; scutum and scutellum densely punctate; metanotum with fine contiguous punctures and weak tubercle; area of propodeum coarsely rugose, propodeum laterally with dense, round punctures; metasoma terga polished, densely punctate except on broad posterior margins; sterna with punctures sparse basally becoming dense apically; pygidial plate not striate, but with fine lineolation like a fingerprint.—Face with small area of dense, plumose, white pubescence between antennal socket and eye margin; frons and clypeus closely covered with weakly- branched, erect, fine, curly hair (Fig 9); vertex sparsely covered with weakly- branched, straight hair; scutum and scutellum closely to densely covered with very short, tightly-branched brown hair, becoming paler and longer around margins; mesepisternum closely covered with long, white, plumose hair; hind tibial scopa white; hind basitarsus with fringe of long white plumose hair on posterior margin; metasomal T1 with dense white hair apicolaterally, T2-4 with close white hair bands basally (usually obscured by preceding tergum) and broad, dense, white, apical hair bands, interrupted medially on T2; prepygidial fimbria brown; S5 with dense, pale brown apical fringe.
Remarks. The sexes were associated by the shape of the labrum. While there was no example of coincident collection, a male and females were taken in pan traps on the same day and in the same area, and specimens from the
Charleville area were collected within days and a few kilometers of each other.
Etymology. The specific name is a Latin adjective meaning curly, referring to the facial hair of the female.
263 Appendices
Leioproctus (Protomorpha) gibber Batley n. sp.
Type. Holotype Ethabuka Station, Queensland (23.756°S 138.474°E), 21
Apr 2012, M. Batley, ex Scaevola parvibarbata in AM (K. 362309)
Specimens examined. Holotype and the following paratypes. Queensland: , 3 Ethabuka Station
(23.748°S 138.474°E & 3.758°S 138.483°E), 14 Apr 2012, M. Batley, ex Scaevola depauperata in AM (K.
362310,11,54,55); Field River (23.922°S 138.119°E), 18 Apr 2012, M. Batley, ex Scaevola depauperata in AM
(K. 362312); , Ethabuka Station (23.738°S 138.467°E), 21 Apr 2012, M. Batley, ex Scaevola depauperata in AM
(K.362356); 2 Field River (23.804°S 138.055°E), 22 Apr 2010, T. Popic, ex Scaevola parvibarbata in AM (K.
362313,14); Kunnamuka Swamp (23.355°S 138.230°E), 23 Nov 2010 in AM (K. 362315).
Diagnosis
Female with large protuberance on vertex and stiff, pale orange setae across frons (Fig 14). Male with slight projection on vertex, hind tibia with two large, almost straight spurs. Both sexes have mandibles with a prominent outer ridge and no preapical tooth.
Figs 10–13. Leioproctus gibber male: (10) hind tibia and basitarsus (11) S7, ventral view (12) S8, ventral view (13) genitalia, ventral view.
264 Appendices
Fig 14. Leioproctus gibber female: vertex of head.
Descriptions
Male.— Head width 1.9 mm, body length 5.7 mm. Relative measurements:
HW 50, HL 44, UID 34, UFW 30, LID 23, HVO 0.7, WOC 19, OOD 9, IAD 8, AOD 6, SL
14, SW 4, FL 34. Vertex of head weakly elevated medially; inner orbits strongly convergent ventrally; clypeus almost flat basally, gently convex on apical ¼; antennal sockets level with front of eye viewed laterally; mandible simple with prominent outer ridge; labrum basally smoothly convex with medial depression, apically concave; flagellum 0.68× as long as head width; metanotum with fine contiguous punctures and weak nodule; area of propodeum subhorizontal basally, separated abruptly from vertical half but not carinate. Hind femur almost flat ventrally; hind tibia moderately broad, maximum width 0.27× length, bearing 8 stiff, apically bent setae, the most distal being particularly large and flattened (Fig
10); hind tibial spurs large, almost straight, inner distinctly pectinate, outer 0.75× as long as inner; hind basitarsus dished, width 0.40× length, with small apical extension of anterior margin. Metasomal T2 with postgradular area weakly depressed; S6 with polished medial ridge; S7,8 and genitalia as in Figs 11-13. —
Integument black except flagellum brown, mandibles orange-brown with dark red tips, anterior face fore tibia, fore tarsi yellow-brown, mid and hind tarsi dark
265 Appendices brown, metasomal terga black or dark brown with posterior margins translucent brown, sterna dark brown or black.—Clypeus densely punctate, supraclypeal area medially impunctate and polished; frons with strong, dense, medium punctures, fovea obscured by tomentum; scutum and scutellum polished and closely punctate; mesepisternum densely punctate; metanotum with fine contiguous punctures and weak tubercle; area of propodeum area coarsely rugose, laterally propodeum densely punctate; metasomal terga densely punctate except for wide impunctate apical margins.—Pubescence: Face densely covered with adpressed, silver-white pubescence, vertex with close, erect, white setae between ocelli, weakly branched laterally, gena with open, white, plumose hair; scutum and scutellum openly covered with white, finely plumose hair, longer around margin of scutellum; similar long hair medially on metanotum; metasomal T1 with open, long, white hair on basal half, T1,2 with white hair apicolaterally (possible basal hair bands hidden), hair sparse elsewhere; S3,4 with long apical fringes, S6 with dense apical tuft.
