Neolucia bollami Eastwood, Braby & Graham, sp. nov. (Lepidoptera: Lycaenidae): speciation of a new allochronic cryptic butterfly from south-western Western Australia
Rodney G. Eastwood
A * , Michael F. Braby B C and Matthew R. Williams D
+ Author Affiliations
- Author Affiliations
A Department of Terrestrial Zoology, Western Australian Museum, 49 Kew Street, Welshpool, WA 6106, Australia.
B Division of Ecology and Evolution, Research School of Biology, The Australian National University, Acton, ACT 2601, Australia.
C The Australian National Insect Collection, National Research Collections Australia, GPO Box 1700, Canberra, ACT 2601, Australia.
D Department of Biodiversity Conservation and Attractions, 17 Dick Perry Avenue, Kensington, WA 6151, Australia.
Invertebrate Systematics 37(8) 552-570 https://doi.org/10.1071/IS23009
Submitted: 16 March 2023 Accepted: 8 August 2023 Published: 6 September 2023
© 2023 The Author(s) (or their employer(s)). Published by CSIRO Publishing.
Abstract
South-western Western Australia is a global biodiversity hotspot renowned for exceptional diversity of plants and animals. The evolutionary processes that have generated this high biodiversity are not always clear, particularly for invertebrates, yet the area supports a very large number of endemic species that have diversified in situ. We use an integrative taxonomic approach based on adult and immature morphology, ecology, behaviour and molecular data to investigate the taxonomic status of a sympatric but seasonally isolated form (Neolucia agricola occidens Waterhouse & Lyell, 1914 form ‘Julimar’) of the polyommatine butterfly Neolucia agricola (Westwood, 1851) in south-western Western Australia. Our molecular dataset comprised 112 samples representing all Neolucia Waterhouse & Turner, 1905 species (100 COI 5′ sequences, 658 bp, plus 12 COI 3′, tRNA Leu, COII and EF1-α sequences, 3303 bp). Maximum likelihood phylogenetic analysis of the combined dataset recovered form ‘Julimar’ and N. agricola as reciprocally monophyletic, with a mean uncorrected ‘p’ pairwise divergence of 5.77% for the ‘barcode’ region of COI. Based on this and other evidence we recognise form ‘Julimar’ as a new species, Neolucia bollami Eastwood, Braby & Graham, sp. nov., sister to N. agricola and endemic to south-western Western Australia. As a result of these findings, we evaluated the evolutionary history of the two Neolucia species in WA and the processes that may have contributed to the diversification in sympatry or allopatry. We conclude that the multiple effect traits associated with a host shift, including host fidelity and temporal divergence, played an important role in the diversification process and in maintaining the reproductive integrity of the nascent allochronic species.
ZooBank: urn:lsid:zoobank.org:act:53D9AD14-9694-4B5E-889C-A8D533E7F57D
Keywords: allochronic speciation, Fabaceae, host fidelity, host plant shift, integrative taxonomy, magic traits, modes of diversification, South-West Australian Ecoregion, sympatric speciation.
References
Akaike, H (1974). A new look at the statistical model identification. IEEE Transactions on Automatic Control AC-19, 716–723.| A new look at the statistical model identification.Crossref | GoogleScholarGoogle Scholar |
Alexander, RD, and Bigelow, RS (1960). Allochronic speciation in field crickets, and a new species, Acheta veletis. Evolution 14, 334–346.
| Allochronic speciation in field crickets, and a new species, Acheta veletis.Crossref | GoogleScholarGoogle Scholar |
Bell, KL, Hamm, CA, Shapiro, AM, and Nice, CC (2017). Sympatric, temporally isolated populations of the pine-white butterfly Neophasia menapia, are morphologically and genetically differentiated. PLoS One 12, e0176989.
| Sympatric, temporally isolated populations of the pine-white butterfly Neophasia menapia, are morphologically and genetically differentiated.Crossref | GoogleScholarGoogle Scholar |
Berlocher, SH, and Feder, JL (2002). Sympatric speciation in phytophagous insects: moving beyond controversy? Annual Review of Entomology 47, 773–815.
| Sympatric speciation in phytophagous insects: moving beyond controversy?Crossref | GoogleScholarGoogle Scholar |
Bethenod, M-T, Thomas, Y, Rousset, F, Frérot, B, Pélozuelo, L, Genestier, G, and Bourguet, D (2005). Genetic isolation between two sympatric host plant races of the European corn borer, Ostrinia nubilalis Hübner. II: assortative mating and host-plant preferences for oviposition. Heredity 94, 264–270.
