Journal Articles

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    Cretaceous Connections Among Camel Cricket Lineages in the Himalaya Revealed Through Fossil-Calibrated Mitogenomic Phylogenetics
    (MDPI (Basel, Switzerland), 2025-07-01) Dorji C; Morgan-Richards M; Trewick SA; Heller K-G
    The nocturnal, flightless camel crickets (Rhaphidophoridae) have a global distribution and are believed to have originated prior to the breakup of Pangea. We investigated the phylogeny and the timing of the radiation of East Asian species with mitogenomic data. Initially we analyzed a large taxon dataset (n = 117) using available partial mitochondrial and nuclear DNA sequences to confirm the monophyly of subfamilies and current taxonomy. Our findings support the monophyly of each genus within the subfamily Aemodogryllinae, with a minor inconsistency between taxonomy and phylogeny resolved by resurrection of the genus Gymnaeta Adelung. Fossil-calibrated molecular clock analysis used 11,124 bp alignment of 13 complete mitochondrial protein-coding genes for 20 species of Rhaphidophoridae, with a focus on the neglected Rhaphidophorinae and Aemodogryllinae lineages. Divergence time estimates suggest that the most recent common ancestor of the family lived during the Early Jurassic (189 Mya ± 23 Mya) before Pangea broke into the supercontinents or possibly during the early stage of breakup when Gondwana and Laurasia were still connected by land. The two subfamilies, Rhaphidophorinae and Aemodogryllinae, that overlap in Asia are estimated to have diverged 138 Mya ± 17 Mya, well before the Late Cretaceous northern connection between America and Asia (the Bering Land Bridge). Thus, our extended sampling of species from East Asia and Oceania refutes the importance of continental drift in the evolution of this wingless orthopteran family.
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    Integrative taxonomic revision of the Australian cave cricket Speleotettix Chopard, 1944 (Orthoptera: Rhaphidophoridae): New species, distribution and conservation implications
    (John Wiley and Sons Australia, Ltd on behalf of Australian Entomological Society, 2025-08-01) Beasley-Hall PG; Trewick SA; Hedges BA; Cooper SJB; Reed EH; Austin AD
    Cave crickets, also called wētā, are an important component of subterranean realms globally, but the true diversity of the group is poorly known. Speleotettix Chopard, 1944 contains two species from southeast Australia, Speleotettix tindalei Chopard, 1944 and Speleotettix chopardi (Karny, 1935). However, the initial description of the genus was poorly characterised, and its taxonomy has remained unclear since. Here, we take an integrative molecular and morphological approach to redescribe Speleotettix and describe three new species: Speleotettix aolae Beasley-Hall, sp. nov. from Victoria and Speleotettix binoomea Beasley-Hall, sp. nov. from New South Wales, both found in limestone caves, and Speleotettix palaga Beasley-Hall, sp. nov. from mineshafts in Victoria. To provide consistency in future work on the group, we also redescribe S. tindalei and treat S. chopardi as a nomen dubium. Finally, we transfer the threatened species Cavernotettix craggiensis Richards, 1974, an island species from Tasmania's Bass Strait, into the genus as Speleotettix craggiensis (Richards, 1974) comb. nov. In so doing, we increase the number of described rhaphidophorid species in Australia to 27 and significantly expand the distribution of Speleotettix, making it the most widespread of the Australian genera currently known. As all members of Speleotettix are short-range endemics at risk of decline, these findings have implications for their future conservation management.
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    A new species of Celatoblatta cockroach that coincides with the distribution of kauri forests in New Zealand
    (Taylor and Francis Group on behalf of the Royal Society of New Zealand, 2025-07-16) Morgan-Richards M; Trewick SA
    A new species of Blattidae cockroach is described from northern Aotearoa New Zealand. Celatoblatta kauri sp. nov. can be distinguished from similar conspecific species by its distinctive facial markings. The recorded distribution of Celatoblatta kauri sp. nov. matches that of the historic distribution of kauri forest (Agathis australis) in northern New Zealand. Zoobank LSID: urn:lsid:zoobank.org:pub:93794424-1A7B-4CCA-A7A1-A303BCE8EACA.
