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Article

Wētā Aotearoa—Polyphyly of the New Zealand Anostostomatidae (Insecta: Orthoptera)

by
Steven A. Trewick
1,*,†,
Briar L. Taylor-Smith
2 and
Mary Morgan-Richards
1
1
Wildlife and Ecology, School of Food Technology and Natural Sciences, Massey University, Private Bag 11-222, Palmerston North 4442, New Zealand
2
Ecology Group, Institute of Agriculture and Environment, Massey University, Private Bag 11-222, Palmerston North 4442, New Zealand
*
Author to whom correspondence should be addressed.
†
zoobank.org:pub:65583178-0930-47F3-A9A8-C887572BC0A8.
Insects 2024, 15(10), 787; https://doi.org/10.3390/insects15100787
Submission received: 12 September 2024 / Revised: 1 October 2024 / Accepted: 2 October 2024 / Published: 9 October 2024
(This article belongs to the Section Insect Systematics, Phylogeny and Evolution)
Browse Figures
Versions Notes

Abstract

:

Simple Summary

Large crickets in the family Anostostomatidae, which are locally known as wētā, are a prominent feature of New Zealand endemic diversity and ecology. However, their systematics are poorly resolved, which reflects the wider uncertainty about the treatment on this orthopteran family. We examined the relationships among the New Zealand genera with representatives of the fauna from nearby Australia and New Caledonia by using a DNA dataset. We found that the New Zealand genera comprised four distinct lineages that were each more closely related to taxa outside New Zealand. We also found that the most speciose genus in New Zealand comprised two paraphyletic taxa.

Abstract

The Anostostomatidae of Aotearoa New Zealand are well-characterized at the genus and species level, but the higher-level systematics of the family as a whole remain poorly resolved. We tested the hypothesis that the New Zealand anaostostomatid fauna consists of a single monophyletic group consistent with a single common ancestor. For phylogenetic analysis, we sampled the genera in Aotearoa New Zealand as well as representatives of the family from Australia and New Caledonia. Maximum likelihood analyses including topological comparison statistics with a DNA alignment of thirteen mitochondrial and four nuclear protein coding genes rejected the monophyly of lineages in New Zealand. We found phylogenetic support for four separate New Zealand lineages; three with their closest relatives in Australia and one in New Caledonia. The New Zealand genus Hemiandrus is paraphyletic and the establishment of a morphologically distinct genus is justified. We determined that six of the valid species previously placed in Hemiandrus form a distinct clade that we designated here as Anderus gen. nov. The putative Hemiandrus that we sampled from Australia was sister to neither of the New Zealand lineages.

1. Introduction

The Anostostomatidae (Orthoptera) comprises taxa with a predominantly Southern Hemisphere distribution [1], and about 302 valid species [2]. The majority of species are flightless nocturnal predators or scavengers that hunt on the ground and burrow in soil to conceal themselves during the day. The family is richly represented in Central and South Africa, Australasia, Central and South America, and is also found in North Africa through Asia including the Himalaya, India, and China. Species of Anostostomatidae are also known in Sri Lanka and on islands near Taiwan and south of Japan. Despite considering many genera of this family insufficiently studied, Gorochov (2021) [3] suggested two subfamilies with Anostostomatinae divided into nine tribes, and Lezininae not divided. The placement of anostostomatid genera within tribes must be considered preliminary, and additional phylogenetic studies are required to understand the deeper evolutionary relationships within this widespread family.
In Aotearoa New Zealand, the family is represented by >40 species (Figure 1) in four informal groups of wētā: tree (Hemideina Walker), giant (Deinacrida White), ground (Hemiandrus Ander), and tusked (Anisoura Ander, Motuweta Johns). All are wingless. Hemideina and Deinacrida comprise most of the species of the small subfamily Deinacridinae [1], or all of the tribe Deinacridini [3], which are distinguished by stridulatory ridges, the musculature of their hind femura, and their predominantly herbivorous diet. These two genera are closely allied, and finding support for the reciprocal monophyly of the species within each of them has proven difficult [4,5,6,7]. Tree and giant wētā will scavenge invertebrate food but they are distinctive among anostostomatids in that they are primarily arboreal herbivores that feed on the foliage, flowers, and fruit of trees and shrubs [8,9]. This contrasts with, for example, the anostostostomatid fauna of Australia (king crickets), most of which forage on the forest floor, eating decaying material or are predatory [10].
The New Zealand ground wētā (Hemiandrus) and tusked wētā (Anisoura, Motuweta) have a predominantly predatory diet and burrowing habit (with one exception) typical of Anostostomatidae elsewhere [11,12] (Figure 2). Hemiandrus and Motuweta were placed by Johns (1997) in the tribe Anostostomatini, along with 13 other genera from Australia, southern Africa, Madagascar, and South America [1]. Reappraisal a few years later left Motuweta in Anostostomatini, with just seven genera from Australia, New Caledonia, and southern Africa [3,13,14]. The tribal affinity of several genera was considered unclear [13], and the taxonomic uncertainty remains with Hemiandrus and Carcinopsis, among others in the subfamily Anostostomatinae but not assigned to a tribe [2]. Phylogenetic analyses based on short DNA sequences suggest the New Zealand tusked wētā (Anisoura, Motuweta) are monophyletic but not closely related to the New Zealand Hemiandrus [15,16]. The three tusked wētā species are united by the presence in adult males of prominent curved tusks protruding from the mandibles (Figure 2b) [17,18]. Salmon (1950) [19] placed the smallest (~20 mm long), and at that time only species, with the ground wētā as Hemiandrus monstrosus. However, unlike New Zealand Hemiandrus, the tusked wētā have auditory pits (tympana) on their fore tibiae. Johns (1997) [1] revised the name of the small tusked wētā, recognizing the precedence of Anisoura nicobarica Ander 1932 and placed it in subfamily Deinacridinae (i.e., with the tree and giant wētā). The name of this species appears to be the result of the mislabeling of the type specimens, as it does not occur in the Nicobar Islands in the Bay of Bengal.
In New Zealand, there are nineteen valid species of Hemiandrus and about six undescribed species in the grey literature referred to by unofficial tag-names [20,21]. In addition, there are undescribed species in Australia that may belong in the genus Hemiandrus [1,15]. Evolutionary relationships among the New Zealand anostostomatid fauna remain unclear, although, in agreement with the current taxonomy with tribal divisions [3], there is evidence that they are not a monophyletic group with respect to taxa outside New Zealand [15,22]. Although previous analyses involved short DNA sequences [5,6,15], these data and morphological examination [21] suggest polyphyly involving lineages in Australia and New Caledonia. Here, we specifically tested the hypothesis that the flightless wētā Aotearoa is a monophyletic group. This includes the possibility that Hemiandrus Ander (ground wētā), as currently applied, comprises two distinct lineages consistent with the resurrection of Zealandosandrus Salmon. We analyzed mitochondrial DNA genomes and nuclear genes from specimens representing the three prominent groups of New Zealand wētā with examples of Anostostomatidae from further afield to resolve these relationships. The current taxonomy is unclear about the tribal affiliations of Hemiandrus, so we included representatives of Anabropsini from Australia and Anostostomatini from New Caledonia as well as a species that represented the Australian putative Hemiandrus lineage. Additional sampling of Hemiandrus diversity in New Zealand was used for morphological examination and COI mtDNA sequencing to clarify membership to the two distinct clades we identified in the course of this work.

