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OCCASIONAL PAPERS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
No. 151
August 13, 2001
MADAGASCAR
Contributions to the Study of the Biodiversity of Madagascar
Number 1
A MONOGRAPH OF THE MIGID TRAP DOOR SPIDERS OF MADAGASCAR AND
REVIEW OF THE WORLD GENERA (ARANEAE, MYGALOMORPHAE, MIGIDAE)
By
Charles E. Griswold and Joel Ledford
California Academy of Sciences
San Francisco, California
MADAGASCAR
Contributions to the Study of the Biodiversity of Madagascar
Number 1
A MONOGRAPH OF THE MIGID TRAP DOOR SPIDERS OF MADAGASCAR AND
REVIEW OF THE WORLD GENERA (ARANEAE, MYGALOMORPHAE, MIGIDAE)
(Abo\e) Female of Poecilomigas abrahami (O.P. Cambridge)
from Dlinza Forest, Zululand, South Africa
(Below) Nest oi Moggridgea teresae Griswold on fallen log
at Kirstenbosch. near Cape Town. South Africa
Photographs by Teresa Meikle
OCCASIONAL PAPERS
OF THE
CALIFORNIA ACADEMY OF SCIENCES
No. 151
August 13,2001
MADAGASCAR
Contributions to tiie Study of tlie Biodiversity of Madagascar
Number 1
A MONOGRAPH OF THE MIGID TRAP DOOR SPIDERS OF MADAGASCAR AND
REVIEW OF THE WORLD GENERA (ARANEAE, MYGALOMORPHAE, MIGID AE)
By
Charles E. Griswold and Joel Ledford
California Academy of Sciences
San Francisco, California
SCIENTIFIC PUBLICATIONS
Alan E. Leviton, Editor
Katie Martin, Managing Editor
© 2001 California Academy of Sciences, Golden Gate Park, San Francisco, California 94118
All rights reser\ed. No part of this publication may be reproduced or transmitted in any form or by
any means, electronic or mechanical, including photocopying, recording, or any information storage
or retrie\al system, w ithout permission in writing from the publisher.
ISSN 0068-5461
Printed in the United States of America
Norcal Printina, Inc., San Francisco, California
Table of Contents
Page
Abstract 7
Introduction 7
Materials and Methods 8
Acknowledgments 9
Material examined 11
Taxonomy 12
Migidae 12
Key to World Genera and Malagasy Species of Migidae 12
Migid genera not from Madagascar 14
Calathotarsiis Simon 14
Hetewmigas Hogg 15
Mallecomigas Goloboff & Platnick 15
Migas L. Koch 15
Moggridgea O. P. Cambridge 16
Poecilomigas Simon 17
Goloboffia Griswold & Ledford, new genus 17
Migidae of Madagascar 18
Paramiginae Petrunkevitch 18
Genus Micwmesomma Pocock 18
Micromesomma cowani Pocock 19
Genus Paramigas Pocock 20
Paramigas alluaiidi (Simon) 20
Paramigas andasibe Raven, new species 21
Paramigas goodmani Griswold & Ledford. new species 22
Paramigas macrops Griswold & Ledford. new species 23
Paramigas manakambus Griswold & Ledford, new species 24
Paramigas milloti Griswold & Ledford, new species 25
Paramigas oracle Griswold & Ledford, new species 26
Paramigas paitliani (Dresco & Canard) 27
Paramigas pectinatiis Griswold & Ledford, new species 28
Paramigas perroti (Simon) 29
Paramigas rothorum Griswold & Ledford, new species 30
Genus Thyropoeiis Pocock 31
Thyropoeus malagasus (Strand) 31
Thyropoeiis mirandus Pocock 33
Phylogenetics 34
Data 34
Outgroups 35
Analysis 35
Preferred tree 35
Character optimization and branch support 36
Character descriptions and interpretations 36
Characters not included in data matrix 43
Phylogeny of Migidae 43
Classification 44
Biogeography 44
Conclusions 47
Literature cited 47
Figures 1-68 51
Appendix 119
5
A MONOGRAPH OF THE MIGID TRAP DOOR SPIDERS OF
MADAGASCAR AND REVIEW OF THE WORLD GENERA
(ARANEAE, MYGALOMORPHAE, MIGIDAE)
Charles E. Griswold (1)* and Joel Ledford (2)
The Malagasy fauna of the spider family Migidae is monographed and a phylogeny for Malagasy
species and exemplars of the world genera is proposed. This phylogenetic analysis corroborates the
monophyly of Migoidea (Actinopodidae plus Migidae) and Migidae, and suggests that the Mala-
gasy migids form a monophyletic group (Paramiginae, sensu Petrunkevitch 1939, but not Raven
1985), that Migas, Moggridgea, and Poecilomigas form a monophyletic group (Miginae Simon
1892), that the Calathotarsinae Simon 1903 is paraphyletic with respect to the Miginae, and that
the South American migids form a monophyletic group. A key to the world genera and 14 species
of Malagasy Migidae is presented. The following new species are described: Paramigas goodmani,
P. macrops, P. manakambus, P. milloti, P. oracle, P. pectinatus, and P. rothorum, all authored by Gris-
wold and Ledford, and Paramigas andasibe Raven. Paramigas subrufus Pocock is a new junior syn-
onym of P. perroti (Simon). The male of Thyropoeus malagasus, the first male known for the genus,
is described. Goloboffia new genus is proposed for Migas vellardi Zapfe 1961, giving Goloboffla vel-
lardi new combination.
The trap door spider family Migidae has long at-
tracted attention because of its strikingly disjunct
southern continent distribution. Migids are known
from Australia, Africa, Madagascar, New Zealand,
New Caledonia and the southern cone of South Amer-
ica: virtually all parts of the former supercontinent
Gondwanaland except the Indian subcontinent and
Antarctica. Migids have figured prominently in discus-
sions of spider biogeography and of southern disjunc-
tions (Pocock 1903; Legendre 1979; Platnick 1981;
Nelson & Platnick 1984; Griswold 1991a). Reasonably
complete modem treatments exist for the African
(Griswold 1987a, 1987b), Australian (Raven 1984;
Main 1991), South American (Schiapelli & Gerschman
1975; Goloboff & Platnick 1987; Goloboff 1991), and
New Zealand faunas (Wilton 1968). The faunas of
Madagascar and New Caledonia have received less at-
tention. Dresco and Canard (1975) briefly reviewed the
Malagasy genera. Nothing has been published on the
migids of New Caledonia since Berland (1924). With
*To whom correspondence should be addressed: (1) Schhnger Cura-
tor of Arachnida, Department of Entomology, Cahfomia Academy
of Sciences, Golden Gate Park, San Francisco California 94118
USA and Research Professor of Biology, San Francisco State Uni-
versity; Fax: (415) 750-7228; Internet: cgriswold@calacademy.org;
(2j California Academy of Sciences Summer Systematics Institute,
Golden Gate Park, San Francisco California 94118 USA and De-
partment of Biology, San Francisco State University; Fax: (415)
750-7228; Internet: jledford(gicalacademy.org.
this paper, the 15th in a series treating the afromontane
spider fauna (Griswold 1985, 1987a, 1987b, 1987c,
1990, 1991b, 1994, 1997a, 1997b, 1998a, 1998b, 2000,
2001 ; Griswold and Platnick 1987), we monograph the
Malagasy migids utilizing all available material.
Madagascar is widely recognized as being of great
conservation importance (National Research Council
1980; Rasoanaivo 1990; Myers et al. 2000) because the
island is known for high rates of endemism and unique
occurrence of primitive members of otherwise wide-
spread taxa (Myers 1988). Ongoing rapid habitat de-
struction, particularly of forests, makes the collection,
description, and study of the evolutionary and biogeo-
graphic significance of the Malagasy biota particularly
urgent.
The spider fauna of Madagascar remains poorly
known. Since the first Malagasy spider {Augusta
glyphica Guerin 1839) was described more than 150
years ago, nearly 150 papers have been published on
Malagasy spiders. The number of spiders recorded
from the whole island only slightly exceeds 450 de-
scribed species (Roth 1992a; Griswold, in press), sig-
nificantly less than the 626 species recorded from the
British Isles (Merrett et al. 1985, Merrett & Millidge
1992). Yet, nearly 400 species (new and described to-
gether) have been collected from a single site (Pare Na-
tional Ranomafana) in the southern part of the island
(Roth 1992b), suggesting a rich fauna yet to be de-
OCCASIONAL PAPERS OF THE CALIFORNIA AC ADEMY OF SCIENCES, NO. 151
scribed. The important afromontane spider families
Cyatholipidae (Griswold 1997a) and Phyxelididae
(Griswold 1990) have only recently been recorded
from the island. Remarkably, a new family (Halidae)
was recently described from Madagascar from hitherto
undescribed species (Jocque 1994). Alderweireldt &
Jocque (1994) suggest that the known component of
the Malagasy spider fauna is around 10%. Our knowl-
edge of the evolutionary relationships of Malagasy spi-
ders is poorer still: of the more than 120 papers that
deal explicitly with the taxonomy of Malagasy spiders,
fewer than 1 present cladograms or other explicit phy-
logenetic hypotheses. Clearly, much remains to be
learned about this rich fauna.
