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  Te Pūtaiao o Tokoriro:

Taxonomy and diversity of New Zealand cave wētā (; )

A thesis presented in fulfillment of the requirements for the degree of

Doctor of Philosophy

In Zoology

at

Massey University, Manawatū New Zealand

Josephine Fitness

2016

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Abstract Species are the fundamental unit for ecology and evolution. , the naming of species, grabbles with the problem of accurately representing these fundamental units. In this research I targeted a group of understudied and undervalued that are common throughout New Zealand. This work focuses on Rhaphidophoridae, a family of Orthoptera found globally, but the diversity in New Zealand is poorly understood and poorly described. I have been the first to use high specimen numbers in order to establish within and between species differences of New Zealand cave wētā. I have established the importance of multiple taxonomic methods. At no stage was the aim to fully resolve all issues, but rather to identify morphological characters that are useful in distinguishing species, and integrating mtDNA sequence data to test species hypotheses.

I focused first on cave wētā specimens that came from a biodiversity studied but had not been identified to genus or species. I was able to identify characters that could distinguish between the taxa present in this sample and developed a method that could be transferred to other locations. Two key findings were that multiple cave wētā species co-exist across a range of habitats and that variation in abundance was species dependent. Of importance was my finding that juveniles cannot be distinguish and placed with their correct adult form due to changes in both subgenital plate shape and apical spines.

From three regions in North Island New Zealand I was able to distinguish and identify fourteen putative cave wētā species. mtDNA sequence data were use to test putative species clusters identified by morphology and allowed me to confidently pair male and female specimens. Combinations of apical leg spines and subgenital plate shape could consistently diagnose most taxa. Many of the species are new to science. Therefore I described three new species in the genus Neonetus. I reviewed our current knowledge of the endemic genera Pleioplectron, Weta and Miotopus and based on evidence from mtDNA sequences and large samples I was able to clarify current species and describe one new Miotopus species. As with many species, male terminalia are the key to distinguishing among species, and species within the same genus have similar female subgenital plates.

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Acknowledgements $-./'4  2*0'  '$&  /* &)*2' "  /#  .0++*-/ )  )*0-" ( )/ *! (4 .0+ -1$.*-. -*! ..*- / 1  - 2$& )  ..*$/  -*! ..*- -4 *-")B $#- .A # 4 #1   ) 3/- ( '4 .0++*-//$1  *! (4 2*-& )  +-*% / 1 ) /#*0"# 2.'$1$)"*0/*!/*2)>) # /2*&$ ./*'**&!/ -A#$.2*0' ) 1 - #1  )+*..$' 2$/#*0//# (A  2*0' '.*'$& /*&)*2' " /# $)$./-4*!0$.) ..)  ))*1/$*)!*-/#  .#*'-.#$+ 0/$* ''*2.#$+A#$./#- 4 -!0) $)"( $/+*..$' /*--4 *0/) *(+' / /#$.+-*% /!-*(./-//*!$)$.#A'.*)&)*2' " ( )//*/#  .0++*-/*!/# )$1 -.$/4) .0+ -1$.*-.2#*'.*+-*1$ !$))'.0++*-//* *(+' / /#$.+-*% /A  # -  -  .* ()4 + *+'  /* &)*2' "  !*- /# $- *)/-$0/$*) )  .0++*-/ /#-*0"#*0/ /#  +-*% /A $./'4  $" /#)&. /* /#  - ./ *! /#  # *)$3 ' 2#* / /$( . $  .*(  ' 2*-& 3/-/$)" )  . ,0 )$)"  !*- 1  29/7 1 ) /#*0"#/# 4#1 ) 1 -( /( A$/#*0//# # '+*!/#$."-*0+.*( /#$)".%0./ 2*0' )*/#1 #++ ) A*-/#*. 2#*# '+ $)!$ ' *'' /$)"*!.+ $( ). $"/#)&./* (0''$1)>#0)$ '.*)>-$-4'*-B($/#> 4*2' >.#> 2$.$/) ..A'.*$"/#)&./*/#*. 2#*#1 +-*1$ 1 29/7.+ $( ). *1 - /#  4 -. /* / 1  )  -4 !-*( 1-$*0. '*/$*). -*0)  /#  *0)/-4> .+ $''4 $&  0.&> *--$)  **&> '$1 - ''> )  / 1  )  -4E. 3/ ).$1  *'' /$)"/# (. '1 .A 2*0' '.*'$& /*/#)&'$1 -''"$)!*-#$.# '+$) +-*1$ $)"'-" .(+' !-*($* $1 -.$/4./0 4!*-(%*-*(+*) )/*!/#$. +-*% /A'.*/* *#)/#)* !- 4!*-#$.# '+$).*( )4. .) *)/-$0/$*) !*-+0'$/$*)A  #1 # .*( #0" .0++*-/!-*(*/#(4!($'4) (4#0.) E.!($'4A# 4 #1 # /* '2$/#(4./- ..*))0( -*0.*..$*).) '**&$)"!/ -(4&$ . 2#$'   #1   ) *0/ $) /#  !$ '  *- / .. 4 )$1 -.$/4A $/#*0/ /# $- # '+  2*0' #1  )'*./A*/#)&./*(4+- )/.-#) #$''$+) (4$)B'2. / +#) -4A

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Table of Contents                         "   " " ) -'$)/-* 0/$*)/*$* $1 -.$/4>/3*)*(4) #+#$ *+#*-$  *) /# "'*') )/$*)'' 1 'A    #  #( #$.$.++ -2-$// )) +0'$.# . *)- . -#'**&$)"/1  29/7!-*(/# -*-/#) #*22 - ' /* $./$)"0$.#.+ $ .A*(  *'*"$' /2.0. .2 '' 0 /*/# *) $/$*).*!/# $)$/$' $* $1 -.$/4.0-1 4!*-/# - A  &((! %+$@$"0- OAR*-- /$*)   &(  **$%*&&%*(!+*!&% &)    $   '" #$.#+/ -'**&.//#- - "$*).-*0) /# *-/# .') ) *(+- . /# .+ $ ./#/- !*0) $)!*- ./$)/# . - "$*).A

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Chapter 6 Figure 6.1 Summary of genera 161

xi

The Rhaphidophoridae

CHAPTER 1

Introduction to Rhaphidophoridae

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1 Chapter 1

‘Takoto kau ana te whānau a Tāne” The descendants of Tāne are laid low – Tāne is the god of the forest. This is a plea for conservation.

Māori view the natural world with a holistic approach, their beliefs and traditions acknowledge their relationships within the natural environment (Harmsworth & Awatere, 2013). Māori traditional whakapapa (genealogy) places all living flora and fauna within the same family tree. Thus they see themselves as part of the environment and ecosystems rather than being separate from it. This whakapapa or the genealogical sequence from the beginning of time to present is central to Māori perspective of the natural world and their link within it. Central to their belief is the creation myth of the separation of Ranginui (Sky father) and papa-ti-a-nuku (Earth mother) which allowed the birth of their children (wind, forest and plants, sea, rivers and ). The key thing here is that humans are not separate to other and plant species but are included and therefore a key component of a balanced ecosystem. This is reflected in their roles as Kaitiaki (guardians) for carrying out Kaitiakitanga (sustainable resource management and guardianship). Viewing all life as being part of the same whakapapa is compatible with our current understanding of the evolution of life on earth, because all species are linked by a common ancestor (Darwin, 1859; Theobald, 2010).

It has become increasingly more common to take a holistic view of conservation and include indigenous people in planning and monitoring local ecosystems. Through traditional ecological knowledge, tangata whenua (people of the land, or original inhabitants of New Zealand) are well placed to be highly involved in current planning, management and conservation of the environment (Ramstad et al., 2007). This thesis brings the importance of whakapapa to the understanding and conservation of the endemic New Zealand insect species known as cave wētā or Rhaphidophoridae.

 ! Taxonomy is the discovery, description, naming and classification of living organisms on earth and the study of their whakapapa (genealogy or evolutionary relationships). Good taxonomy is important in all biological research (Khuroo et al., 2007). As

2 The Rhaphidophoridae researchers we need to know what taxonomic units we are considering, whether we study conservation, ecology, phylogenetics, physiology, biochemistry or biomedicine. For the most part we know and have described the large megafauna on earth and have a general understanding of species (see below), but our knowledge of the much greater diversity of smaller life continues to expand (Troia & McManamay, 2016). The diversity of unicellular organisms is only just being realised (Nee, 2004). The discovery of reproductively isolated but morphologically similar taxa (cryptic species) reveals underestimated species diversity. Tools to study the shape and form of plants and animals have resulted in a greater appreciation of the morphological variation that exists. Detailed assessment of morphological differences among specimens can sometimes reveal examples of very subtle patterns of morphological variation, yielding recognition of “pseudo-cryptic” species (Knowlton, 1993). In other situations, morphological variation is found to be within a single lineage, thus for some groups we have overestimated total biodiversity (Bickford et al., 2007). Problems of both underestimation and overestimation are seen in New Zealand cave wētā taxonomy; species diversity has been partitioned into too many genera (Cook et al., 2010), and new species are discovered by combining better sampling and genetic data. For example Talitropsis specimens from the Chatham Islands were originally thought to belong to a highly variable single species but are now recognised as two distinct species (Steven A Trewick, 1999).

Taxonomy is limited by poor sampling so that the diversity is not fully represented. But it can also suffer from an excessive number of descriptions where later taxa are recognised as synonymies (Gaston & Mound, 1993). This creates a problem for the biologists who follow, as historical taxonomy must be traced and corrected, before new taxa can be described. In addition, the definition of “species” differs among researchers and this too can lead to conflicts of understanding the literature. The subject of what a species is has been extensively discussed e.g Darwin, 1859; De Queiroz, 2007, 2011; Mallet, 1995; Mayr, 1942. Here I take the view that a ‘species’ is a taxonomic description of an arbitrarily delineated segment of an evolutionary lineage in time (see Fig. 1 Vaux et al., 2016). However, although a species is artificial, it remains a hypothesis based on empirical observations of an evolutionary lineage and as a hypothesis provides a set of important predictions. In practice, Mallet’s genotypic

3 Chapter 1 cluster definition provides a suitable method for identification of “species” using either (or both) morphological and genetic data (Mallet, 1995).

variation (unlabelled lineages following the designation of a species). The assignment of these taxonomic categories is arbitrary as the size and scale of segments varies. Not all lineage-splits are classified as speciation (cyan asterisk), and species classifications based on ancestral and derived difference without evidence of lineage-splitting (e.g. chronospecies) do not invoke a discrete speciation event. Species Vaux et al. 2016. (Figure 1) may be described based on limited Yarn as a metaphor for evolutionary fossil evidence (blue Sp), because lineages: lineage-splitting and variation is novel or of interest, even splitting hairs. A, a piece of yarn when there is limited knowledge of represents an evolutionary lineage the lineage to which it belongs. C, (yellow L). Like evolutionary depending upon the scale of lineages, yarn is continuous and observation (limited by the consists of many fibres. In both, availability of data such as zoom splits can be identified (red and resolution in photography or asterisks). The origin of each piece sampling in biology) further lineages of yarn in the tangled ball of wool (fibres) and splits (lineage-splits) represents the unknown common can be identified. Many lineages do ancestry of lineages as we move not persist for a significant length of backward in time. Many lineage- time and either go extinct or splits are also missed due to hybridize with the original lineage. extinction (orange asterisks). B, D, lineages are made up of particular segments of lineages can individuals within populations and be classified as species (green Sp, hybridization can unite populations purple lines representing temporal (pieces of yarn that split may boundaries of segments), relative to recombine soon thereafter). the studied organism, the availability Differentiating lineages (fibres) is of data and the hypotheses under easier when divergence has investigation. Segments of lineages followed a lineage-split. can also be classified as subspecies or varieties (green Ssp), or consolidated as intraspecific

4 The Rhaphidophoridae

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5 Chapter 1

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6 The Rhaphidophoridae

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7 Chapter 1

" "                                #!""  Subfamily Genera Species Range Asia- Aemodogryllinae 12 93 tropical/temperate, (Walker, 1871) Europe, NA Anoplophilinae (Storozhenko & Paik 2 28 Korea 2010) Ceuthophilinae 24 71 North America Dolichopainae South Europe, Asia (Brunner von 1 47 Minor, Caucasus, Wattenwyl 1888) Transcaucasia Gammarotettiginae 1 6 California Macropathinae 29 95 Austral (Karny 1929) Protroglophilinae 2 4 Extinct from Europe (Gorochov 1999) Rhaphidophorinae Eat Asia, Australia 6 84 (Walker 1871) and Samoa Troglophilinae South Europe to 1 17 (Krauss 1879) Syria Tropidischiinae 1 1 California (Scudder 1869)