| Genetic isolation between two sympatric host plant races of the European corn borer, Ostrinia nubilalis Hübner. II: assortative mating and host-plant preferences for oviposition.Crossref | GoogleScholarGoogle Scholar |
Braby MF (2000) ‘Butterflies of Australia. Their Identification, Biology and Distribution.’ (CSIRO Publishing: Melbourne, Vic., Australia)
Braby MF (2016) ‘The Complete Field Guide to Butterflies of Australia’. 2nd edn. (CSIRO Publishing: Melbourne, Vic., Australia)
Braby, MF, and Wurtz, GE (2018). A new subspecies of Neolucia hobartensis (Miskin, 1890) (Lepidoptera: Lycaenidae) from Mainland Southeastern Australia, with a Review of Butterfly Endemism in Montane Areas in this Region. Records of the Australian Museum 70, 423–433.
| A new subspecies of Neolucia hobartensis (Miskin, 1890) (Lepidoptera: Lycaenidae) from Mainland Southeastern Australia, with a Review of Butterfly Endemism in Montane Areas in this Region.Crossref | GoogleScholarGoogle Scholar |
Bush, GL (1994). Sympatric speciation in animals – new wine in old bottles. Trends in Ecology & Evolution 9, 285–288.
| Sympatric speciation in animals – new wine in old bottles.Crossref | GoogleScholarGoogle Scholar |
Byrne, M, Steane, DA, Joseph, L, Yeates, DK, Jordan, GJ, Crayn, D, Aplin, K, Cantrill, DJ, Cook, LG, Crisp, MD, Keogh, JS, Melville, J, Moritz, C, Porch, N, Sniderman, JMK, Sunnucks, P, and Weston, PH (2011). Decline of a biome: evolution, contraction, fragmentation, extinction and invasion of the Australian mesic zone biota. Journal of Biogeography 38, 1635–1656.
| Decline of a biome: evolution, contraction, fragmentation, extinction and invasion of the Australian mesic zone biota.Crossref | GoogleScholarGoogle Scholar |
Clement, M, Posada, D, and Crandall, KA (2000). TCS: a computer program to estimate gene genealogies. Molecular Ecology 9, 1657–1659.
| TCS: a computer program to estimate gene genealogies.Crossref | GoogleScholarGoogle Scholar |
Cocroft RB, Rodríguez RL, Hunt RE (2009) Host shifts, the evolution of communication, and speciation in the Enchenopa binotata species complex of treehoppers. In ‘Specialization, Speciation, and Radiation: The Evolutionary Biology of Herbivorous Insects’. (Ed. K Tilman) pp. 88–100. (Oxford University Press)
Coyne, JA (2007). Sympatric speciation. Current Biology 17, R787–R788.
| Sympatric speciation.Crossref | GoogleScholarGoogle Scholar |
Coyne JA, Orr HA (2004) ‘Speciation.’ (Sinauer Associates: MA)
Crisp, MD, and Cook, LG (2003). Phylogeny and Evolution of Anomalous Roots in Daviesia (Fabaceae: Mirbelieae). International Journal of Plant Sciences 164, 603–612.
| Phylogeny and Evolution of Anomalous Roots in Daviesia (Fabaceae: Mirbelieae).Crossref | GoogleScholarGoogle Scholar |
Darriba, D, Taboada, GL, Doallo, R, and Posada, D (2012). jModelTest 2: more models, new heuristics and parallel computing. Nature Methods 9, 772.
| jModelTest 2: more models, new heuristics and parallel computing.Crossref | GoogleScholarGoogle Scholar |
Dolman, G, and Joseph, L (2012). A species assemblage approach to comparative phylogeography of birds in southern Australia. Ecology and Evolution 2, 354–369.
| A species assemblage approach to comparative phylogeography of birds in southern Australia.Crossref | GoogleScholarGoogle Scholar |
Drès, M, and Mallet, J (2002). Host races in plant-feeding insects and their importance in sympatric speciation. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences 357, 471–492.
| Host races in plant-feeding insects and their importance in sympatric speciation.Crossref | GoogleScholarGoogle Scholar |
Dröllner, M, Barham, M, Kirkland, CL, Danišík, M, Bourdet, J, Schulz, M, and Aspandiar, M (2023). Directly dating Plio-Pleistocene climate change in the terrestrial record. Geophysical Research Letters 50, e2023GL102928.
| Directly dating Plio-Pleistocene climate change in the terrestrial record.Crossref | GoogleScholarGoogle Scholar |
Eastwood RG (2006) Ant Association and Speciation in Lycaenidae (Lepidoptera): Consequences of Novel Adaptations and Pleistocene Climate Changes. PhD thesis, Australian School of Environmental Studies, Griffith University, Qld, Australia.