Our studies of the Malagasy migids benefit from
two recent developments: (1)3 well supported cladis-
tic hypothesis of migid affinities and (2) new migid
material collected in the course of general sui"veys of
Madagascar biGdi\'ersiry. Raven (1985) and Goloboff
(1993a) proposed mygalomorph cladograms that con-
finned a Migoidea (sensii Platnick & Shadab 1976)
comprising Migidae and Actinopodidae and suggested
Ctenizidae and Idiopidae as close reiati\es. Our phylo-
genetic analysis builds on this hypothesis. After initial
collections of Malagasy migids in the late 19th century
k\\ of these spiders were collected for the next 100
years. Rene\\ed interest in the Malagasy biota and con-
cern for impending extinctions led to new efforts to
survey the island. In particular, the efforts of Steve
Goodman and Brian Fisher in the 1990s to collect ter-
restrial arthropods and small vertebrates produced as
many migid specimens as collected in the pre\ious 100
years. Nevertheless, our attempts to monograph the
Malagasy migids and understand their phylogen>' are
hindered by absent and ambiguous data. Of the ten
world genera two (Mallecomigas and Micromesomma)
are still known only from females. Malagasy migids re-
main ver)' rare in collections: we ha\e examined only
113 specimens, adults and ju\eniles alike. Many
species are known from single specimens, and collect-
ing data may be absent or ambiguous. Of the 14 Mala-
gasy species that we recognize, only one (Thyropoeiis
malagasiis) is known with certaint\' from both sexes.
Indeed, only once ha\ e a male and female of a Mala-
gasy migid species been collected in association (in
1996 b\' Goodman in a pitfall trap at Vohimena)!
Recognizing these shortcomings in the data, we at-
tempt to understand the phylogenetic position of Mala-
gasy migids. re\iew the biology of the family, describe
the Malagasy fauna, and pro\ide a key to the world
genera and Malagasy species. We are confident that
continued exploration of the arthropod faunas of the
southern continents will complete the picture of the
Migidae and provide tests of the hypotheses presented
herein.
MATERIAL AND METHODS
Conventions — Throughout the text, figures cited
from previous papers are listed as "fig." and those ap-
pearing in this paper as "Fig." All illustrations are by
Jenny Speckels (JS), Joel Ledford (JL), Virginia Kirsch
(VK) or Charles Griswold (CG) and are attributed in
the figure captions. When referring to size, 'small' =
<10mm, 'medium'= 10-20 mm, and Targe' = >20 mm.
In the discussion of biogeographic history "ma" refers
to millions of years before the present. Records of dis-
tribution in Madagascar presented in Figure 68 include
localities for the juveniles listed in Table 1 .
Descriptions — Each genus is treated, a brief tax-
onomic history is provided, major diagnostic features
are listed, and published reports on the biology are
summarized. For Malagasy and Afrotropical genera
the description summarizes all included species: for
other genera the description reflects the exemplars (see
Table 2) supplemented by the literature. Species de-
scriptions follow the format in Griswold (1987a). Each
description is of a single individual of each sex, which
is identified either as a type or by the locality at which
it was collected. Illustrations cited in the description
may not refer to the specimen described but serve to
describe the features reported. For each sex there is
also a section reporting the variation in the most con-
spicuous and variable features. For variation in quanti-
tative features three to five adult individuals represent-
ing the full range in overall size were chosen to sample
the species: the number (N) is reported at the beginning
of each section. All measurements are in millimeters.
Abbreviations used in the text and figures are listed in
Table 3.
Eyes are measured from above. The sternal sigilla
are the concave regions near the sternal margin as
viewed in oblique lighting, not the discolored area as-
sociated with these structures. Spines are recorded as
arising from the dorsal (d), prolateral (p), ventral (v) or
retrolateral (r) surface of the segment and are recorded
from proximal to distal. For example, tibia p 0-0-1, d
1-0-0. r 0-1-1 signifies that the tibia has a proximal
spine on the dorsum, a distal spine on the prolateral
side, and a median and distal spine on the retrolateral
side. The convention r 0-0- la signifies that the retro-
lateral spine is at the apex of the segment (a = apical).
Especially on the lateral surfaces of the tibiae-tarsi of
the female pedipalpus and legs I and II spines may be
in diagonal series oriented in a proximo-dorsal to disto-
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS 9
Table 1: Juvenile Paramigas from Madagascar (ail specimens are in CASC)
Antsiranana Province:
Reserve Speciale Manongarivo. 10.8 km SW Antanambao (13°57.7'S, 48°26'E) elev. 400 m, rainforest, sifted litter, 8 No-
vember 1998, B.L.Fisher
Reserve Speciale d'Anjanaharibe-Sud, 12.2 km WSW Befingotra (14°45'S, 49°26'E) elev. 1985m, montane rainforest, sift-
ed litter. 25 November 1994, B.L. Fisher
Toamasina Province;
Presquil de Masoala, 5.3 km SSE Ambanizana, Andranobe (15°40'S, 49°58'E) elev. 425 m, rainforest, sifted litter, 21 No-
vember 1993. B.L. Fisher
SF Tampolo, 10 km NNE Fenoarivo Atn. (17°17'S, 49°26'E) elev 10 m, littoral rainforest, sifting litter, 4 April 1997, B.L.
Fisher
Fianarantsoa Province:
Reserve Speciale Ivohibe, 8 km E Ivohibe (22°29'S, 46°58.rE) elev. 1200 m, forest, sifting leaf litter, 15-21 October 1997,
B.L. Fisher
Reserve Andringitra, 38 km S Ambalavao (22°12'S, 46°58'E) elev. 1680 m, montane rainforest, sifted litter, 23 October 1993,
B. L. Fisher
Reserve Andringitra, 38 km S Ambalavao (22°12'S, 46°58'E) elev. 1875 m, montane rainforest, sifted litter, 29 October 1993,
B. L, Fisher
Reser\'e Andringitra, 8.5 km SE Antanitotsy (22°10'S, 46°58'E) elev. 1990 m, rainforest, sifting litter, 6 March 1997, B. L.
Fisher
Toliara Province:
Mahafaly near Eloetse, by Lac Tsimanampetsoa (24°10'S, 43°45'E) 15-16 September 1992, V. & B. Roth
Somhem Isoky-Vohimena Forest (22°4rS, 44°50'E) elev. 730 m, tropical dry forest, sifted litter, 21 January 1996, B.L. Fish-
er
Foret de Petriky, 12.5 km W Tolagnaro (25°03.73'S, 46°52.16'E) elev. 10 m, littoral rainforest, sifting litter, 22 November
1998, B.L. Fisher
ventral direction, with the smallest spines most dorsal
and the largest most ventral (Figs. 1, 24A): in this case
the convention P 1-1-2-4-3 signifies that on the prolat-
eral surface there are two proximal spines in a row fol-
lowed by diagonal series of 2, 4 and 3 spines. We de-
fine "thorns" as socketed setae that taper to the apex
and that are less than 5x as long as the diameter at the
base (Fig. 25D; Griswold 1987a, figs. 23-26). Thorns
have a nearly smooth surface. Cuspules are socketed
setae that are narrow at the base, widest medially, and
blunt to pointed at the apex, and deeply grooved (Gris-
wold 1987a, figs. 358, 359) or smooth (Figs. 43C-D).
Localities in Madagascar may be hard to trace due
to many identical or similar place names, abbreviation
of place names on labels, illegible labels, and name
changes over time. We have tried our best to decipher
difficult labels. The recent compilation of field stations
where insects have been collected in Madagascar (Vi-
ette 1991) has been very helpful.
Specimen Preparation — Vulvae were examined
in lactic acid. In some cases they were cleaned by ex-
posure to trypsin or cleared with Clorox® bleach. Ex-
aminafion was via Wild M5Apo, Olympus SZHIO,
Leica MZApo and Leitz Ortholux II microscopes.
Prior to examination with a Hitachi S-520, JEOL T-
200 or Cambridge Stereoscan Scanning Electron Mi-
croscope (SEM), large structures were air-dried and
delicate structures (spinnerets, whole mounts of juve-
niles) were critical-point-dried.
Phylogenetics — On the preferred cladogram
(Fig. 65) all nodes were assigned a letter (e.g.. A, B).
Throughout the discussions of the phylogenetic analy-
sis and relationships among taxa these letters are used
to refer both to nodes on the cladogram (e.g., node N)
and clades distad to that node (e.g., clade N, compris-
ing Migas, Moggridgea and Poecilomigas). For fijrther
discussion see 'phylogenetic methodology' under
'Phylogenetics' below.
ACKNOWLEDGMENTS
Support for this project came from National Sci-
ence Foundation grants BSR-9020439 and DEB-
9020439 to Griswold and DEB-0072713 to Brian Fish-
er and Griswold. Initial participation by Ledford was
made possible by the California Academy of Sciences
(CASC) Summer Systematics Institute, supported by
NSF grant BIR-9531307. Additional space and equip-
ment support was provided to Ledford by the Ento-
10 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 1 5 1
Table 2: Exemplars and other taxa examined for migid phylogenetic analysis (note: M=male; F=female).
Outgroups:
Actinopus rufipes (Lucas, 1834): F from Brazil, Barra de Pirai, Rio de Janeiro, det. Bucherl (AMNH)
Actinopus sp.: F from Tobago, Grafton Estate (AMNH)
Actinopus sp.: FF from Colombia, Meta, Puerto Lleras, Lomalinda, elev. 300m., lyiT'^N, 3°18'N, 15 April 1986, B. T. Car-
roll (CASC), MM , same locality, March 1987 (CASC)
Actinopus sp.: M from Argentina, Corrientes, 29 April 1964, E. S. Ross (CASC)
Bothriocyrtum califomicum (O.P. Cambridge, 1874): M from USA, California, Los Angeles Co., campus of University of
California at Los Angeles, November 1955, M, Maher (CASC); F from Los Angeles Co., Eagle Rock, July 1991, C.E.
Griswold (CASC)
Idiops sp.: ME from South Africa, Natal, Pietermantzburg, 1-15 October 1985, C. & T. Griswold (CASC)
Non-Malagasy Migidae:
Calathotarsus coronatus Simon, 1903: specimens from Chile, Valparaiso, Parque Nacional La Campana, Cerro La Campana,
M from elev. 980m.. 1 1 November 1986, F from 1080 m., 26 August 1981, R. Caderon (AMNH); also M from Santia-
go Prov.. Quilicura. September 1979, Pena (AMNH) and F from El Canelo, elev. 950 m., 8 September 1966, E.