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8 The Rhaphidophoridae

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9 Chapter 1

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10 The Rhaphidophoridae

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11 Chapter 1

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12 The Rhaphidophoridae

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13 Chapter 1

.-$+/$*). )  /-4$)" /* !$)  /# .  #-/ -.>  #1  !*0)  /#/ $/ $. *!/ ) $!!$0'/ /* $)/ -+- / 2#/ * 4 +-/ *- .#+  /#  2-$/ - $. - ! --$)" /*A ) +-/$0'- .+$)  *0)/. -  #-  /* !*''*2 $! .+$) . -  )*/ 0-/ '4 )  *).$./ )/'4 - ! --  /*A !/ ) (*-  /#) *)  )(  #.  ) "$1 ) /* /#  .(  ./-0/0- >2#$# ./*/# *)!0.$*)A  . 2 '' . /#  ($.$ )/$!$/$*) +-*' (.> /# -  $. '.* /#  +-*' ( 2# -  .+ $( ).))*/ $ )/$!$ /*.+ $ .A# - - ()4 3(+' .$)+0'$.#  2*-& 2# -  -#+#$ *+#*-$ . -  $ )/$!$  /* /#  " )0. *- !($'4 ' 1 'A *- 3(+' >+0--)  - )JOMMQK'$./.+ $ .*!29/7!-*(*/# -./0 $ .2# -  /# 4 #1   ) - *-  . /$)" $/.A   ) .  /#/ *)'4 *)  *! /#  $"#/ .+ $( ).'$./ #. )$ )/$!$ /*.+ $ .' 1 '>/# - .//*" )0.) *) . F0)$ )/$!$ GA )!*-(/$*) '$&  /#$. #. /#  +*/ )/$' /*   /* /#  - + -/*$-  *! &)*2' " !*-/#/.+ $ .>.0#.2# - .+ $ .$.!*0) ) /#  !! /.$/. #1 *)+*+0'/$*).>0/*)'4$!$/) ++-*+-$/ '4..$") A  *- /#  " ) -' +0'$ /# -  $. 1 -4 '$//'  $)!*-(/$*) *) /#  .-$+/$*). )  *'*"4*!.+ $ .$)'0 $)".*( .$$ )/$!$/$*)*!#-/ -./#/(4# '+ +' .+ $ .$)/*" ) -A!0/0- "*'2$'' /*- / 2 .$/ /#/*)/$).. (0# $)!*-(/$*) *) /# .  - /0- . . $. 1$'' A  /" / J#//+@HH2 /" /A(.. 4AA)5HK $.  . -#'  2 .$/  /#/ * . %0./ /#$. 0/ *)'4*)/$)./# $)!*-(/$*)2 #1 /* / A 

14 The Rhaphidophoridae

Table 1.2 Science literature relating to New Zealand cave wētā. Author Name Topic Allegrucci et al., 2010 Macropathus filifer Systematics Pachyrhamma edwardsii Pallidoplectron turneri Pleioplectron simplex Talitropsis sedilotti

Bolivar, 1882 Talitropsis sedilotti

Brunner von Neonetus variegatus Systematics Wattenwyl, 1888 Pachyrhamma

Chopard, 1923 Neonetus variegatus Systematics Neonetus huttoni Weta thomsoni Pachyrhamma fascifer Pleioplectron cavernae

Cook et al., 2010 Gymnoplectron longipes Systematics Macropathus sp. Pachyrhamma acanthoceras P. delli P. edwardsii P. fuscum P. giganteum P. longicaudum P. ngonogtahaense P. spinosum P. tuarti P. uncatum P. waipuense P. waitomoensis Pleioplectron cavernae P. simplex Turbottoplectron cavernae T. unicolor Weta thomsoni

Gorochov, 1988 Tallitropsini Systematics

Hutton, 1897 Talitropsis sedilotti Systematics; Hutton T. crassicruris attempted to define T. irregularis genera and a number Ischyroplectron isolatum of species. Many he Gymnoplectron longipes described himself Pachyrhamma speluncae others are P. novae seelandiae redefinitions of P. fascifer existing taxa Pleioplectron simplex P. hudsoni P. pectinatum P. diversum Neonetus variegatus N. pilosus Isoplectron armatum I. calcaratum

15 Chapter 1

Pharmacus montanus Macropathus filifer M. edwardsii Johns 2014 Maotoweta virescens Systematics

Karny, 1930 Talitropsis apoduroides Systematics; also refers to higher taxonomic levels

Karny 1935 Pachyrhamma chopardi

Karny, 1937 Isoplectron aciculatum Systematics Talitropsis chopardi

Kirby, 1906

Milligan 1926

Richards, 1954a Macropathus filifer Ecology Observation

Richards, 1954b Macropathus filifer Observation

Richards, 1954c Macropathus filifer Systematics Macropathus acathocera Ecology

Richards, 1955 Ecology of general cave weta

Richards, 1958a Talitropsis crassicruris Systematics Novoplectron serratum Ecology

Richards, 1958b Macropathus filifer Systematics

Richards, 1958c Pachyrhamma longipes Systematics Pachyrhamma waitomoensis Pallidoplectron turneri

Richards, 1959a Pachyrhamma fusca Systematics Pachyrhamma uncata

Richards, 1959b Pleioplectron simplex Systematics P. hudsoni P. pectinatum P. diversum P. edwardsii P. cavernae Richards, 1959c Pachyrhamma longicauda Systematics

Richards, 1960 Pachyrhamma waipuensis Systematics Pallidoplectron peniculosum

Richards, 1961a Pachyrhamma waitomoensis Ecology Pallidoplectron turneri

Richards, 1961b Observations

Richards, 1961c Gymnoplectron spinosa Systematics G. ngongotahaensis

16 The Rhaphidophoridae

G. tuarti

Richards, 1961d Gymnoplectron Systematics and Macropathus synonym Pachyrhamma

Richards, 1962a Gynmoplectron waitomoensis Ecology Pallidoplectron turneri

Richards, 1962b Gymnoplectron giganteum Systematics

Richards, 1964 Dendroplectron aucklandense Systematics Notoplectron campbellense

Richards, 1965a

Richards, 1965b Gymnoplectron waitomoensis Ecology Pallidoplectron turneri

Richards, 1965c Pallidoplectron subterraneum Systematics

Richards, 1970 Insulanoplectron spinosum Systematics

Richards, 1971 Gymnoplectron Biogeography/ Talitropsis Relationships with Paraneonetus Australia Dendroplectron Novoplectron Turbottoplectron Pleioplectron Insulanoplectron

Richards, 1972 Pharmacus montanus Systematics P. chapmanae P. brewsterensis P. dumbletoni Petrotettix serratus P. spinosus P. cupolaensis P. nigripes Setascutum ohauensis S. pallidum

Salmon, 1948 Turbottoplectron unicolor Systematics

Thomas et al., 1998 Pleioplectron simplex Ecology

Trewick, 1999 Talitropsis crassicruris Systematics T. megatibia T. sedilotti

Ward, 1997 Generic key to NZ family

17 Chapter 1

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18 The Rhaphidophoridae

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'' "-0$>A>-0#$>A>:*- *)$>AJOMNNKA (+*) (* *!.+ $ . $1 -.$!$/$*)$)*'$#*+* 1 -$& /.J-/#*+/ ->#+#$ *+#*-$  KA &#+#( /#& %*!)%,&#+*!&%>A;JNK>NMUIONA

#())> A> ))5 //$>A>:*- *)$>AJNVVQKA#-/ -$5/$*)*!.+ $ .B .+ $!$./ ''$/ !($'4*!*'$#*+* .#$155$$J-/#*+/ -> #+#$ *+#*-$  K1 -$& /.A &+(%#&&#+#(,&#+*!&%>>DJPK>OTQI OUNA

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19 Chapter 1

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20 The Rhaphidophoridae

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21 Chapter 1

'' /> AJNVVRKA.+ $ . !$)$/$*)!*-/# (* -).4)/# .$.A(%)!%&#& /2 ,&#+*!&%><;JTK>OVQIOVVA

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22 The Rhaphidophoridae

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23 Chapter 1



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24 The Rhaphidophoridae

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03>A>- 2$&>AA>:*-")B$#- .>AJOMNSKA $) " .>.+'$/.) 

25 Chapter 1

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26 Te Paki

CHAPTER 2

Improved resolution of cave wētā diversity

(Orthoptera: Rhaphidophoridae): ecological

implications for Te Paki, Far North, New Zealand.

   

27 Chapter 2



28 Te Paki



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29 Chapter 2

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

30 Te Paki

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

31 Chapter 2

 Figure 2.1. Te Paki Ecological District, Far North, New Zealand, showing location of pitfall trap sites that provided cave wētā specimens.

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

 

32 Te Paki

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

33 Chapter 2

( .0- ( )/. 2 -  /& ) !-*( /#  .$  /#/ #  /#  (*./ *(+' /  ' ". ./$'' //# /*/# * 4/* ).0- *-- /''*/$*)A-*)*/0(' )"/#2.+- ! -'  /* * 4 ' )"/#> 2#$# $. 1-$'  $) '$1$)" )  +- . -1  .+ $( ).A #  .0B " )$/' +'/ . *! (' . )  ! (' . 2 -  3($)  )  '..$!$  $)/* -  $'4 - *")$. /4+ .A$"( )//$*)) +$'*.$/4J#$-$) ..K2 - *).$ - 0/*/# - '/ - *- $!!$0'//*.. ..$)2 /"'4*'+- . -1 .+ $( ).A 

34 Te Paki

Table 2.1. Apical spines used for diagnosis of species and a summary of the presence and absence on each of the four putative species of cave wētā from Te Paki Ecological District: 1= present 0 = absent

   Pro. Pro. Pro. Retro. Retro. Retro. Mid Femur Pro. Mid Fore Femur Pro. Hind Femur Pro. Femur Pro. Hind Mid Femur Retro. Femur Retro. Mid Fore Femur Retro. Hind Femur Retro. Femur Retro. Hind Mid Tibia Inferior Prol. Prol. Inferior Tibia Mid Fore Tibia Inferior Pro. Inferior Fore Tibia Hind Tibia Inferior Pro. Inferior Tibia Hind Mid Tibia Superior Pro. Superior Tibia Mid Hind Tibia Inferior Sub. Hind For Tibia Inferior Retro. Retro. Inferior For Tibia Mid Tibia Inferior Retro. Retro. Inferior Tibia Mid Hind Tibia Inferior Sub.l Sub.l Inferior Tibia Hind Fore Tibia Superior Pro. Superior Fore Tibia Hind Tibia Superior Pro. Superior Pro. Tibia Hind Hind Tibia Superior Sub. Tibia Hind Superior Sub. Tibia Hind Hind Tibia Inferior Retro. Inferior Tibia Hind Mid Tibia Superior Retro. Retro. Superior Tibia Mid Fore Tibia Superior Retro. Superior Fore Tibia Hind Tibia Superior Retro. Superior Retro. Tibia Hind

Spine Code S1 S2 S3 S4 S5 S6 S7 S8 S9 S10 S11 S12 S13 S14 S15 S16 S17 S18 S19 S20 S21 S22 TPA 1 1 1 1 0 0/1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 TPB 1 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 TPC 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 TPD 1 0 0 1 0 1 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1

 

35 Chapter 2

),*")' .,#. ),)'*,.#0 (&3-#-1 , ).#(  ,)'-/-'*& )  -* #' (-."., *, - (. ." ,(! ) -"* -<-#4 -(- 2) -* #' (-?)-. ) ." - 1 , /&.-),&,! $/0 (#& -(1 (). ." *, - ( ) '(3'& - ( '& - .". 1 ,  (). /&. /. 1 ,  ( 0 ,." & -- &,!  )'*,  .) ."  0 ,! -#4 ) 0 17.5#()/,-'*& ?&&-..#-.#&(&3- -/- 0O?LH),   'NLMOI?2*&),.),3(&3-#-/- ,#(#*&)'*)( (.(&3-#-HI#(."  )!*%! H%-( ( .&?NLMOI?"#-(&3-#-#(&/ ." . !),#&*#& -*#( (' .,#..)" &* . ,'#( 1" ." ,.2)/& - *,. )(."  -#-) ." - ",. ,-?&-) 2'#( 1" ." ,$/0 (#& -)/& '."  .)." #,/&.)/(. ,*,.-/-#(!."#-'),*")&)!#&#( ),'.#)(<(1"#"# (3 ) ." - .,#.-),, -*)( )('& -( '& -?  #." . ,)' /&.- ( &,!  $/0 (#& - H#? ? ( , /&.-I #( ."  -  )  )(  '),*").3* I<.@. -.-1 , /- .))'*, -#4 # , ( - .1 (." - 2 -( ." */..#0 -* # -?1#."/% 3 . -.1-/- .))'*, # , (. ' -/, .,#.-')(!*/..#0 .2? **&# .)." ' .,#.H,)< < I)(." #,)1(.)-- --'),*")&)!#&0,#.#)(')(!#(#0#/&-1#."#(."  &,!  -* #' ( - .?   # (.# #   )'#(.#)( )  ",. ,- .". #!()-   (/' ,) */..#0 -* # -?&#!(' (.) ."  %#.2.)."  2#-.#(! -,#  0 17.5 /(#(0)&0 $/##)/-/- )  2#-.#(!&#. ,./, - .#&  &)1?  (!! ).& )/(.- )  && # (.# #&  ( /(# (.# #  $/0 (#&  0  17.5 ,)' " -'*&  (&  /- .) (&3-  ." #, , &.#0  /((  )0 , -*  ( .#' ? && -..#-.#&(&3- -/- 0O?LH),  'NLMOI? /-  ( ,&#-  #( , ) &&#(!H I--/'#(!)#--)(#-.,#/.#)() )/(...))'*, 0  17.5/(( )0 ,.#' ?#. ("#.., ( -. 0,#& -?.,#.&3D-#. E#- ,()' . /. 1- (). #..  #( ."#- 13  /-  1  1(.  .)  . ,'#(  1" ." , ." ,  1- #.#)(& -#.  G -#.  0,#.#)( ( -) 1    . ,'- - +/ (.#&&3#(.)." ') &? (! ( ,&." (/' ,) .,*@3-1-)(-.(. ), && -#. - #(  !#0 ( ')(." ( 0,#.#)( ')(! ')(."- 1- )'').  3 #(&/#(!')(."- .),#(." ') &-? ,))/(.-1 , #(- ,. 1" , .,*- 1 , #(/- /./!".()0 17.5<1"#& .,*-().#(/-  ),*,.#/&,')(."

