Evolutionary and biogeographical history of an ancient and

39 Pages·2011·1.69 MB·English

Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••. With 10 figures
Evolutionary and biogeographical history of an ancient
and global group of arachnids (Arachnida: Opiliones:
Cyphophthalmi) with a new taxonomic arrangement
GONZALO GIRIBET1*, PRASHANT P. SHARMA1, LIGIA R. BENAVIDES2,
SARAH L. BOYER3, RONALD M. CLOUSE1,4, BENJAMIN L. DE BIVORT1,5,
DIMITAR DIMITROV6, GISELE Y. KAWAUCHI1, JEROME MURIENNE1,7 and
PETER J. SCHWENDINGER8
1Museum of Comparative Zoology, Department of Organismic and Evolutionary Biology, Harvard
University, 26 Oxford Street, Cambridge, MA 02138, USA
2Department of Biological Sciences, The George Washington University, 2023 G Street NW,
Washington, DC 20052, USA
3Biology Department, Macalester College, 1600 Grand Avenue, Saint Paul, MN 55105, USA
4Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th
Street, New York, NY 10024, USA
5Rowland Institute at Harvard, Harvard University, 100 Edwin Land Boulevard, Cambridge, MA
02142, USA
6Center for Macroecology, Evolution and Climate, Zoological Museum, University of Copenhagen,
Department of Entomology, Universitetsparken 15, DK-2100, Copenhagen, Denmark
7UMR 7205 CNRS, Département Systématique et Évolution, CP 50, Muséum national d’Histoire
naturelle, 45, rue Buffon, 75005 Paris, France
8Muséum d’histoire naturelle, Département des Arthropodes et d’Entomologie I, 1, Route de
Malagnou, Case Postale 6434, CH-1211 Genève 6, Switzerland
Received 27 May 2011; revised 8 July 2011; accepted for publication 8 July 2011
bij_17741..39
We investigate the phylogeny, biogeography, time of origin and diversification, ancestral area reconstruction and
large-scale distributional patterns of an ancient group of arachnids, the harvestman suborder Cyphophthalmi.
Analysis of molecular and morphological data allow us to propose a new classification system for the group;
Pettalidae constitutes the infraorder Scopulophthalmi new clade, sister group to all other families, which are
divided into the infraorders Sternophthalmi new clade and Boreophthalmi new clade. Sternophthalmi includes
the families Troglosironidae, Ogoveidae, and Neogoveidae; Boreophthalmi includes Stylocellidae and Sironidae,
the latter family of questionable monophyly. The internal resolution of each family is discussed and traced back
to its geological time origin, as well as to its original landmass, using methods for estimating divergence times
andancestralareareconstruction.TheoriginofCyphophthalmicanbetracedbacktotheCarboniferous,whereas
the diversification time of most families ranges between the Carboniferous and the Jurassic, with the exception
of Troglosironidae, whose current diversity originates in the Cretaceous/Tertiary.Ancestral area reconstruction is
ambiguousinmostcases.SternophthalmiistracedbacktoanancestrallandmassthatcontainedNewCaledonia
and West Africa in the Permian, whereas the ancestral landmass for Neogoveidae included the south-eastern
USA and West Africa, dating back to the Triassic. For Pettalidae, most results include South Africa, or a
combination of SouthAfrica with theAustralian plate of New Zealand or Sri Lanka, as the most likely ancestral
landmass, back in the Jurassic. Stylocellidae is reconstructed to the Thai-Malay Penisula during the Jurassic.
Combination of the molecular and morphological data results in a hypothesis for all the cyphophthalmid genera,
although the limited data available for some taxa represented only in the morphological partition negatively
affects the phylogenetic reconstruction by decreasing nodal support in most clades. However, it resolves
*Corresponding author. E-mail: [email protected]
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–•• 1
2 G. GIRIBET ETAL.
the position of many monotypic genera not available for molecular analysis, such as Iberosiro, Odontosiro,
Speleosiro, Managotria or Marwe, although it does not place Shearogovea or Ankaratra within any existing
family. The biogeographical data show a strong correlation between relatedness and formerly adjacent
landmasses, and oceanic dispersal does not need to be postulated to explain disjunct distributions, especially
when considering the time of divergence. The data also allow testing of the hypotheses of the supposed total
submersion of New Zealand and New Caledonia, clearly falsifying submersion of the former, although the data
cannot reject the latter. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society,
2011, (cid:129)(cid:129), ••–••.
ADDITIONAL KEYWORDS: biogeography – distribution modelling – Gondwana – Laurasia – MAXENT –
New Caledonia – New Zealand – Pangea.
INTRODUCTION Giribet, 2005; Boyer & Giribet, 2007; Clouse &
Giribet, 2007; Giribet & Kury, 2007; Boyer etal.,
The harvestman suborder Cyphophthalmi (Fig.1) 2007b; Boyer & Giribet, 2009; Clouse, de Bivort &
constitutes an ancient lineage of arachnids and was Giribet, 2009; Karaman, 2009; Murienne & Giribet,
probably one of the earliest inhabitants of terrestrial 2009; Sharma & Giribet, 2009a; Clouse & Giribet,
ecosystems. Currently distributed on all continental 2010; de Bivort & Giribet, 2010; Murienne, Karaman
landmasses (with the exception ofAntarctica) and on & Giribet, 2010b; Clouse etal., 2011). These include
most large islands of continental origin, the group is some recent and more general debates on the total
considered to have been in close association to these submersion of large fragment islands such as New
landmasses since its origins (Juberthie & Massoud, Caledonia and New Zealand (Sharma & Giribet,
1976; Boyer etal., 2007b). The fact that deep genetic 2009a; Giribet & Boyer, 2010; Giribet etal., 2010).