Schlinger & M. Irwin (CASC)
Calathotarsus simoni Schiapelli & Gerschman, 1975: F from Argentina, Buenos Aires, Cerro Negro, Sierra de la Ventana,
April 1974, Cesari (AMNH)
Heteromigas dovei Hogg, 1902: MF from Australia. Tasmania. Patersonia. 29 January 1926, V. V. Hickman (QMB)
Migas vellardi Zapfe, 1961 : holotype M from La Herradura, Coquimbo Province. Chile (UCS); F from Chile. Coquimbo Re-
gion. Guanaqueros. 10 January 1984. E. Maury & P Goloboff (MACN 8338)
Mallecomigas schlingeri Goloboff & Platnick, 1987: holotype F from Chile, Araucania Region, elev. 610m in Cordillera de
Nahuelbuta, 18 km W Angol, 10 February 1967, E.I. Schlinger (UCS at CASC). Note: This specimen has been trans-
ferred to UCS.
Migas exemplars:
Migas gatenhyi Wilton. 1968: M (fragmentary) from New Zealand, Wellington, Town Belt, Oriental Bay, 41°17'S, 174°46'E,
1 1 January 1995, L.J. Boutin (CASC), F from same locality, 17 April 1995, L.J. Boutin (CASC)
Migas taierii Todd, 1945: M from New Zealand, Patearoa (AMNH)
Other Migas:
Migas affinis Berland, 1 924: holotype M. from New Caledonia, Foret du Mt. Panie, 27 May 1911. Roux & Sarrasin (MNHN
AR4132)
Migas distincius O.P. Cambridge, 1879: F from New Zealand, Portobello (AMNH)
Migas giveni Wilton, 1968: F from New Zealand. N. Island. Waipoua Forest. Yakas Track, 2 February 1994, E.I. Schlinger
(CASC)
Migas nitens Hickman. 1927: F from Cornelian Bay, Tasmania, 28 August 1930, V. V. Hickman (MNHN AR4131)
Migas paradoxus L. Koch. 1873: F from New Zealand. C. L. Wilton (AMNH)
Migas variapalpus Raven. 1984: M from .Australia. Queensland, O'Reilly's, Lamington National Park, elev. 935m, 28°13'S,
153°08'E, 12 July 1992, C.E. Griswold (CASC)
Migas sp.: M from Tauranga, New Zealand (OMD)
Moggridgea crudeni Hewitt, 1913: F from Alicedale, South Africa. F Cruden (AMSA)
Moggridgea dyeri O. P. Cambridge, 1875: M from Uitenhage, South Africa, F. Cruden (AMSA)
Moggridgea intermedia Hewitt, 1913: ME from South Africa, Western Cape, Diepwalle Forest Station, 21 km. N. Knysna,
33°57'S, 23°09'E, elev. 540 m, 12 December 1996, C.E. Griswold (CASC)
Moggridgea pseudocrudeni Hewitt, 1919: M from Alicedale. South Africa (AMSA)
Moggridgea peringueyi Simon. 1903: M from South Africa, Cape. Oudtshoom, 29 October 1949, B. Malkin (CASC); E
(holotype) from South .■Africa, Cape, Matjiesfontein (MNHN 19274)
Moggridgea rupicola Hewirt, 1913: MMF from South Africa, Alicedale (AMSA)
Moggridgea tingle Main, 1991: paratypes, from Walpole National Park in Western Australia, M from Big Tingle Tree, 16
July 1908 (WAM 89'330), EE from Valley of the Giants, 14 January 1990 (WAM 90/1 1 12) and Deep River crossing at
Manjinup, 14 January 1990, B.Y. Main (WAM 90/1 113)
Poecilomigas abrahami (O.P. Cambridge, 1889): F from South Africa, Natal, Sordwana Bay, elev. m, 50 mi. E of Ubom-
bo, 5 April 1958, E. S. Ross (CASC); E from South Africa, Uitenhage (AMSA); MM from South Africa, Natal, Pieter-
maritzburg. Town Bush, elev. 3000 ft., 29=33 'S, 30^1 E, 8 January 1984, T. Meikle (CASC), same data (NMSA).
Poecilomigas basilleupi Benoit, 1962: ME from Tanzania, Tanga Region. West Usambara Mts., Mazumbai, 4°49'S, 38°30E,
elev. ca. 1400 m, 10-20 November 1995. C. Griswold, D. Ubick, & N. Scharff, (CASC)
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
11
Table 3 : List of Anatomical and Other Abbreviations
Used in the Text and Figures.
a
apical
AER
anterior eye row
ALE
anterior lateral eyes
AME
anterior median eyes
d
dorsal
HS
spermathecal head
ITC
inferior tarsal claws
OAL
ocular area length
OAW
ocular area width
P
prolateral
PER
posterior eye row
PLE
posterior lateral eyes
PME
posterior median eyes
r
retrolateral
SS
spermathecal stalk
STC
superior tarsal claws
V
ventral
mology Department, University of California, Davis.
Additional support came from the Exline-Frizzell and
In-house Research Funds of the CASC and the
Schlinger Foundation. The 1998 CASC expeditions to
Ranomafana, Madagascar were supported by grants
from the Oracle and McBean Foundations.
Robert Raven provided the specimens and de-
scriptive notes on Paramigas andasibe, which new
species is attributed to his authorship. Specimens and
observations essential to the completion of this study
were obtained through fieldwork in Australia, Mada-
gascar, and New Zealand. This fieldwork was made
possible through permissions granted by a variety of
government bodies. Permits to do research in Australia
were granted by the Queensland National Parks and
Wildlife Service, facilitated by Dr. R. Raven of the
Queensland Museum. Permits to do research in and ex-
port specimens from Madagascar were granted by the
Association Nationale pour le Gestion des Aires Pro-
tegees (ANGAP) and Direction des Eaux et Forets of
the Ministre d'Etat a L' Agriculture et au Development
Rural, under Accordes de Collaboration of the Xerces
Society (facilitated by Dr. C. Kremen, Mr. C. Ramili-
son, and Ms. B. Davies of that organization) and Insti-
tute for Conservation of Tropical Environments
(ICTE) (facilitated by Drs. R. van Berkum and Ben-
jamin Andriamihaja). A permit to collect specimens in
New Zealand protected areas was granted by Te Papa
Atawhai (Department of Conservation). Research in
Tanzania was made possible through a Research Per-
mit from the Tanzania Commission for Science and
Technology (COSTECH) and Residence Permit Class
C from the Tanzanian Department of Immigration, and
export of specimens made possible by a CITES Ex-
emption Certificate from the Wildlife Division of the
United Republic of Tanzania, facilitated by Professor
Kim M. Howell of the University of Dar-es-Salaam.
Research at Mazumbai was made possible by Dr. S. A.
O. Chamshama, Dean of Forestry, Sokoine University,
Morogoro, and Mr. Modest S. Mrecha, Officer in
Charge. Mazumbai Forest Reserve. Research in South
Africa was made possible through a permit from the
Department van Waterwese en Bosbou.
The following are thanked for their hospitality:
(Australia) Drs. Robert Raven and Val Davies; (Bel-
gium) Dr. Rudy Jocque; (France) Drs. Christine Rol-
lard and the late Jacqueline Heurtault; (New Zealand)
Drs. Lyn and the late Ray Forster; (UK) Mr. John and
the late Mrs. Frances Murphy, Mr. Paul Hillyard;
(USA) Drs. Norman Platnick and Petra Sierwald. For
generous hospitality and able assistance in Madagas-
car, especially in fieldwork, I thank the following: Mr.
Rija Andriamasamanana, Ms. Marie Jeanne Raheri-
lalao, Mr. Samuelson Randrianarisoa, Mr. Christophe
Roland and Ranomafana Park Ranger Mr. Emile Raje-
riarison.
Dr. David Kavanaugh deserves special thanks for
help in the field as do the members of the California
Academy of Sciences/Pare Botanique et Zoologique
Tzimbazaza (CAS/PBZT) 2000 Ant-Spider course:
Daniella Andriamalala, Balsama Rajemison, Jean-
Jacques Rokotoarison, Jean Claude Rakotonirina and
Helian Ratsirarson. Assistance with Scanning Electron
Microscope examination was provided by Mr. Darrell
Ubick (CASC), and Mr. Vijay Bandu, Mrs. Connie
Bowland, and Mr. Antony Bruton (University of
Natal). Darrell Ubick and Mr. Keith Dabney assisted in
ways too numerous to mention. A draft of the manu-
script was critically read by Fred Coyle, Pablo
Goloboff, Norm Platnick, Robert Raven and David Ka-
vanaugh. Agreement with all of our conclusions is not
implied
This is contribution #14 from the CASC Center for
Biodiversity Research and Information (CBRI).
MATERIAL EXAMINED
The following institutions and individuals lent or
otherwise provided specimens: Ms. Lisa Joy Boutin,
Tasmania, Australia; Dr. Ariel Camousseight, Museo
Nacional de Historia Natural, Universidad de Chile,
Santiago (USC); Mr. Peter Croeser, Natal Museum,
Pietermaritzburg, South Africa (NMSA); The late Dr.
Ray Forster, Otago Museum, Dunedin, New Zealand
(OMD); The late Dr Maria Elena Galiano, Museo de
Ciencias Naturales "Bernardino Rivadivia", Buenos
Aires, Argentina. (MACN); Dr. Fred Gess, Albany
12
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Museum, Grahamstown, South Africa (AMSA); Dr.