Edwards ED, Newland J, Regan L (2001) ‘Lepidoptera: Hesperioidea, Papilionoidea. Zoological Catalogue of Australia. Vol. 31.6.’ (CSIRO Publishing: Melbourne, Vic., Australia)
Egan, SP, Ragland, GJ, Assour, L, Powell, THQ, Hood, GR, Emrich, S, Nosil, P, and Feder, JL (2015). Experimental evidence of genome-wide impact of ecological selection during early stages of speciation-with-gene-flow. Ecology Letters 18, 817–825.
| Experimental evidence of genome-wide impact of ecological selection during early stages of speciation-with-gene-flow.Crossref | GoogleScholarGoogle Scholar |
Eliot, JN (1973). The higher classification of the Lycaenidae (Lepidoptera): a tentative arrangement. Bulletin of the British Museum (Natural History) Entomology 28, 371–505.
| The higher classification of the Lycaenidae (Lepidoptera): a tentative arrangement.Crossref | GoogleScholarGoogle Scholar |
Feder, JL, Opp, SB, Wlazlo, B, Reynolds, K, Go, W, and Spisak, S (1994). Host fidelity is an effective premating barrier between sympatric races of the apple maggot fly. Proceedings of the National Academy of Sciences 91, 7990–7994.
| Host fidelity is an effective premating barrier between sympatric races of the apple maggot fly.Crossref | GoogleScholarGoogle Scholar |
Fitzpatrick, BM, Fordyce, JA, and Gavrilets, S (2008). What, if anything, is sympatric speciation? Journal of Evolutionary Biology 21, 1452–1459.
| What, if anything, is sympatric speciation?Crossref | GoogleScholarGoogle Scholar |
Forbes, AA, Hood, GR, and Feder, JL (2010). Geographic and ecological overlap of parasitoid wasps associated with the Rhagoletis pomonella (Diptera: Tephritidae) species complex. Annals of the Entomological Society of America 103, 908–915.
| Geographic and ecological overlap of parasitoid wasps associated with the Rhagoletis pomonella (Diptera: Tephritidae) species complex.Crossref | GoogleScholarGoogle Scholar |
Forbes, AA, Devine, SN, Hippee, AC, Tvedte, ES, Ward, AKG, Widmayer, HA, and Wilson, CJ (2017). Revisiting the particular role of host shifts in initiating insect speciation. Evolution 71, 1126–1137.
| Revisiting the particular role of host shifts in initiating insect speciation.Crossref | GoogleScholarGoogle Scholar |
Forest, F, Grenyer, R, Rouget, M, Davies, TJ, Cowling, RM, Faith, DP, Balmford, A, Manning, JC, Procheş, Ş, van der Bank, M, Reeves, G, Hedderson, TAJ, and Savolainen, V (2007). Preserving the evolutionary potential of floras in biodiversity hotspots. Nature 445, 757–760.
| Preserving the evolutionary potential of floras in biodiversity hotspots.Crossref | GoogleScholarGoogle Scholar |
Graham, AJ, Bollam, HH, and Williams, M (1996). An unusual temporally isolated population of ‘Neolucia agricola’ Waterhouse and Turner in Western Australia (Lepidoptera: Lycaenidae). Australian Entomologist 23, 111–114.
| An unusual temporally isolated population of ‘Neolucia agricola’ Waterhouse and Turner in Western Australia (Lepidoptera: Lycaenidae).Crossref | GoogleScholarGoogle Scholar |
Guindon, S, Dufayard, JF, Lefort, V, Anisimova, M, Hordijk, W, and Gascuel, O (2010). New Algorithms and Methods to Estimate Maximum-Likelihood Phylogenies: Assessing the Performance of PhyML 3.0. Systematic Biology 59, 307–321.
| New Algorithms and Methods to Estimate Maximum-Likelihood Phylogenies: Assessing the Performance of PhyML 3.0.Crossref | GoogleScholarGoogle Scholar |
Heath, A (1997). Myrmecophily and the male genitalia of African Lycaenidae: a preliminary discussion. Metamorphosis 8, 89–97.