divergences in cytochrome c oxidase subunit I (COI) However, to use a system for biogeographical infer-
have been reported within one species (Boyer, Baker ences, a sound systematic hypothesis of the group is
& Giribet, 2007a), and are suspected for many others required. The taxonomy of Cyphophthalmi has ben-
(R. Clouse & P. Sharma, unpubl. data), corroborates efitted from the contributions of many studies, espe-
theobservationsthatindividualsmaylivealongtime cially the synthetic work of Hansen & Sørensen
(Juberthie, 1960b) and do not disperse far during the (1904), who produced the first and still best mono-
course of life history. These, together with the old graph on the group, and established the first classi-
history of the group [a Burmese amber specimen fication system of the suborder Cyphophthalmi with
probablybelongingtoStylocellidaeisknownfromthe one family, Sironidae, and two subfamilies, Stylocel-
EarlyCretaceous(Poinar,2008)andtheoriginsofthe lini(includingthegeneraStylocellusWestwood,1874,
group has been estimated to have taken place during Ogovia Hansen & Sørensen, 1904, which was pre-
the Devonian or Carboniferous using molecular occupiedandbecameOgoveaRoewer,1923,andMiop-
datingtechniques(Giribetetal.,2010)]haveresulted salis Thorell, 1890) and Sironini (including Pettalus
in a broad use of Cyphophthalmi for biogeographical Thorell, 1876, Purcellia Hansen & Sørensen, 1904,
inferences and zoogeographical discussions (Rambla, Siro Latreille, 1796, and Parasiro Hansen &
1974;Juberthie&Massoud,1976;Boyer,Karaman& Sørensen, 1904). Another major contributor was
(cid:2)
Figure1. Habitus.A, Karripurcellia harveyi (Pettalidae) from Warren National Park, WesternAustralia, July 2004. B,
Pettalusthwaitesi(Pettalidae)fromPeradeniyaBotanicalGardens,CentralProvince,SriLanka,October2007.C,Rakaia
pauli (Pettalidae) from Kelcey’s bush, near Waimate, North Island, New Zealand, February 2008. D, Aoraki longitarsa
(Pettalidae) from Governor’s bush, Mt Cook, South Island, New Zealand, January 2006. E, male Pettalus thwaitesi
(Pettalidae) from Peradeniya Botanical Gardens, Central Province, Sri Lanka, June 2004. F, Ogovea cameroonensis
(Ogoveidae) from Ototomo Forest, Central province, Cameroon, June 2009. G, Parogovia sp. (Neogoveidae) from Mt.
Koupé,South-WestProvince,Cameroon,June2009.H,twospeciesofParogoviafromCampoReserve,LittoralProvince,
Cameroon, June 2009; upper left, adult specimen of Parogovia n. sp.; lower right, juvenile specimen of Parogovia cf.
sironoides.I,juvenilespecimenofParamiopsalisramulosus(Sironidae)fromP.N.PenedaGerés,Portugal,May2008.J,
Paramiopsalis ramulosus (Sironidae) from P.N. Peneda Gerés, Portugal, May 2008. K, Parasiro minor (Sironidae) from
Monte Rasu, Sardinia, Italy, March 2008. L, Suzukielus sauteri (Sironidae) from Mt. Takao, Tokyo Prefecture, Honshu,
Japan,April2005.M,juvenilespecimenofLeptopsalissp.(Stylocellidae)fromBantimurung-BulusaraungN.P.,Sulawesi
Selatan, Indonesia, June 2006. N, female Leptopsalis sp. (Stylocellidae) from Bantimurung-Bulusaraung N.P., Sulawesi
Selatan, Indonesia, June 2006.
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
EVOLUTIONARY AND BIOGEOGRAPHICAL HISTORY OF ARACHNIDS 3
AA B
C D E
F G
H
I J K
L M N
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
4 G. GIRIBET ETAL.
Juberthie, who described and monographed many and subsequently included in Neogoveidae (Giribet,
genera(e.g.Juberthie,1956,1958,1960a,1961,1962, 2007b);FangensisRambla,1994wastransferredfrom
1969, 1970a, b; Juberthie & Muñoz-Cuevas, 1970; Sironidae to Stylocellidae (Schwendinger & Giribet,
Juberthie, 1979) in addition to his contributions to 2005); Metasiro was transferred from Sironidae to
the biology of the group; the regional work of Forster Neogoveidae (Giribet, 2007b); Meghalaya Giribet,
(1948,1952)inNewZealand,thatofLawrence(1931, Sharma&Bastawade,2007wasincludedinStylocel-
1933, 1939, 1963) in South Africa, that of Rambla lidae (Clouse etal., 2009); and Shearogovea mexasca
(Rambla & Fontarnau, 1984, 1986; Rambla, 1991, (Shear, 1977) was excluded from Neogoveidae
1994)intheIberianPeninsulaandsoutheastAsia,to (Benavides&Giribet,2007;Giribet,2011)–thefami-
mention just a few of them. More recently, Shear has lies are currently considered to be stable.
contributed with descriptions of numerous species in Recent phylogenetic analyses based on nucleotide
almost all cyphophthalmid families (Shear, 1977, sequence data have resolved the relationship among
1979a,b,1985,1993a,b,c;Shear&Gruber,1996).He some of these families, providing strong support for
also proposed the bases of modern cyphophthalmid a relationship of Troglosironidae and Neogoveidae
systematicsinaseminalfirstcladisticanalysisofthe (Boyer etal., 2007b; Boyer & Giribet, 2009; Sharma
group(Shear,1980),withfivefamilies,threeofwhich & Giribet, 2009a; Giribet etal., 2010), a result also
were new, and two infraorders, equivalent to Hansen obtained in a recent analysis of morphometric char-
and Sørensen’s subfamilies (Table1). A sixth family, acters(deBivort,Clouse&Giribet,2010).Monophyly
Troglosironidae, was also proposed a few years later of Pettalidae is well supported both by discrete and
(Shear, 1993b). continuous morphological characters (Giribet &
Two decades after Shear’s classification system Boyer,2002;Giribet,2003a;Boyer&Giribet,2007;de
appeared, Giribet (2000) compiled all the cyphoph- Bivortetal.,2010;deBivort&Giribet,2010),aswell
thalmid literature to date, recognizing 113 species in asadiversityofmolecularanalyses(Boyer&Giribet,
26 genera.Asubsequent analysis including represen- 2007, 2009; Boyer etal., 2007b; Giribet etal., 2010).