Mark Harvey, Western Australian Museum, Perth,
Australia (WAM); Mr. Paul Hillyard and Mrs. Janet
Beccaloni, The Natural History Museum, London, UK
(BMNH); Dr. Rudy Jocque, Musee Royal de L' Afrique
Centrale, Tervuren, Belgium (MRAC); Dr. Torbjom
Kronestedt, Naturhistorika Riks-museet, Stockholm,
Sweden (NRS); Dr. Norman I. Platnick, American Mu-
seum of Natural History, New York, USA (AMNH);
Dr. Robert J. Raven, Queensland Museum, South Bris-
bane, Australia (QMB); Dr. Christine Rollard and the
late Dr. Jacqueline Heurtault, Museum National d'His-
toire Naturelle, Paris, France (MNHN); Mrs. Barbara
and the late Mr. Vincent D. Roth, Portal, Arizona; Dr.
Petra Sierwald and Mr. Phil Parillo, Field Museum of
Natural History. Chicago, USA (FMNH); The other
specimen depository listed is ZFMK (Staatliches Mu-
seum fiir Naturkunde, Karlsruhe, Germany).
TAXONOMY
60A). STC with tooth or single row of teeth, ITC with
or without teeth. Trichobothrial pattern: pedipalpal tar-
sus with a dorsal row of 4—6, row may divide proxi-
mally, tibiae with dorsobasal converging rows of 3—6,
metatarsi with group of several near dorsoapex, poste-
riad of this point trichobothria form a single dorsal row
that may extend to metatarsal base, tarsi with numerous
trichobothria (10-25) on their dorsal surface (Fig.
27D); trichobothrial base distally embedded, with
proximal hood, smooth or weakly corrugiform (Figs.
24C, 42D, 55C; Griswold 1987a, figs. 41-42). Tarsal
organ in apical half of segment (Fig. 27D), oval to
round, weakly domed, with concentric ridges (Figs.
24D, 42D, 55D; Griswold 1987a, figs. 43^4). Posteri-
or lateral spinnerets 3- segmented, basal segment
longest, median and distal approximately equal in
length, distal segment domed, length less than 2/3 di-
ameter, median segment with 12^0 spigots, apical
with 30-50; anterior lateral spinnerets single segment-
ed, separated at base by diameter, 0.5 x length PLS,
with ventroapical group of 4— 20 spigots (Figs. 3A— B);
Migidae male with rectangular field of epiandrous spigots ante-
^. . ^. . . , , ^ „ , , riad of epigastric furrow, spigots evenly spaced, sepa-
Diaenosis. — Distmguished from all other mygalo- ,, ,,,,•, , ^ ,
, , , . , ... ^ , , rated by approximately V2 their length. Spermathecae
morphs by having the combination 01 characters rastel- ,_. ,-,.v. -.^. t-^ ■ j ■ , , . ■
, ^. „„ ^„ __, ,, . . (Figs. 17A-C, 35A-E) paired, simp e, unbranched,
lum absent (Figs. 2B, 5B, 9B), tangs quadrate and ^ ® , ,. ' , ', ■ . J
■ ■ ^- .,^ ..-.. ■ ■ ^ • , straight to distally curved, with pores throughout, un-
keeled (Fiss. 2C, 43 A), thoracic fovea straight to re- , • , , ■ . » , , ■ ,
, ,r- , ^ A X J T,r r. -1 • J- sclerotized or sclerotized. Male with or without clasp-
curved (Fies. 1, 2A), and PLS with spigots on median ,.,,_. -„„^ ., ,„r
," . , , ,-r- , . r^. ing processes on legs, with (Fig. 30B) or without (Fig.
and distal articles only (Figs. 3A-B . ,°J^ ■ . c t j- T
^ r,,. , J , 1 ID) megaspine at retroapex of metatarsus I, pedipal-
Descnption. — Small to large sized mygalo- -.u u lu ■ 1 r /c- ta ^ 1 , a r-
^ ,^ . , , „ , „ pus with bulb simple, pinform (Figs. 7A-C, llA-C,
morphs. 5-45 mm in length. Sparsely setose. Carapace , ita r» ^M^_n\
smooth in temale (Fig. 1), rugose in males (Figs. 6,
52), length 1-1.25 x width; caput low to arched. Ocu-
lar area 0.40-0.65 x width carapace; AER straight to KEY TO WORLD GENERA AND
slightly recur%'ed (Figs. 2A, D). Thoracic fovea re- MALAGASY SPECIES OF MIGIDAE
curved, in some specimens straight (Figs. 29, 52),
0. 12-0.40 X width carapace. Chelicerae lacking rastel- ' Females 2
lum, fangs quadrate, with keels (Fig. 43 A), in most ~ Males 19
species obliquely oriented (Fig. 2C), pro and retromar- -,,,t ., , ,, , ,. ,r- ,,^ ,,^ ,,„
. r^ n L f.T- -.^r^ ^^T^^ o 2( 1 ) Tarsi I aud Illacking spiucs (Figs. 1 3C, 1 5C, 16B,
gins of fang fijrrow with teeth (Figs. 24E, 43B). Ster- ^ &f \ & ' -
num length 1.0-1.50 x width, with 1 pair sigilla oppo- _' ' '. ' ' ' ' 'tt ■'. • /t-- i ^ a<^\ c
„ TTT .T- ^r^. ^ , , 1 J ,, -Tarsi I and II With spines (Figs. 1, 14C) 5
site coxae Il-III (Fig. 5B). Cuspules (socketed, blunt ^ v & ,
macrosetae) present on labium and pedipalpal coxae at 3(2) Patellae I, II, and IV with ventral setae normal, ta-
least of females (Figs. 2C, 43C, D); serrula absent (Fig. pe^ng from base to apex (Fig. 13C); fang with basal
43C). Leg formula 4123 or 1423; legs I and II of fe- jqqij^ (pjg 59j)\ 4
males with strong series of spines (Figs. 1, 57) on pro- _Patellae I, II, and IV with ventral patches of lamellate
and retrolateral margins of tibiae, metatarsi, and, in ^^^^ (pigs. 15C, 16B, 6 ID); fang without basal tooth
some species, tarsi, these spines in most species form- Moggridgea
ing diagonal rows; legs III and IV with dense patterns
of spinules and stout setae, especially dorsally and pro- 4(3) Dark dorsal and lateral maculations forming trans-
laterally on patellae and on metatarsus III (Figs, verse bands or annuli on tibiae and metatarsi (Fig. 18);
25A-C, 44D); scopulae absent from females, present metatarsus IV preening comb absent. . . Poecilomigas
beneath at least some tarsi in males (Figs. 30B, 3 IF, - Tibiae and metatarsi lacking such dorsal and lateral
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
13
maculations (Fig. 13C); metatarsus IV preening comb
present in some species (Fig. 59C) Migas
5(2) Tibia III cylindrical, convex dorsally at base (Figs.
4, 8, 14D, 59A); caput setose, with rows or at least a
few setae posteriad of the ocular area (Figs. 8, 12A) .
6
-Tibia III concave dorsally at base (Figs. 1, 25A-C);
caput lacking setae except some species with pair of
prefoveal setae (Figs. 2A, 23A, 50) 9
6(5) Patellae lacking ventral lamellate setae (Figs. 12C,
D); fang unarmed (Figs. 5B, 12B) 7
— Patellae with ventral lamellate setae (Fig. 14C); fang
with basal tooth (Fig. 14B)
Goloboffia vellardi (Zapfe)
7(6) Pedipalpal tibia cylindrical, tapering to apex, un-
modified, without cluster of spines (Fig. 8); OAW less
than 0.5 x width of caput (Figs. 8, 12A) 8
— Pedipalpal tibia swollen ventrally, with dorsoapical
cluster of spines (Figs. 4, 5C); OAW greater than 0.5 x
width of caput (except C. pihuychen) (Fig. 4)
Calathotarsus
8(7) Cuspules on pedipalpal coxae restricted to base
near labium (Fig. 12B)
Mallecomigas schlingeri Goloboff & Platnick
— Cuspules on pedipalpal coxae extending for length of
coxae (Fig. 9B) Hetemmigas
9(5) Sternal sigilla deeply excised, lunate (Figs. 5 IB,
58B) 10
— Sternal sigilla flat (Figs. 23B, 34B) 11
10(9) Fang unarmed (Fig. 58B); eyes reduced, PLE di-
ameter less than 0.4 x ALE (Fig. 57); total length
greater than 30mm. . . . Thyropoens mirandus Pocock
— Fang with basal tooth (Fig. 5 IB); eyes normal, PLE
diameter greater than 0.5 x ALE (Fig. 50); total length
less than 20mm Thyropoeus malagasus (Strand)
11(9) Leg coxae with only ordinary, slender setae (Fig.
34B); carapace lateral margin evenly rounded posteri-
orly (Fig. 33) 12
— Leg coxae II and III with thorns (Figs. 23B, 25D);
carapace lateral margin narrowed posteriorly (Fig. 22)
Micromesomma cowani Pocock
12(11) Dorsally legs I and II with slender procumbant
setae at apices of tibiae and bases of metatarsi (Figs.