2. Materials and Methods

We sampled representatives of Anostostomatidae from Australia, New Zealand, and New Caledonia, seeking to assess the monophyly of the endemic New Zealand taxa (Table 1, Figure 1 and Figure 2). We included representation of New Zealand tusked (Motuweta), tree (Hemideina), giant (Deinacrida), and ground (Hemiandrus) wētā.
We used a skim-sequencing high-throughput next-generation sequencing approach [32,33,34] to generate DNA data for our samples, targeting the whole mitochondrial genome and nuclear histones. Insect DNA was extracted using a high salt method [6,35] and quantified using Qubit fluorometry (Life Technologies, Thermo Fisher Scientific Inc., Waltham, MA, USA). Genomic DNA samples were paired-end sequenced with high-throughput sequencing on an Illumina HiSeq 2500 (either BGI Genomics, Tai Po, Hong Kong or Macrogen Inc., Seoul, Republic of Korea) following fragmentation and indexing using the Illumina TruSeq Nano DNA Kit. The resulting 100 or 150 bp paired-end reads were filtered and edited to remove the sample barcodes and assembled in Geneious v9.1.4 [36].
Mitochondrial genomes were obtained from each specimen using an iterative reference mapping approach, starting with available short sequence data. Paired reads were iteratively mapped to the reference sequence in Geneious, generating a novel consensus sequence that was then used as a reference to remap the raw sequence reads. This process was repeated until all alignment gaps were filled by extension with the new sequence data and ambiguities resolved. Subsequent assemblies began with the more similar reference templates from our first anostostomatid mtDNA genomes. This approach has been proven to be fast and efficient for other Orthoptera [32,34,37]. Mitochondrial assemblies were uploaded as raw FASTA files for protein coding regions, rDNAs and tRNAs were identified using MITOS [38] for a comparison with the published data and detailed examination of the amino acid translations. Annotations were transferred and individually cross-checked through a comparison of the reading frames, amino acid translation, and RNA structure. A similar approach was used to assemble histone sequences using available Ensifera histone sequence and iterative assembly to extend across four exons (H2A, H2B, H3, and H4).
DNA sequence alignments were analyzed using maximum likelihood (ML) implemented in IQ-Tree v2.2 through IQ-Tree tools [39,40] utilizing model selection [41]) and ultrafast bootstrapping [42]. Partition models [43] were applied in ML analyses. Initially, gene and codon partitions were applied but these were optimized in IQTree v2.2 using the partition model test and merge functions to reduce overparameterization. A third codon RY coding scheme was also assessed using binary coding (0,1). Final trees used the optimal model scheme and 1000 bootstrap replicates. We repeated analyses using an unrooted tree approach to further assess the reciprocal monophyly of Australian and New Zealand anostostomatids in our sample.
We used tree constraints to test support for alternative topologies and making statistical comparisons of the fit of the data to the resulting ML topologies using the bootstrap proportion RELL (bp-RELL) approximation [44], (KH) Kishino–Hasegawa test [45], (SH) Shimodaira–Hasegawa test [46], (c-ELW) expected likelihood weights [47], and (AU) approximately unbiased test [48].
Topology constraint tests in Newick format:
Hypothesis 1.
Geographic monophyly but considering the Motuweta/Carcinopsis lineage as sister to core Anostostomatidae, as implied by preliminary analysis [15] ((Carcinopsis, Motuweta), (((‘Rakiura’, H. brucei, H. focalis, H. pallitarsis), (H. crassidens, D. connectens)), (E. ornata, ‘Hemiandrus’, P. flavocalceata, T. laudatum))).
Hypothesis 2.
Geographic monophyly (Carcinopsis, ((Motuweta, (‘Rakiura’, H. brucei, H. focalis, H. pallitarsis)), (H. crassidens, D. connectens)), (E. ornata, ‘Hemiandrus’, P. flavocalceata, T. laudatum)).
Hypothesis 3.
Unconstrained tree (Carcinopsis, Motuweta, ((((‘Rakiura’, H.brucei), (‘Hemiandrus’, T. laudatum)), ((P. flavocalceata, E. ornata), (H. focalis, H. pallitarsis))), (H. crassidens, D. connectens)).
We identified two clades within our sampling of New Zealand Hemiandrus species, so we collected morphological information by examining additional species (Table 2). Thirteen species of Hemiandrus were sampled to encompass the known variation in traits (Table 2) [1,26,49,50,51]. Specimens were examined and anatomical features photographed using a SZX7 Zoom Stereomicroscope with SC100 digital camera and Cellsens v4.2 software from Olympus Corp., Tokyo, Japan. Amplification and sequencing were carried out using primers for mtDNA cytochrome oxidase subunit I (COI) C1-J–2195 [52] and mtd12_wetaR [53]. Previously published sequences [15,54] were obtained from GenBank. Sanger sequences were aligned and analyzed in Ugene [55].