36 Te Paki

1 , ) -? ') && 0 17.5/(( 1#."(1#.")/.. ), ." /(# (.# # $/0 (#& -- # .".2)(?" , -*)(- 0,#& 1-." )/(. ), "-* # -?"  2*&(.),30,#& -1 , -#. <"#...3* H(.#0  ), -.<*#(  ),-",/&(I(.#' H')(.") ." 3 ,.,** I<(&&*)--#& #(. ,.#)(- ) ." - 0,#& -1 , )(-# , ?

  -#(! ."  &,! , 0  17.5 .". 1 ,  #." , /&.- ), ( , /&.-< 1  # (.# #  )/, '),*").3* - ,)' ." #, -*#(.#)(< )0 ,&& -#4  ( ** ,( ? & - ( '& - )/&   , &#&3 *#,  -  )( -*#(  )/(.- ( )0 ,&& -#4 ?  -*#.  ." #,-#4 <#.1-**, (..".." &,! -.#(#0#/&-<&&) 1"#"1 , --#!(&  .).2)(<#(&/ ()/&.-? '& -< ),#(-.( <", &.#0 &3-) .<*& ( &/(. )0#*)-#.),- 1" , - /&. "*"#)*"),#  )0#*)-#.),- ,  -& ,).#- < ,%<(*)#(. ) . (1#." #( - ,,.#)(-?" -#4 ,(! ) ." -* #' (- 1- !, . , ."( ."  /&.- )   -/!! -.#(! .".  (/' , )  #(-.,- 1 ,  #(&/ ')(!-.." -'*& ?(1 , *& ,."((<( ""/(#+/ -*#( )'#(.#)(H& N?MI?1--#'#&,#(-#4 .) (1-#(#.#&&3)0 ,&))%  /- #.-*&&#)&)/,.#)(( &#.  ),''  #. &))% &#%   $/0 (#&  )  ()." , -* # -? "  )/,." *" ().3* < < 1- ."  -'&& -. )  && )  ."  */..#0  .2 ( ."#- -* # - &-) "  *& < $/0 (#& @&#%  ** ,( ? , /& 2'#(.#)( )  ."  )0#*)-#.),- #( ."  '& - )   -* #' (- , 0 &  )." ,1#- ? "  /(#+/  -*#(  )'#(.#)(- )  && )/, .2 , #( ), )/,"3*)." -#-.".." 31 , # , (.-* # -(().$/-.0,#(.-)  ."  #,-.'),*").3* - ()/(. , H#!/, N?OI?  &% !'! "%#"',#& (#.#& -- --' (. )  ."  , &##&#.3 )  &--# 3#(! $/0 (#& - #(.) )(  )  ."  )/, '),*").3* -#(#. .".#.1-*,/ (..) 2&/ -'&&$/0 (#& -HWJML''I ,)' -/- +/ (. '),*")&)!#& (&3-#-? " ,  1- () & , #-,#'#(.#)( )  $/0 (#& -- )(-#4 #(.))( ) ."  )/,*/..#0 -* # -H#!/, N?PI? &-) )/(.".*, - ( F- ( ) -)' -*#( -.) '), 0,#& ')(!." -'&& $/0 (#& -."(/&.-<-/!! -.#(!()(.)! ( .#)'*)( (..)-*#(  0 &)*' (.? /0 (#&  '& -)/&/-/&&3 #-.#(!/#-" , #&3 ,)'." #,, -* .#0 /&.-

37 Chapter 2

3 ." #, *& < -) . ( &/(. )0#*)-#.),- &%#(! - ,,.#)(-? /0 (#&  '& - 1 ,  & -- , #&3 #-.#(!/#-"  - &.  #(-.,- 1 ,  0 ,3 -#'#&, .) /&.-? )1 0 ,< $/0 (#& -. ( .) -'&& ,<*& ,<1#." 1 ,*#&(&. ,&-*#( -)(."  & !-< ( ."  ! (#.&# 1 ,  /( , 0 &)*  ( ) . ( # #/&. .) - ? #-.#(!/#-"#(!* (/&.#'. (&-.#(-.,#(#0#/&-1-." ')-.# #/&.(#( '(3 - - #'*,.#&< /. &-) '3 ().   (  --,3 2 *. 1" ( - )(,3 - 2/& ",. ,- ,  2*, --  )(&3 #( ."  #(& #(-.,? ( ), , ), -* # - #!()-#-.) -,)/-.-*)--#& 1 , ')0 &&-'&&$/0 (#& (/( ,.#( #(#0#/&- ,)'." -'*& - .- ), "'),*").3* < 2 *.#(!.")- &,! /. ()./&.#(#0#/&-) <1"#")/&&13- , &#&3#-.#(!/#-"  ,)'."  )." ,'),*").3* -?  & - ( '& - 1#."#( ."  -'  -* # - " 0 ,3 # , (. -/@! (#.& *&. - --)#. 1#."." #,# , (.! (#.&-.,/./, ( /(.#)(?,)' 2'#(.#)()  /&.-* #' (-H(( ,@/&.#(#0#/&-I)&& . #(."#--./31 1 ,  & .)# (.# 3 #!".#-.#(.-/@! (#.&*&. -"* -= )/,, &.#(!.) '& -( )/,.)'& -H#!/, N?QI?)*#,." '& -( '& -) ." -' -* # -1  /- ." *#&& !-*#( -?  ( ."   -* #' (-< '& - "  -/@! (#.& *&.  .". 1- -'&& ( .,#(!/&,<1#."().".." * 2<1"#&-.#('& -#.1-/,0 #-.&&31#." )/&  % & - *,.  3  -"&&)1 !,))0 ?  '& - "  .,#(!/&, -/@ ! (#.&*&. /.." '& -"&)(!<(,,)1.)(!/ @&#% -.,/./, #( (. )( ."  0 (.,& -/,  ? "   '& -C -/! (#.& *&.  1- ,) 1#." *,&& & -# -(.1)#-.&&) -<1"#& #('& -#.1--&#!".&3/,0 /*1,-#-.&&3 1#."()% &?"  '& -/@! (#.&*&. 1-,)(-"),.<1#."." #-.& ',!#(.))." <(#(." '& #.1-(,,)1.#.-- /.1# (#(!-#./,0  /*1,-H#!/, N?QI?  )/,'#(* ,'/..#)(-) *#&-*#( -1 , # (.# # H& N?MI? (#.#)(1  )/( .". ."  "#( '/, , .,)&. ,& *#& -*#(  1- -)' .#' - *, - (. ( -)' .#' - - (. #( -* #' (- )  ? "#- 0,#.#)( -")1  () )(-#-. (. *.. ,(#(-* ),.#' <(1-." , ), )(-# , .) *)&3'),*"#1#."#(

38 Te Paki

."#- -* # -? "  .",  )." , *#& -*#(  )'#(.#)(- 1 ,  /(0,3#(! ')(! #(#0#/&-) ." -' '),*")!,)/*#(&/#(!'& -( '& -<1"#"&&)1  ." ' .)   '." ? , - ( F- (  )  -#2 *#& -*#( - 0,#  )(-#-. (.&3 ')(!."  )/,-* # -= ), @ '/,, .,)&. ,&*#&-*#( <'#@ '/,*,)&. ,& *#& -*#( < ), @.## -/* ,#), *,)&. ,& *#& -*#( < '#@.## -/* ,#), *,)&. ,& *#& -*#( < "#(@.## #( ,#), -/@*#& *,)&. ,& -*#( < ( "#(@ .###( ,#),-/@*#&, .,)&. ,&-*#( H& N?M9#!/, N?NI?)! ." ,." -  *,)0# )'#(.#)() ",. ,-.. -.".)(-#-. (.&3(, &#&3#!()-  /&.-(( ,/&.-) ."  )/,.2 ()/(. , ?  '%! ,&&

. ,)'OMU/&.<(<MLP&,! *, @/&.-(MQP)." , /(# (.# #  -'&& $/0 (#& - 1 ,  /-  #( ' .,# (&3-#- .) )'*,  -"*  ( -#4   .1 ( - 2 - ( ')(! ."  )/, .2? (&3 ),  # 1  #( (3 -#!(# #(.-#4 # , (  .1 (." ' (-) /&.'& -( '& -H.@. -.< WL?LLLM<& N?NI?&.")/!"'& -( '& -) ." )." ,*/..#0 -* # -# (). # , -#!(# #(.&3 #( -#4  1  ().  .". ." ,  1 ,  +/(.#..#0  # , ( -  .1 ( - 2 -? ), #(-.( < ."  "#(@ '/, )  '&  -* #' (- 1- &)(! , ."( ."  '& -C H#!/,  N?OI? /&.  '3 1 && # , /. ."#- )/& ().   . -. 1#."." *, - (.-'*& ? Table 2.2. Comparisons using t-tests of male and female mean sizes for three morphological traits in four species of cave wētā (cave wētā) from Te Paki Ecological District. Bold = P<0.05. Standard errors for each mean value are in brackets. Mean size in Neonetus Pachyrhamma n.sp Pallidoplectron Talitropsis n.sp mm. variegatus (TPA) (TPB) n.sp (TPC) (TPD) (Standard Male Female Male Female Male Female Male Female Error) n=115 n=182 n=53 n=51 n=5 n=2 n=5 n=6

Pronotum 2.690 2.980 3.906 4.358 3.190 3.275 2.450 2.384 (0.023) (0.023) (0.185) (0.198) (0.107) (0.215) (0.088) (0.084)

Hind femora 8.630 8.927 12.610 13.293 16.700 10.040 7.120 7.409 (0.090) (0.074) (0.717) (0.614) (0.175) (2.15) (0.362) (0.467)

Hind tibia 9.346 9.422 13.750 14.508 12.500 11.720 6.880 7.665 (0.099) (0.078) (0.740) (0.638) (0.196) (3.45) (0.423) (0.679)

39 Chapter 2



 Figure 2.2. Dorsal view of cave wētā showing the location of apical spines. S1) Fore femur prolateral, S2) Fore femur retrolateral, S3) Mid femur prolateral, S4) Mid femur retrolateral, S5) Hind femur prolateral, S6) Hind femur retrolateral, S7) Fore tibia superior prolateral, S8) Fore tibia superior retrolateral, S9) Fore tibia inferior prolateral, S10) Fore tibia inferior retrolateral, S11) Mid tibia superior prolateral, S12) Mid tibia superior retrolateral, S13) Mid tibia inferior prolateral, S14) Mid tibia inferior retrolateral, S15) Hind tibia superior prolateral, S17) Hind tibia inferior prolateral, S19) Hind tibia inferior sub-apical prolateral, S21) Hind tibia superior sub-apical prolateral. Spines S16, S18, S20 and S22 match and appose to S15 S17, S19 and S21 on the retrolateral spine.

40 Te Paki

 Figure 2.3. Four species of cave wētā identified from Te Paki Ecological District, Females on left, males on right.



41 Chapter 2

/% 3 )'*,#-)(. -.--")1 ." , 1 , -#!(# #(.# , ( -#(-#4 )  &&.", ' .,#0,#& -' -/,  .1 ((H@0&/ WL?LLLMI<(  .1 ((H@0&/  WL?LLLMI? ( && - -  1- ."  &,! -. 0 ( .")/!"()/&.-) ."#-.2)(1 , *, - (.#(." -'*& ? )1 0 ,<-#4 &)( 1- ()., &#&  ),#-.#(!/#-"#(!&&.2(." /- ) -*#( )'#(.#)(",. ,- (-/@! (#.&*&. -"* 1- -- (.#&H& N?OI?