27A-C, 41B) 13
— Dorsally legs I and II with thorns at apices of tibiae
and bases of metatarsi (Fig 33)
Paramigas goodmani, new species
13(12) Dense vestiture of long, curved, silky hairs be-
neath at least tibiae and metatarsi I and II (Figs. 33,
41B, 47C) 14
— Long, curved, silky hairs sparse or absent from be-
neath tibiae and metatarsi I and II (Figs. 1, 28B) .15
14(13) Spermathecae elongate, length greater than 1.67
X head diameter, head diameter less than 2.30 x diam-
eter stalk (Figs. 35A-C, 47 G-H)
Paramigas perroti (Simon)
— Spermathecae short, length less than 1 .67 x head di-
ameter, head diameter greater than 2.14 x diameter
stalk (Figs. 40A, B, 41C, D)
Paramigas oracle, new species
15(13) Tibia I with fewer than 30 ventrolateral spines
(Fig. 46B); prefoveal setae small (Fig. 46A) to absent
(Fig. 48); spermathecae short, length less than 1 .20 x
head diameter, head diameter greater than 3.50 x di-
ameter stalk (Figs. 45A-C) 16
— Legs I and II densely spinose, tibia I with more than
40 ventrolateral spines (Fig. 1); prefoveal setae large
(Fig. 39A); spermathecae long, length greater than
1.63 X head diameter, head diameter less than 1.81 x
diameter stalk (Figs. 35E, 39C)
Paramigas milloti, new species
16(15) Carapace with conspicuous prefoveal setae
(Figs. 44A, 46A) 17
— Prefoveal setae minute to absent (Figs. 28A, 48). 18
17(16) Legs I and II dorsally with patches of procum-
bant setae at apices of tibiae and bases of metatarsi
sparse (<10) (Fig. 44C)
Paramigas pauliani (Dresco & Canard)
— Legs I and II dorsally with patches of conspicuously
serrate, procumbant setae at apices of tibiae and bases
of metatarsi dense (>20) (Fig. 46B)
Paramigas pectinatus, new species
18(16) Labium with fewer than 10 cuspules; dense net-
work of reticulate striae on lateral margins of caput
(Fig. 28A) and longitudinal banding patterns on legs
(Fig. 28B) Paramigas alhtaudi (Simon)
— Labium with more than 15 cuspules (Fig. 49B); lat-
eral margins of caput without striae and legs not longi-
tudinally banded (Fig. 48)
Paramigas rothorum, new species
19(1) Fang with basal tooth (Fig. 53B) 20
— Fang lacking basal tooth (Figs. 30A, 37B) 24
20(19) Tibia I uiunodified, cylindrical; sternal sigilla
flat (Fig. 30A); pedipalpal tarsus with apical spinules
(Figs. 20A-B, 37E-G) 21
14
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
— Tibia I swollen prolaterally (Figs. 52, 55A); sternal
sigilla deeply incised (Fig. 53B); pedipalpal tarsus
lacking apical spinules (Figs. 53E-G, 54B-C)
Thyropoeus malagasus (Strand)
21(20) OAW <0.41 X carapace width; embolus longer
than bulb (Fig. 20A) 22
— OAW >0.44 X carapace width; embolus length equal
to bulb (Legendre & Calderon Gonzalez 1984, fig. X,
8) Goloboffia vellardi (Zapfe)
22(21) Scopulae present beneath tarsi III and IV (Fig.
60A); femora with dorsal spines short or absent (Gris-
wold 1987b, fig. 70) 23
— Scopulae absent fi-om tarsi III and IV (Fig. 60B);
femora with dorsal spines long (Griswold 1987b, fig.
69) Migas
23(22) Dark dorsal and lateral maculations or annuli on
tibiae and metatarsi; embolus straight (Figs. 20A-B).
Poecilomigas
— Tibiae and metatarsi lacking such dorsal and lateral
maculations or annuli; embolus reflexed (Raven 1984
fig. 6) Migas variapalpus Raven
24(19) Femur I venter rounded Figs. 7D, 30B); apical
lobes of pedipalpal tibia subequal (Figs. 7B, 1 IC, 32C)
25
— Femur I venter carinate (Fig. 60C); ectal apical lobe
of pedipalpal tibia longer than mesal (Figs. 16A, 32D)
Moggridgea (Africa)
25(24) Caput low, not highly arched, height less than
2.0 X height at thoracic fovea; retroapex of tibia I with
megaspine (Figs. 7D, 30B) 26
— Caput highly arched, height greater than 2.3 x height
at thoracic fovea; retroapex of tibia I lacking
megaspine (Fig. 1 ID) Heteromigas
26(25) Metatarsus I straight, cylindrical or slightly
swollen retroapically (Figs. 29, 36) 27
— Metatarsus I swollen for apical half (Figs. 6, 7D) . .
Calathotarsus
27(26) Metatarsus I cylindrical, unmodified (Figs.
37C, 38C) 28
— Metatarsus I pale and swollen retrolaterally (Figs. 29,
SOB, 31D-E)
Paramigas andasibe Raven, new species
28(27) Pedipalpal tarsus with apical spinules (Figs.
37E-G); embolus length less than or equal to bulb
length (Figs. 38E-G) 29
— Apex of pedipalpal tarsus lacking spinules; embolus
length much longer than bulb (Fig. 16A)
Moggridgea (Australia)
29(28) Tarsi III and IV cylindrical, with weak ventral
scopulae (Fig. 36); tibia I with 3—4 retroventral spines
(Fig. 37C) Paramigas macrops, new species
— Tarsi III and IV swollen, curved, sausage shaped,
with dense ventral scopulae (Fig. 38D); tibia I with
only retroapical megaspine (Fig. 38C)
Paramigas manakambus, new species
MIGID GENERA NOT FROM
MADAGASCAR
Calathotarsus Simon, 1903
(Figs. 4, 5A-E, 6. 7A-F. 62A-B, 63A, 65, 66)
Calathotarsus (type species C. coronatus Simon,
1903) comprises three species from Argentina and
Chile. The genus was described by Simon (1903), the
Argentinean fauna was reviewed by Schiapelli and
Gerschman (1973), the Chilean fauna was reviewed by
Legendre and Calderon Gonzalez (1984), Goloboff and
Platnick (1987) and Goloboff (1994), and new species
were described by Schiapelli and Gerschman (1975)
and Goloboff (1991). Possible synapomorphies for the
genus are the modified female pedipalpal tarsus bear-
ing a ventral expansion and dorsal group of cuspules
(Figs. 5C) and bent male metatarsus 1 (Figs. 6, 7D) that
is swollen for the apical half The very wide ocular area
(Fig. 4), mentioned as a synapomorphy by Goloboff
(1991) may relate "Migas' vellardi to Calathotarsus.
Calathotarsus are medium sized (12—20 rrmi) with an
arched caput (Fig. 5A) in the female, wide ocular area
(except in C. pihuychen Goloboff) and with rows of
setae on the caput. The thoracic fovea is simple or may
have a weak posterior extension (Fig. 4). The chelicer-
al fang fiirrow has denticles between the tooth rows
(Figs. 5D, 7E) and intercheliceral tumescence in the
male, the cuspules of the pedipalpal coxae are clustered
near the base (Fig. 5B), the sternal sigilla are flat, and
there are no thorns on the leg coxae. Female tarsi I and
II have spines, patellae III and IV and tibiae and
metatarsi III are densely spinulose, tibiae III are cylin-
drical, metatarsi III and IV have broad retrolateral
preening combs of long and short setae (Figs. 62A— B),
and the ITC have one to a few teeth (Fig. 5E). The
spermathecae are short, cylindrical and without easily
distinguished head and stalk (Fig. 63A). Males have a
megaspine on metatarsus I, the pedipalpal tibia lacks
spinules, and there are scopulae beneath at least tibiae
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
15
III and IV. Biology has been reported by Schiapelli and
Gerschman(1973, 1975) and Goloboff( 1991). All Ca-
lathotarsiis appear to be burrowers rather than nest
builders. Golobofif reports that Calathotarsus corona-
tus and C. pihuychen make burrows closed with a thick
and rigid trapdoor (Goloboff 1991 figs. 9-1 1 ) that has
several small pits on the iimer surface that presumably
mark where the spider inserts its fangs to hold the door
shut.
Heteromigas Hogg, 1902
(Figs. 8, 9A-D, 10, llA-F, 63B, 65, 66)
Heteromigas (type species Heteromigas dovei
Hogg, 1902) comprises two species from eastern Aus-
tralia. Hogg described H. dovei from Tasmania (Hogg
1902), Raven (1984) described H. terraereginae from
Queensland and Raven and Churchill (1989) added
notes on the morphology and biology of//, dovei. Pos-
sible synapomorphies for the genus are long pedipalpal
tarsi of males (Fig. 1 0) and loss of the male retrolater-
al tibia I megaspine (Fig. 1 ID). Heteromigas are small
to medium sized (8—18 mm) with an arched caput (Fig.
9A) in the female, moderately broad ocular area (ocu-
lar area width 0.38-0.44 x carapace width) (Fig. 8) and
with rows of setae on the caput. The thoracic fovea has
a posterior extension in most specimens. The chelicer-
al fang furrow has denticles between the tooth rows
and the male lacks intercheliceral tumescence (Figs.
9C, 1 IE). The cuspules of the pedipalpal coxae extend
to the apex, the sternal sigilla are flat, and thorns are
absent from the leg coxae (Fig. 9B). Female tarsi I and
II have spines, patellae III and IV and tibiae and
metatarsi III are densely spinulose, tibiae III are cylin-
drical and preening combs are absent. Tarsal claws of
the female have simple or divided teeth (Fig. 9D) and
those of the male have multiple teeth (Fig. IIF). The
spermathecae of H. dovei are long and have the head
narrower than the stalk (Fig. 63B) whereas those of//.
terraereginae are short and cylindrical. Male metatar-
sus I lacks a megaspine and the pedipalpal tibia lacks
spinules, and there are scopulae beneath at least
metatarsi III and FV. Biology has been reported by
Raven (1984) and Raven and Churchill (1989). Het-
eromigas terraereginae lives in the ground in burrows
that are fitted with circular trapdoors impregnated with
soil. Heteromigas dovei also makes burrows in the soil
that are up to 10 cm deep with a soil-impregnated lid.