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    Molecular phylogenetics illuminates the evolutionary history and hidden diversity of Australian cave crick ets (Orthoptera: Rhaphidophoridae)
    (John Wiley and Sons Ltd on behalf of Royal Entomological Society., 2025-06-23) Beasley-Hall PG; Trewick SA; Eberhard SM; Zwick A; Reed EH; Cooper SJB; Austin AD; Blaimer B
    Cave crickets (Orthoptera: Rhaphidophoridae) are a globally distributed group of insects found in dark, humid microhabitats including natural caves, alpine scree, and forest litter. Ten extant subfamilies are currently recognised, of which Macropathinae, which comprises the entirety of the fauna in South America, South Africa, Australia, and New Zealand, is thought to be the most ancient. New Zealand comprises high phylogenetic diversity of Rhaphidophoridae throughout its mesic zone, with most species occurring above ground. In contrast, the Australian fauna is poorly known and contains an apparently greater relative proportion of species utilising caves as refugia. A robust phylogenetic framework is needed to underpin future taxonomic work on the group and uncover potentially contrasting patterns of taxonomic diversity. Here, we performed fossil-calibrated phylogenetic analysis using whole mitochondrial genomes and nuclear markers to reconstruct the evolutionary history of Macropathinae with a focus on the Australian fauna. By dramatically increasing taxon sampling relative to past studies, we recovered the Australian fauna as rampantly polyphyletic, with the remaining Macropathinae nested among six distinct Australian lineages. Deep divergences between major clades imply additional Australian lineages remain undetected, either due to extinction or sampling bias, and have likely confounded past biogeographic signal. We inferred the radiation of Macropathinae began during the Lower Cretaceous prior to the fragmentation of Gondwana with a potential Pangaean origin for Rhaphidophoridae. Finally, we found evidence for several undescribed species and genera of Australian Macropathinae, all of which qualify as short-range endemics, and discuss the conservation implications of these restricted distributions.
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    The impact of experimental warming on bumblebees: Higher temperatures induce behavioral changes in Bombus terrestris queens
    (Schweizerbart Science Publishers, 2025-05-22) Kim HS; Yoon HJ; Kim BY; Lee KY; Trewick SA; Lee KS; Jin BR
    Bumblebees (Bombus terrestris) are important pollinators in most temperate and boreal ecosystems, but these regions are increasingly affected by heatwaves that are a symptom of anthropogenic climate change. Bumblebee queens may be particularly vulnerable to elevated temperatures during their solitary period following overwintering. Using an experimental system with numerous replicates, we investigated the oviposition behavior of overwintered spring queens, the resulting colony characteristics, and the mating behavior of newly emerged fall queens at three different temperatures: optimal (27 °C; C27), moderately high (32 °C; C32), and high (35 °C; C35). Spring queens exposed to higher temperatures exhibited unusual behavior, with spring queens in the C35 group more frequently digging up and destroying egg cells, ultimately leading to colony failure. Compared to C27, spring queens in the C32 treatment displayed more frequent ovipo sition, a higher colony formation rate, a higher number of offspring per colony, a shorter colony development period, and lower body sizes in their adult offspring. These results indicate that elevated temperatures lead to behavioral changes in spring queens and affect the characteristics of resulting colonies. Fall queens exhibited mating avoidance behavior at higher temperatures, resulting in a lower mating rate. Our results demonstrate that bumblebee queens respond to thermal condi tions that can lead to colony failure when thermal limits are exceeded. In nature, behavioral changes induced in bumblebee queens exposed to persistent heatwaves during the solitary period may be seriously deleterious to bumblebee populations in many regions.
