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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    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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    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.
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    New Caledonian rovers and the historical biogeography of a hyper-diverse endemic lineage of South Pacific leaf beetles
    (John Wiley & Sons Ltd on behalf of Royal Entomological Society, 2024-10-01) Platania L; Cardoso A; Anderson M; Fikáček M; Gauthier J; Hendrich L; Mille C; Morii Y; Reid CAM; Seidel M; Morgan-Richards M; Trewick SA; Toussaint EFA; Gómez-Zurita J
    South Pacific archipelagos are central in the biogeographic debate on the relative importance of vicariance and dispersal in shaping the distribution of species. However, each taxonomic group was subject to different processes and histories, and here, we reveal the historical biogeography of the diverse Eumolpinae leaf beetles, widely distributed in the region. Extensive taxon sampling focusing on South Pacific Eumolpinae was used to infer the first molecular phylogeny of the group using three single-copy protein-coding nuclear and two mitochondrial markers. Upon assessing the clade of interest for lineage-specific variation in substitution rates, the age of the most recent common ancestors was estimated using out-group calibration and multi-gamma site models (MGSMs). Biogeographic analyses used standard event-based inferences also incorporating phylogenetic uncertainty. Zealandian Eumolpinae are monophyletic and appear to have split from their global relatives in the transition from the Cretaceous to the Paleogene. Variation in the rates of molecular evolution affected the in-group stem branch, with a significant drop in the substitution rate, and the MGSM correction recovered the crown age of Zealandian Eumolpinae during the Late Eocene–Oligocene transition. Biogeographic inference resolved the origin of the radiation in New Caledonia, favouring a null model without island age constraints, and repeated dispersal events to the other islands, including three independent but synchronous colonisations of New Zealand during the Miocene. New Caledonia, with a highly diverse Eumolpinae fauna of uncertain origin, acted as a hub and pump of biodiversity of these beetles in the entire South Pacific region, sending migrants to other islands through long-distance dispersal with lineages establishing when land became available.
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    Mwhitiwhiti Aotearoa: Phylogeny and synonymy of the silent alpine grasshopper radiation of New Zealand (Orthoptera: Acrididae)
    (Magnolia Press, 2023-12-11) Trewick SA; Koot EM; Morgan-Richards M
    Aotearoa New Zealand has a fauna of endemic alpine grasshoppers, consisting of thirteen species distributed among four genera. The many re-classifications of species within this group and the presence of species complexes highlight the uncertainty that surrounds relationships within and between these genera. High-throughput Next Generation Sequencing was used to assemble the complete mitochondrial genomes, 45S ribosomal cassettes and histone sequences of New Zealands four endemic alpine genera: Alpinacris, Brachaspis, Paprides and Sigaus. Phylogenetic analysis of these molecular datasets, as individual genes, partitions and combinations returned a consistent topology that is incompatible with the current classification. The genera Sigaus, Alpinacris, and Paprides all exhibit paraphyly. A consideration of the pronotum, epiphallus and terminalia of adult specimens reveals species-specific differences, but fails to provide compelling evidence for species groups justifying distinct genera. In combination with phylogenetic, morphological and spatial evidence we propose a simplified taxonomy consisting of a single genus for the mwhitiwhiti Aotearoa species radiation.
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    Standard metabolic rate variation among New Zealand Orthoptera
    (Elsevier B V, Amsterdam, 2024) Bulgarella M; Haywood J; Dowle EJ; Morgan-Richards M; Trewick SA
    Standard metabolic rates (SMR) of ectotherms reflect the energetic cost of self-maintenance and thus provide important information about life-history strategies of organisms. We examined variation in SMR among fifteen species of New Zealand orthopteran. These species represent a heterogeneous group with a wide geographic distribution, differing morphologies and life histories. Gathering original data on morphological and physiological traits of individual species is a first step towards understanding existing variability. Individual metabolic rates of ectotherms are one of the first traits to respond to climate change. Baseline SMR datasets are valuable for modeling current species distributions and their responses to a changing climate. At higher latitudes, the average environmental temperature decreases. The pattern that cold-adapted ectotherms display higher SMR at colder temperatures and greater thermal sensitivity to compensate for lower temperatures and the shorter growing and reproductive seasons is predicted from the metabolic cold adaptation (MCA) hypothesis. We predict higher SMR for the orthopteran species found at higher latitudes. We further compared the index of thermal sensitivity Q10 per species. We used closed-system respirometry to measure SMR, at two test temperatures (4 °C and 14 °C), for the fifteen species acclimated to the same conditions. As expected, we found significant differences in SMR among species. The rate of oxygen consumption was positively correlated with body mass. Our findings do not support the MCA hypothesis. In fact, we found evidence of co-gradient variation in SMR, whereby insects from higher elevations and latitudes presented lower SMR. We discuss our findings in relation to life histories and ecology of each species. The novel physiological data presented will aid in understanding potential responses of these unusual species to changing climatic conditions in Aotearoa/New Zealand.
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    No barrier to fertilisation when different sexual populations of the mānuka stick insect are crossed
    (Taylor and Francis Group, 2023-01-01) Morgan-Richards M
    The mānuka stick insect Clitarchus hookeri (White) is facultatively parthenogenetic, but females from sexual populations that have mated with males from their own population do not produce any offspring via asexual reproduction. In contrast, females from parthenogenetic populations of C. hookeri mate with males (in captivity) but show a partial barrier to fertilisation with more than 90% of their offspring resulting from asexual reproduction post mating. Captive crossing experiments with parthenogenetic females require the mating of individuals from different populations (sexual and parthenogenetic), thus potential intraspecific differences bring a confounding element to these experiments. Experiments mating sexual females with males from different sexual populations were undertaken to determine whether offspring resulting from such a cross would be the result of sexual or parthenogenetic reproduction. Virgin females and males were collected from two sexual populations known to represent distinct genetic lineages (Waikato and Whanganui). Eleven adult females were caged with non-local males and eggs collected post-mating. Approximately equal numbers of sons and daughters hatched (168 female; 210 male) suggesting all offspring were the result of sexual reproduction. In these intraspecific crosses no barriers to fertilisation were detected, suggesting that in the absence of males the decay of some sexual trait in Phasmids can occur in fewer than 100 generations.