Mallecomigas Goloboff «& Platnick, 1987
(Figs. 12A-F, 63C-D, 65, 66)
This monotypic genus (type species Mallecomigas
schlingeri Goloboff & Platnick, 1987), known only
from Chile, was proposed by Goloboff and Platnick
(1987) for a unique female specimen. A possible
synapomorphy for Mallecomigas is spermathecae that
are long and have the head narrower than the stalk
(Figs. 63 C— D). Mallecomigas are large (22 mm) with
an arched caput, narrow ocular area (width 0.43 x
carapace width) (Fig. 12 A) and with rows of setae on
the caput. The thoracic fovea is simple with a faint,
shallow posterior extension. The cheliceral fang fur-
row lacks denticles between the tooth rows (Fig. 12E),
the cuspules of the pedipalpal coxae are clustered near
the base, the sternal sigilla are flat, and thorns are ab-
sent from the leg coxae (Fig. 12B). Female tarsi I and
II have spines (Fig. 12C), patellae III and IV and tibi-
ae and metatarsi III are densely spinulose, tibiae III are
cylindrical (Fig. 12D) and metatarsi III and IV have
preening combs of setae that are uniformly long and
separated at the base by distances greater than their di-
ameter (Goloboff & Platnick 1987:9). The comb on
metatarsus IV is inconspicuous, being formed of very
thin setae. The spermathecae are characteristic: they
are long and have the head narrower than the stalk
(Figs. 63C— D). Tarsal claws have few teeth (Fig. 12F).
The male is unknown and nothing has been published
on the biology. The collector indicated that the speci-
men had been dug from a deep hole in the ground with-
out a door (E. Schlinger, pers. commun.).
Migas L. Koch, 1873
(Figs. 13A-E, 14A-F, 19C, 59A-C, 60B, 63E-F, 65. 66)
With 34 described species from Australia, New
Caledonia, and New Zealand, Migas (type species
Migas paradoxus L. Koch, 1873 from New Zealand) is
the largest migid genus. It comprises a disparate as-
semblage of species united by the presence of a tooth
near the base of the fang, preening combs in most
species, and lacking the synapomorphy (tibial banding)
of Poecilomigas . We do not know the genus in detail
but our species examined do include the type species
and comprise species from the flill geographic range of
the genus: Migas affinis Berland from New Caledonia,
M. distinctus O. P. Cambridge, M. gatenbyi Wilton, M
giveni Wilton, M. paradoxus L. Koch, and M. taierii
Todd from New Zealand, and M. nitens Hickman and
M. variapalpus Raven from Australia. Most of our
other data come from the literature. Our analysis sug-
gests that the Chilean species Migas vellardi is mis-
placed in Migas and is more closely related to the other
South American genera Calathotarsus and Mallecomi-
gas (see Goloboffia). The composition and affinities of
Migas constitute the greatest remaining problem in
migid taxonomy. The species from Australia, New
16
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Caledonia and New Zealand are similar in being small
to medium sized (6-1 7mm) with a nearly flat caput no
higher than the thoracic region (Fig. 13 A), ocular area
moderately wide (OAW 0.37-0.49 x carapace width)
(Fig. 13B) and with well developed rows of setae on
the caput leading to a pair of enlarged prefoveal setae.
The thoracic fovea is simple. The cheliceral fang flir-
row lacks denticles between the tooth rows (Fig. 13D)
and intercheliceral tumescence in the male may be
present (some CASC and AMNH specimens deter-
mined as M. variapalpis, although these may be a dis-
tinct species [Goloboff 1991:71]) or absent (holotype
of M variapalpis [Raven in Goloboff 1991:71]), the
cuspules of the pedipalpal coxae extend to the apex, the
sternal sigilla are flat, and thorns are absent from the
leg coxae. Female tarsi I and II lack spines (Fig. 13C),
patellae III and IV and tibiae and metatarsi III are
sparsely spinulose and tibia III is cylindrical (Fig.
59A). A dense vestiture of long, curled hairs beneath
legs I and II may be present (e.g., M. gatenbyi) or ab-
sent (e.g., M giveni), leg 1 and II dorsal procumbant
setae are absent (M affinis has retrodorsal patches of
thorns at the bases of metatarsi I and II that resemble
those of the Malagasy Paramigas goodmani but it
lacks procumbant setae on the tibiae) and preening
combs of widely spaced or mixed long and short setae
are present in most species (Fig. 59C), though absent at
least from M. affinis and M. distinctus. Tarsal claws of
the female are simple or have a few teeth (Fig. 13E).
Males typically lack scopulae from tarsi 111 and IV
(Fig. 60B) but have a metatarsus I megaspine and spin-
ules on the pedipalpal tarsus. The spermathecae of at
least some New Zealand species are folded so that they
appear mushroom-like (Fig. 19C): this morphology
may prove to be a synapomorphy uniting at least some
Migas species. Males have spinules at the apex of the
pedipalpal tarsus.
The biology of several species of Migas has been
reported. Species from New Zealand and Australia ap-
pear to make nests or burrows, and some species (e.g.,
M distinctus) are reputed to make both (Wilton 1968:
77). The biology of New Zealand species was summa-
rized by Wilton (1968). He reports that Migas con-
struct unbranched trap-door nests that comprise a "flat-
tened silk tube with a hinged lid with exposed surfaces
camouflaged and strengthened with material gathered
from the surrounding terrain. Nests may be on the
trunks of trees, in moss on banks, under stones, in
crevices ... or in burrows in sand or clay banks"
(Wilton 1968:77, figs. 221, 412). Nests have one or in
rare cases two doors. Todd (1945) studied the biology
of Migas. Raven and Churchill (1989) report that
Migas nitens makes burrows directly in the soil with a
thin door whereas M. plomleyi makes a camouflaged
silken, sac-like nest.
Moggridgea O. P. Cambridge, 1875
(Figs. 15A-F, 16A-B, 17A-C, 32D, 60C,
61A-D, 62C-D, 65, 66)
Moggridgea (type species Moggridgea dyeri O. P.
Cambridge, 1875) comprises 33 species from Africa
and some surrounding islands and from Australia. This
genus was described in 1 875 by Cambridge for Mog-
gridgea dyeri from South Africa, placed in the Migeae
by Simon (1892) and transferred from the Miginae to
Paramiginae by Raven (1985). Moggridgea are record-
ed from mainland Africa by Benoit (1962), Cambridge
(1875), Griswold (1987a), Hewitt (1913a, 1913b,
1913c, 1914, 1915a, 1916, 1919), Lawrence (1928),
Purcell (1903), and Simon (1903), from the surround-
ing islands of Bioko (Griswold & Ubick 1999), the Co-
moros (Griswold 1987a), Principe (Simon 1907), and
Socotra (Griswold 1987a). Griswold (1987a) mono-
graphed the African species and Main (1991) described
two new species from Australia. Our analysis suggests
that Moggridgea should be returned to the Miginae
(sensu Simon 1903) (placement supported by a low
caput and loss of spines from female leg tarsi) and that
the dorsobasal depression on female tibia III, consid-
ered by Raven (1985) to be a synapomorphy of the
Paramiginae, has arisen in parallel in Moggridgea.
Possible synapomorphies for the genus are the lamel-
late setae beneath at least patellae I, II and IV (with ho-
moplasy in Goloboffia vellardi), a dorsobasal depres-
sion on female tibia III (with homoplasy in the Mala-
gasy migids), and a median sclerotized band across the
spermathecae. Thorns on the leg coxae were postulat-
ed to be a synapomorphy uniting Moggridgea and Mi-
cromesomma (Griswold 1 987a: 109); our new analysis
suggests that these have evolved in parallel. Mog-
gridgea are small to large (5— 25nmi) with a low caput
(Fig. 15 A), moderately broad ocular area (OAW
0.42-0.86 X caput width) (Fig. 15B) and with at least a
few setae on the caput. Prefoveal setae may be absent,
small, or enlarged. The thoracic fovea is simple or may
have a weak posterior extension. The cheliceral fang
fijrrow lacks denticles between the tooth rows (Fig.
15E) and intercheliceral tumescence in the male may
be present (M tingle) or absent (most African species).
The cuspules of the pedipalpal coxae extend to the
apex, the sternal sigilla are flat, and thorns on the leg
coxae may be present or absent. Female tarsi 1 and II
lack spines (Fig. 15C), patellae III and IV and tibiae
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
17
and metatarsi III are densely spinulose, tibiae III have
a weak (most species) to moderate (A/, breyeri Hewitt)
dorsobasal depression (Figs. 15 D, 61 A— B) and
metatarsus IV has a preening comb (Figs. 62C— D) of a
few, closely spaced setae, in most species on a com-
mon base (some lack this comb, e.g., M. anactenidia
Griswold). Peculiar, lamellate setae are present beneath
at least patellae I, II and IV, occurring in both sexes of
most species (Figs. 16B, 6 ID). Tarsal claws may have
simple (Fig. 15F) or multidentate (Griswold 1987a fig.
37) teeth. The spermathecae are short to long, cylindri-
cal to sinuate, with the head slightly wider than the
stalk (M ampullata has the head narrower than the
stalk: Griswold 1 987a fig. 80) and have a median trans-
verse sclerotized band (Figs. 17A-C). Males lack tibia
I megaspines (Fig. 163) and spinules on the pedipalpal
tarsus (Fig. I6A), most species have scopulae beneath
metatarsi III and IV and at least most of the African
species have male femur I carinate ventrally (Fig. 60C)
and the apical lobes of the pedipalpal tarsus unequal
(Figs. I6A, 32D). The biology of African species was
summarized by Griswold (1987a) and that of Aus-
tralian species by Main (1991). The majority of
African species for which biology is known make a
bag or sac-like nest, oval to pear shaped, constructed of
silk mixed with bits of the surrounding substrate, pro-
vided with a single wafer or cork trap door, and placed
in a niche or crevice in the substrate (Griswold 1987a,
figs. 2—3). Most nest building species have been taken
from the trunks of trees though some build their nests
on rock faces or stone walls. The Australian M tingle
constructs similar nests (Main 1991, fig. 1). Some
species make a true silk-lined tubular burrow, excavat-
ed in the substrate and usually terrestrial. At least the
African M. mordax, M. peringueyi. M. terrestris and
M. terricola and the Australian M. austmlis are bur-
rowing species.