tatives of most genera and based on a numerical Stylocellidaeisalsowellsupportedbasedonmorphol-
cladistic analysis of 32 morphological characters ogy (Giribet & Boyer, 2002; Clouse etal., 2009) and
(Giribet & Boyer, 2002) recognized most of the fami- molecules (Schwendinger & Giribet, 2005; Clouse &
lies erected by Shear (1980) but also challenged some Giribet, 2007; Boyer etal., 2007b; Clouse etal., 2009;
of his systematic propositions because the root of Clouse&Giribet,2010;Giribetetal.,2010).However,
the tree, based on a limited molecular data set also monophylyofSironidae,especiallythemembershipof
published in the same study, was placed between the Mediterranean genus Parasiro and the Japanese
stylocellids and the rest (rendering Tropicophthalmi Suzukielus Juberthie, 1970, remains controversial,
paraphyletic) or between pettalids and the rest (ren- both based on morphology (Giribet & Boyer, 2002; de
deringTemperophthalmiparaphyletic).Shear(1993b) Bivort&Giribet,2004;deBivortetal.,2010),aswell
had also proposed the new family Troglosironidae as on molecular analyses (Boyer etal., 2005; Boyer &
as sister to (Pettalidae + Sironidae) and this result Giribet, 2007; Giribet etal., 2010).
was refuted by Giribet & Boyer (2002), who found it Inadditiontotheuncertaintyaboutthemonophyly
nestedwithinanunresolvedNeogoveidae.Aftersome of Sironidae, which we approach here by including
minor familial reassignments – Huitaca Shear, 1979 an expanded taxon sampling in problematic genera
wasremovedfromOgoveidae(Giribet&Prieto,2003) previously represented by a single species (Parasiro),
we include a much larger diversity of Neogoveidae,
both from the Neotropics (29 species versus six used
Table1. Classification system of Shear (1980, 1993)
in Boyer etal., 2007b; including data on the new
Brazilian genus Canga DaSilva, Pinto-da-Rocha &
Suborder Cyphophthalmi
Giribet, 2010) and from theAfrotropics (12 terminals
Infraorder Tropicophthalmi Shear, 1980
versussevenusedinBoyeretal.,2007b).Mostimpor-
SuperfamilyStylocelloideaHansen&Sørensen,1904
tantly, we include the first molecular data on the
Family Stylocellidae Hansen & Sørensen, 1904
family Ogoveidae, from specimens collected in Cam-
Superfamily Ogoveoidea Shear, 1980
eroon in 2009. In total, we provide novel sequence
Family Ogoveidae Shear, 1980
data for 34 species (of a total of 162 molecular ter-
Family Neogoveidae Shear, 1980
minals), include 27 genera and a family previously
Infraorder Temperophthalmi Shear, 1980
unsampled, and include new landmasses (Mindanao,
Superfamily Sironoidea Simon, 1879
the eastern Neotropics, the westernmost distribution
Family Sironidae Simon, 1879
of the Afrotropics) not considered in previous phylo-
Family Pettalidae Shear, 1980
genetic analyses. The present study also provides
Family Troglosironidae Shear, 1993
the first total evidence analysis of molecules and
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
EVOLUTIONARY AND BIOGEOGRAPHICAL HISTORY OF ARACHNIDS 5
morphology for the whole suborder Cyphophthalmi molecular and morphological study. Museum speci-
and new data on the timing of diversification and mens have also been used for the morphological
cladogenesis of the group, aiming to revisit interest- studies. A detailed discussion of the specimen col-
ing biogeographical topics. Finally, we provide an lecting effort is provided in the Supporting informa-
estimate of the ancestral area for each lineage and tion (AppendixS1).
presentthefirsthabitatsuitabilityanddistributional The study includes the first molecular data for the
patterns analysis for this dispersal-limited, yet family Ogoveidae (Ogovea cameroonensis Giribet &
globally-distributed group of arthropods. Studying Prieto,2003)andincludesadditionalsamplingwithin
macroecologicalpatternsinCyphophthalmiiscompli- all other families, building upon previous studies on
cated as a result of the scarce occurrence data for the phylogenies of Pettalidae (Boyer & Giribet, 2007;
most species. Thus, species-level assessment of large- Boyer etal., 2007b). Stylocellidae (Clouse etal., 2009;
scale distributional patterns and their primary eco- Clouse & Giribet, 2010; Clouse etal., 2011), Troglo-
logical and evolutionary drivers is difficult. Recently, sironidae (Sharma & Giribet, 2009a), and Sironidae
theoretical and practical arguments for the utility of (Boyer etal., 2005; Giribet & Shear, 2010; Murienne
modelling distributional patterns or even ecological etal., 2010b). However, data for Neogoveidae were
niche characteristics above the species level have restricted to a single previous study (Boyer etal.,
been proposed (Heino & Soininen, 2007; Hadly, 2007b), and the family was poorly sampled. For the
Spaeth & Li, 2009; Diniz, De Marco & Hawkins, African diversity, we are now able to add data on
2010). Despite some obvious limitations (Diniz etal., anotherdescribedspecies,Parogoviagabonica(Juber-
2010), this approach may be very useful to evaluate thie, 1969), from near its type locality (Ipassa
patterns in groups with limited distributional data Reserve, Makokou, Gabon). We also add new data on
such as insects and other arthropods including two new species from Mount Koupé and the Campo
Cyphophthalmi. reserve, in Cameroon, and an additional specimen of
Parogovia cf. sironoides also from the Campo reserve
in Cameroon. The third African species, Parogovia
MATERIAL AND METHODS
pabsgarnoni Legg, 1990, is known only from its type
SPECIMENS locality in Sierra Leone, resulting in a large biogeo-
Most specimens used in the molecular part of graphical gap from the known distribution of species
this study (Fig.2) were collected by one or more in the Gulf of Guinea, and differs morphologically
of the authors through direct sifting of leaf litter from the other species in the genus in many key
and transferred to approximately 95% ethanol for characters, showing very different spermatopositor
Figure2. Distribution map of the sampled specimens for the molecular study (Table2). Pettalidae are represented in
red, Troglosironidae in purple, Ogoveidae in cyan, Neogoveidae in green, Sironidae in orange, and Stylocellidae in navy
blue.