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

Table 2.3. Adjusted P values from Tukey HSD test comparing mean lengths of pronota, hind femura, and hind tibiae of four species of cave wētā from Te Paki Ecological District pitfall traps. Significant differences are underlined (P<0.05) or in bold (P<0.0001). TPA (Neonetus variegatus), TPB (Pachyrhamma sp.), TPC (Pallidoplectron sp.), TPD (Talitropsis sp).  Pronotum Length Hind Femur Hind Tibia

TPA vs TPB <0.001 <0.001 <0.001

TPA vs TPC 0.543 0.297 0.025

TPA vs TPD 0.174 0.095 0.014

TPB vs TPC 0.014 0.073 0.284

TPB vs TPD <0.001 <0.001 <0.001

TPC vs TPD 0.089 0.023 <0.001

42 Te Paki





Figure 2.4.

Principal component analysis of Te Paki cave wētā morphology. A) All taxa and juveniles using spine and metric data, B) Four diagnosed taxa using spine and metric data (data from unidentified juveniles excluded), C) Four diagnosed taxa using only metric data. Juveniles (Juv.) were specimens less than 10mm long.

43 Chapter 2

 Figure 2.5. Subgenital plates of the four cave wētā species identified from the Te Paki area. Row A: TPA (Neonetus variegatus) female ventral (left), male ventral (centre), male lateral (right). Row B: TPB (Pachyrhamma sp.) female ventral (left), male ventral (centre), male lateral (right). Row C: TPC (Pallidoplectron sp.) female ventral (left), male ventral (centre), male lateral (right). Row D (Talitropsis sp.) female ventral (left), male ventral (centre), male lateral (right).



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44 Te Paki

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#!(# #(.#(. ,.#)(-) -* # -1#."')(."("#..1 ,  . . <(1  ().  -#!(# #(. 0,#.#)( ')(! -#. - -/" .". 0  17.5 /((  1#."#( "#..-1-#( &/ ( 3-#. ?), 2'*& <')-.) ." . '*),&0,#.#)(#( (/' ,- ( # , ( - ')(! "#..- ** ,- .)    ,#0  ,)' ."  *#(  ), -.-#. -. ,,)#(.("%*/%/?"/-0 17.5/(( . %# , +/#, -# (.# #.#)((- *,. ., .' (.) ."  )/,-* # -?

45 Chapter 2

 Figure 2.6. Mean monthly counts of four cave wētā species in pitfall traps in Te Paki Ecological District between July 2006 and June 2007. A) TPA, B) TPB, C) TPC, D) TPD. Dashed line = pine forest, dotted line = native forest and full line = shrubland. Note that y axis scales differ.



+"!" ,  , 0# 1  2#-.#(! -,#*.#)(-)  1 &("*"#)*"),# #(, &.#)( .) ."  *#& -*#(  .< -/@! (#.& *&.  -"* < )." , '),*")&)!#& ..,#/. - H)&)/,<*.. ,(<"#,- .I(&).#)(#( ),'.#)(?"#- (& /-.))( #,'."  ! (/-.)1"#" "'),*").3*  &)(! <(#()( - ." ')-.&#% &3-* # - (' ? "/- < ."  ')-. /((. -* # - #( ."  -'*& < 1- ')-. &#% &3 "!'(&)%'(&,/(( ,0)(.. (3&<MTTT),#(!.)."  -,#*.#)()  ")*, HMUNOI? "  &,! -. -* # - < )  1"#" () /&.- 1 ,  )/( #( ."  *#. &&.,*-<(, #&3 --#!( .),% )(-#-. (.1#."(/&.*#, ) ,% *, 0#)/-&3)&& . .)! ." ,#(")&&)1&)!.(/1")H))% . &?NLMLI?&.")/!"." ,%  ,)'." *, - (.-./31 , ()./&.-<." #, -*#( )/(.-1 , )(-#-. (.1#."%()1(/&.,% -* # - ,)',)/(  1  &(?  " (/' ,)/- -'&& , .,) ( *,) &. ,& -*#( - )( ."  "#( '/,-#'#&,.) "# '%"!#",-<MUQT? )1 0 ,<1-'/"-'&& , -* # -."(." .",  "# '%"!/,, (.&3 -,# H#",-<MUQT<MURLI? 1-')-.-#'#&,.) '%"#&&)�,<MTTN<"0#(!)'#(.#)() -*#( -<

46 Te Paki

)&)/,(!&)--3** ,( 0 ,3&#%  '%"#&&& "')�,< MTTN /. ."  -/@! (#.& *&. - )  '& - ( '& - & ,&3 #-.#(!/#-"  #. ,)' .". 1# -*, (/((.'#(&(-* # -0

 Table 2.4. Generalised linear model (Poisson, link: log) to explain the variation in cave wētā abundance (response) captured in pitfall traps in Te Paki, New Zealand. Terms added sequentially (first to last). There were eight sites situated in one of three habitats (native forest, pine forest and scrub). Four cave wētā species were identified and juveniles were excluded.

Df Deviance Resid. Resid. Pr (> Chi) Df Dev NULL 431 6461 Species 3 3177 428 3284 < 0.001 Month 11 336 417 2948 < 0.001 Habitat 2 1260 415 1688 < 0.001 Site 6 448 409 1240 < 0.001 Species:Month 33 260 376 979 < 0.001 Species:Habitat 6 91 370 888 < 0.001 Month:Habitat 22 105 348 783 < 0.001 Species:Month:Habitat 66 86 282 697 0.049

  

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47 Chapter 2

(0#,)(' (..",)/!")/.." #,&# 3& <) . ()'#(!)/.#(.)."  ), -..) ),!  )( -/,,)/(#(! 0 ! ..#)( . (#!". H))% . &? NLML= #",- MUQPI? /,." ,'), <#(0 -</&.-($/0 (#& -"0  ()- ,0 .)! ." ,H#",- MURNI? "  *, - (. -./3 1- &#'#.  3 ."  -#(!&  )&& .#)( ' .") /- ? #-)0 ,3)  /&&0 17.5#0 ,-#.3<*)*/&.#)(-.,/./, ( "0#)/,&-"# .- 1)/&*,)&3  -."# 0 /-#(!)'#(.#)() -'*&#(!-.,. !# -?

.1-*)--#& .)*& ."  )/,0 17.5-* # -#(.)! ( , 0 (.")/!"'(3 2#-.#(! ,"*"#)*"),#  -,#*.#)(- ,   ## (.? "!'(& 1- -.&#-"  3 ,/(( ,0)(.. (13&#(MTTT</."#-),#!#(& -,#*.#)(#-, )!(#- - #(! ,." ,0!/ 1#.", -* ..)-* # -#!()-#-H")*,MUNOI?  -,#*.#)() 0 )%'(&3")*,HMUNOI" &* , )(#& # , ( - .1 (,/(( ,0)( .. (13&C- ),#!#(&  -,#*.#)( ( .". )  /..)( HMTUSI ( ."  -* #' (- 2'#(  3 ")*, HMUNOI "#'- & ? /..)(C-  -,#*.#)( )  0 )%'(& #- -  )( -* #' (- ,)' /%&(< 1" , - ")*,  &# 0  .". ,/(( ,C-  -,#*.#)( )  0 )%'(& 1- &)- , .) /..)(C- 0 # "&(& ,)'  &&#(!.)(? )&&)1#(!")*,C- -,#*.#)(<0)%'(&#-." ')-.&#% &3(#.  ),."  -* # - ,)' %#)&)!#&#-.,#.?

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48 Te Paki

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

,#.#)( #( ."  *, - ( F- (  )  -*#( - ( ."  -#4  ( -"*  )  ."  0  17.5 , 0 &  ")1 "&& (!#(! # (.# #.#)( ), ."#- '#&3 (  ? (#.#&&3 #(#0#/&-) -#'#&,-#4 1 , **, (.&3#(#-.#(!/#-"& /.&)- ,#(-* .#)( )  (/' ,)/- 0  17.5 #( ."    %# -'*&  , 0 &  .". -*#(  )/(.- &)(  1 ,  -/ ## (. .) #-.#(!/#-" )/, -* # - H1" ( /&. ), ( , /&.I? " (  &#(! 1#." ."  **,)2#'. &3 QQ  -,#  -* # - (.#)(1# < *#& -*#(  )'#(.#)( 1#&& &#% &3 (  .)   )'#(  1#."  .#&- )  ! (#.&#< -#4  (

49 Chapter 2

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50 Te Paki

#&(!!

. & -. )/, -* # - )  ( '# 0  17.5 ,  -3'*.,# #( ."    %# , !#)(? /&.- )  "!'(& )%'(& 1 ,  /!". #( && .",  "#.. .3* - -/,0 3 < &.")/!" ." 3 1 ,  ()/(. ,  '),  ) . ( #( *#(  ), -. ."( (.#0  ), -. ), -",/&(? ()(.,-.</&. '%"#&&1 , ()./!".#((3) ." -",/&( .,*-&.")/!"." -'&&-'*& -#4 ) ."#-.2)(*, &/ - )&)!#&#( , ( -?

,* *./,  ,. - '3 (). /,. &3 , & . .,/  /(( -< - # , ( - #(&/#(!.#0#.3& 0 &-<.,**#(!0/&( ,#&#.3("#.." . ,)! ( #.3, &#% &3 .) #(. ,. .) 3# & #(& .,**#(! ,. -? )1 0 ,< *./,  ,. - ,  -* # -@  * ( (.<(-* # -*./, ,. -1 ,  )/(.) * ()().""#..( .#' ) 3 ,?(#(. , -.#(!)- ,0.#)(#-.".)(&3#(,% -*?HI# 1  -   -*,#(!.#'  #(, -  #( ."  *./,  ,. ? "#- -/!! -.- .". 1" ( "*"#)*"),# , )'#( #(.)-#(!& /(#./,#(!((&3-#-<#'*),.(. #( ),'.#)(#-&#% &3.) &)-.?), 2'*& <-./# -#' ., ),#(!."  .- )  *, .), )(.,)& )( 0  17.5 /((  '#!". *,)/  '#-& #(! , -/&.- 1" ((/' ,)/--* # -, ., . -)( H/-)  .&?NLMOI?

( )(-# ,#(! , !#)(& #)#0 ,-#.3 ( #.- )(- ,0.#)( ."  +/&#.3 )  ')# #  "#..- ( ." #, *#.3 .) -/**),. (.#0  -* # - #- #'*),.(.? ( ')-. #(-.( -< (.#0  -* # - 1)/&   --/'  .)   '),  -/ -- /& #( (.#0  0 ! ..#)(< /. -./# - ,  #(, -#(!&3 -")1#(! .". '(3 (.#0  #(0 ,. ,. - (.",#0 #(#-./, (')# # "#..-#(&/#(!/,(, -H ?!?&(")( .&?NLMM=,)% ,")  .&?NLLMI?#."#(!(&%'*&(..#)(-. %#1  )&& . ." -'  )/,-* # -/. . . -#!(# #(.&3"#!" ,(/' ,-) .1) 0  17.5 -* # - )'*,  .) ."  (.#0  ), -. ), -",/&( "#..< *,)0##(! 0# ( ) ." #'*),.(.,)& *#(  ), -.("0 #('#(.#(#(!*)*/&.#)(-)  ( '##(0 ,. ,. -?







51 Chapter 2

 

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



#."#'*,)0 -3-. '.#, -)&/.#)(<." .,**#(!.*,)0# #(-#!".-#(.)."  #-.,#/.#)() -* # -(#(#. --* # --* # #-*.#&<. '*),&("#.. # , ( - #( , &.#0  /(( -? "#- #(#. - .". /./,  -./# - )  #)#0 ,-#.3 1#&&   '),  #( ),'.#0  ( ' (#(! /&  /-  #-.#(. .2 1#." #-.#(. )&)!# -1#&& ")(.,#/. .)(&3- -?#-./,  (0#,)(' (.-** , ()..)#'*  ." #-.,#/.#)() "!'(&)%'(&.(*#(  ), -.** ,-.) *,)0# 0&/& "#.. ),.& -.-)'  ( '##(0 ,. ,. -? .#-<")1 0 ,< & ,.".-#(!& .,**#(!, !#' 1#&&()./,. &3*./, &&#(0 ,. ,. -), 0 (&&,"*"#)*"),##0 ,-#.3?