Female—Head width 2.3 mm, body length 7.2 mm. Relative measurements:
HW 50, HL 43, UID 36, UFW 34, LID 26, HVO 4, WOC 17, OOD 10, IAD 8, AOD 8, SL
16, SW 3, FL 23. Vertex of head with large mesial protuberance (Fig 10), inner orbits convergent ventrally; clypeus flat except at extreme ventrolateral corners, apical marginal straight with a tiny medial projection; mandible with prominent outer ridge and no preapical tooth; labrum strongly carinate, basally and apically concave; flagellum less than ½ as long as head width; propodeum as for male. Hind basitibial plate carinate, bluntly acute apically; inner hind tibial spur gently curved with ca10 strong teeth; pygidial plate emarginate apically.—Integument black except labrum and flagellum brown, mandibles amber with dark red tips,
266 Appendices metasomal terga with posterior margins translucent brown; sterna dark brown.—
Clypeus dull, densely punctate, except ventrolateral corner impunctate and polished; interantennal area with small medial impunctate area; frons and vertex densely punctate, fovea slightly narrower than base of scape not reaching lateral ocellus; scutum and scutellum densely punctate; metanotum with fine contiguous punctures and weak tubercle; area of propodeum irregularly rugose, propodeum laterally with dense round punctures; metasomal terga polished and densely punctate except on posterior margins; sterna with punctures sparse basally becoming dense apically; pygidial plate coarsely striate.—Face with small area of dense, white pubescence in paraocular areas to just above antennal sockets, vertex and frons with close, stiff, pale orange setae, clypeus with close, fine simple hair; scutum and scutellum closely covered with short, densely-branched, erect, cream hair, becoming dense around margins; metanotum with slightly longer hair near tubercle; hind tibial scopa white; hind basitarsus with open fringe of long white plumose hair on posterior margin; metasomal T1 with dense white hair apicolaterally, T2-4 with broad, dense white hair bands apically, interrupted medially on T2 and close white hair bands basally; prepygidial fimbria white; S5 with dense, white apical fringe.
Remarks. The sexes were associated by coincident collection and the protruding vertex.
Etymology. The specific name is a Latin noun in apposition meaning hump, referring to the vertex of the female.
Leioproctus (Protomorpha) gurneyi Batley n. sp.
267 Appendices
Type. Ethabuka Station, Queensland (23.753°S 138.493°E), 20 Apr 2012,
M. Batley, ex Lechenaultia divaricata in AM (K.362316).
Specimens examined. Holotype and the following paratypes. 3 same data as holotype, in AM (K.
362317-9).
Diagnosis
Both sexes with face convex viewed laterally. Female vertex with a single row of stiff, curved, pale orange setae. Male hind tibia slender with weak setae, and large, almost straight spurs.
Figs 15-18. Leioproctus gurneyi male: (15) hind tibia and basitarsus (16) S7, ventral view (17) S8, ventral view (18) genitalia, ventral view.
Fig 19. Leioproctus gurneyi female: vertex of head.