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
6 G. GIRIBET ETAL.
from the species in the Gulf of Guinea (Legg & Pabs- for 170 terminals. The 28S rRNA fragment was
Garnon,1989;Legg,1990).Apossiblecloserelativeto divided into ten regions and was available for 169
this species is included here, based on data from a terminals. 16S rRNA was divided into eight frag-
female collected in Ivory Coast. The South American ments and was available for 127 terminals. All the
sampling has also been enriched considerably. The ribosomal genes were submitted to direct optimiza-
monotypic genus Huitaca is now represented by data tion or to multiple sequence alignment for homology
from seven species all from Colombia, including the assignment. The 143 COI sequences, unlike in many
nominal Huitaca ventralis Shear, 1979. Metagovea is other organisms, show clade-specific considerable
now represented by 13 specimens in 12 putative new sequence length variation, and hence the gene was
species, including one from Guyana. These include divided into seven fragments and analyzed under
the Colombian species identified as Neogovea in our dynamic homology (Wheeler, 2005) or submitted to
previous studies, although they appear to be related multiple sequence alignment. The histone H3 data
to Metagovea. A large biogeographical gap in our were available for 108 terminals and were treated as
previous studies was the easternmost distribution of prealigned in all analyses because they show no
the genus Neogovea (type species Neogovea immsi length variation.
Hinton, 1938, from Amapá, Brazil). We now include
putative representatives of this clade based on speci-
menscollectedinFrenchGuianaandGuyana,includ- MORPHOLOGICAL DATA MATRIX
ing specimens from the recently described N.virginie Amorphologicalmatrixof62characterswascompiled
Jocqué & Jocqué, 2011. With the exception of one for 161 taxa based in part on our previous studies
specimen from Guyana clustering with Metagovea, (Giribet & Boyer, 2002; Giribet, 2003a; de Bivort &
these specimens group with the unidentified juvenile Giribet, 2004; Boyer & Giribet, 2007; de Bivort &
from Venezuela used in Boyer etal. (2007b) and with Giribet, 2010), direct observation of specimens,
a species from the ‘Tepuis’ in Colombia, which we mostly through scanning electron microscopy, and
reassign here to the genus Brasilogovea Martens, complemented by some new literature sources
1969, previously considered a synonym of Neogovea (Karaman, 2009). All 19 multistate characters were
(Shear, 1980; Giribet, 2000). Finally, we were able to unordered. Spermatogenesis in Cyphophthalmi is a
include data on the monotypic genus Canga based on promising source of phylogenetic characters, as
a specimen from its type locality (DaSilva etal., recentlyoutlinedbyAlberti,Giribet&Gutjahr(2009;
2010). However, large areas of the Neotropics with seealsoJuberthie&Manier,1976;Juberthie,Manier
known specimens of Neogoveidae remain unsampled & Boissin, 1976; Juberthie & Manier, 1978; Alberti,
in our molecular phylogeny, and they should be 1995, 2005), although taxon sampling is still sparse
includedinfuturestudies(Benavides&Giribet,2007: andthesecharacterswerenotconsideredinthisdata
fig.1). set(G.Alberti&G.Giribet,unpubl.data).Wedidnot
have access to specimens of a few species that were
included in the data matrix based entirely on litera-
MOLECULAR DATA ture sources. These have missing data for several
DNA extraction, amplification, and sequencing were characters, especially those observed through scan-
performed as described in several of our previous ning electron microscopy, such as the prosomal
studies on molecular systematics of Cyphophthalmi sternal characters. These species include Ankaratra
using the same markers (Schwendinger & Giribet, franzi Shear & Gruber, 1996, Manangotria taolanaro
2005; Boyer etal., 2007b; Boyer & Giribet, 2009; Shear & Gruber, 1996, and Odontosiro lusitanicus
Sharma & Giribet, 2009a; Clouse & Giribet, 2010). Juberthie, 1961. Similarly, several Cyphophthalmus
Weusedthefivemarkersasinthesepreviousstudies, Joseph, 1868 species, included in our molecular
including the nuclear ribosomal 18S and 28S rRNA, matrix, were not scored for several morphological
the nuclear protein-encoding histone H3, the mito- characters because males were never available for
chondrialribosomal16SrRNA,andthemitochondrial examination, and their published descriptions do not
protein-encoding COI genes. For outgroups, we used includescanningelectronmicrographsoftherelevant
seven noncyphophthalmid Opiliones from the subor- charactersandtheirdescriptionsandillustrationsare
ders Eupnoi, Dyspnoi, and Laniatores (Table2). Pub- inadequate for scoring those features. Finally, a few
lishedsequencedatafromthesestudiesandthenovel species scored in the matrix are not known for one
data presented here are deposited in GenBank and gender, and therefore the corresponding scorings are