Hebert, PDN, Penton, EH, Burns, JM, Janzen, DH, and Hallwachs, W (2004). Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences 101, 14812–14817.
| Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator.Crossref | GoogleScholarGoogle Scholar |
Helbig, AJ, Knox, AG, Parkin, DT, Sangster, G, and Collinson, M (2002). Guidelines for assigning species rank. Ibis 144, 518–525.
| Guidelines for assigning species rank.Crossref | GoogleScholarGoogle Scholar |
Hirowatari, T (1992). A generic classification of the Tribe Polyommatini of the Oriental and Australian regions (Lepidoptera, Lycaenidae, Polyommatinae). Bulletin of the University of Osaka Prefecture, Series B 44, 1–102.
Hopper, SD (1979). Biogeographical aspects of speciation in the southwest Australian flora. Annual Review of Ecology and Systematics 10, 399–422.
| Biogeographical aspects of speciation in the southwest Australian flora.Crossref | GoogleScholarGoogle Scholar |
Hopper, SD (2009). OCBIL theory: towards an integrated understanding of the evolution, ecology and conservation of biodiversity on old, climatically buffered, infertile landscapes. Plant and Soil 322, 49–86.
| OCBIL theory: towards an integrated understanding of the evolution, ecology and conservation of biodiversity on old, climatically buffered, infertile landscapes.Crossref | GoogleScholarGoogle Scholar |
Hopper, SD, and Gioia, P (2004). The Southwest Australian floristic region: evolution and conservation of a global hot spot of biodiversity. Annual Review of Ecology, Evolution, and Systematics 35, 623–650.
| The Southwest Australian floristic region: evolution and conservation of a global hot spot of biodiversity.Crossref | GoogleScholarGoogle Scholar |
Hurvich, CM, and Tsai, C-L (1989). Regression and time series model selection in small samples. Biometrika 76, 297–307.
| Regression and time series model selection in small samples.Crossref | GoogleScholarGoogle Scholar |
Jaenike, J (1990). Host specialization in phytophagous insects. Annual Review of Ecology and Systematics 21, 243–273.
| Host specialization in phytophagous insects.Crossref | GoogleScholarGoogle Scholar |
Jiggins, CD (2006). Sympatric speciation: why the controversy? Current Biology 16, R333–R334.
| Sympatric speciation: why the controversy?Crossref | GoogleScholarGoogle Scholar |
Kisdi, É, and Priklopil, T (2011). Evolutionary branching of a magic trait. Journal of Mathematical Biology 63, 361–397.
| Evolutionary branching of a magic trait.Crossref | GoogleScholarGoogle Scholar |
Klots AB (1970) Lepidoptera. In ‘Taxonomist’s Glossary of Genitalia in Insects’, 2nd edn. (Ed. SL Tuxen) pp. 115–130. (Munksgaard: Copenhagen, Denmark)
Korol, A, Rashkovetsky, E, Iliadi, K, Michalak, P, Ronin, Y, and Nevo, E (2000). Nonrandom mating in Drosophila melanogaster laboratory populations derived from closely adjacent ecologically contrasting slopes at ‘Evolution Canyon’. Proceedings of the National Academy of Sciences 97, 12637–12642.
| Nonrandom mating in Drosophila melanogaster laboratory populations derived from closely adjacent ecologically contrasting slopes at ‘Evolution Canyon’.Crossref | GoogleScholarGoogle Scholar |
Larsson, A (2014). AliView: a fast and lightweight alignment viewer and editor for large data sets. Bioinformatics 30, 3276–3278.
| AliView: a fast and lightweight alignment viewer and editor for large data sets.Crossref | GoogleScholarGoogle Scholar |
Lukhtanov, VA, Kandul, NP, Plotkin, JB, Dantchenko, AV, Haig, D, and Pierce, NE (2005). Reinforcement of pre-zygotic isolation and karyotype evolution in Agrodiaetus butterflies. Nature 436, 385–389.
| Reinforcement of pre-zygotic isolation and karyotype evolution in Agrodiaetus butterflies.Crossref | GoogleScholarGoogle Scholar |
Mallet, J, Meyer, A, Nosil, P, and Feder, JL (2009). Space, sympatry and speciation. Journal of Evolutionary Biology 22, 2332–2341.
| Space, sympatry and speciation.Crossref | GoogleScholarGoogle Scholar |
Marchese, C (2015). Biodiversity hotspots: a shortcut for a more complicated concept. Global Ecology and Conservation 3, 297–309.