3. Results

Seventeen protein coding genes comprising thirteen mtDNA and four nuclear histones were extracted from data for each of twelve species representatives, then concatenated, aligned, and trimmed, giving an alignment of 12,612 bp of DNA sequence, which formed the basis of the primary analysis (Table 1).
Phylogenetic relationships were inferred from the combined mitochondrial and nuclear genes. A consistent topology was returned regardless of the partition models used including AA, 3rd codon exclusion from CDS, and RY (0,1) coding the 3rd codon position. Only the placement of the Deinacridini lineage (represented by Deinacrida connectens and Hemideina crassidens) varied, which is consistent with the short internal branch returned from the analyses (Figure 3). In all instances, the lineage comprising the New Zealand tusked wētā Motuweta riparia and the New Caledonian Carcinopsis was sister to the other sampled Anostostomatidae. This effectively resulted in the position of this lineage as the outgroup to the ingroup comprising four Australian and six other New Zealand taxa.
Among the Australian taxa in this set were two fully-winged species (Figure 2i,l), but they were not each other’s closest relatives. Instead, the two Australian representatives of the tribe Anabropsini were sister to one another, and the wingless Australian ‘Hemiandrus’ sp. was sister to the winged species Transaevum laudatum in all analyses (Figure 3).
We assessed the possibility that the monophyly of the New Zealand taxa was statistically as likely as this by constraining the tree topology to enforce monophyly (Figure 4) and used 1000 RELL replicates. These analyses confirmed the best fit of the molecular data to the unconstrained phylogeny (Figure 4c, Table 3).
We then removed the Motuweta and Carcinopis sister lineage and repeated the ML analysis with optimal partition models and found improved support for internal nodes in the resulting trees (Figure 5). Without other information, the root could fall on any edge of this unrooted topology (Figure 5), but all possible placements of the root nevertheless resulted in paraphyly of the Australian and New Zealand taxa in the analysis. Furthermore, no placement of a root resulted in the monophyly of Australian and New Zealand Hemiandrus or the monophyly of Hemiandrus in New Zealand, which comprises two independent lineages.
Having identified compelling phylogenetic evidence that the New Zealand representatives of the genus Hemiandrus were not monophyletic, we extended our sampling to determine which of the 19 valid species were part of the Hemiandrus clade and which were not. Using mitochondrial COI sequences, we identified seven species within the genus Hemiandrus and six species that formed a separate clade (Table 2, Figure 6). We propose a new genus to align with these evolutionary relationships.