are shown in Table2. missing. The total number of missing cells was thus
Allsequencefilesforeachgenewerepreparedwith 1798 (17% of cells). In the present study, we were not
MacGDE (Linton, 2005). 18S rRNA sequence data able to use morphometrics, as we have done in pre-
were divided into six fragments and it was available vious studies (Clouse, 2010; de Bivort etal., 2010; de
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
EVOLUTIONARY AND BIOGEOGRAPHICAL HISTORY OF ARACHNIDS 7
3680 025 7868 23456767901231 94 4701532 19136
0590 609 4447 74933773344475 45 8556556 98898
7167 176 1116 61611661111161 11 6111111 66666
3833 833 8883 38388338888838 88 3888888 33333
7177 177 1117 71711771111171 11 7111111 77777
6566 566 5556 65655665555565 55 6555555 66666
3 UQUU QUU QQQU UQUQQUUQQQQQUQ QQ UQQQQQQ UUUUU
H EDEE–DEE––DDDE–EDEDDEEDDDDDED–DD–EDDDDDD––EEEEE
OI U673667Q518120Q992309Q992203 Q518122 Q992313Q992318Q992319Q518111Q518112Q518110Q518108Q133745Q133746Q518106Q825638Q518098Y639582U673666Q825636Q518100Q518102Q518103Q518104Q518105U673665Q518115Q992331Q518113Q518118U673668Q992333Q518125Q518114U673670Q518119Q518116Q518123Q518117U673671Q992344 Q992349Q992322Q992348
C EDDD–D–DDDDDDDDDDDDAEDDDDDDEDDDDEDDDEDDDDED–DDD
A 084517 455426 39 6890122731154235980 9 316
RN 807806358358807806 806806805805806806 805356 805805805806806805807807808808357357807808807807807807808 357 357358357
r 117711 111111 17 1111111111771111111 7 777
16S DQ5DQ5EU6EU6DQ5DQ5––––––DQ5DQ5DQ5DQ5DQ5DQ5–DQ5EU6–DQ5DQ5DQ5DQ5DQ5DQ5DQ5DQ5DQ5DQ5EU6EU6DQ5DQ5DQ5DQ5DQ5DQ5DQ5–EU6–EU6EU6EU6
A 83469282 93421349098326734591357600629658875792
RN 803804365365803804365365 365802802802802373373801365800800363363801801801801801362803803802802363364803802803802803803802363364364363364364
r 11771177 71111331711771111171111771111111777777
28S DQ5DQ5EU6EU6DQ5DQ5EU6EU6–EU6DQ5DQ5DQ5DQ5DQ1DQ1DQ5EU6DQ5DQ5EU6EU6DQ5DQ5DQ5DQ5DQ5EU6DQ5DQ5DQ5DQ5EU6EU6DQ5DQ5DQ5DQ5DQ5DQ5DQ5EU6EU6EU6EU6EU6EU6
18SrRNA EU673626DQ518000EU673618EU673620DQ517999DQ518002EU673622EU673613EU673614EU673624DQ517984DQ517985DQ517983DQ517982DQ133721DQ133722DQ517980DQ517998DQ518973AY639494EU673592DQ825537DQ517974DQ517976DQ517977DQ517978DQ517979EU673589DQ517988DQ517990DQ517986DQ517995EU673596EU673601DQ517996DQ517987DQ517992DQ517997DQ517993DQ517989DQ517994EU673599EU673608EU673606EU673597EU673610EU673603
*
30646622 907776377 1168566875418477438
04964272 06012632263366996046699024558309447436
91040842 75229635516399449367983810858093972121
64464329*31467187557366999046060274164900316232
37376110714642629383911114723991016294621414090
nates 387,172.404,175.283,171.408,171.478,175.673,174.686,177.666,170.7,175.63708,174.610,145.560,145.151,145.458,145.833,72.-666,73.-500,115.780,169.347,29.3421,31.1251,80.5483,80.8483,80.8517,80.8517,80.8359,80.8551,80.3294,18.4145,171.163,174.258,172.327,168.000,170.225,172.478,175.641,175.073,170.852,175.967,167.177,169.327,168.083,173.595,175.166,169.869,173.181,174.757,175.
ers Coordi 41.78-37.53-42.09-43.03-39.93-41.08-38.79-43.73-39.66-41.08-16.16-17.24-16.06-16.59-41.50-39.66-34.49-44.10-29.05-28.74-7.277.387.386.926.926.826.5533.98-43.25-41.28-40.83-46.89-45.92-42.33-39.93-39.41-44.70-41.46-46.10-46.58-46.89-41.22-40.97-45.90-43.80-40.85-41.15-
b
m
u
n
on aliaaliaaliaalia
nBankaccessi Locality NewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandQueensland,AustrQueensland,AustrQueensland,AustrQueensland,AustrChileChileWesternAustraliaNewZealandSouthAfricaSouthAfricaSriLankaSriLankaSriLankaSriLankaSriLankaSriLankaSriLankaSouthAfricaNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealandNewZealand
e
G
h
t
wi 2989555941406606 2651474546139003398685238283858687888757439528086292916894697044129707589354
12998998 19999443933222222239921892292999828929
ndata Voucher DNA101DNA101DNA100DNA100DNA101DNA100DNA100DNA101DOC094DNA101DNA100DNA100DNA100DNA100DNA100DNA100DNA101DNA100DNA100DNA100DNA101DNA101DNA101DNA101DNA101DNA101DNA101DNA100DNA100DNA100DNA101DNA101DNA101DNA100DNA101DNA101DNA100DNA101DNA100DNA100DNA100DNA101DNA101DNA101DNA100DNA101DNA100
o
ti
c
e
oll a
able2.Specimenandc AMILYPETTALIDAEAorakicalcarobtusawestlandicAorakicryptaAorakidenticulatadenticulataAorakidenticulatamajorAorakigranulosaAorakihealyiAorakiinermaAorakilongitarsaAorakicf.tumidataAorakisp.AustropurcelliaarcticosaAustropurcelliadaviesaeAustropurcelliaforsteriAustropurcelliascopariaChileogoveaoedipusChileogoveasp.KarripurcelliaharveyiNeopurcelliasalmoniParapurcelliamonticolaParapurcelliasilvicolaPettalusthwaitesiPettalussp.Pettalussp.Pettalussp.Pettalussp.Pettalussp.sp.PettalusPurcelliaillustransRakaiaantipodianaRakaiadorotheaRakaiaflorensisRakaialindsayiRakaiamacraRakaiamagnaaustralisRakaiamediaRakaiaminutissimaRakaiapauliRakaiasolitariaRakaiasorensenisorenseniRakaiasorensenidigitataRakaiastewartiensisRakaiaunilocaRakaiasp.Rakaiasp.Rakaiasp.Rakaiasp.Rakaiasp.