Descriptions
268 Appendices
Male.— Head width 1.9 mm, body length 5.6 mm. Relative measurements:
HW 50, HL 41, UID 32, UFW 32, LID 23, HVO 1, WOC 20, OOD 7, IAD 10, AOD 6, SL
13, SW 4, FL 34. Inner orbits strongly convergent ventrally; face convex, antennal sockets in front of eyes viewed laterally; clypeus transversely convex; labrum basal part convex with small medial depression, apical part concave, less than half as long as basal part; flagellum 0.68× as long as head width; metanotum with fine contiguous punctures and weak tubercle; area of propodeum subhorizontal basally, vertical apically with a sharp change of slope but no carina. Hind tibia slender, maximum width 0.22× length, with 6 stiff setae; hind tibial spurs large, almost straight, outer 0.85× as long as inner; hind basitarsus dished, width 0.30× length, with small spathulate apical projection of anterior margin. Metasomal T2 with postgradular area depressed; S6 with very weak medial ridge; S7,8 and genitalia as in Figs 16-18.—Integument black except flagellum brown, mandibles orange-brown with dark red tips, labrum dark brown basally, orange-brown apically, metasomal terga with posterior margins translucent brown, sterna dark brown, anterior face fore tibia and fore tarsi yellow-brown, mid and hind tarsi brown.—Clypeus densely punctate; supraclypeal area dull, punctate; frons with strong, dense, medium punctures, fovea obscured by tomentum; scutum and scutellum polished, closely to openly punctate; mesepisternum densely punctate; metanotum with fine contiguous punctures and weak tubercle; area of propodeum area coarsely rugose, laterally propodeum densely punctate; metasomal terga densely punctate except for wide impunctate apical margins.—Face densely covered with adpressed, silver-white pubescence; vertex with close, erect, plumose, white hair; gena with similar hair; scutum and scutellum openly covered with long, finely plumose, white hair, longer and denser around margins;
269 Appendices metasomal T1 with open, long, white hair on basal half, T1,2 with dense, white hair apicolaterally, T3,4 with sparse, white apical hair bands, T2-4 with bands of close white hair basally; S3,4 with long apical fringes; S6 with weak apical hair tuft.
Female—Head width 2.3 mm, body length 7.5 mm. Relative measurements:
HW 50, HL 42, UID 34, UFW 35, LID 25, HVO 1, WOC 21, OOD 7, IAD 9, AOD 8, SL
14, SW 3, FL 25.—Inner orbits convergent ventrally; face convex, antennal sockets in front of eyes viewed laterally, interantennal area somewhat bulbous; clypeus convex with convex apical margin; labrum strongly carinate, basally convex with a large medial depression, apically concave; mandible narrow with preapical tooth and weak, rounded outer ridge; flagellum ½ as long as head width; propodeum as for male. Hind basitibial plate strongly carinate, apically rounded; inner hind tibial spur gently curved with 5 well-spaced teeth; pygidial plate striolate, apex rounded, entire.—Integument black except flagellum brown, mandibles amber with dark red tips, labrum brown basally, apically amber; metasomal terga with posterior margins translucent brown, sterna black or dark brown, anterior face fore tibia yellow-brown.—Clypeus, interantennal area, frons and vertex densely punctate; fovea strongly impressed, impunctate, ca 2/3 width base of scape, curving towards but not reaching lateral ocellus; scutum and scutellum densely punctate; metanotum with fine contiguous punctures and weak tubercle; area of propodeum irregularly rugose, propodeum laterally densely punctate; metasomal terga polished and densely punctate except on posterior margins; sterna with punctures sparse basally becoming dense apically; pygidial plate striolate.—Face with dense, adpressed, white pubescence in paraocular areas from bottom of eye to near its top; vertex with a row of ca 12 pale orange, erect setae curved forward; frons and clypeus openly covered with smaller, fine unbranched setae; scutum and scutellum
270 Appendices closely covered with short, densely-branched, erect, white hair, becoming dense around margins; metanotum with slightly longer hair near tubercle; hind tibial scopa white, sparsely branched; hind basitarsus with open fringe of long white plumose hair on posterior margin; metasomal T1,2 with dense white hair apicolaterally, T3,4 with broad, dense white apical hair bands, T2-4 basally with close white hair bands, usually obscured by preceding tergum; prepygidial fimbria pale brown; S5 with dense, white apical fringe.