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    Origins and diversity of invasive brushtail possums (Trichosurus vulpecula) in New Zealand surveyed with mtDNA haplotype and nuclear microsatellite data
    (Taylor and Francis Group on behalf of the Royal Society of New Zealand, 2025-05-07) Pattabiraman N; Morgan-Richards M; Trewick SA; Su J
    The brushtail possum Trichosurus vulpecula is native to Australia where six subspecies exist in distinct regions. A composite invasive population is established in Aotearoa New Zealand, which has since been subject to localised bouts of culling. We surveyed population genetic structure across New Zealand to identify the scale of genetic diversity introduced to New Zealand and the resulting pattern of admixture. Australian brushtail possums have high mitochondrial diversity (17%) and prominent spatial structure. Thirty-eight haplotypes among 25 New Zealand population samples (n = 465) were closely related to 45 haplotypes sampled from Victoria and Tasmania in Australia (n = 120), but just one was shared. High haplotype diversity is consistent with multiple successful introductions and rapid population expansion in New Zealand. Nuclear diversity of microsatellite loci screened in 18 New Zealand population samples (n = 374) comprised five genotypic clusters (K = 5), but these groups did not correlate with geography. An overall signal of genetic partitioning within the invasive population suggests limited mixing but ongoing management towards eradication will influence patterns of population recovery, migration and evolution of traits including toxin resistance. We consider the implications of mixed ancestry of the invasive population in terms of variation in toxin tolerance detected in New Zealand.
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    Revision of the New Zealand cave wētā genus Isoplectron Hutton (Orthoptera: Rhaphidophoridae), with synonymy of Petrotettix Richards and Setascutum Richards, and the description of a new genus
    (EJT Consortium, 2024-12-10) Hegg D; Morgan-Richards M; Trewick SA; Robillard T; Tan MK; Pepe F
    The New Zealand cave wētā genus Isoplectron was first described by Hutton (1896) and included two species, Isoplectron armatum and I. calcaratum. Two more species, I. cochleatum and I. aciculatum, were described by Karny in 1935 and in 1937, respectively. The species I. cochleatum was later moved to the genus Pharmacus Pictet & de Saussure, 1893. Here we clarify the status and appearance of all known species of Isoplectron, and of other closely related cave wētā. Based on morphology and mtDNA sequences we determine that the genera Petrotettix Richards, 1972 and Setascutum Richards, 1972, belong in Isoplectron, and are thus synonymised with the latter. We also infer that Isoplectron aciculatum Karny, 1937 does not hold species rank, and that it should be considered a subspecies of Isoplectron armatum instead. The male of Isoplectron pallidum (Richards, 1972) comb. nov. and the female of Isoplectron armatum aciculatum Karny, 1937 are described here for the first time. Using molecular data, we also establish that two new species that are morphologically similar to Isoplectron are in fact more closely related to Pharmacus. These species are assigned to a new genus, Praecantrix gen. nov. We describe seven new species and two new subspecies of New Zealand rhaphidophorids belonging to the genera Isoplectron Hutton, 1896 and Praecantrix: Isoplectron bicolor sp. nov., I. maculatum sp. nov., I. virgatum sp. nov., I. ferratum sp. nov., I. parallelum sp. nov., Praecantrix silvatica silvatica gen., sp. et subsp. nov., P. silvatica lutea gen., sp. et subsp. nov. and P. saxicola gen. et sp. nov. Species demoted to subspecies rank: Isoplectron aciculatum Karny, 1937 becomes Isoplectron armatum aciculatum Karny, 1937. New combinations: Setascutum pallidum Richards, 1972 becomes Isoplectron pallidum (Richards, 1972) comb. nov.; Petrotettix serratus Richards, 1972 becomes Isoplectron serratum (Richards, 1972) comb. nov. New synonyms: Isoplectron calcaratum Hutton, 1896 = Isoplectron armatum Hutton, 1896 syn. nov.; Setascutum ohauensis Richards, 1972 = Isoplectron armatum Hutton, 1896 syn. nov.; Petrotettix spinosus Richards, 1972 = Isoplectron serratum (Richards, 1972) syn. nov.; Petrotettix cupolaensis Richards, 1972 = Isoplectron serratum (Richards, 1972) syn. nov.; Petrotettix nigripes Richards, 1972 = Isoplectron serratum (Richards, 1972) syn. nov.