 

52 Te Paki

    &(")(</-. ,# <&,#."<"),* NLMM?'*&#(!#(#! ()/-!,)/(@ �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

53 Chapter 2

 -<.. <#!&#()<,/( NLMO?,.")*. ,-* # - #& )(&#( <  ,-#)(Q?LFQ?L<HNPFLRFMOI?"..*>FF,.")*. ,?* # -#& ?),!?  ,",NLML?0)&/.#)() ! (#.&#>." ),# -< 0# ( <(( 1#, .#)(-? !'MOT>QGMT? #(!,( NLLO?#)#0 ,-#.3<)(- ,0.#)(<(." K2)()'##'* #' (.K? $('"!&%)'"!1 %!!%&*'%"&,&' &MO>PQMGPQU? #-MUUU?)/,( 1-* # -) !#(.17.5<!%H,.")*. ,> ()-.)-.)'.# > #(,##( I ,)' 1 &(? "(%! "'",  "',"* !NU>OLSGONP? )& ,! <, 1#%?NLMM?2*&),#(!*"3&)! )!,*"#)(!,/ ( #( )(.#( (.&#-&(-3-. '? !&'&N>ORUGOUU? ),)(NLML0* ! !)!'"%,"")%&',?)&/' N> #(!)' (#'&#" .)(."<23-)4)< "() )--#&-0(. ,/,3(#0 ,-#.3 , --<",#-."/,"< 1 &(? #&.3 < , (& ( ,NLLL?/(&#(#.),.2- & .#)( ),')(#.),#(! )-3-. '" &."?" " "!&%)'"!UN>MTQGMUS? ) '((<( ,- (NLLO? -*)(- -) (.-.)#-./,( #(/-.,&#< 1#."*,.#/&,, , ( .) /(.#)(&!,)/*-0(&'% " ",NT>PPPGPRP? /..)(MTUS?" . ()* &'.# )  1 &(?%!&'"!&"*  ! !&''('NU>NNOGNPL? , ' (<)&1 && <,1#( </,*"3<)--<($3(MUUO?  ,, -.,#&,.",)*)-- '&! ->." #,/- #()(- ,0.#)(*&((#(!? "!&%)'"!" ",S>SURGTLT? /( ,<"1<# , NLLQ? (#! ()/-#)#0 ,-#.3(&(/- @1".) 2).#*&(..#)( ), -.-)(.,#/. A* ! "(%! ""%&'%,PU> NLGNR? ) < - MUUN?-/,0 3) #(0 ,. ,. -#(-,/&(-( ), -. 1% K-3< 1 &(?* !!'" " "&'MQ>ROGSM? %-( ( </#&&/' < #(.< ! (, <#("#(<K ,<#'*-)( < )&3')-<. 0 (- <!( , NLMO? !(>)''/(#.3 )&)!3 *%! ?#)%&"!6041540''#/2201 #%"'0"%2#7)!?

54 Te Paki

#",-MUQP?). -)( ))(((#&#-'#( %"#'(& %&% , MTRUH"*"#)*"),# <,.")*. ,I?%!&'"!&"'", "'," * !TN>SOOGSOS? #",-MUQT? 0#-#)() ." "*"#)*"),# H,.")*. ,I)  1  &(?,. ?"  (/- %"#'(&&% ,#(." ,#.#-"/- /' H.? #-.?I)&& .#)(?%!&'"!&"'", "',"* !TQ> PRQGPSL? #",-MUQT? 0#-#)() ." "*"#)*"),# H,.")*. ,I) ( 1  &(*,. ?" ! ( ,,% ,/(( ,( "# '%"!(?!? %!&'"!&"'", "',"* !TQ>RUQGSLR? #",-MUQU? 0#-#)() ." "*"#)*"),# H,.")*. ,I)  1  &(@,.?"  (/- "# '%"! /..)(<MTUS?%!&'"!&"' ", "',"* !TS>OMUGONS? #",-MUQU? 0#-#)() ." "*"#)*"),# H,.")*. ,I)  1  &(@,. ? 1* # -K &)(!#(!.)." ! (/-,%  ,/(( ,<MTTT?%!&'"!&"'", "',"* !TS>ONUG OON? #",-MURL? 0#-#)() ." "*"#)*"),# H,.")*. ,I)  1  &(,. ?" "*"#)*"),# ) ." #*/0 -?%!&'"!&" '", "',"* !TT>NQUGNRP? #",-MURN? #(! "0#)/,( ( '# -) "*"#)*"),#  H,.")*. ,I ,)'#.)')0 -< 1 &(?%!&'"!&"'",  "',"* !."" ",N>MNMGMNU? /-) <1 .**&  < ,,3</((NLMO? .-) -*.#&&3 2. (-#0  )(.,)&) #(0-#0 ,.-)(/(( ) (.#0 #(0 ,. ,. -#(#(&(  1 &(), -.-?"!&%)'"!" ",NS>SPGTN? &')( MUPT? 1! ( ,<-* # -<(, ),-) ,.")*. , ,)'." ",  #(!- -&(-< 1 &(?"%&"'( ! !&''('- (&( O> OLMGOLS? " ,& 3MUUR?),*")&)!#&0,#.#)(#(." -" &&-)  "&', (&-* # - H-.,)*)>/&#'/&# I#( 1 &((#'*&#.#)(- ),." #, )(- ,0.#)(?* ! "(%! """ ",NO>SOGTN? ),  'NLMO?>&(!/! ( (0#,)(' (. ),-..#-.#&)'*/.#(!?

55 Chapter 2

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

56 Te Paki

    

  "  )0  #!/,  #- ( ' (' (. )  #!/,  N?Q .". #- #( ."  "*. , ( 1- */&#-"  #( ."  $)/,(& ,.#& ?  . , */&#.#)(  '#-.%  "-  ( *#%  /* 1#." *").)-  #(! *&  #( ."  #!/,  #( ."  1,)(! ), ,? "  ( 1 0 ,-#)( ),, .&3*& -." *").)-) '& &. ,&(0 (.,&0# 1 ),(#(." ,#!". *& -?" - " (-1#." ,)/(#(." ),#!#(&0 ,-#)(?  

57 Chapter 2



58 Te Paki

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                   )      & &       &        *  "       %    " !    '               !      %            $   ! %)Josephine Fitness   *%!# "&"#'$!#)Dr Steve Trewick   !&$$#&%"&% &## ) Fitness, J. L., Morgan-Richards, M., Ball, O.-P., Godfrey, A. J. R., & Trewick, S. A. (2015). Improved resolution of cave wďtă diversity (Orthoptera: Rhaphidophoridae): ecological implications for Te Paki, Far North, New Zealand. New Zealand Journal of Zoology, 42(1), 1–16.  ("%#$%&$!#)Chapter 2    # x          !  # 90% & x            #

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59 Chapter 3

60 Three Locations

Chapter 3

Comparing species level diversity of cave wētā among

three locations of New Zealand



























 

61 Chapter 3

  

62 Three Locations



 

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63 Chapter 3

-* # -<1"#", %()1( #,&31 &&?".#(&/ --#.2)()'3) ." !,)/*.)  /- H0(! &#-.# .&?<NLMRI? 0 &)*#(!13-.)#-.#(!/#-" .1 (-#'#&, -* # - "-  (  '$), !)& ), '(3 #)&)!#-.- )0 , ."  *-. 1   -? ),*")&)!#& ",. ,- )'#(  1#." ! ( .# . ,  ()1 1# &3 /-  .) #-.#(!/#-"  .1 ( &)- &3 , &.  -* # -? ( .2 .". ,  '),*")&)!#&&3 # (.#&<',% ,-(  -* #&&30&/& (."#-**,)"#-)'')(&3 , ,,  .) -  ,)#(! H ?!?  ,.< . &?< NLLP< /. -  ),#.4 9 # ,)< NLLP=, 1#%<NLLTI?

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64 Three Locations

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

,   2. ( ."  1),% )  ."  *, 0#)/- "*. , H#.( -- . &?< NLMQI &))%#(! . #!()-#-(# (.# #.#)() -* # -? (." *, 0#)/-"*. ,  0 &)* -)'  -# '),*")&)!#& ' -/, - .) -- -- ."  '),*")&)!#& # , (   .1 ( # , (. !,)/*- )  #(#0#/&- (  . ,'#(  1" ." , -* # - )/&   #-.#(!/#-" /-#(!'),*")&)!3?2*(#(!."#-**,)".)# , (., .". '#!".)(.#('), (# , (.-* # -< /- '#.)")(,#&- +/ ( -.) . -.-* # -"3*)." - -(*,)0# #.#)(&#( ),'.#)(.) 2*&), #)#0 ,-#.3 ( -.&#-".2)()'3? (."#-- -'*&#(! ),.1#&&.,! ..", ),." -&(  1 &(, !#)(-1#."-#'#&,! ( ,& (0#,)(' (.&)(#.#)(-.".1#&&&&)1 )'*,#-)() -* # -& 0 &#0 ,-#.3')(!&).#)(-?

65 Chapter 3

 ),  ."  ,,#0& )  "/'(-< &)1&( ), -. 1- 2. (-#0  #( ),." -&(  1  &(<1#." .#0 &3)(.#!/)/-"#...".)/&"0  (& #(#0#/&-)  '(3-* # -.)"0 ,(! - 2. ( .",)/!")/.),." -&(?(& --." , 1- &)& *..#)( ), 2., ' &3 &)1 ,. - )  ! (  &)1 1  1)/& 2* . &#..&  , !#)(& '#,) ( '#-'< ( -* # - 1)/&   1# -*, ? "#- *, #.- .". $ (. , !#)(- #( ),." -&( 1#&& -",   - . )  .2? &. ,(.#0 &3< ()/(. ,#(!-- '&! -/(#+/ .)-* # #, !#)(-1#&&-/!! -.." -*.#&-&  )0 , 1"#" *,.#.#)(#(! )/,- H#!/,  O?MI? 0 ( 1" ,  # , (. , !#)(- "0  #-.#(. .2< #. 1)/&   2* .  .". $ (. , !#)(- 1)/&"0 , &.  -* # -?). -.*, #.#)(-)/.." , &.#)(-"#* .1 (-* (#0 ,-#.3  /- )'#(.#)() '),*")&)!#&(! ( .##( ),'.#)( ),-'*& ) 0  17.5 ,)'.", , !#)(-) -)/." ,(),." -&( 1 &(?



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

66 Three Locations

0  17.5 -* #' (- 1 ,  ).#(  /-#(!  )'#(.#)( )  "( )&& .#(! . (#!".< 2.,.#)( ,)',))-.")& -/,#(!." 3(*#. &&.,*-?#. &&.,**#(! 1-,,# )/./-#(!-.(,' .")1#.".,*-)(-#-.#(!) *&-.#,)/( )(.#( ,HMML''#' . ,(UL'' *I/,# #(.)." !,)/(?" ,#')  ."  )(.#( , 1- &/-" 1#." ."  !,)/(? ,)*3& (  !&3)& 1- /-  - ."  *, - ,0.#0 (." )(.#( ,-1 , "&  #&& 1#."#.?.#()0 ,1-*& )0 , ." .,*)/.O')0 ." !,)/(?"#-&&)1 #(0 ,. ,. -.)')0 /( , (#(.)." .,*1#.")/. #(!#'*  /.*, 0 (. &,! ,(#'&-<,#((  ,#- ,)' (. ,#(! #(.) ."  .,*? #. && .,*- 1 ,  )&& .  ( , - . 0 ,3 ')(."/,#(!." -/'' ,) NLMMFNLMN<1"#& ." #(- .-1 , .#0 ?, - ,0  0  17.5 1 ,  -),.  )/. )  ."  !&3)& ( *&  #( MLLV ."()&? 3.#'  )&& .#(! #(0)&0  &# .#(!  3#(! &)!- 1" ,  0  17.5 ) . ( ,))-. /,#(! ."  3 ), - ,"#(! ")&&)1- #( ., - ( (%-? ,.# ##& .,  17.5 0#.# - H17.5 '). &-I1 , - ," 1" , 0#&& H, 1#%9),!(@#",-<NLLLI?

67 Chapter 3

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68 Three Locations

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

'+'%'"!. # '"!!&$(!!  ")&  ! ()'#  2.,.#)(- 1 ,  * , ),'  /-#(!  B-&.#(! )/.C *,).))& H/((/%- 9 & -< MUURI  -#!(  ), , -" .#--/ < /. /-  -/ -- /&&3 ), *, - ,0  ,.")*. ,( .#--/  H, 1#% 9 ),!(@#",-< NLLPI?), " -'*& <  JMQLL -  *#, H*I ,!' (. -*((#(! ')-. )  ."  3.)",)'   )2#-  H I! ( ) ." '#.)")(,#&! ()' 1-'*&# # /-#(!*)&3' ,-  "#( , .#)( HI (  )'#(.#)( )  /(#0 ,-& #(0 ,. ,.  *,#' ,-> MPULH)&' ,<&%< ) "< /.4<MUUPI<M@ @MSMT<M@@NMUM<M@ @ NMUQ( N@@OLMPH#')( .&?<MUUPI?" , *)--#& , *, - (..#0 )  " *" ().3*  ,)' "&).#)(1-- +/ ( /.().&&-* #' (-*,)0#  .".)/& -/ -- /&&3'*&# # ?  / -- /&*,)/.-1 , *, *, /-#(!." FM#! -.*,).))&H ),*?< & 0 &(<  I ( - +/ (  1#." #!3  " '#-.,3 ( (   OSOL