| Biodiversity hotspots: a shortcut for a more complicated concept.Crossref | GoogleScholarGoogle Scholar |
Mayr E (1942) ‘Systematics and the Origin of Species.’ (Columbia University Press: New York, NY, USA)
Meier, R, Zhang, G, and Ali, F (2008). The use of mean instead of smallest interspecific distances exaggerates the size of the ‘barcoding gap’ and leads to misidentification. Systematic Biology 57, 809–813.
| The use of mean instead of smallest interspecific distances exaggerates the size of the ‘barcoding gap’ and leads to misidentification.Crossref | GoogleScholarGoogle Scholar |
Meyer, CP, and Paulay, G (2005). DNA barcoding: error rates based on comprehensive sampling. PLoS Biology 3, e422.
| DNA barcoding: error rates based on comprehensive sampling.Crossref | GoogleScholarGoogle Scholar |
Munday, PL, van Herwerden, L, and Dudgeon, CL (2004). Evidence for sympatric speciation by host shift in the sea. Current Biology 14, 1498–1504.
| Evidence for sympatric speciation by host shift in the sea.Crossref | GoogleScholarGoogle Scholar |
Múrias dos Santos, A, Cabezas, MP, Tavares, AI, Xavier, R, and Branco, M (2016). tcsBU: a tool to extend TCS network layout and visualization. Bioinformatics 32, 627–628.
| tcsBU: a tool to extend TCS network layout and visualization.Crossref | GoogleScholarGoogle Scholar |
Nosil, P, Crespi, BJ, and Sandoval, CP (2002). Host-plant adaptation drives the parallel evolution of reproductive isolation. Nature 417, 440–443.
| Host-plant adaptation drives the parallel evolution of reproductive isolation.Crossref | GoogleScholarGoogle Scholar |
Ogden, R, and Thorpe, RS (2002). Molecular evidence for ecological speciation in tropical habitats. Proceedings of the National Academy of Sciences 99, 13612–13615.
| Molecular evidence for ecological speciation in tropical habitats.Crossref | GoogleScholarGoogle Scholar |
Orr AG, Kitching RL (2010) ‘The Butterflies of Australia.’ (Allen and Unwin: Sydney, NSW, Australia)
Peters JV (1971) ‘A catalogue of the type specimens of the Hesperioidea and Papilionoidea (Lepidoptera) in the Australian Museum.’ (Australian Entomological Press: Sydney, NSW, Australia)
Ragland, GJ, Egan, SP, Feder, JL, Berlocher, SH, and Hahn, DA (2011). Developmental trajectories of gene expression reveal candidates for diapause termination: a key life-history transition in the apple maggot fly Rhagoletis pomonella. Journal of Experimental Biology 214, 3948–3960.
| Developmental trajectories of gene expression reveal candidates for diapause termination: a key life-history transition in the apple maggot fly Rhagoletis pomonella.Crossref | GoogleScholarGoogle Scholar |
Rix, MG, Edwards, DL, Byrne, M, Harvey, MS, Joseph, L, and Roberts, JD (2014). Biogeography and speciation of terrestrial fauna in the south-western Australian biodiversity hotspot. Biological Reviews 90, 762–793.
| Biogeography and speciation of terrestrial fauna in the south-western Australian biodiversity hotspot.Crossref | GoogleScholarGoogle Scholar |
Rosser, N, Kozak, KM, Phillimore, AB, and Mallet, J (2015). Extensive range overlap between heliconiine sister species: evidence for sympatric speciation in butterflies? BMC Evolutionary Biology 15, 125.
| Extensive range overlap between heliconiine sister species: evidence for sympatric speciation in butterflies?Crossref | GoogleScholarGoogle Scholar |
Rundle, HD, and Nosil, P (2005). Ecological speciation. Ecology Letters 8, 336–352.
| Ecological speciation.Crossref | GoogleScholarGoogle Scholar |
Santos, H, Burban, C, Rousselet, J, Rossi, J-P, Branco, M, and Kerdelhué, C (2011). Incipient allochronic speciation in the pine processionary moth (Thaumetopoea pityocampa, Lepidoptera, Notodontidae). Journal of Evolutionary Biology 24, 146–158.
| Incipient allochronic speciation in the pine processionary moth (Thaumetopoea pityocampa, Lepidoptera, Notodontidae).Crossref | GoogleScholarGoogle Scholar |
Schluter, D (2009). Evidence for ecological speciation and its alternative. Science 323, 737–741.