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    Comparison of growth, relative abundance, and diet of three sympatric Hemiandrus ground wētā (Orthoptera, Anostostomatidae) in a New Zealand Forest
    (Pensoft Publishers for the Orthopterists' Society, 2025-01) Chikwature N; Morgan-Richards M; Vereijssen J; Trewick SA
    Ecology is an essential discipline for understanding the biology and behavior of organisms. This study increased knowledge of three sympatric species of ground wētā (Hemiandrus spp.). Hemiandrus ground wētā are nocturnal Ensifera that live in burrows during the day, and for these reasons, there is limited knowledge of their activity, development, and diet. We examined the diet of Hemiandrus electra, Hemiandrus ‘disparalis’, and Hemiandrus nox by examining the crop contents of specimens caught in malaise traps set in a native forest (St Arnaud) over seven months during two spring-summer-autumn seasons (2004/2005 and 2005/2006). The three species investigated varied in the plant and invertebrate fragment proportions in their diets. Hemiandrus electra and H. ‘disparalis’ were predominantly herbivores, while H. nox was primarily carnivorous, although plant matter constituted 20% of its diet. We identified the species and sex of 966 Hemiandrus wētā that were intercepted by the same malaise traps. Our results showed that H. electra was the most abundant species, with 701 individuals, while 157 and 109 were identified as H. ‘disparalis’ and H. nox, respectively. Surprisingly, the species with maternal care (H. electra) was the only one of the three sympatric Hemiandrus to have an even sex ratio; for the other two species, more males were caught in the traps. Using hind leg dimensions, we categorized each female H. electra specimen using naïve Gaussian mixture model clustering, which identified five size clusters (putatively corresponding to instars). Based on the month of collection and the growth category, we observed no seasonality in the development of this species of ground wētā, as almost all instars were found in each month of the sampling period. This study found no evidence that these nocturnal forest species synchronize their growth or reproduction with seasons.
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    Forecasting Range Shifts in Terrestrial Alpine Insects Under Global Warming
    (John Wiley and Sons Incorporated, Hoboken, New Jersey, 2025-01) Meza-Joya FL; Morgan-Richards M; Trewick SA
    Anthropogenic planetary heating is disrupting global alpine systems, but our ability to empirically measure and predict responses in alpine species distributions is impaired by a lack of comprehensive data and technical limitations. We conducted a comprehensive, semi-quantitative review of empirical studies on contemporary range shifts in alpine insects driven by climate heating, drawing attention to methodological issues and potential biotic and abiotic factors influencing variation in responses. We highlight case studies showing how range dynamics may affect standing genetic variation and adaptive potential and discuss how data integration frameworks can improve forecasts. Although biotic and abiotic factors influence individual species responses, most alpine insects studied so far are shifting to higher elevations. Upslope shifts are often accompanied by range contractions that are expected to diminish species genetic variation and adaptive potential, increasing extinction risk. Endemic species on islands are predicted to be especially vulnerable. Inferences drawn from the responses of alpine insects, also have relevance to species in other montane habitats. Correlative niche modelling is a keystone tool to predict range responses to planetary heating, but its limited ability to consider biological processes underpinning species' responses complicates interpretation. Alpine insects exhibit some potential to respond to rising temperatures via genetic change or phenotypic plasticity. Thus, future efforts should incorporate biological processes by using flexible hybrid niche modelling approaches to enhance the biological realism of predictions. Boosting scientific capability to envisage the future of alpine environments and their associated biota is imperative given that the speed and intensity of heating on high-mountain ecosystems can surpass our ability to collect the empirical data required to guide effective conservation planning and management decisions.