Remarks. The sexes were associated by coincident collection and the convex face, viewed laterally.
Etymology. The species is named after Alex Gurney, creator of the comic strip "Bluey and Curley".
Leioproctus (Protomorpha) latifrons Batley n. sp.
Type. Holotype Kunnamuka Swamp, Queensland (23.353°S 138.260°E),
22 Nov 2010, T. Popic ex Scaevola parvibarbata in AM (K. 362320)
Specimens examined. Holotype and the following paratypes. Queensland: , Kunnamuka Swamp
(23.353°S 138.260°E), 15 Apr 2012, M. Batley, ex Scaevola depauperata in AM (K. 362321); 5 , Ethabuka
Station (23.756°S 138.474°E), 21 Apr 2012, M. Batley, ex Scaevola depauperata in AM (K. 362322-26);
Ethabuka Station (23.740°S 138.466°E), 14 Nov 2010, T. Popic ex Scaevola parvibarbata in AM (K362327).
Diagnosis
Female with vertex broadly elevated, bigibbous bearing stiff orange setae, frons with lateral patches of similar setae, clypeus flat. Male vertex weakly bigibbous, hind tibia with hooked setae, basitarsus broad (width 0.32× length) with moderately long spurs, inner recurved near base, outer 0.7× as long as inner.
271 Appendices
Figs 20-23. Leioproctus latifrons male: (20) hind basitarsus (21) hind tibia
(22) S7, ventral view (23) S8, ventral view (24) genitalia, ventral view.
Fig 25. Leioproctus latifrons female: vertex of head.
Descriptions
Male—Head width 1.9 mm, body length 5.7 mm. Relative measurements:
HW 50, HL 42, UID 32, UFW 33, LID 23, HVO 3, WOC 20, OOD 8, IAD 7, AOD 7, SL
15, SW 4, FL 38.—Inner orbits strongly convergent; clypeus gently convex apically, basal ¾ almost flat; labrum strongly carinate, basal part smoothly convex, apical part obscured; flagellum 0.75× as long as head width; area of propodeum subhorizontal basally, vertical apically with a sharp change of slope but no carina.
Hind tibia broad, maximum width 0.32× length, with 7 stiff, apically hooked setae and two large spurs, inner recurved near base, outer 0.7× as long as inner; hind basitarsus weakly concave, width 0.34× length, with a short apical extension of anterior margin. Metasomal T2 with postgradular area depressed; S6 with strong
272 Appendices medial ridge; S7,8 and genitalia as in Figs 22-24.—Integument black except flagellum dull orange-brown, mandibles orange-brown with dark red tips, labrum orange-brown, metasomal terga with posterior margins translucent brown, sterna dark brown, anterior face fore tibia, distal ends of all tibiae and all tarsi orange- brown, S6 brown.—Clypeus, supraclypeal area and frons densely punctate, fovea obscured by tomentum; scutum and scutellum polished, closely punctate; mesepisternum densely punctate; metanotum with fine contiguous punctures and weak tubercle; area of propodeum coarsely rugose, laterally propodeum with large, contiguous punctures; metasoma strongly punctate except for wide impunctate apical margins.—Face below mid frons densely covered with adpressed, silver-white pubescence; vertex with open, erect, plumose pale brown hair; gena with open, white, plumose hair; scutum and scutellum openly covered with long, white, finely plumose hair; metasomal T1 with open, long, white hair on basal half, T1,2 apicolaterally with weak white hair patches, T2-4 basally with bands of close white hair; S3,4 with long apical fringes; S6 with dense apical hair tuft.