Taxonomy

  • Class Insecta
  • Order Orthoptera
  • Suborder Ensifera
  • Superfamily Stenopelmatoidea
  • Family Anostostomatidae Saussure (1859)
  • Anderus gen. nov.
  • zoobank.org:pub:65583178-0930-47F3-A9A8-C887572BC0A8
Nocturnal anostostomatids lacking wings. Small to medium (body length approximately 8 to 15 mm) in size and pigmentation varied. Males and females similar in size or females slightly larger. Leg, head, and mandible dimensions are similar in the two sexes. All have maxillary palps with dense small hairs extending through the 5th, 4th, and 3rd segments (c.f. Hemiandrus in which pilosity is on the 5th and distal half of the 4th segments only). Some longer setae scattered on the 3rd and 4th segments. Adult females of all species have prominent, long, narrow curved, or almost straight ovipositors. Adult males have a pair of relatively long and straight, pointed falci on the 10th tergite meeting or overlapping (c.f. Hemiandrus typically with short, hooked, or knob-shaped falci) (Figure 7). Foretibia lack tympana.
Designated type species: Anderus brucei (Formerly Hemiandrus brucei: [26]) (Figure 7). Anderus gen. nov. also includes the species A. fiordensis [19] (NB: the species H. nitaweta Jewell is a synonym of this species); A. maculifrons [28]; A. luna [26]; A. subantarticus [19]; A. nox [26]. The undescribed species included in our main analysis (Figure 5) as ‘Rakiura’ from Stewart Island (=‘saxatilis’ [20]) also belongs to Anderus gen. nov. and awaits formal description.

4. Discussion

Recent subdivisions of the family Anostostomatidae into tribes based on morphological characters have left a number of genera unplaced due to the lack of suitable morphological evidence [1,2,3]. Two of these unclear genera, Hemiandrus and Transaevum, are represented in our phylogenetic analysis. Based on our sampling, we concluded that both genera are more closely related to representatives of the tribe Anabropsini than either Deinacridini or Anostostomatini. In the present analysis, we tested the hypothesis that the extant, flightless Anostostomatidae of Aotearoa New Zealand form a monophyletic group, and found that this could be rejected. Instead, we identified four lineages of wētā Aotearoa that are each more closely related to taxa in either Australia or New Caledonia than to other wētā Aotearoa.
We found that the tusk wētā lineage (Motuweta) was more closely related to New Caledonian taxa, represented here by Carcinopsis, than to other New Zealand genera. Together, Motuweta and Carcinopsis form a lineage that appears to be sister to the core Anostostomatidae, at least as represented in the present sample. Although adult males of both these genera have exaggerated mandible structures that are probably associated with male–-male sexual competition, the structures involved differ in form [56]. In the New Zealand Motuweta (and Anisoura), each mandible is relatively small but bears a long, curved projection (tusk) whereas in males of Carcinopsis that have exaggerated structures, it is the mandibles themselves that are enlarged [23,57]. These are among the paraphyletic diversity of secondary sexual head structures displayed by male Anostostomatidae around the world [58]. Analysis including wider representation of global ansostostomatid diversity to resolve the deeper systematic relationships and evolution of these interesting traits requires a similar scale of DNA sequence data as presented here, as short DNA sequence data are not sufficient [15,16]. The New Zealand genus Hemiandrus emerges as paraphyletic, comprising two separate lineages of ‘ground’ wētā with independent ancestry among Australasian Anostostomatidae. The undescribed Australian species that were proposed as belonging to Hemiandrus [1] belong to another independent lineage that is not closely allied to either of the New Zealand Hemiandrus lineages. In the context of the present sampling, the undescribed Australian ‘Hemiandrus’ species is sister to the Australian winged Transaevum laudatum. It is notable that both taxa bear a single prolateral tympanum on each fore tibia (Figure 8), whereas none of the Hemiandrus in New Zealand have any tympana. The tympanum was noted in the description of Transaevum but not in Australian putative ‘Hemiandrus’ discussed in the same paper [1].
Ander (1838) [59] proposed the name Hemiandrus for relatively small New Zealand anostostomatids in which the females have minute ovipositors, making it difficult to distinguish the sexes (hence half male). Salmon (1950) [19] proposed the name Zealandosandrus for New Zealand ground wētā species where the females have a long ovipositor, and retained Hemiandrus for species where females have very short ovipositors. However, ovipositor length has emerged as a paraphyletic trait. Although one lineage of New Zealand ground wētā includes a distinctive radiation of species with minute ovipositors such as H. bilobatus Ander, 1938 [51], it also includes some species with long ovipositors such as H. focalis (Hutton) and H. jacinda Trewick as well as others with intermediate length ovipositors (H. electra, H. maia [49]) (Figure 6). Salmon (1950) [19] also confounded species of different lineages by confusing H. focalis with H. maculifrons, making the name Zealandosandrus unclear and unusable [1]. Thus, we propose the name Anderus gen. nov., masculine, in honor of Kjell Ernst Viktor Ander (1902–1992), a Swedish entomologist who contributed to the systematics of Ensifera and established the genus Hemiandrus. Anderus gen. nov. is readily distinguished from Hemiandrus by pilosity on the maxillary palps that that extends over all of MP4 and part of MP3 (Figure 7, Table 2) [20,21].
There remain many questions about the higher-level systematics of the family Anostostomatidae and related Orthoptera [16], and identifying the polyphyly of the living representatives of the family in Aotearoa New Zealand indicates that inferences from biogeography should not overly influence this work. Although not ubiquitous in the Australian Anostostomatidae fauna, several fully winged species exist, and partial or fully developed wings also occur in some Ansostostomatidae in Asia and the Americas (e.g., [14,60,61]). While none of the extant wētā of Aotearoa New Zealand have wings, it is now apparent that this endemic fauna is not derived from a single common ancestor in New Zealand. Given the abundant evidence of long-distance dispersal of Rhaphidophoridae in the region [37,62], there is no reason to exclude either active dispersal (flying) or passive dispersal (rafting) of Anostostomatidae to explain the ancestral relationships inferred here. This mirrors the situation in some birds, where island faunas often comprise high numbers of independently derived endemic flightless species (e.g., [63,64,65]). Most profoundly, the large moa (Dinornithiformes) that lacked all bones of the forelimb, which, prior to recent appropriate phylogenetic analysis, were presumed to have a flightless ancestor along with other members of the order. It is now clear that moa and other ‘ratite’ lineages most likely had flying ancestors with that critical trait lost after colonization of separated lands [66]. The absence, in extant species, of traits (such as wings) that are ecologically and taxonomically influential, challenges biogeographic interpretation [67,68], but reemphasizes the dynamism of evolutionary biology.