Poecilomigas Simon, 1903
(Figs. 18, 19A-B, 20A-C, 59D, 60A, 65, 66)
This genus (types species Poecilomigas pul-
chripes Simon 1903) comprises three species from
southern and eastern Africa. The genus was proposed
by Simon (1903), revised by Griswold (1987b), and
Griswold (1998b) provided additional data on behavior
and taxonomy. Possible synapomorphies for the genus
are annulate tibiae (Fig. 18) and, implied by our analy-
sis, the loss of preening combs from metatarsus IV.
Poecilomigas are small to large sized (females
6.&-26.0 mm) with a low, nearly flat caput, moderate-
ly wide ocular area (OAW 0.5—0.65 width caput) and
with rows of setae on the caput and enlarged prefoveal
setae (Fig. 18). The thoracic fovea is simple. The che-
liceral fang fiirrow lacks denticles between the tooth
rows (Fig. 20C) and intercheliceral mmescence occurs
in the male, the cuspules of the pedipalpal coxae ex-
tend to or near to the apex, the sternal sigilla are flat,
and thoms are absent from the leg coxae. Females are
with or without dense, long silky hairs beneath legs I
and II, tarsi I and II lack spines (Fig. 18), patellae III
and IV and tibiae and metatarsi III are weakly spinu-
lose, and preening combs are absent. The spermathecae
are short to long, cylindrical, uniformly sclerotized and
have a well-defined head and stalk (Figs. 19A-B).
Males are with or without a retroapical megaspine on
tibia I, have spinules at the apex of the pedipalpal tar-
sus (Figs. 20A, B), and have scopulae beneath tarsi III
and IV (Fig. 60A). Biology has been reported by Cam-
bridge (1889), Pocock (1895), Hewitt (1915b), Gris-
wold ( 1 987b) and Griswold ( 1 998b). Both P abrahami
and P. basilleupi appear to build nests, usually on the
trunks of trees. Although Cambridge, Pocock, and He-
witt report P. abrahami making nests with one or two
doors, Griswold found only 2-door nests for this
species and it is likely that the 1-door nests were made
by other sympatric migids. The nests of P. abrahami
are vertical, situated in a crevice and woven of silk and
bits of the substrate, and are provided with a wafer type
door at the upper and lower end (Griswold 1987b, figs.
16, 17). Poecilomigas basilleupi makes a similar nest
with a single door at the upper end (Griswold 1998b,
figs. 2-3).
Goloboffia, new genus
(Figs. 14A-F, 63E-F)
Type species. — Migas vellardi Zapfe 1961. male
holotype from La Herradura, Coquimbo Province,
Chile, deposited in Museo Nacional de Historia Natur-
al, Universidad de Chile, Santiago, examined.
Etymology. — ^Named in honor of Pablo GolobofF,
smdent of phylogenetic systematics and collector of
many new and interesting South American mygalo-
morphs. Gender feminine.
Diagnosis. — Distinguished from migids other than
Calathotarsus by having a very wide ocular area
(OAW >0.6 X carapace width) and from migids other
then Moggridgea by having ventral lamellate setae on
the patellae. Goloboffia is distinguished from Calatho-
tarsus by lacking a ventral expansion and dorsal group
of cuspules on the female pedipalpal tarsus and bent
male metatarsus I and from Moggridgea by lacking a
dorsal excavation on female tibia III and having tarsal
spines and spinules on the tibia of the male pedipalpus.
18
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Note. — We transfer Migas vellardi because it does
not belong in Migas. It shares with the other South
American genera Calathotarsus and Mallecomigas the
synapomorphies of pedipalpal coxa cuspule distribu-
tion that is concentrated at the proximal edge near the
labium, slender spinuies densely situated on patella-
tibia III of females, and preening combs that consist of
many setae that extend part way around the ventral
apex of the segment and that occur on both metatarsi
III and IV. We place Migas vellardi in the new genus
Goloboffia because it differs from both Calathotarsus
and Mallecomigas in several features both diagnostic
and synapomorphic for these genera. Synapomorphies
for Goloboffia are the low caput, basal tooth on the
fang, ventral lamellate setae on the patellae, apical
spinuies on the male pedipalpal tibia and loss of the
male metatarsus I megaspine.
Description. — The male was described by Zapfe
(1961: 153) and redescribed by Calderon and Calderon
Gonzalez (1983) and Legendre and Calderon Gonzalez
(1984) and the female was described by GoloboflFand
Platnick (1987). Small (6— 8mm); caput raised above
thoracic region (though not highly arched), the ocular
area very wide (OAW >0.6 x carapace width); with
few setae on the caput and small prefoveal setae (Fig.
I4A); thoracic fovea simple, nearly straight; chelicerae
with anteromedian brush of stout setae (like Calatho-
tarsus), fang with basal tooth and fang furrow with few
(female) or many (male) denticles between tooth rows
(Fig. 14E); male with intercheliceral tumescence; cus-
pules of pedipalpal coxae restricted to base (Fig. 14B);
sternal sigilla flat; female tarsi 1 and II with spines (Fig.
I4C), patellae I, II and IV with ventral lamellate setae
(Fig. 14C), patellae III and IV and tibiae and metatarsi
III densely spinulose (Fig. 14D); metatarsi III and IV
with broad preening combs; female tarsal claws slen-
der with short teeth (Fig. I4F); spermathecae short and
cylindrical (Figs. 63E— F); male lacks tibia I
megaspine; pedipalpal tarsus with apical spinuies;
scopulae beneath male metatarsi III and IV.
Natural History. — Goloboff (1991) reports that
Migas vellardi is terrestrial and makes burrows closed
with a thin and rigid trapdoor lacking beveled edges
(Goloboff 1991. figs. 12-13). The door fits over the
burrow entrance and has two mesal holding marks on
the inner surface that presumably mark where the spi-
der inserts it fangs to hold the door shut.
Composition. — One species.
Distribution. — ^Northern Chile (Fig. 66).
MIGIDAE OF MADAGASCAR
Paramiginae Petrunkevitch 1939
Myrtaleae Simon 1892: 84 (unavailable through homonymy
of type genus)
Paramiginae Petrunkevitch 1939: 154 (nomen novum).
Roewer 1942: 192. Bonnet 1958: 3329. Raven 1985: 144.
Diagnosis. — Migids with reduced caput setation,
with prefoveal setae only or lacking setae altogether
posteriad of ocular area (Figs. 1, 2A, 33, 50), femur III
with a ventral membrane that extends at least 4/5 the
length of the femur (Fig. 42C), and tibia III with a deep
dorsobasal excavation (Figs. 25A, C, 42A— B) and an-
terior diagonal ridge (Fig. 25 A, 42A), and most species
with an angular and strongly tripartite thoracic fovea
(Figs. 2A, 22, 46A).
Genus Micromesomma Pocock 1895
Micromesomma Pocock 1895:190 (type species, by mono-
typy, Micromesomma cowani Pocock 1895). Roewer 1942:
192. Bonnet 1957:2886. Dresco & Canard 1975:783-788.
Raven 1985:145. Griswold 1987a:109-l 17. Platnick 1989:
71. Dippenaar-Schoeman & Jocque 1997:77. Platnick
2001.
Diagnosis. — ^Distinguished ft-om all other migid
genera, except some Moggridgea, by having thorns on
the ventral surfaces of coxae II and III (Figs. 23B,
25D), and from Moggridgea by having tarsal spines in
the female (Fig. 22), a group of dorsal thorns at the
apices of tibiae I and II (Figs. 24A-B), and lacking
lamellate setae beneath the patellae.
Note. — A possible synapomorphy for the genus is
the thorns on leg coxae.
Description. — Small to large, 6.0-22.0 in length.
Sparsely setose. Carapace smooth, length 1.14-1.21 x
width, posteriorly narrowed (Fig. 22); caput with me-
dian ocular seta, prefoveal setae present. Ocular area
0.43-0.52 X width caput; AER straight to slightly re-
curved, ALE equal to or smaller than AME; PER
slightly recurved, PLE equal to or smaller than PME;
ocular quadrangle narrowed anteriorly. Thoracic fovea
tripartite, recurved, width 0.19—0.30 x width carapace.
Sternum length 1.30-1.53 x width, margin sinuate; se-
tose along margin, sparsely setose on surface; sigilla
oval to irregular, shallowly excavated, length 2.25—3.0
X sternum width (Fig. 238). Thorns present at least on
coxae II and IIL Leg formula 4123; tibiae, metatarsi,
and tarsi I and II ventrally with sparse distribution of
short setae; femur III with ventral membrane extending
to base, tibia III with deep dorsobasal excavation and
anterior ridge (Figs. 25A— B). Spermathecae with broad
head and narrow stalk (Figs. 23D-E, 26A-C). Male
unknown.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
Natural History. — Unknown.
Composition. — One species.
Distribution. — Central Madagascar.
Micromesontma cowani Pocock 1895
(Figs. 22, 23A-E, 24A-E, 25A-B, D, 26A-C, 65, 68)
Miavmesomma cowani Pocock, 1895:190-191 (two speci-
mens from Central Madagascar, purchased of Rev. Deans
Cowans, lectotype female, here designated in order to en-
sure the name's proper and consistent application, and
paralectoiype female, BMNH 82.26, in BMNH, exam-
ined). Roewer 1942:192. Bonnet 1957:2886. Dresco & Ca-
nard 1975: 783-788. Raven 1985:145. Griswold
1987a: 109-1 17. Plamick 1989:71. Platnick 2001.