| Evidence for ecological speciation and its alternative.Crossref | GoogleScholarGoogle Scholar |
Schmidt, DJ, Grund, R, Williams, MR, and Hughes, JM (2014). Australian parasitic Ogyris butterflies: east-west divergence of highly specialized relicts. Biological Journal of the Linnean Society 111, 473–484.
| Australian parasitic Ogyris butterflies: east-west divergence of highly specialized relicts.Crossref | GoogleScholarGoogle Scholar |
Servedio, MR, Doorn, GSV, Kopp, M, Frame, AM, and Nosil, P (2011). Magic traits in speciation: ‘magic’ but not rare? Trends in Ecology & Evolution 26, 389–397.
| Magic traits in speciation: ‘magic’ but not rare?Crossref | GoogleScholarGoogle Scholar |
Smadja, CM, and Butlin, RK (2011). A framework for comparing processes of speciation in the presence of gene flow. Molecular Ecology 20, 5123–5140.
| A framework for comparing processes of speciation in the presence of gene flow.Crossref | GoogleScholarGoogle Scholar |
Stireman, JO, Nason, JD, and Heard, SB (2005). Host-associated genetic differentiation in phytophagous insects: general phenomenon or isolated exceptions? Evidence from a goldenrod-insect community. Evolution 59, 2573–2587.
| Host-associated genetic differentiation in phytophagous insects: general phenomenon or isolated exceptions? Evidence from a goldenrod-insect community.Crossref | GoogleScholarGoogle Scholar |
Su, YN (2016). A simple and quick method of displaying liquid-preserved morphological structures for microphotography. Zootaxa 4208, 592–593.
| A simple and quick method of displaying liquid-preserved morphological structures for microphotography.Crossref | GoogleScholarGoogle Scholar |
Swofford DL (2002) PAUP* ‘Phylogenetic Analysis Using Parsimony (* and Other Methods)’, ver. 4. (Sinauer Associates: Sunderland, MA, USA)
Sword, GA, Joern, A, and Senior, LB (2005). Host plant-associated genetic differentiation in the snakeweed grasshopper, Hesperotettix viridis (Orthoptera: Acrididae). Molecular Ecology 14, 2197–2205.
| Host plant-associated genetic differentiation in the snakeweed grasshopper, Hesperotettix viridis (Orthoptera: Acrididae).Crossref | GoogleScholarGoogle Scholar |
Talavera, G, Lukhtanov, V, Pierce, NE, and Vila, R (2021). DNA barcodes combined with multi-locus data of representative taxa can generate reliable higher-level phylogenies. Systematic Biology 71, 382–395.
| DNA barcodes combined with multi-locus data of representative taxa can generate reliable higher-level phylogenies.Crossref | GoogleScholarGoogle Scholar |
Taylor, RS, and Friesen, VL (2017). The role of allochrony in speciation. Molecular Ecology 26, 3330–3342.
| The role of allochrony in speciation.Crossref | GoogleScholarGoogle Scholar |
Waterhouse GA, Lyell G (1914) ‘The Butterflies of Australia. A Monograph of the Australian Rhopalocera.’ (Angus and Robertson: Sydney, NSW, Australia)
Western Australian Herbarium (2023a) Daviesia angulata Lindl. In ‘Florabase’. (Department of Biodiversity, Conservation and Attractions) Available at https://florabase.dpaw.wa.gov.au/browse/profile/3793
Western Australian Herbarium (2023b) Daviesia daphnoides Meisn. In ‘Florabase’. (Department of Biodiversity, Conservation and Attractions) Available at https://florabase.dpaw.wa.gov.au/browse/profile/3803
Western Australian Herbarium (2023c) Daviesia chapmanii Crisp. In ‘Florabase’. (Department of Biodiversity, Conservation and Attractions) Available at https://florabase.dpaw.wa.gov.au/browse/profile/14199
Williams, MR, Williams, AAE, Lundstrom, TD, Hay, RW, Bollam, HH, and Graham, AJ (1995). Range extensions and natural history notes for some Western Australian butterflies. Victorian Entomologist 25, 94–96.
Williams AAE, Williams MR, Powell R, Walker G (2012) ‘Rare butterflies of the south-west.’ (Department of Environment and Conservation: Perth, WA, Australia)
Wood TK (1993) Speciation of the Enchenopa binotata complex (Insecta: Homoptera; Membracidae). In ‘Evolutionary Patterns and Processes’. (Eds DR Lees, D Edwards) pp. 299–317. (The Linnean Society of London and Harcourt Brace Co: London, UK)