5. Conclusions

With a small number of representative taxa and rich multigene mitochondrial and nuclear DNA sequence data, we found a convincing signal of paraphyly among Anostostomatidae from New Zealand, Australia, and New Caledonia. In doing so, we found that the genus Hemiandrus actually comprises two separate genera that are both endemic to New Zealand. We propose a new genus (Anderus gen. nov) to accommodate this.

Author Contributions

Conceptualization, S.A.T., M.M.-R. and B.L.T.-S.; Methodology and formal analysis, S.A.T.; Writing—original draft preparation, S.A.T. and M.M.-R.; Writing—review and editing, S.A.T., M.M.-R. and B.L.T.-S.; Visualization, S.A.T. All authors have read and agreed to the published version of the manuscript.

Funding

This research was supported by the Massey University College of Sciences REaDi Fund (RM25248).

Data Availability Statement

The original data presented in the study are openly available in the NCBI GenBank database https://www.ncbi.nlm.nih.gov/genbank/. See Table 1 and Table 2 for accession numbers.

Acknowledgments

We thank the iNaturalist observers for use of the photographs (accessed on 1 October 2024): Exogryllacris ornata Reiner Richter https://inaturalist.ala.org.au/observations/186384142; Penalva flavocalceata https://inaturalist.ala.org.au/observations/204264514; Transaevum laudatum https://inaturalist.ala.org.au/observations/188303308; ‘Hemiandrus’ Australia Nick Lambert https://inaturalist.nz/observations/97002596; Motuweta riparia Sam Purdie https://inaturalist.nz/observations/71114059; Carcinopsis Emilie Ducouret https://inaturalist.nz/observations/83746504; Hemiandrus ‘Rakiura’ Danilo Hegg https://inaturalist.nz/observations/9593056. We also thank Geoff Monteith for providing essential Australian Anostostomatidae material. Collecting permits were provided by the New Zealand Department of Conservation (#NM15823-RES), and New Caledonia Direction du Développement Économique et de L’environement (No. 60912-873-2012/JJC).