Female—Head width 2.2 mm, body length 6.8 mm. Relative measurements:
HW 50, HL 40, UID 35, UFW 34, LID 26, HVO 2, WOC 21, OOD 7, IAD 8, AOD 8, SL
15, SW 3, FL 22.—Inner orbits convergent ventrally; clypeus flat, except ventrolateral corners strongly convex, apical margin gently sinuous area with small medial projection; labrum strongly carinate, basal and apical areas concave; mandibles narrow with no preapical tooth (Fig 1) and weak, rounded outer ridge; flagellum <½ as long as head width; propodeum as for male. Hind basitibial plate strongly carinate; inner hind tibial spur gently curved with 8 strong teeth; pygidial plate striate with small apical emargination.—Integument black, except labrum
273 Appendices and marginal area of clypeus amber, flagellum brown, mandibles amber with dark red tips, metasomal terga with posterior margins translucent brown, sterna dark brown, anterior face fore tibia yellow-brown.—Clypeus densely punctate except ventrolateral corner impunctate and polished; frons and vertex densely punctate; fovea impressed, impunctate and dull, about as wide as base of scape, almost reaching lateral ocellus; scutum and scutellum densely punctate; metanotum with fine contiguous punctures and small tubercle; area of propodeum irregularly rugose, propodeum laterally densely punctate; metasomal terga polished and densely punctate except on posterior margins; sterna with punctures sparse basally becoming dense apically, pygidial plate striate.—Face with dense, adpressed, white pubescence covering a triangular part of paraocular area from bottom of eye to level of mid frons; vertex and upper frons laterally with stiff orange setae; clypeus and frons medially with close, fine simple hair; scutum closely covered with short, densely-branched, erect, white hair, dense around margins, open anteriorly; scutellum with pale brown hair; hind tibial scopa white, plumose; hind basitarsus with open fringe of long white plumose hair on posterior margin; metasomal T1,2 with dense white hair apicolaterally, T3,4 with broad, dense white apical hair bands, T2-4 basally with close white hair bands; prepygidial fimbria white; S5 with dense, white apical fringe.
Remarks. The sexes were associated on the basis of their biggibous vertex.
A male and female were collected within days from the same plant species in neighbouring locations.
Etymology. The specific name is a Latin noun in apposition meaning broad brow.
274 Appendices
Leioproctus (Protomorpha) nix Batley n. sp.
Type. Holotype Field River, Queensland (23.808°S 138.058°S) 19 Nov
2010, T. Popic, ex pan trap in AM (K. 362330).
Specimens examined. Holotype and the following paratypes. Queensland: Field River (23.804°E
138.055°S), 24 Jun 2010, T. Popic in AM (K. 362353); , 6 Ethabuka Station (23.745°S 138.470°E), 12-14 Nov
2010, T. Popic, pan trap and ex Trachymene glaucifolia in AM (K. 362331-37); Field River (23.808°S
138.058°S), 17 Nov 2010, T. Popic, ex Dicrastylis costelloi in AM (K. 362338); Field River (23.808°S
138.058°S), 19 Nov 2010, T. Popic, ex Brunonia australis in AM (K. 362339); 13 Kunnamuka Swamp
(23.361°S 138.264°E), 22-24 Nov 2010, T. Popic, ex Brunonia australis, Dicrastylis costelloi, Trachymene glaucifolia, Trianthema pilosa in AM (K.362340-52).
Diagnosis
Female with hair on clypeus and frons simple, pygidial plate striolate, fovea ca as wide as base of scape, but obscured by pubescence. Male hind femur with broad ventral projection of anterior face, inner hind tibial spur flattened and strongly recurved, outer spur short, 0.3× as long as inner (Fig 3).
Figs 26-30. Leioproctus nix male: (26) hind basitarsus outer view (27) hind femur anterior view (28) S7, ventral view (29) S8, ventral view (30) genitalia, ventral view.
275 Appendices
Fig 31. Leioproctus nix female: vertex of head.
Descriptions
Male—Head width 2 mm, body length 6.9 mm. Relative measurements: HW
50, HL 48, UID 33, UFW 33, LID 24, HVO 4, WOC 19, OOD 8, IAD 8, AOD 7, SL 15, SW
4, FL 41.—Inner orbits convergent ventrally; clypeus transversely convex; interantennal area raised; labrum strongly carinate, basal part convex with medial depression, apical part concave; lower gena concave; flagellum 0.8× times as long as head width; metanotum with distinct tubercle; area of propodeum subhorizontal basally, vertical apically with a sharp change of slope but no carina.