69 Chapter 3

! ( .# (&3- , H**&#  #)-3-. '- (?< ,&-< I? /& ).#  - +/ ( - 1 ,  -- '&  ( &#!(  /-#(!  ( #)/- 0T H ,-  . &?< NLMNI? ) #(- ,.#)(-F & .#)(-1 ,  . . (- +/ ( -1 , .,(-&. .))( #,'.". ." ,  1 ,  () -.)* ))(- ), ,'  -"# .- .". 1)/& #(#.  ."  *, - (  )  (/& ,*,&)!-?  *&).3* -1 , &/-. , - )(- +/ ( -#'#&,#.3H *#,1#- #-.( -I /-#(! ."   #!"),@ )#(#(! &!),#."' H#.)/ 9  #< MUTSI #'*& ' (.  #(  ( #)/- 0T?  1  &( 0  17.5   - +/ ( - --)#.  1#." *, 0#)/- -./# -1 , 0#&&  ),."  )&&)1#(!-* # -> '%"#&&& "''H)& ,! . &?< NLLTI<  '%"#&& '. 0 %&&%(%& H, 1#%< NLLLI< %"#'(&-*? H))% . &?< NLMLI= %"#'(&  %.  "# '%"! '(%!%.  "# '%"! & # +H&& !,/# .&?<NLMLI?  * # - "3*)." - - -  )( '),*")&)!3 H-I 1 ,  . -.  1#." ."  '. .?#,1#- ! ( .##-.( -1 , &/&. (/- .)-- --*). (.#& # (.# #  3 '),*")&)!3?  * # -  &#'#..#)( .))&- #(  ( #)/- 1 ,  /-  .) &/&.  ."  0 ,!  ! ( .# #-.(  1#."#(  &/-. , H& I< ( ."  ' ( ! ( .# #-.(   .1 ( ."  ( , -. "*&).3*  & ? "  ,.#)) 1#."#((  .1 (&/-. ,#-.( -H (.,F (. ,I( /- .)# (.# 3D!*-E .1 (."   - +/ ( -#'#&,#.3.".'#!".),, -*)(.)-* # -& 0 &# , ( -H  ,.< .) %& < . &?< NLLPI? 3 ,#. ,# ), )( #,'.#)( )  -* # - , +/#,  *#,1#-  ! ( .##-.( -.) &)1H& --."(JPVI()(),( ) '.&/-. ,- ),-* #' (- ,)'." -' '),*")&)!#&? ()'),*")&)!#&# , (  )/&   # (.# # < '. - +/ ( - # , (  )( #.- )1( 1- (). ()/!" .) -.&#-"/(#+/ ? '),*")&)!3# , /.'.- +/ ( -1 , -#'#&, ."#- ' (. '3 -* # - "3*)." -#- 1- (). -/**),. ?  - +/ ( - ), )/, */..#0 -* # -."." ()&& .  ,)'&&.", , !#)(-1 , /- .)#( , "*&).3* ( .1),%-/-#(!." '#(#'/'-*((#(!**,)"H( &. .&?<MUUUI /-#(! ."  -) .1,   H #!" 9 ,3(.< NLMQI ( )  3 )&& .#)( &).#)(? 

70 Three Locations

   (.).&<MRO#(#0#/&/&.0 17.51 , #(&/ #(." (&3-#-?* #' (- 1 ,  .&)!/  #(.) ."  ") (#2 )&& .#)( #( ."  )&)!3 ,)/*< -- 3 (#0 ,-#.3? "  -'*&  #(&/  QS /&. 0  17.5 ,)' 1% C- 3< QR ,)' (1.8 ( QL ,)' ,(%#? "  /-  )  *#. && .,*- #( (1.8 ( ,(%##(, - ." -'*& -#4  ),." - , !#)(-<")1 0 ,'(3) ." 0  17.5 /!". 1 ,  $/0 (#& -? ( ."  *, 0#)/- "*. , H."#- ." -#-I $/0 (#& - 1 ,  -")1(.) # #/&..)-- --'),*")&)!#&&3#(.)-* # -(-), ().#(&/  #(."#--./3?                       $%    "   

Location Number of Number of Phenotypes OTUs

Hawke’s Bay 18 8 Manawatū 26 11 Taranaki 28 10 

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

71 Chapter 3

#,#(! '& - 1#." '& - 1- -# , ), -)'  .2 ."( )." ,- /  .) '#(), *#&-*#( 0,#.#)(?" ().3* -1#."-*#( .3* -ML(MM, &13- )/( 1#."*&. -O),OMH- 2 * ( (.=& O?NI?))." ,*" ().3* "- ." -  -*#(  .3* -< , #( ),#(! ."  #  .". ." -  ",. ,- ,  ), '&  ( '&  )  ."  -'  -* # -? ( -)'  - - ."  -*#(  .3*  H#? ? OI #- )/( #(  (/' ,) .2 ),)."'& -( '& -'%#(!." /- ) -*#( -#'*)--#& .) #-.#(!/#-" -* # - &)( ? *#(  .3*  O #- )/( #( .2 .". ** , .)   ,)' #-.#(.! ( ,<#(&/#(! "'"#(&("!'(&.-1 &&-.2)(-# , .)  ."  -'  ! (/-< H-  "*. , P3? " ,  '."#(! 1- (). *)--#& < '& - ( '& -* #' (-1 , & .#(- *,. -?"/-< ),-)' .2)(&3)( - 21- , *, - (. #(." .).&-* #' (-'*& - .H#? ?> <<<9I?

                 #

72 Three Locations

                         " "  "!     

ate S1 S2 S3 S4 S5 S6 S7 S8 S9 Sex S10 S11 S12 S13 S14 S15 S16 S17 S18 S19 S20 S21 S22 OTU Pl Location Spine type Subgenital Phenotype

P1 M HB, SB22 T11 0 0 0 1 0 1 1 0 1 0 1 0 1 1 1 1 1 1 0 0 1 1 A T, M P2 M HB, SB22 T22 0 0 0 1 0 1 1 0 1 1 1 0 1 1 1 1 1 1 0 0 1 1 A T P3 M T SB22 T17 0 0 0 1 0 0 1 0 1 1 1 0 1 1 1 1 1 1 0 0 1 1 A P4 M T SB22 T14 0 0 0 1 0 0 1 0 1 0 1 0 1 1 1 1 1 1 0 0 1 1 A P5 F HB, SB16 T11 0 0 0 1 0 1 1 0 1 0 1 0 1 1 1 1 1 1 0 0 1 1 A T, M P6 M M SB15 T1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 C P7 M M SB15 T20 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 C P8 M M SB15 T16 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 C P9 M M SB15 T19 0 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C P10 F M SB19 T1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 C P11 F HB SB19 T21 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 C P12 F T SB23 T3 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 B P13 M HB, SB3 T9 1 0 0 1 0 1 1 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 G T, M P14 M HB, SB3 T10 1 0 0 1 0 0 1 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 G M P15 M T SB3 T5 1 0 0 1 0 0 1 0 1 1 1 0 1 1 1 1 1 1 0 0 1 1 G P16 F T SB21 T9 1 0 0 1 0 1 1 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 G

73 Chapter 3

P17 F HB, SB21 T10 1 0 0 1 0 0 1 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 G T, M P18 M T SB2 T26 1 1 1 1 0 0 1 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 D P19 M HB, SB2 T21 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 D T, M P20 M HB, SB2 T3 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 D T, M P21 M M SB2 T7 1 0 0 1 0 0 1 0 1 1 1 1 1 1 1 1 1 1 0 0 1 1 D P22 F T SB9 T1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 E P23 F T SB9 T20 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 E

P24 M T SB10 T1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 E

P25 M T SB10 T20 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 E P26 M M, T SB11 T3 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 H P27 F HB, SB14 T3 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 H T, M P28 F T SB14 T21 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 H P29 M HB, SB12 T4 1 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 F T, M P30 M HB, SB12 T24 1 0 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 F M P31 F HB, SB6 T4 1 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 F T, M P32 F T SB6 T24 1 0 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 F P33 M HB, SB4 T3 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 B M P34 M HB SB4 T21 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 B P35 F HB SB20 T21 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 D P36 F T SB26 T3 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 D P37 M T SB1 T16 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 L

74 Three Locations

P38 F T SB1 T16 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 L P39 F T SB13 T6 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 K P40 M M SB5 T12 0 0 0 1 0 0 1 0 1 0 1 0 1 0 1 1 1 1 1 1 1 1 I P41 M HB SB5 T13 0 0 0 1 0 1 1 0 1 0 1 0 1 0 1 1 1 1 1 1 1 1 I P42 F M SB17 T12 0 0 0 1 0 0 1 0 1 0 1 0 1 0 1 1 1 1 1 1 1 1 I P43 F M SB19 T25 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 C P44 F M SB24 T14 0 0 0 1 0 0 1 0 1 0 1 0 1 1 1 1 1 1 0 0 1 1 P P45 F M SB19 T16 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 C P46 F T SB26 T18 1 0 0 1 0 0 0 1 1 1 0 1 1 1 1 1 1 1 0 0 1 1 M P49 F HB SB16 T22 0 0 0 1 0 1 1 0 1 1 1 0 1 1 1 1 1 1 0 0 1 1 A

P51 F M SB19 T8 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 C

P53 F M SB23 T21 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 U

75 Chapter 3

Table 3.3 Line drawings of the different types of cave wētā subgenital plates represent by specimens from three North Island New Zealand locations. Subgenital Plate Ventral View Lateral View Type

SB1

SB2

SB3

SB4

SB5

SB6

SB9

76 Three Locations

SB10

SB11

SB12

SB13

SB14

SB15

SB16

SB17

77 Chapter 3

SB18

SB19

SB20

SB21

SB22

SB23

SB24

SB26

SB27



78 Three Locations

$() )- ' $"%'( $ ) !() ),$ '#) %#$).%( ,)  "! # "! $"# )$ "  # $%')$#! )-$#$" *#) :;7 #!. > $' ? #+*!('%'(#)($"$)%#$).%(7#)$#!.'#),# (%"#(,('#)(%#).%*)$(#!%!(%#'#()( ,1)0,'*(*!!.%!#)("(*#)!%!)")7 $,+'4 ($"#+*!(#*#*(*!%#$).%,'$#')',(*#&*7$' -"%!4 ACC # FB= :$) "!(4 '$" '#  # ) #,)24 '(%)+!.;    ($') ()*"%. $+%$()$' *#!  #. $)' + ,1)0 (%"#-"#7(),$(%"#(:ACC4FB=;(!)!.'#) (*#)!%!)(%#'#)(%#$"#)$#(:>E4>E#>A4?A '(%)+!.; # )'$' ' $#(' ( (%') (4 :# '(%)+!.;7$")#"!(%"#$')'$',($*##". ("%!7





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

79 Chapter 3

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





80 Three Locations

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87 Chapter 3

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88 Three Locations

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93 Chapter 4



94 Neonetus

 

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95 Chapter 4

   

96 Neonetus

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97 Chapter 4

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98 Neonetus

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99 Chapter 4

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100 Neonetus

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101 Chapter 4

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102 Neonetus

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103 Chapter 4

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104 Neonetus

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105 Chapter 4

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106 Neonetus

107 Chapter 4

Figure 4.2 Phylogenetic relationships within the New Zealand cave wētā genus Neonetus inferred from MtDNA sequences (COI) using maximum likelihood (numbers above nodes are posterior probabilities, below nodes are bootstrap values). Main clades are labelled with either existing species name, Te Paki codes from Chapter 2, or letters referenced in the text. Clade G: Neonetus CIRRATUS sp. nov.; Clade C Neonetus UNCINUS sp. nov.; Clade B Neonetus PATERLONGIPES sp. nov.

[Following page]

108 Neonetus

109 Chapter 4



$%   #  ' Class: Insecta

Order: Orthoptera

Suborder:

Superfamily: Tettigoniodea

Family: Rhaphidophoridae

Genus: Neonetus

Type species: variegatus Brunner von wattenwyl 1888

Neonetus Brunner von Wattenwyl 1888 (Type species variegatus) variegatus Brunner von wattenwyl 1888 (syntypes: MHNG, Geneva Museum; NMW, Vienna Museum) pilosus [syn.] Hutton 1896 huttoni Chopard 1923 (holotype: missing)

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111 Chapter 4

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112 Neonetus

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114 Neonetus

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115 Chapter 4

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116 Neonetus

Figure 4.5 Lateral view off NeNeonetusonetus huttonihuttoni ((Left)Le male, (right) female. Scale bars 10mm.

Figure 4.6 Subgenital plates of Neonetus huttoni (Left) male lateral view, (centre) male ventral view, (right) female venteral view.

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119 Chapter 4

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120 Neonetus

  

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121 Chapter 4

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122 Neonetus

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125 Chapter 4

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126 Neonetus

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Reference

Chopard, L. (1923). On some New Zealand Cave Orthoptera. Transaction of the Royal Society of New Zealand, 54, 230–239.

Folmer O, Black M, Hoeh W, Lutz R, V. R. (1994). DNA primers for the amplification of mitochondrial cytochrome c oxidase subunit I from diverse metaxoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.

Hutton, F. W. (1897). The Stenopelmatidae of New Zealand. Transaction and Proceedings of the New Zealand institue, 29, 208–242.

Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C. (2012). Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28(12), 1647–1649.

Richards, A. M. (1959). Revision of the Rhaphidophoridae (Orthoptera) of New Zealand - Part V. The Genus Pleioplectron Hutton 1897. Transaction of the Royal Society of New Zealand, 87, 319–327.

Saitou, N., & Nei, M. (1987). The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4(4), 406– 425.

Salmon, J. T. (1948). New Genera Species and Records of Orthoptera from the Three Kings Islands New Zealand. Records of the Auckland Institute and Museum, 3(4/5), 301–307.

Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H., & Flook, P. (1994). Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87(6), 651–701.

Sunnucks, P., & Hales, D. F. (1996). Numerous transposed sequences of mitochondrial cytochrome oxidase I-II in aphids of the genus Sitobion (Hemiptera: Aphididae). Molecular Biology and Evolution , 13(3), 510–524.

129 Chapter 4

Trewick, S. A., & Morgan-Richards, M. (2004). Phylogenetics of New Zealand’s tree, giant and tusked weta (Orthoptera: Anostostomatidae): evidence from mitochondrial DNA. Journal of Orthoptera Research, 13(2), 185–196.

130 Pleioplectron

Chapter 5

Clarification of the genera Pleioplectron, Weta,

Miotopus and the species within.

   : !", !)-+ "'%., + - -1('(&" ,+")-"(', (  -!+ )+()(,'0,)",; ('(-!(0/+(',"+-!",-!,",!)-+-((',-"-.- (+&%).%"-"('80!"!0"%%0"- .+-!+)++/"0').%"-"('"'' ))+()+"-'0"%2,,"%#(.+'%;-,.!-"&-!)+()(,'&,0"%%  (+&-- ))+()+"-%2 ' &.,.& -%( . (, (+ /(.!+ ,)"&', 0"%%))'-8"'(+'0"-!+-"%R( -! '-+'-"('%(&&",,"(' ('((%( "%(&'%-.+(( ((%( "%(&'%-.+;

131 Chapter 5



132 Pleioplectron

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133 Chapter 5

   .--('KRSSC2),)",:2&('(-2)D   .--('KRSQ

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134 Pleioplectron

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135 Chapter 5

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

136 Pleioplectron

-( -!+0"-!,. '"-%)%-,!)( &%,-(",-"' .",!,)",( -!-!+ ).--"/ '+!  ' ; 

 "!/,(!*!&%&+#*,-*% #'!(%*!*)'!)4**('#&* & *  #% *  & '(&%&*+$ % !% $+( & @? )'!$%) &     %  4'!## )'!%&$!%*!&%)( !,%!)*!% +!) !% &#&+()3       6=1 (" #+72    6? 8 =D1 #+ 8 (%72      6<1 (72   6>8, (+" "'% -+-&'- (  0,/%"',!(.%+,.++-; .--('CKRSQD0,(++- -(,)+- #"'-(-!'0 '.,80!"!",",-"' .",! +(&  2 ",-"'-"/ )"% ,)"' (.'-8 ,. '"-% )%- ,!) ' %+  ,"3( .%-,; '-!(+-! ,%'8,)"&',!/-0()"%,)"',(' -!"+ (+ &(+0!"% "'"/".%,!/#.,-('C%O;KD; ! +%-"/%2 0",)+ (+-! ,%' ,)", ", (',",-'- 0"-! .--('>,

137 Chapter 5

!''0(.-! ,%',)",( !,'"'-" "; ",+ "'''0,)",,+"%(0;

 '('-+,--(-!,"-.-"('0"-!8-!,)"&',( -!&('(-2)" '.,   ! &- !)%(-2), -!- %.,-+ 0"-!"' -! "/+,"-2 (   (%%- +(& -! (.-! ,%'#    ", %,( &(+)!(%( "%%2 &(+ ,"&"%+ -(  -!'  "' -! ,' (  -! "' +"(+ ,.)"% ,)"',('-!!"'-""; '"-"('8-!&%,. '"-%)%-(   ", %-'",<%"$8,"&"%+-( #,  ' +,!(0'-(  &(+)!(%( "%%2 ' '-"%%2 ,( ,"&"%+ ' ,  !, !+('(%( "% )+' (/+  8 -!", ,)", ,!(.% '(0   "' , ;%-!(. ! .--(',+"-0( ,)", +(& '$,'"',.%+  (.''(/"' +(&1&"'-"('( &(+)!(%( 2(+&- /+"-"(' "' -! /"%% ,&)% -( "'"- -!- '$, '"',.%+ ' '+2 '-+.+2('-"'&(+-!'(',)",(  ;

138 Pleioplectron

Table 5.1 Specimens representing Pleioplectron and Miotopus from which morphological data were obtained. Measurements are in mm. Spine codes are as given in Figure 4.1. Pro = Pronotum length, HF = Hind femur length, HT = Hind tibia length, Location codes T = Taranaki, M = Manawatu, HB = Hawkes Bay      $ % & #' $"     4736      4: 4 3 4 4 3 7,35 45,64 47,:8 466      6 4 3 4 4 4 7,6< 46,;: 48,;: 476<      6 4 3 4 4 4 8,:: 4:,67 4<,:< 465:      6 4 3 4 4 4 8,74 48,6; 4: 4;6:      6 4 3 4 4 4 8,59 48,34 49,98 4;74      6 4 3 4 4 4 7,7< 46,39 47,56 793      6 4 3 4 4 4 8,57 47,<: 49,7; 795      6 4 3 4 4 4 8,77 48,46 49,48 :;6      6 4 3 4 4 4 7,86 45,4: 45,: . 6.      6 4 3 4 4 4 8,5< 48,6: 4:,49 .4<.      58 4 3 3 4 4 8,48 47,:6 49,96 478<      6 4 3 4 4 4 8,89 49,:: 53,79 479:      6 4 3 4 4 4 9,47 4:,;4 54,99 794      6 4 3 4 4 4 7,;: 48,47 4:,4< '798      6 4 3 4 4 4 7,65 45,4; 46,8; 4

139 Chapter 5

.4<.      6 4 3 4 4 4    678    #!, &,   8 4 4 4 4 4 7,< 4:,7: 55,89 635    #!, &,   8 4 4 4 4 4 8,48 49,6; 4<,<9 594:    #!, &,   8 4 4 4 4 4 8,78 4<,54 54,:5 589:    #!, &,   8 4 4 4 4 4 8,7: 54,:9 59,95 4773          5< 3 3 3 3 3 6,4 :,9; 43,55 4775          5< 3 3 3 3 3 6,6; 43,;4 45,49 4797          5< 3 3 3 3 3 6,65 <,8 43,:< 493<          7 3 3 4 3 3 6,6< 43,38 44,45 4;:          7 3 3 4 3 3 6,98 43,4 44,5 93          7 3 3 4 3 3 6,39 ;,<: 43,88 4

140 Pleioplectron

464          7 3 3 4 3 3    494          7 3 3 4 3 3    639          5 3 3 4 4 4 7,96 45,:; 47,9< 65;          5 3 3 4 4 4 6,66 43,<; 45,4< 694          5 3 3 4 4 4 7,7: 45,3; 45,<5 4;::       4 4 3 4 3 3 8,97 54,44 55,:: 4;:;       4 4 3 4 3 3 8,8: 4;,75 54,74

141 Chapter 5

Table 5.2 Summary of species and their authority, spine and subgenital plate description.

              

         

       

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142 Pleioplectron

#&  3 2'#(.#)( )  "*"#)*"),#  -* #' (- ,)' ."  ),." -&( "- # (.# # )'')(-* # -) -'&&:,%: ), -.1 &&#(!0 16.5="#--* # - #- )(-#-. (. #( '),*")&)!3 . && &).#)(-= /&. '& - ( '& - ,  ) . ( )/(.)! ." ,(." 3"0 ." -' -*#( )/(.D& P=LE=" -*#( - ,)' ." ),." -&(-* # -, -#'#&,.) /..)(A-),#!#(& -,#*.#)() ." ! (/-= " # , (  #(!.".2& # +"- )/,*#,-) "#(.##*#&-*#( -( ." ),." -&(-* # -D2(&"!E"-)(&3.", *#,-= '& -/! (#.&*&. - 1#."#(." ! (/- "# '%"!, -#'#&,:"0#(!&,! *&. >&#% -.,/./, .". '3 "0   .,#>* 2 ), &. &))%#(! * 2= &  -/! (#.& *&. - "0  -* # - #-.#(.#)(-=.- +/ ( -) 16.5 ,)'0,#)/-),." -&(-* #' (-D V E ,  -#-. , .)  "# '%"! & # + ,)' )/." -&( &).#)(- #(&/#(! (. ,/,3 D** (#2 ME=  "/-  )(-# , .". ."  1# -*,  ),." -&( '),*", *, - (.- /..)(A--* # - "# '%"!(&"!2 /- ) ." */#.3)  /..)(A- -,#*.#)() 2(&"!D(&%)  '& -E "0 ,  #( #. &)1<   "# '%"!(&"!#-&#..& -'&& ,."(2& # +D#!/, P=LE(#-&#..&  ,% ,#()&)/,=/! (#.&*&. )  '& -"-.", *)#(.-:1"#", ( ,&3#( &#( :()0#*)-#.),) /&. '& -"-)/.P- ,,.#)(-)(." &)1 ,',!#( .." .#*= ()(.,-.:'& -/! (#.&*&. -, -&#!".&3&)(! ,."(,)#( /-*#. -"* =" , #--&#!".% &)1(."  (., ) ." *&. = "# '%"! & # +-/! (#.&*&. ) /&.'& -"-.", *#&/&! -:." '#& , -/&.#(! ,)'% &=  HHH  

143 Chapter 5

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144 Pleioplectron

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

 " !<"#(3:,)1()&)/, ="),.- . )0 ,#(!)."./,!#. -(-. ,(#. -; -. ,/'*& ,)1()&)/,= & D#!/, P=NE< ,#&)(!:,)/(;,)1()&)/,;&)." #(- . ;-.3&#-"),.:(). 2. (#(! 3)(."  () ." -/>! (#.&*&. ;-/>! (#.&*&. #--1# - &)(!#(.,#(!/&,-"* ;. ,'#(. -.)&/(.*)#(. .1 (." -.3&#;! (.&  % &)1(." '#& ) ." *&.  (#(!#-&/.." * 2=  '& D#!/, P=NE</>! (#.&*&. #-.,#&) ;)/. ,.1)&) -"#!" ,."(."  '#& &) ;* 2,)/( (&/(.;)0#*)-#.),,)1(1#." 1- ,,.#)(-. ." .#*)(." )..)' ! =  &'%('"!1< &&#(!.)(:#,*:(!1 %:(1./:,(%#: 1% A- 3:"(!(/#: 1 %(! -:#%.)D#!/, P=SE=  ,# '% 1  ).3* < /&. '&  D#!/,  P=ME< )&& .  LPBKSBMKLK /,#. : (1./

145 Chapter 5

DOKWMPIOS=QOJ < LRPWOKIMO=KKJ E )&& .  3 == , 1#%= /- /'  --#)( (/' , D222  ( LOOME= &-) 2'#( < /&. '&  D#!/,  P=ME< )&& . LTBLLBMKLK/,#. :(1./DOKWMPIOS=QOJ<LRPWOKIMO=KKJE3== , 1#%=/- /' --#)((/' ,D222(LOQPE=

Figure 5.2 Pleioplectron hudsoni. Lateral view, (left) male, (right) female. Scale bars 10mm

Figure 5.3 Subgenital plate of Pleioplectron hudsoni. (Left) male lateral view, (centre) male ventral view, (right) female ventral view.

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146 Pleioplectron

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

147 Chapter 5

."( -/* ,#), -*#( ; *&  1#." ,% .#*; "#( '), 1#." K>L , .,)&. ,& *#& -*#( :-'&&;-&#!".&3,%;"#(.## &)(! ,."( '),;-'&&,)1(&. ,(.  -*#( - ,/((#(! /* ."  & (!." )  ."  .## )( -/* ,#), -/,  ; "#( *#& .## -*#( - - )&&)1-< M -/* ,#), -/*#& -*#( -: L *,)&. ,& ( L , .,)&. ,&; M -/* ,#), *#& -*#( -: L *,)&. ,& ( L , .,)&. ,&; -*#( - .1#  - &)(! - -/* ,#),-/*#&-*#( -;M#( ,#),*#&-*#( -:L*,)&. ,&(L, .,)&. ,&; MBN & (!." )  -/* ,#), *#& -*#( - )0 ; () #( ,#), -/*#& -*#( -; .,-/- 1#."O- !' (.-:L-.(M(- !' (.1#."*#,) -*#( )(#-.& (:L-.- !' (. "-N-'&&-*#( -/* ,)'."  (#(&. ,(.  -"#)(;)(." /( ,-# ) ." L-. - !' (.- '#(/.  ,)1( -*#( - ,/( /* ."  & (!." )  ."  - !' (. )( ."  , .,)&. ,&(*,)&. ,&-# ;." M(- !' (."-L*#,) '#(/. -*#( -."  /( ,-# ) ." .,-#;O."- !' (."& ." & (!.") ." L-.- !' (.=

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Figure 5.4 Lateral view of Miotopus diversus. (Left) male, (right) female.