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Diversity and approximate species ranges of wētā (Orthoptera: Anostostomatidae) in Aotearoa New Zealand.
Figure 1. Diversity and approximate species ranges of wētā (Orthoptera: Anostostomatidae) in Aotearoa New Zealand.
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Figure 2. Species of Anostostomatidae included in this phylogenetic analysis. (a) Carcinopsis sp. male; (b) Motuweta riparia male; (c) Deinacrida connectens female; (d) Hemideina crassidens female; (e) Hemiandrus brucei male; (f) Hemiandrus focalis female; (g) Hemiandrus ‘rakiura’ female; (h) Hemiandrus pallitarsis female; (i) Transaevum laudatum female; (j) Australian ‘Hemiandrus’ sp. pair; (k) Penalva flavoclceata female; (l) Exogryllacris ornata, not to scale.
Figure 2. Species of Anostostomatidae included in this phylogenetic analysis. (a) Carcinopsis sp. male; (b) Motuweta riparia male; (c) Deinacrida connectens female; (d) Hemideina crassidens female; (e) Hemiandrus brucei male; (f) Hemiandrus focalis female; (g) Hemiandrus ‘rakiura’ female; (h) Hemiandrus pallitarsis female; (i) Transaevum laudatum female; (j) Australian ‘Hemiandrus’ sp. pair; (k) Penalva flavoclceata female; (l) Exogryllacris ornata, not to scale.
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Figure 3. Maximum likelihood topologies inferred from 17 protein coding genes for 12 Anostostomatidae from New Zealand (pink), Australia (blue), and New Caledonia (yellow). Node support above 60% from 10,000 ML bootstrap replicates is shown. Two traits (presence/absence of tympana on the foretibia (P = prolateral; R = retrolateral), presence/absence of wings), and current higher-level classification [2] (Cigliano et al. 2024) are indicated.
Figure 3. Maximum likelihood topologies inferred from 17 protein coding genes for 12 Anostostomatidae from New Zealand (pink), Australia (blue), and New Caledonia (yellow). Node support above 60% from 10,000 ML bootstrap replicates is shown. Two traits (presence/absence of tympana on the foretibia (P = prolateral; R = retrolateral), presence/absence of wings), and current higher-level classification [2] (Cigliano et al. 2024) are indicated.
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Figure 4. Maximum likelihood topologies inferred from 17 protein coding genes for 12 Anostostomatidae from New Zealand (pink), Australia (blue), and New Caledonia (yellow) with constraints: (a) hypothesis 1; (b) hypothesis 2; (c) hypothesis 3 (no constraint).
Figure 4. Maximum likelihood topologies inferred from 17 protein coding genes for 12 Anostostomatidae from New Zealand (pink), Australia (blue), and New Caledonia (yellow) with constraints: (a) hypothesis 1; (b) hypothesis 2; (c) hypothesis 3 (no constraint).
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Figure 5. Evolutionary relationship of 10 Anostostomatidae species from Australia (blue) and New Zealand (pink) inferred with unrooted ML phylogeny with 1000 bootstrap replicates for DNA sequences from 13 mtDNA and 4 nuclear protein coding genes (12,612 bp).
Figure 5. Evolutionary relationship of 10 Anostostomatidae species from Australia (blue) and New Zealand (pink) inferred with unrooted ML phylogeny with 1000 bootstrap replicates for DNA sequences from 13 mtDNA and 4 nuclear protein coding genes (12,612 bp).
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Figure 6. Neighbor-joining trees of available mitochondrial DNA COI (720bp) sequences for recognized species of New Zealand ground wētā that are currently placed in the genus Hemiandrus. The two lineages resolved as separate clades by the analysis of 17 protein coding genes (Figure 3) are presented here as (a) Hemiandrus Ander and (b) Anderus gen. nov. Silhouettes represent the approximate shape and size of adult female ovipositor profile of each species (10 mm scale bar). GenBank accession numbers in Table 2.
Figure 6. Neighbor-joining trees of available mitochondrial DNA COI (720bp) sequences for recognized species of New Zealand ground wētā that are currently placed in the genus Hemiandrus. The two lineages resolved as separate clades by the analysis of 17 protein coding genes (Figure 3) are presented here as (a) Hemiandrus Ander and (b) Anderus gen. nov. Silhouettes represent the approximate shape and size of adult female ovipositor profile of each species (10 mm scale bar). GenBank accession numbers in Table 2.
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Figure 7. Anderus brucei comb. nov. (a) Adult female; (b) Adult male; (c) Dorsal male terminalia showing long falci in contact that is typical of Anderus gen. nov.; (d) Maxillary palp in situ and in silhouette showing the distribution of fine hairs along the full length of MP4, which is typical of Anderus gen. nov. (a, photocredit Uwe Schneehagen). Full description of Anderus brucei comb. nov. is available in Taylor-Smith et al. (2016) [26]. Falci are, as defined by Johns 2001, are the sclerotized projections on the tenth tergite of males [20]. Although the etymology was not given, the term most probably derives from the sickle-like shape of the structures in many taxa (Falx in Latin).