Hind femur swollen ventrally on anterior face (Fig. 27); hind tibia moderately broadened, maximum width 0.27× length, with 8 stiff, curved setae and two spurs, inner flattened and strongly recurved, outer only 0.3× as long as inner; hind basitarsus dished with short, curved apical extension of anterior margin, width
0.43× length. Metasomal T2,3 with postgradular area depressed; S6 with strong medial ridge; S7,8 and genitalia as in Figs 28-30.—Integument black except flagellum mostly orange-brown, labrum and mandibles orange-brown, metasomal terga with posterior margins translucent brown; sterna dark brown, tarsi dull orange-brown.—Clypeus densely punctate; supraclypeal area polished and impunctate on upper half; frons densely punctate, fovea obscured by pubescence;
276 Appendices scutum and scutellum polished, closely punctate; metanotum with fine contiguous punctures and distinct tubercle; area of propodeum coarsely rugose, propodeum laterally with contiguous punctures; metasomal terga closely punctate except for wide impunctate apical margins.—Face below mid frons densely covered with adpressed, silver-white pubescence; vertex with open, long, erect, plumose, white hair; gena with open, white, plumose hair; scutum and scutellum openly covered with long, white, finely plumose hair; metasomal T1 with open, long, white hair on basal half, T2-6 basally with broad bands close white hair; S6 with relatively weak apical tuft; S3,4 with long, even fringes.
Female—Head width 2.4 mm, body length 7.3 mm. Relative measurements:
HW 50, HL 47, UID 33, UFW 34, LID 27, HVO 3, WOC 18, OOD 8, IAD 8, AOD 9, SL
16, SW 3, FL 27.—Inner orbits convergent ventrally; clypeus weakly convex except at ventral margin; labrum strongly carinate, basally convex with oval medial depression, apically concave; flagellum ca ½ as long as head width; mandible narrow, with rounded outer ridge and subapical tooth; propodeum as for male.
Hind basitibial plate strongly carinate; inner hind tibial spur gently curved with 9 strong, well-spaced teeth; pygidial plate with irregular striae.—Integument black except flagellum dorsally dark brown with 4 apical flagellomeres orange-brown, ventrally orange-brown, labrum and mandibles orange-brown, metasomal T1,2 brown on apical half, all terga with posterior margins translucent brown.—Clypeus and frons densely punctate with polished interspaces; fovea impressed, about as wide as scape, upper end curving towards lateral ocellus; scutum and scutellum polished, closely punctate; propodeum area rugulose; lateral parts polished with close, even punctures; metanotum with fine contiguous punctures and distinct tubercle; metasomal terga closely punctate except in apical marginal areas; sterna
277 Appendices with punctures sparse basally becoming dense apically, pygidial plate striolate.—
Face with dense white, pubescence in paraocular areas extending almost to top of eye and across lower ¼ of frons; frons and clypeus with open to sparse simple hair; scutum and scutellum closely covered with short, densely-branched, white hair becoming dense around margins; metasomal T1-4 with dense white apical hair bands, interrupted medially on T1,2; T2,3 with close, white, basal hair bands; prepygidial fimbria pale brown.
Remarks. The sexes were associated by the shape of the clypeus and coincident collection.
Etymology. The specific name is a Latin noun in apposition meaning snow, referring to the colour of the facial hair of the female.
Pollen examination.
Pollen was removed from the scopae of four female specimens: two L. latifrons collected from Scaevola parvibarbata, one L. gibber collected from
Scaevola depauperata and one L. gurneyi collected from Lechenaultia divaricata.
Bees collected from Scaevola species carried tricolporate pollen indistinguishable by light microscopy from pollen taken from the flowers. The L. gurneyi specimen was carrying pollen tetrads indistinguishable from that taken from flowers and that illustrated in the Australasian Pollen and Spore Atlas (APSA 2013).
Discussion
All five new species have the major characteristics that distinguish
Protomorpha from other Leioproctus: small size (5-9 mm long); metasomal terga with strong, dense punctures and apical hair bands; females with impressed facial
278 Appendices foveae and sparse, weakly branched hair on the metasomal sterna; and above all males with elaborately shaped seventh sterna and modified hind tibiae and basitarsi. The additional species will help refine the diagnostic characteristics of the subgenus. Although the carinate labrum and preapical tooth on the mandibles are common features, they are absent in one or more species, just as it was previously demonstrated (Maynard 1991) that the sculpture of the female pygidial plate varies. The newly described males consistently have strong apical hair fringes on S3 and S4, which suggests that this feature may be more common than previous thought.