T F
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
8 G. GIRIBET ETAL.
1
H3 ––AY639444–AY639446–AY639447AY639449AY639450AY639451 AY639453–AY639454– AY639456JF786415––JF786416AY918882–––––––––AY639457.DQ518166DQ825520 JF786417 DQ518164–DQ518165––JF786340–
COI FJ946438AY639556AY639557AY639559–FJ946433AY639561AY639563AY639565AY639567 AY639569FJ946432AY639571AY639572 AY639574EU638288DQ825641JF786389JF786390DQ825642JF786391DQ825643DQ513112–AY639579DQ513115–DQ513111–AY639580DQ513108DQ825640 JF786392 AY639584EU887047DQ825639EU887043DQ518128EU887040EU887061
16SrRNA FJ946364AY639526AY639527AY639529AY639532FJ946358AY639534AY639536AY639537– AY639539FJ946357AY639544– AY639546EU638281AY639550JF935023JF935024AY918877JF935025AY639551–––––––AY639552DQ518086DQ825615 JF935026 AY639555EU887077DQ518084EU887074DQ518055EU887073EU887075
* * *
28SrRNA FJ946415DQ513120DQ825593DQ825587DQ825592FJ946409AY639504DQ825589DQ825591DQ825594 DQ825588FJ946408DQ513118DQ513119 AY639515EU638287DQ513121JF934990JF934991DQ513122JF934992DQ513128DQ513125DQ513133DQ825585DQ513134JF934993DQ825584DQ513136DQ513123DQ513116DQ825583 JF934994 DQ825580EU887121DQ825582EU887116DQ825581EU887119EU887125
18SrRNA FJ946390AY639461AY639462AY639464AY639467FJ946384AY639469AY639471AY639473AY639475 AY639477FJ946383AY639482AY639483 AY639485EU638284AY639489JF934956JF934957AY918872JF934958AY639490DQ513139DQ513146AY639491DQ513147JF934959AY428818DQ513149AY639492DQ513138DQ825541 JF934960 AY639497DQ825540DQ518089EU887101DQ518088EU887102EU887107
13 77 5
*44 07 4 26
47 47 7 3977 6772 665 27 0 26 9
46 46 6 3866 59289864030506 11 7 9767076
96 26 6 3366 36023418337143*43 0 5636010
Coordinates 39.61056,20.3346.01667,14.6643.83333,20.0541.96667,21.3542.45,19.2666743.41344,16.9141.41667,22.2642.4,18.7666742.65,18.6666743.01667,18.51 42.16667,19.3343.27917,22.0642.23333,19.0642.23333,19.16 42.93333,18.543.41718,8.06-42.31580,8.48-41.56944,8.14-43.31968,6.87-42.15251,1.93040.43047,9.00944.5833,123.5-46.99221,121.-38.27744,120.-39.4833,79.43-47.74646,116.-46.86777,117.-44.08338,3.58141.06367,122.-45.9833,9.866635.63440,139.234.83333,138.9 3.64621,11.29 21.1833,165.30-22.0500,166.46-22.3527,166.97-21.6000,165.71-21.7500,166.15-22.1945,166.50-22.1770,166.51-
A
Locality GreeceSloveniaSerbiaMacedoniaMontenegroCroatiaMacedoniaMontenegroMontenegroBosnia&HerzegovinaMontenegroSerbiaMontenegroBosnia&HerzegovinaMontenegroSpainSpainPortugalSpainSpainSardinia,ItalyOregon,USAWashington,USCalifornia,USAMaryland,USAIdaho,USAIdaho,USAFranceCalifornia,USAItalyJapanJapan Cameroon NewCaledoniaNewCaledoniaNewCaledoniaNewCaledoniaNewCaledoniaNewCaledoniaNewCaledonia
11879998959997949339 92981038 077859382483358814238911135722614350 17 45446780771024
1444404440 4090 984563546646646455 6 3385573
her 102100100100100102100100100101 100102100101 100101100103104101103100101101100101101100101100101101 104 100100100101100101102
uc NANANANANANANANANANA NANANANA NANANANANANANANANANANANANANANANANANA NA NANANANANANANA
Vo DDDDDDDDDD DDDD DDDDDDDDDDDDDDDDDD D DDDDDDD
able2.Continued AMILYSIRONIDAECyphophthalmuscorfuanusCyphophthalmusduricoriusCyphophthalmusereCyphophthalmuscf.gjorgjeviciCyphophthalmusgordaniCyphophthalmushlavaciCyphophthalmusmarkoiCyphophthalmusmartensiCyphophthalmusminutusCyphophthalmusognjenovici CyphophthalmusrumijaeCyphophthalmusserbicusCyphophthalmusteyrovskyiCyphophthalmustrebinjanus CyphophthalmuszetaeParamiopsaliseduardoiParamiopsalisramulosusParamiopsalisramulosusParamiopsalissp.ParasirocoiffaitiParasirominorSiroacaroidesSiroboyeraeSirocalaverasSiroexilisSirocf.kamiakensisSirokamiakensisSirorubensSiroshastaSirovalleorumSuzukielussauteriSuzukielussauteri AMILYOGOVEIDAEOgoveacameroonensis AMILYTROGLOSIRONIDAETroglosiroaelleniTroglosirojuberthieiTroglosirolongifossaTroglosiromonteithiTroglosironinquaTroglosirourbanusTroglosirowilsoni