Figure 7. Anderus brucei comb. nov. (a) Adult female; (b) Adult male; (c) Dorsal male terminalia showing long falci in contact that is typical of Anderus gen. nov.; (d) Maxillary palp in situ and in silhouette showing the distribution of fine hairs along the full length of MP4, which is typical of Anderus gen. nov. (a, photocredit Uwe Schneehagen). Full description of Anderus brucei comb. nov. is available in Taylor-Smith et al. (2016) [26]. Falci are, as defined by Johns 2001, are the sclerotized projections on the tenth tergite of males [20]. Although the etymology was not given, the term most probably derives from the sickle-like shape of the structures in many taxa (Falx in Latin).
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Figure 8. Anostostomatid tympana on fore tibiae. Views of paired legs provides a view of both pro- and retro-lateral surfaces: (a) Transaevum laudatum and (b) Australian ‘Hemiandrus’ have shallow prolateral tympana (pink arrows); (c) Hemideina crassidens; (d) Motuweta riparia; (e) Carcinopsis sp.; (f) Penalva flavocalceata and (g) Exogryllacris ornata have prominent paired prolateral and retrolateral tympana.
Figure 8. Anostostomatid tympana on fore tibiae. Views of paired legs provides a view of both pro- and retro-lateral surfaces: (a) Transaevum laudatum and (b) Australian ‘Hemiandrus’ have shallow prolateral tympana (pink arrows); (c) Hemideina crassidens; (d) Motuweta riparia; (e) Carcinopsis sp.; (f) Penalva flavocalceata and (g) Exogryllacris ornata have prominent paired prolateral and retrolateral tympana.
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Table 1. Anostostomatidae sampled for whole mitochondrial genomes and nuclear loci. Voucher material is in the Phoenix Collection at Massey University, Palmerston North (MPN). Current classification within Anostostomatidae, after Cigliano et al. (2024) [2]. * Undescribed but proposed as synonymous with New Zealand Hemiandrus. Images of vouchers for undescribed taxa can be found at: MPN_GW1392 https://inaturalist.nz/observations/9593056, MPN_ORT190 https://inaturalist.nz/observations/223875743.
Table 1. Anostostomatidae sampled for whole mitochondrial genomes and nuclear loci. Voucher material is in the Phoenix Collection at Massey University, Palmerston North (MPN). Current classification within Anostostomatidae, after Cigliano et al. (2024) [2]. * Undescribed but proposed as synonymous with New Zealand Hemiandrus. Images of vouchers for undescribed taxa can be found at: MPN_GW1392 https://inaturalist.nz/observations/9593056, MPN_ORT190 https://inaturalist.nz/observations/223875743.
TaxonClassificationLocationMPN CodeGenBank mtDNAGenBank
Histone		Collector
Carcinopsis sp. Brunner 1888 [23]AnostostomatinaeCol d’Amieu,
New Caledonia		ORT388PQ442192PP965128, PP965140, PP965152, PP965164E.X.M. Trewick
Deinacrida connectens (Ander, 1939) [24]DeinacridiniMount Peel,
South Island,
New Zealand		Dco2011PQ442198PP965131, PP965143, PP965155, PP965167S.A. Trewick
Exogryllacris ornata Willemse, 1963 [25]AnabropsiniBartle Frere,
Queensland,
Australia		ORT193PQ442190PP965130, PP965142, PP965154, PP965166G. Monteith
Hemiandrus ‘Rakiura’UnplacedTin Range,
Rakiura,
New Zealand		GW1392PQ442194PP965139, PP965151, PP965163, PP965175D. Hegg
Hemiandrus brucei Taylor-Smith, 2016 [26]UnplacedWhanganui,
North Island,
New Zealand		Hbr2011PQ442197PP965138, PP965150, PP965162, PP965174B.L. Taylor-Smith
Hemiandrus focalis (Hutton, 1897) [27]UnplacedLake Alta,
South Island,
New Zealand		GW262PQ442196PP965134, PP965146, PP965158, PP965170M. Morgan-Richards
Hemiandrus pallitarsis (Walker, 1869) [28]UnplacedPalmerston North,
North Island,
New Zealand		Hpa2012PQ442195PP965133, PP965145, PP965157, PP965169S.A. Trewick
‘Hemiandrus’ sp. [1]UnplacedBellenden Ker,
Queensland,
Australia		ORT190PQ442193PP965137, PP965149, PP965161, PP965173G. Monteith
Hemideina crassidens (Blanchard, 1851) [29]DeinacridiniMaitai Valley,
South Island,
New Zealand		Hcr2011PQ452770PP965132, PP965144, PP965156, PP965168S.A. Trewick
Motuweta riparia (Gibbs, 2002) [30]AnostostomatiniRaukumura Range,
North Island,
New Zealand		TW29PQ423746PP965129, PP965141, PP965153, PP965165E. Dowle
Penalva flavocalceata (Karny, 1929) [31]AnabropsiniBartle Frere,
Queensland,
Australia		ORT200PQ442191PP965136, PP965148, PP965160, PP965172G. Monteith
Transaevum laudatum (Johns, 1997) [1]UnplacedMt Finnigan,
Queensland,
Australia		ORT178PQ442189PP965135, PP965147, PP965159, PP965171G. Monteith
Table 2. Aotearoa New Zealand ground wētā (Hemiandrus and Anderus gen. nov.) used for morphological and mtDNA COI examination.
Table 2. Aotearoa New Zealand ground wētā (Hemiandrus and Anderus gen. nov.) used for morphological and mtDNA COI examination.