The presence of so many species in a relatively limited area was unexpected. In addition to the five new species, L. (Protomorpha) alloeopus and L.
(P.) tarsalis females were collected at about 5 sites in sand dune country with a total area of approximately 3,000 km2. Together with the known distributions of previously described species, this suggests that the subgenus is strongly adapted to life in arid environments with highly variable rainfall.
Females of four species have distinctive setae on the vertex or frons, a feature not previously found in the subgenus. Modified facial hairs have been suggested (Müller 1996) as adaptations to pollen collection from flowers that deposit pollen on dorsal areas of visitors. The small number of specimens and absence of behavioural observations (Müller 1996, Gonzalez & Chavez 2004) prevent the drawing of similar conclusions for the new species, but the limited information available is consistent with this explanation for the facial setae. (1)
Only females carry the modified setae. (2) Flowers visited by L. gibber, L. gurneyi and L. latifrons, the species with stiff facial setae, are members of the Goodeniaceae
279 Appendices family which present pollen to the backs of visitors. Pollen removed from the scopae of all three species was indistinguishable by light microscopy from the pollen of the flowers they were visiting. (3) Flowers visited by L. crispus do not have specialised pollen presenters, but are zygomorphic and the anthers are where insect visitors would frequently have their backs. (4) The flowers visited by
L. (Protomorpha) species without special setae are more varied and less specialised (Table 1). L. tarsalis specimens were collected from Tribulus within minutes and a few meters from where L. gurneyi were collected from Lechenaultia.
Rainfall in the Lake Eyre Basin, in which these bees were found, is more variable and dry periods last longer than in any other part of the World (McMahon et al. 2008). Study of these morphologically similar species of L. (Protomorpha) could provide interesting insights into the effect of such "boom-bust" conditions on the evolution of and interactions between species.
Table 1. Flowers visited by Leioproctus (Protomorpha) species. Leioproctus Females with Flowers visited species modified setae alloeopus no Grevillea stenobotrya, Scaevola depauperata, Senna pleurocarpa, Euphorbia drummondii, Tribulus terrestris crispus yes Eremophila gilesii, Stemodia glabella, Petalostylis labicheoides gibber yes Scaevola depauperata, Scaevola parvibarbata gurneyi yes Lechenaultia divaricata latifrons yes Scaevola depauperata nix no Trachymene glaucifolia, Dicrastylis costelloi, Brunonia australis tarsalis no Tribulus terrestris
Acknowledgements
We thank Professors Glenda Wardle and Chris Dickman, Dr Yvonne Davila, the Desert Ecology Research Group and volunteers for valuable assistance in the
280 Appendices
Simpson Desert and to Bush Heritage Australia and Carlo Station for granting access to the Simpson Desert sites. Funding for the project that allowed collections in the Simpson Desert was provided by the Australian Research Council.
References
Alqarni, A.S., M.A. Hannan, V.H. Gonzalez, and M.S. Engel, 2012. A new species of Chalicodoma from Saudi Arabia with modified facial setae
(Hymenoptera, Megachilidae). ZooKeys 204: 71-83.
ASPA, 2013. http://apsa.anu.edu.au/sample/282-5-
2?order=desc&q=lechenaultia. Accessed 1/4/2013.
Gonzalez V.H., and F.C. Chavez, 2004. Nesting biology of a new high Andean bee, Anthophora walteri Gonzalez (Hymenoptera: Apidae: Anthophorini). Journal of the Kansas Entomological Society 77: 584–592.
Maynard, G.V., 1991. Revision of Leioproctus (Protomorpha) Rayment
(Hymenoptera: Colletidae) with Descriptions of Two New Species. Journal of the
Australian Entomological Society 30: 67-75.
McMahon, T.A., R.E. Murphy, M.C. Peel, J.F. Costelloe, and F.H.S. Chiew, 2008.
Understanding the surface hydrology of the Lake Eyre Basin: Part 1—Rainfall.