T F F F
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
EVOLUTIONARY AND BIOGEOGRAPHICAL HISTORY OF ARACHNIDS 9
1
– AY639460– – GQ488114GQ488121GQ488123 – GQ488128–GQ488119GQ488132GQ488133GQ488134– –GQ488136GQ488135–GQ488127 – –GQ488118GQ488117 ––GQ488115GQ488116– DQ825508.– ––HQ593871 JF786430–
GQ488181 AY639583GQ488153 – GQ488178GQ488144GQ488146 GQ488137 GQ488182GQ488184–GQ488167GQ488168GQ488171GQ488170 GQ488175GQ488176GQ488174GQ488155GQ488158 DQ825632 GQ488154–– ––GQ488180GQ488142– GQ488179– GQ488139–HQ593870 JF786414–
– GQ488195– GQ488230 DQ825611GQ488206GQ488226 GQ488221 DQ825610DQ518087GQ488204GQ488211GQ488212GQ488188– GQ488190GQ488213–GQ488192– – ––GQ488219 GQ488196–GQ488228GQ488193– –– GQ488194–– JF935028DQ825617
DQ825551 DQ825554GQ488439 GQ488451 GQ488435GQ488382GQ488385 GQ488402 GQ488454GQ488456GQ488380GQ488422GQ488359GQ488361GQ488428 GQ488433GQ488434GQ488432GQ488394GQ488352 GQ488398 GQ488390GQ488379GQ488377 GQ488396GQ488392GQ488436GQ488375GQ488444 DQ825564GQ488425 GQ488367GQ488371HQ593869 JF935011DQ825600
1
GQ488337 DQ133712DQ133717 GQ488322 GQ488317GQ488264GQ488267 GQ488283 DQ825532DQ518095GQ488262GQ488298GQ488299GQ488308GQ488307 GQ488250GQ488314GQ488312GQ488278DQ825534 DQ825530 GQ488276GQ488261GQ488233 GQ488280GQ488239DQ825526GQ488260GQ488328 DQ825527.GQ488305 GQ488252GQ488257HQ593868 JF934963DQ825545
733 61 8 97 1 9 3*9 2 33
85,98.43694 9972,98.983061667,134.5* 333,134.0333* 6667,110.31669833,107.01585694,106.5233 7,101.15* 9694,102.43051639,101.73864,107.012789028,125.1527425,119.735560583,100.66384611,100.0691 4583,100.54362111,98.497227722,100.35386667,98.413898528,98.43722 1806,98.94278 7,100.10*0778,90.231678833,98.27778 6667,98.413895,101.98*4,117.09*1,116.24*4,117.09* 6667,117.83330583,100.6638 1,116.53*5833,110.32978,125.09* 7972,72.6233-5000,67.6333-
8 27 8 777 6 3774013 92478 9 956 72686 01 074 16
7. 4.2. 0. 1.6.6. 6. 4.3.6.1.5.0.0. 0.3.0.9.7. 9. 6.5.7. 9.4.0.5.0. 1.0. 6.1.6. 0.5.
1- - -- - -- - - 22 1 -
aaaa aaa a
Thailand ThailandNewGuinea,IndonesiaNewGuinea,IndonesiaBorneo,MalaysiaJava,IndonesiaJava,Indonesia Thailand MalaysiaMalaysiaJava,IndonesiaSulawesi,IndonesiSulawesi,IndonesiSumatra,IndonesiSumatra,Indonesi Sumatra,IndonesiSumatra,IndonesiSumatra,IndonesiThailandThailand Thailand ThailandIndiaChina ThailandThailandBorneo,IndonesiaBorneo,MalaysiaBorneo,Indonesia Borneo,IndonesiaSumatra,Indonesi Borneo,MalaysiaBorneo,MalaysiaMindanao,Philippines ColombiaVenezuela
3 5 6 8 0 6,5 3,4 0 3,8
A100388,NA10106A100669A101064 A101510 A101514A101932A101944,NA10194A101093,NA10049A101483A101489A101930A101937A101938A102032A102033,NA10204A102039A102042A102061A103250A101094,NA10150A101494,NA10150NA10176A101767A102051A103242,NA10324NA10324A103251A103265A101519A103259A101468,NA10195A101517A102032,NA10205NA10205A103249A103254A104981 A101665A100869
NDNN N NNNDNDNNNNNNNDNNNNNDNDDNNNDDNNNNNDNNDDNNN NN
D DD D DDD D DDDDDDD DDDDD D DDD DDDDD DD DDD DD
E
Table2.Continued FAMILYSTYLOCELLIDAFangensisinsulanus FangensisspelaeusLeptopsalislydekkeri Leptopsalisnovaguinea Leptopsalissp.Leptopsalissp.Leptopsalissp. Leptopsalissp. Leptopsalissp.Leptopsalissp.Leptopsalissp.Leptopsalissp.Leptopsalissp.Leptopsalissp.Leptopsalissp. Leptopsalissp.Leptopsalissp.Leptopsalissp.Leptopsalissp.Meghalayasp. Meghalayasp. Meghalayasp.Meghalayasp.Meghalayasp. Meghalayasp.Meghalayasp.Miopsalissp.Miopsalissp.Miopsalissp. Miopsalissp.Miopsalissp. Miopsalissp.Miopsalissp.Miopsalissp. FAMILYNEOGOVEIDAEBrasilogoveasp.‘Brasilogovea’sp.Tobogan