148 Pleioplectron

Figure 5.5 Subgenital plate of Miotopus diversus. (Left) male lateral view (centre) male ventral view, (right) female ventral view.  #-.,#/.#)(<< &&#(!.)(:(1./:,(%#: 1% A-3D#!/, P=SE=  ,# % 1 )&).3* </&.'& D#!/, P=O:P=PE<LN=QP'':)&& . MPBKTBKQ: /3A-/&&3: #.% (! :!')(..#)(&,%=D!,#, :>NT=LOTKO<LRN=TOMPOE)&& .  3B=/- /' --#)((/' ,D222(OQLE= ).3* /&. '& D#!/, P=O:P=PE<LM=OS'':)&& . MKKT: "/.,10&& 3-., ' : (1./ D!,# , : >OK=ORNLS< LRP=QLQKOE )&& .  3 !,/.  -./ (.= /- /' --#)((/' ,D222(LNMRE=

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149 Chapter 5

         !"&&1' #/'-#4 0 16.5 )/(#( ), -. , -) ." )/." -&(:  1 &(1#."0,# !. )&)/,*.. ,(=#'#&,.) "'"#(&)%&(&-  )(*#&-*#( -1#."."  2 *.#)() ." *, - ( ) "#( '/,*,)&. ,&*#& -*#( =#'#&, '& -/! (#.&*&. /.# ,-#('& ! (#.&. ,'#(&#D#!/,  P=Q8P=RE=  ', " ",<'  ),)&#",-1")-./#  1 &(0 16.5( */&#-" '(3#'*),.(.-3-. '.#-** ,- ,)'LTPO/(.#&LTRM=

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150 Pleioplectron

1#."L,.#/&. &)(!*,)&. ,&*#&-*#( ;L,.#/&. &)(!, .,)&. ,&*#& -*#( ;'#.##1#."M*#,-) &)(!-*#( -*)-#.#)(#(." '#.)&)1 ,*),.#)()  ." .##;*,)&. ,&-# -*#( -&)(! ,."(." , .,)&. ,&-*#( ;'#.## *#& -*#( - - )&&)1-< L '# .## *,)&. ,& -/* ,#), *#& -*#( ; L '# .## , .,)&. ,&-/* ,#),*#&-*#( ;*& 1#.",%.#*;L'#.##*,)&. ,&#( ,#), *#& -*#( ; L '# .## , .,)&. ,& #( ,#), *#& -*#( ; #( ,#), -*#( - &)(! , ."( -/* ,#), -*#( ; *&  1#." ,% .#*; "#( '), 1#." L , .,)&. ,& -*#( -: -'&&;-&#!".&3,%;"#(.## &)(! ,."( '),;-'&&,)1(&. ,(. -*#( - ,/((#(!/*." & (!.") ." .##)(-/* ,#),-/,  ;O*#,-, -*/,&#% (N .#' - ."  & (!." )  ."  )." , -'&& -*#( -; "#( *#& .## -*#( - - )&&)1-< M -/* ,#),-/*#&-*#( -:L*,)&. ,&(L, .,)&. ,&;M-/* ,#),*#&-*#( -: L*,)&. ,&(L, .,)&. ,&;-*#( -.1# -&)(!--/* ,#),-/*#&-*#( -;M #( ,#),*#&-*#( -:L*,)&. ,&(L, .,)&. ,&;MBN& (!.") -/* ,#),*#& -*#( - )0 ; () #( ,#), -/*#& -*#( -; .,-/- 1#." O - !' (.-: L-.(M( - !' (.1#."*#,) -*#( )(#-.& (:L-.- !' (."-N-'&&-*#( -/* ,)' ."  (#(&. ,(.  -"#)(;)(." /( ,-# ) ." L-.- !' (.-'#(/. ,)1( -*#( -,/(/*." & (!.") ." - !' (.)(." , .,)&. ,&(*,)&. ,&-# ;."  M(- !' (."-L*#,) '#(/. -*#( -." /( ,-# ) ." .,-#;O."- !' (."&  ." & (!.") ." L-.- !' (.=

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151 Chapter 5

,# '% < )&).3* < /&. '&  D#!/,  P=Q: P=RE< -#4 : )&& .  ( MKLO: & / && 3: #),&( D!,# , : >OO=RKNSN OQ=MKMOK

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Figure 5.6 Holotype of Miotopus RICHARDSI (Left) male, (right) female. Scale bars 10mm.

Figure 5.7 Subgenital plate of Miotopus RICHARDSI. (Left) male lateral view, (centre) male ventral view, (right) female ventral view.

152 Pleioplectron

Figure 5.8                     

153 Chapter 5



Chopard, L. (1923). On some New Zealand Cave Orthoptera. Transaction of the Royal Society of New Zealand, 54, 230–239. Cook, L., Trewick, S., Morgan-Richards, M., & Johns, P. (2010). Status of the New Zealand cave weta (rhaphidophoridae) genera pachyrhamma, gymnoplectron and turbottoplectron. Invertebrate Systematic, (1906), 131–138. Hutton, F. . (1900). Notes on some New Zealand Orthoptera. Transaction and Proceedings of the New Zealand Institue, 32, 19–21. Hutton, F. W. (1897). The Stenopelmatidae of New Zealand. Transaction and Proceedings of the New Zealand Institue, 29, 208–242. Hutton, F. W. (1899). Supplement to the Stenopelmatidae of New Zealand. Transaction and Proceedings of the New Zealand Institue, 31, 40–43. Richards, A. M. (1959b). Revision of the Rhaphidophoridae (Orthopetera) of New Zealand - Part V. The Genus Pleioplectron Hutton 1897. Transaction of the Royal Society of New Zealand, 87, 319–327. Team, R. (2015). Rstudio: Integrated Development for R. Retrieved from http://www.rstudio.com/. Ward, D. F. (1997). A new generic key to the New Zealand cave weta genera (Orthoptera : Rhaphidophoridae). New Zealand Natural Sciences 23:13-17   

154 Ahu kit te matauranga

       Chapter 6

A step towards understanding New Zealand

Rhaphidophoridae

                  

155 Chapter 6



156 Ahu kit te matauranga



157 Chapter 6

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158 Ahu kit te matauranga

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159 Chapter 6

*, - ( B- ( ) ." - *#&-*#( -#-(). -#&3#(. ,*, . =#-.#(!/#-"#(! ." -  .2 #- ')-.&3  * ( (. )( ./, - )  ."  ! (#.&#= & - ( '& - # ,D) )/,- E(*#,#(!." .1)- 2 -.)! ." ,#-) . (# #/&.= )1 0 ,:."  -/! (#.& *&.  # , ( -  .1 ( '& - )  " -* # - D"*. , N .&  N=NE ,  +/#.  *,)()/(  ( #'*),.(. ), #-.#(!/#-"#(! -* # -=  '& - )  ."  # , (. "!'(& -* # - . ( .) "0  -#'#&, -"*  -/! (#.& *&. -: #= = "!'(& '& - . ( .) "0  #>&)  -.,/./, -= #.)")(,#&  - +/ (  # #(*#,#(!'& ( '& -(. -. */..#0 -* # -)/(,# -/.1- &-) "/! &3 #'*),.(. #( )/, /( ,-.(#(! )  ." #, 1"%** D! ( &)!3 ), 0)&/.#)(,3 , &.#)(-"#*- . ."  -* # - ( ! (/- & 0 &E= . - +/ (#(! *,)0# - #,-.-./3) ." 1"%**) '(3) ." ),." -&(0 16.5=  )/(.".-* # -."., *,.) ." ! (/- "# '%"! , . 3#",- DLTPSE #.(# &3#(.)." ! ( .##0 ,-#.3)  "!'(&=" ! (/- "!'(&"- "/!  '),*")&)!#& #0 ,-#.3 1#."#( #. ( #. '% - #. &')-. #'*)--#&  .) -3 .". ."  ! (/-  "# '%"!#- (). #( . "!'(&2 )- ,0 .".)( ) ."  '$),  #(#(! ",. , 1#."  "# '%"! ,)' )." , "!'(& -* # - #- #.- -#4  /. .". )/& +/&&3   ( *..#)( .) 0  1 &&#(!=  #",- DLTQPE )/(()." ,-* # -)  "# '%"! ,)'0 -3-. '1 -.)  /(.& 3="#- -* # -"-(). (# (.# # -#( =" -#'#&,#.# - .1 (." ( 1&3 -,#  -* # - #( "*. , O "!'(&    (  "# '%"! '(%!% ,  ().& D&.")/!"'.#(#. -." 3, ().-#-. ,.2:- "*. ,OE=)." , &)(!& !! :-'&&)# /.)( #-0 1 && ,(." -'&& ,)( #- )/( #( ), -.=  -/!! -. .". ."  ! (/-  "# '%"! -")/&   -3()(3'#4  #(.) "!'(&-0 1 &&#(!-* # -=  "  ! (/- "!'(& "-  (  &,!  *,. )  ."#- , - ," *,)$ .= . #-  1# -*,  ! (/- ( "- !)(  ,)' N  -,#  -* # - .) P  -,# : N )  1"#" ,  ( 1&3  -,#  #( ."#- , - ,": ( "!'(& # "&(&  -,#  3 /..)(DLSTRE"- (-3()(3'#4 1#." "!'(&)%'(&=" , , .& -. )/,'),  "!'(&-* # -3 ..)  -,# (-." - -* # -, *,)&3 #,&3 1# -*,  D#  (). &)&&3 )'')(E= /- /'- *,)&3 ")& '. ,#& )  ." -  -* # - .". (   /-  - #.#)(& -* #' (- ),  -,#*.#)(- #( ."  /./, =

160 Ahu kit te matauranga

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

161 Chapter 6

Figure 6.1 A developing whakapapa of New Zealand rhaphidophorid genera.   

162 Ahu kit te matauranga

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

163 Chapter 6

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

164 Ahu kit te matauranga

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

165 Chapter 6

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

166 Ahu kit te matauranga

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

167 Chapter 6

 

168 Appendix

Appendix I %" # $ ) "% "! # !%*# !"%*# ,,,                                 &! %  &! % %%*# # 446        44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 476;      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4933      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;46      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;47      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;49      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;4:      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;4;      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;5<      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;65      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 476:      44 4  55 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 7:5      47 7  55 3 3 3 4 3 3 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 ;4:      4: 6  55 3 3 3 4 3 3 4 3 4 4 4 3 4 4 4 4 4 4 3 3 4 4 4;69      55 5  55 3 3 3 4 3 4 4 3 4 4 4 3 4 4 4 4 4 4 3 3 4 4 ;39      55 5  55 3 3 3 4 3 4 4 3 4 4 4 3 4 4 4 4 4 4 3 3 4 4 4;48      55 8  49 3 3 3 4 3 4 4 3 4 4 4 3 4 4 4 4 4 4 3 3 4 4 4;5;      44 7<  49 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4;66      44 7<  49 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4:3;      55 8  49 3 3 3 4 3 4 4 3 4 4 4 3 4 4 4 4 4 4 3 3 4 4 4;4<      44 7<  49 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4

169 Appendix

;49      44 7<  49 3 3 3 4 3 4 4 3 4 3 4 3 4 4 4 4 4 4 3 3 4 4 4937      55 8  49 3 3 3 4 3 4 4 3 4 4 4 3 4 4 4 4 4 4 3 3 4 4 4;:3      54 67  7 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4    4657      6 66  7 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;7<      6 66  7 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 <4;      6 66  7 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;6<      6 66  7 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 :43      6 45  56 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 75<       4 43  4< 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 :;;     4 43  4< 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 :;8     4 43  4< 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 764     4 9  48 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 5<4     53 :  48 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 :;9     53 :  48 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 <7;     49 ;  48 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 3 4 4 :;:     4< <  48 3 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 46;<     58 76  4< 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 4 4 4 76<     ; 84  4< 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 4 4 4;97     54 44  4< 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 5<3     6 45  4< 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;96       54 4<  5 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;98     54 4<  5 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;9<     54 4<  5 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 79:     54 4<  5 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 79;   544<54444334444444444444444 7:4     54 4<  5 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4

170 Appendix

9;:    544<54444334444444444444444 :88     54 4<  5 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;88     54 4<  5 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 46<:     6 53  5 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4:53     6 53  5 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;84     6 53  5 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;8:    65354444344444444444444444 7:6     6 53  5 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 8;4    65354444344444444444444444 :8:     6 53  5 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 ;48     6 53  5 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 495     59 4;  5 4 4 4 4 3 3 4 3 4 4 4 4 4 4 4 4 4 4 3 4 4 4 4:96     54 68  53 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4;8<     54 68  53 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 :8;     6 69  59 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 44-43-4     6 69  59 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4 4. 4:48    546854444334444444444444444 4;-:-44        4 55  43 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 . :<3      53 56  43 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 54-:-44      53 56  43 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 . 43-:-44      4 57  < 4 4 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 . 6/70. '      53 58  < 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 ;. '    5358<4444334444444444443344 44-:-44      53 58  < 4 4 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 .

171 Appendix

4773      57 63  45 4 3 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4    4775       57 63  45 4 3 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 4797       57 63  45 4 3 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 4:8<       57 63  45 4 3 4 4 3 3 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 493<       7 5<  45 4 3 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 4;:       7 5<  45 4 3 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 93       7 5<  45 4 3 4 4 3 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 4 4 4

172 Appendix

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173 Appendix

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174 Appendix

175 Appendix

Appendix II

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176 Appendix

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