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Michener, C. D. 2007. The Bees of the World. Second edition. Baltimore and
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281 Appendices
Michener, C.D., and A. Fraser, 1978. A Comparative Anatomical Study of the
Mandibular Structure in Bees (Hymenoptera: Apoidea). The University of Kansas
Science Bulletin 51: 463-482.
Müller, A., 1996. Convergent evolution of morphological specializations in
Central European bee and honey wasp species as an adaptation to the uptake of pollen from nototribic flowers (Hymenoptera, Apoidea and Masaridae). Biological
Journal of the Linnean Society 57: 235-252.
Popic, T.J., G.M. Wardle, and Y.C. Davila, 2013. Flower-visitor networks only partially predict the function of pollen transport by bees. Austral Ecology 38: 76–
86.
Rightmyer, M.G., M. Deyrup, J.S. Ascher & T. Griswold, 2011. Osmia species
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282 Appendix 8
Plant species in this study and their authorities. Family Plant species Authority Aizoaceae Trianthema pilosa F.Muell. Amaranthaceae Ptilotus atriplicifolius (A.Cunn. ex Moq.) Benl Amaranthaceae Ptilotus latifolius R.Br. Amaranthaceae Ptilotus polystachyus F.Muell Araliaceae Trachymene glaucifolia (F.Muell.) Benth. Asteraceae Calotis erinacea Steetz Asteraceae Calotis plumulifera F.Muell. Asteraceae Helipterum molle (A.Cunn. ex DC.) Paul G.Wilson Asteraceae Pterocaulon sphacelatum (Labill.) F.Muell. Boraginaceae Halgania cyanea Lindl. Boraginaceae Trichodesma zeylanicum (Burm.f.) R.Br. Brunoniaceae Brunonia australis Sm. Chenopodiaceae Salsola kali L. Euphorbiaceae Adriana tomentosa Gaudich (isonym) Euphorbiaceae Euphorbia drummondii Boiss. Fabaceae Acacia bivenosa DC. Fabaceae Acacia coriacea DC. Fabaceae Acacia dictyophleba F.Muell. Fabaceae Acacia ligulata A.Cunn. ex Benth. Fabaceae Acacia stenophylla A.Cunn. ex Benth. Fabaceae Crotalaria cunninghamii R.Br. Fabaceae Crotalaria eremaea F.Muell. Fabaceae Crotalaria smithiana A.T.Lee Fabaceae Psoralea eriantha Benth. Fabaceae Senna artemisioides (Gaudich. ex DC.) Randell Fabaceae Senna pleurocarpa (F.Muell.) Randell Fabaceae Swainsona phacoides Benth. Fabaceae Tephrosia rosea F.Muell. ex Benth. Frankeniaceae Frankenia gracilis Summerh. Goodeniaceae Goodenia cycloptera R.Br. Goodeniaceae Lechenaultia divaricata F.Muell. Goodeniceae Scaevola depauperata R.Br. Goodeniceae Scaevola parvibarbata Carolin Goodeniceae Scaevola parviflora K.Krause Haloragaceae Haloragis gossei F.Muell. Lamiaceae Dicrastylis costelloi F.M.Bailey Lamiaceae Dicrastylis lewellinii (F.Muell.) F.Muell. Lamiaceae Newcastelia cephalantha F.Muell. Lamiaceae Newcastelia spodiotricha F.Muell. Malvaceae Abutilon otocarpum F.Muell. Malvaceae Rulingia loxophylla F.Muell. Malvaceae Sida fibulifera Lindl. Malvaceae Sida trichopoda F.Muell. Malvaceae Triumfetta winneckeana F.Muell. Myrtaceae Eucalyptus pachyphylla F.Muell. Portulacaceae Calandrinia balonensis Lindl. Proteaceae Grevillea juncifolia Hook. Proteaceae Grevillea stenobotrya F.Muell. Rubiaceae Oldenlandia pterospora (F.Muell.) F.Muell. Sapindaceae Atalaya hemiglauca (F.Muell.) F.Muell. ex Benth. Scrophulariaceae Eremophila macdonnellii F.Muell. Violaceae Hybanthus aurantiacus (Benth.) F.Muell.
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