SpeciesMPN CodeLocationYearGenBank
Accession
Hemiandrus brucei
(Anderus brucei nov. comb.)		GW218Raurimu2007EU676796
GW126Pureora Forest2005EU676793
GW49APuketi Forest1990EU676765
GW04Manganuku1998EU676798
GW93APelorus Bridge2005EU676791
Hemiandrus luna
(Anderus luna nov. comb.)		GW104Sky Farm2006EU676742
GW143Lewis Pass2006EU676784
GW1385Tongariro2021PP34546
GW916BArthurs Pass2013PP34545
Hemiandrus fiordensis
(Anderus fiordensis nov. comb.)		GW1481Stuart Mountains2022PP34550
FD3(nitaweta)Sinbad Valley2013PP34541
GW70Lake Roe2004PP34547
GW1516Shy Lake2022PP34553
Hemiandrus nox
(Anderus nox nov. comb.)		GW834Hokitika2012PP34552
GW76Awakiri Valley1997EU676766
GW896aDenniston Plateau2012PP34549
GW899Denniston Plateau2013PP34548
GW1542St Arnaud2005PP34551
Hemiandrus maculifrons
(Anderus maculifrons nov. comb.)		GW119Catlins Coast2006EU676770
GW201Takitimu Mountains2006EU676772
GW68Mount Fyffe2004EU676787
GW217Kahurangi2007EU676776
GW150Franz Josef2006EU676786
Hemiandrus subantarcticus
(Anderus subantarcticus nov. comb.)		GW988The Snares2010MW463359
GW792The Snares2010PP34544
GW989The Snares2010MW463360
Hemiandrus electraGW138St Arnaud2005EU676783
GW1028Mount Richmond2013PP34562
GW1029Mount Richmond2013PP34561
Hemiandrus focalisGW1212Takitimu Mountains2019PP34554
GW1211Takitimu Mountains2019PP34555
FD7Eyre Mountains1999PP34542
GW206Obelisk2006EU676774
GW08Harris Saddle1999EU676773
Hemiandrus superbusFD5Sinbad Valley2013PP34543
FD10Sinbad Valley2010MW463358
FD8Sinbad Valley2011MW463357
GW1198Skippers Range2019PP34556
Hemiandrus jacindaGW1376Pouakai2021PP34560
GW1486Pukeiti 2022PP34559
GW208Whareorino2006MW463354
GW1328Thames2020MW463352
GW62Moehau1990MW463353
Hemiandrus maiaGW125Portabello2006EU676744
GW1067Mount Kyeburn2013PP34557
GW118Hampden2006EU676795
GW136Blue Mountains2006EU676780
Hemiandrus meropeGW674Kapiti Island2011MT6323126
GW682Kapiti Island2011MT623127
Hemiandrus palllitarsisGW227Hauturu2007JF895541
GW226Hauturu2007JF895542
KA90Kauaeranga Valley2012JF895543
KA88Kauaeranga Valley2012F895547
MI98Middle Island2012JF895548
CO145Moehau2012JF895551
GW87Pohaninga Valley2004JF895554
Hemiandrus celaenoGW120Banks Peninsular2005EU676771
GW127Foggy Peak2006EU676778
GW129Foggy Peak2006EU676779
Hemiandrus steropeGW1033Lewis Pass2014MT623110
GW1047Cable Bay2014MT623103
GW717Manaroa2012MT623102
GW602Te Rua Bay2010MT623101
GW54Whites Bay1990EU676788
Hemiandrus taygeteGW491Kaikoura2009MT623112
GW1031Mount Richmond2013MT623115
GW871Upper Clarence Valley2012MT623113
GW1231Youngman Stream2019PP34558
Hemiandrus bilobatusGW586Awatere Valley2010MT623085
GW25Wellington1999EU676794
GW240Wellington2007JF895562
GW657Mana Island2010MT623095
GW122Marfells Beach2000EU676777
GW55Marfells Beach1990EU676789
GW193Muritai2006JF895564
Table 3. Support for alternative evolutionary topologies among Anostostomatidae. Comparison of trees that constrained the relationships based on geography (Hypotheses 1 and 2) or unconstrained (Hypothesis 3). Results of 1000 RELL replications for 12 Anostostomatidae comparing the ML fit of data (12,612 bp DNA sequence) to each of three hypotheses (see Figure 4).
Table 3. Support for alternative evolutionary topologies among Anostostomatidae. Comparison of trees that constrained the relationships based on geography (Hypotheses 1 and 2) or unconstrained (Hypothesis 3). Results of 1000 RELL replications for 12 Anostostomatidae comparing the ML fit of data (12,612 bp DNA sequence) to each of three hypotheses (see Figure 4).
HypothesislogLdeltaLbp-RELLp-KHp-SHc-ELW
1−89,689.86177679.860−0−0−3.53 × 10−231−
2−90,220.767411210.80−0−0−0−
3−89,010.0065901+1+1+1+
deltaL: logL difference from the maximal logl in the set. Bp-RELL: bootstrap proportion using RELL method [44]. P-KH: p-value of one-sided Kishino–Hasegawa test [45]. P-SH: p-value of Shimodaira–Hasegawa test [46]. C-ELW: expected likelihood weight [47]. Plus signs denote the 95% confidence sets. Minus signs denote significant exclusion.
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MDPI and ACS Style

Trewick, S.A.; Taylor-Smith, B.L.; Morgan-Richards, M. Wētā Aotearoa—Polyphyly of the New Zealand Anostostomatidae (Insecta: Orthoptera). Insects 2024, 15, 787. https://doi.org/10.3390/insects15100787

AMA Style

Trewick SA, Taylor-Smith BL, Morgan-Richards M. Wētā Aotearoa—Polyphyly of the New Zealand Anostostomatidae (Insecta: Orthoptera). Insects. 2024; 15(10):787. https://doi.org/10.3390/insects15100787

Chicago/Turabian Style

Trewick, Steven A., Briar L. Taylor-Smith, and Mary Morgan-Richards. 2024. "Wētā Aotearoa—Polyphyly of the New Zealand Anostostomatidae (Insecta: Orthoptera)" Insects 15, no. 10: 787. https://doi.org/10.3390/insects15100787

APA Style

Trewick, S. A., Taylor-Smith, B. L., & Morgan-Richards, M. (2024). Wētā Aotearoa—Polyphyly of the New Zealand Anostostomatidae (Insecta: Orthoptera). Insects, 15(10), 787. https://doi.org/10.3390/insects15100787

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