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••
10 G. GIRIBET ETAL.
H3 JF786420–JF786421DQ518167JF786422JF786423–JF786424–––DQ825514DQ518168JF786425–––JF786426–––DQ825513JF786418JF786419––––DQ518169 DQ825519 AY639459DQ825517DQ825518JF786428DQ825515DQ825516JF786427JF786429– EF108592EF108593EF108594–EF108595EF108596
COI JF786395JF786399JF786396DQ518129JF786397––JF786398JF786400–––GQ912860JF786401JF786402JF786403–DQ825646–JF786404JF786408DQ825645JF786393JF786394JF786405JF786406JF786407JF786411DQ518131 DQ825650 ––DQ825650JF786409DQ825648–JF786410JF786412JF786413 EF108586EF108587EF108588EF108589EF108590EF108591
16SrRNA JF935029––DQ518050JF935032JF935033JF935030JF935031JF935036JF935037JF935038DQ825618JF935034JF935035–––DQ825619JF935042–JF935041DQ825616–JF935027–JF935039JF935040JF935047DQ518051 JF935043 –DQ825623DQ825624JF935045DQ825620DQ825621JF935044JF935046JF935048 EF108581EF108582EF108583EF108584–EF108585
28SrRNA JF934997JF935014JF935016DQ825596JF935015JF935012JF935017JF935013JF934999JF935003JF935004DQ825598DQ825597JF935000JF935001JF934998JF935002DQ825599JF935006JF935005JF935010DQ825595JF934995JF934996JF935007JF935008JF935009JF935019DQ825606 DQ825607 DQ825603DQ825604DQ825605JF935022DQ825601DQ825602JF935021JF935020JF935018 EF028096EF108576EF108577EF108578EF108579EF108580
5
18SrRNA JF934964JF934980JF934979DQ518090JF934977JF934981JF934982JF934978JF934972JF934988JF934989DQ825543DQ518091JF934970JF934971JF934973JF934984–DQ825544JF934986JF934987JF934983DQ825542JF934961JF934962JF934974JF934975JF934976JF934969DQ518092 DQ825550 AY639493DQ825548DQ825549JF934967DQ825546DQ825547JF934966JF934968JF934965 EF028095EF108573AF124942EF108574U37003EF108575
9387488358897 71778103 602 5 *
1377977377776 66167616 3235 03 3 0
3323522302293 684328515 95628 0 346147610
2385088398887 180788659847452 2 018818280
3454455435590 051055997016998 5 810111315
Coordinates 6.41055,50.--7.41667,72.-3.55833,76.-5.77956,73.-5.09956,75.-3.55833,76.-3.55833,76.-7.41667,72.-5.09542,75.-3.55833,76.-3.55833,76.-4.04495,69.--1.28500,78.-1.25,78.25*-5.48583,76.-3.56889,76.-1.61639,76.-1.28500,78.-3.55833,76.-3.56889,76.-1.33655,58.-30.56489,84.-35.06231,82.-32.18923,81.-4.19511,52.-4.08813,52.-1.38803,58.-0.50448,12.73.72570,8.83 3.70284,8.87 2.18305,9.1.65815,10.31.44858,9.782.74108,9.881.65815,10.32.13119,9.874.80084,9.662.74108,9.885.83333,7.3-
AA
SS
Locality BrazilColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaColombiaGuyanaFlorida,USASouthCarolina,USouthCarolina,UFrenchGuianaFrenchGuianaGuyanaGabonBioko,EquatorialGuineaBioko,EquatorialGuineaEquatorialGuineaEquatorialGuineaEquatorialGuineaCameroonEquatorialGuineaEquatorialGuineaCameroonCameroonIvoryCoast California,USASI,NewZealandOregon,USAJapanAustraliaMalaysia
04371061078804150926624534509 1 236927581 332871
87808547844015587048234422225 6 655155117 381102
66646166666446166466856688860 0 400600666 073363
her 105101101101101102104101101104104101101101102101101101101101105101105105104105105104101 101 100101101104101101104104105 101100100100100100
uc NANANANANANANANANANANANANANANANANANANANANANANANANANANANANA NA NANANANANANANANANA NANANANANANA
Vo DDDDDDDDDDDDDDDDDDDDDDDDDDDDD D DDDDDDDDD DDDDDD s.
e
at
n
di
or
o
m ec
Table2.Continued CangarenataeHuitacaventralisHuitacasp.Huitacasp.Huitacasp.Huitacasp.Huitacasp.Huitacasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.Metagoveasp.MetasiroamericanusMetasiroamericanusMetasiroamericanusNeogoveavirginieNeogoveavirginieNeogoveasp.ParogoviagabonicaParogoviasironoides Parogoviasironoides Parogoviacf.sironoidesParogoviacf.sironoidesParogoviacf.sironoidesParogoviacf.sironoidesParogoviasp.Parogoviasp.Parogoviasp.Parogoviasp.Parogoviasp. OUTGROUPSProtolophussingularisMegalopsalissp.HesperonemastomamodestuDendrolasmaparvulumEquitiusdoriaeSandokanmalayanus Asterisksindicateapproximat
© 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, (cid:129)(cid:129), ••–••