Note. — The types are from "Central Madagascar"
(Pocock 1895: 191). There are no precise locality data
with any specimen, so the distribution within Mada-
gascar remains a mystery.
Diagnosis. Same as for genus.
Description. — Female (lectotype): Total length
19.1. Carapace dark yellow-brown; caput and ocular
area light yellow-brown with dark striae extending
along lateral margins and from PME to thoracic fovea,
black surrounding each AME and extending between
each ALE and posterior eyes; chelicerae dark yellow-
brown, coxae and trochanters yellow-brown, sternum
light yellow-brown except darker at margin, pedipalpal
coxae yellow-brown shading to pale yellow along an-
terior margin; legs and pedipalpi dorsally dark yellow-
brown, light yellow-brown ventrally on patellae and
tibiae; abdomen faded to yellow-white. Carapace 6.0
long, 5.1 wide, height at thoracic fovea 0.29 x carapace
width; smooth. Caput inclined (Fig. 23 A); height 1.33
X that at thoracic fovea, 0.74 x carapace width; medi-
an ocular setae situated behind and between AME, two
pairs of setae positioned anteriad to this; clypeus 0.55
X length OAL, margin procurved. Thoracic fovea re-
curved and tripartite, width 0.19 x that of carapace,
1.33 X wider than long, with pair of prefoveal setae.
Ocular area width 0.52 x caput, 2.22 x wider than long;
AER 2.0 wide, 1.14 x width PER. Ratio of eyes: AME:
ALE: PME: PLE: 1.0: 2.13: 1.4: 1.2, diameter AME
0.27; AME separated by 0.45 x their diameter, PME by
5 X their diameter. Ocular quadrangle 1.43 x wider
than long, posterior width 1.64 x anterior. Sternum 4.1
long, 3.1 wide, widest behind coxa II and narrowed an-
teriorly, setose laterally and sparsely setose on surface;
single thorn situated posteriad to labial suture; coxae I
with 3-5, II with 17-20, and III with 35-37 thorns; sig-
illa2.2 long, 1.0 wide, shallow, irregularly shaped, ad-
jacent to coxa II, width 0.31 x width sternum, distance
between 0.5 x distance from margin. Labial sigilla dis-
tinct, swollen; labium with 27 and pedipalpal coxae
with 34—36 cuspules; labium 0.85 long, 1.15 wide,
pedipalpal coxae 1.95 long, 1.1 wide, apex produced to
a sharp point. Chelicerae 2. 1 long, promargin of fang
fiirrow with 3 teeth, retromargin with 1 large basal
tooth, pro-and retromargin interspersed with 7 denti-
cles, with pale swelling at base of tooth rows. Femur I
0.84. tibia I 0.49, femur IV 0.83, and tibia IV 0.52 x
carapace width. Spination: pedipalpus: tibia v 1-0-0,
tarsus p 1-1-1-1, r 0-1-1-0; leg I: tibia p 2-4-2-4, r 3-4-
3-3-3, d 14 thorns at apex of tibia, metatarsus p 5-4-la,
r 4-4-3- la, tarsus p 1-1-1, r 1-1-1; leg II: tibia p 2-1-3-
1-3, r 1-2-2-1-2, d 27 thorns at apex of tibia, metatar-
sus p 3-3-2-la, r 0-2-2-2, tarsus p 1-1-1-1, r 1-0-0; leg
III: patella with approximately 60, tibia with approxi-
mately 110, metatarsus with approximately 130, and
tarsus with approximately 70 spinules; leg IV: patella
with approximately 70 spinules and tibia with approx-
imately 80 stout setae, metatarsus p 0-0-1-1 a. Femur II
with proximal ventral rows of 10-15 stout setae. STC
teeth (pro-retro): I, II, III (2-2), IV (1-1); ITC simple,
pedipalp claw with 1 tooth (Fig. 23F). Leg measure-
ments (Femur + Patella + Tibia + Metatarsus + Tarsus
= [Total]): I: 4.3 + 2.3 + 2.5 + 2.1 + 1.1 = [12.25]; II:
3.75 + 2.25+2.2+ 1.8+ 1.5 = [11.25]; III; 3.05 + 1.9
+ 2.15 + 1.9 + 1.5 = [10.5]; IV: 4.25 + 2.45 + 2.7 + 2.15
+ 2.15 = [13.7]; pedipalpus: 2.9+ 1.25 + 1.4 + (absent)
+ 1.25 = [6.25]. Abdomen 1 1.0 long, 7.0 wide, sparse-
ly covered with short setae. Spermathecae not removed
from type specimens, but glandular covering visible
through cuticle suggests they are like those described
from other specimens below (Figs. 23D-E, 26A— C).
Variation (N = 4). — Total length 16.5-19.4; height
at fovea 0.2-0.29 x carapace width. Caput width
0.73-0.78 X carapace width, height 1.33-2.08 x height
at thoracic fovea; width ocular area 0.43—0.52 x caput
width, PLE 0.4-0.75 x PME; clypeus length 0.33-0.55
X OAL; thoracic fovea width 1.33-2.12 x length. Che-
liceral fang furrow interspersed with 4—8 denticles
(Figs. 23C, 24E). Sternum with or without cuspules,
sigilla oval to irregularly shaped; labium with 19—39,
pedipalpal coxae with 40—45 cuspules; coxa I with 0-6,
coxa II with 24—37, and coxa III with 35^5 thorns.
Tibia I (Fig. 24A) with 11-19, metatarsus I with 7-14
retroventral, tibia II with 6—15 proventral spines. STC
with 1—2 teeth. Spermathecae (N=6 specimens) length
0.55-0.84 X distance between them and 1.12-1.35 x
head diameter, diameter head 3.12-4.0 x diameter
stalk, head length 2.14-^.0 x length stalk (Figs.
23D-E, 26A-C).
Material Examined. — Madagascar: (no other lo-
cality) 1954 A. Verdier (3 females, MRAC 147.158);
Le Barbier No. 29-1920, "Page det. Revu par Dresco
20
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
en 1974" (19 females, MNHN AR4136. 1 female,
CASC); Central Madagascar, Rev. Deans Cowans (lec-
totype and paralectotype females, BMNH).
Genus Paramigas Pocock 1895
Paramigas Pocock 1895:188 (type species, by monotypy,
Paramigas subrufus Pocock 1895 [=Myrtale perroti Simon
1895]). Simon 1892:32. Roewer 1942:193. Dresco & Ca-
nard 1975:783-788. Raven 1985:144. Platnick 1989:73.
Dippenaar-Schoeman & Jocque 1997:77. Platnick 2001.
Legendrella Dresco & Canard 1975:786 (type species, by
monotypy, Legendrella pauliani Dresco & Canard, 1975).
Brignoli 1983:119. First synonymized by Raven 1985:144.
Diagnosis. — Distinguished from all other migid
genera by the presence of retrodorsal procumbant setae
or thorns at the apices of tibiae and bases of metatarsi
I and II (Figs. 27A— C, 46B), from Moggridgea also by
having spines on female tarsi I and II (Figs. 1, 28B),
and from all other genera except some Moggridgea and
the Malagasy Micromesomma and Thyropoeus by hav-
ing tibia III with a deep dorsal basal concavity (Figs.
36, 42A— B). Males have spinules at the apex of the
pedipalpal tarsus (Figs. 30E, 32C, 37F), a retroapical
megaspine on tibia I (Figs. 30B, 37C), and a tripartite
thoracic fovea (Figs. 29, 36, 38A).
Note. — Possible synapomorphies for the genus are
procumbant and stout dorsal setae at the base of
metatarsi I and II and apical spinules on the male pedi-
palp tarsus.
Description. — Small to large sized mygalo-
morphs, 6-22 mm in length. Carapace smooth in fe-
male, rugose in male, length 1—1 .3 x width in female,
0.98—1.02 X width in male, posteriorly narrowed in fe-
male, rounded in male; caput with median ocular seta,
prefoveal setae present (Fig. 1) or absent (Fig. 48). Oc-
ular area 0.41-0.55 x width caput; AER straight to
slightly recurved, ALE equal to or smaller than AME;
PER straight to slightly recurved, PLE equal to or
smaller than PME; ocular quadrangle narrowed anteri-
orly. Thoracic fovea recurred, tripartite, 0. 1 1-0.25 x
width carapace. Fang furrow of females with pale
swelling at base of tooth rows (Fig. 43B). Sternum
length 1.10—1.53 x width, margin sinuate; setose along
margin, sparsely setose on surface; sigilla oval to ir-
regularly shaped, shallow, length 0.21-0.42 x sternum
width. Labial suture distinct; cuspules present on labi-
um and pedipalpal coxae of female, absent from male
(Fig. 30A); thorns absent from coxae. Leg formula
4123 (female) or 1423 (male); coxae, legs and pedi-
palpi setose; legs I and II with strong series of spines
on pro-and retrolateral margins (Figs. 1, 27A);
retrodorsal procumbant setae or thorns at the apices of
tibiae and bases of metatarsi I and II (Figs. 27A-C);
tibiae, metatarsi, and tarsi I and II ventrally with sparse
(Figs. 44C, 48) or dense (Figs. 27A, 33, 47C) distribu-
tion of long filiform setae; legs III and IV with dense
patterns of spinules and stout setae pro-and retrodor-
sally gradually changing to slender setae distally in fe-
male, spination weak in male; femur III with ventral
membrane extending to base (Fig. 42C), tibia III with
deep dorsobasal excavation and anterior ridge; scopu-
lae absent from female, present in male. Male with
retrolateral tibia I megaspine, spinules at apex of pedi-
palpal tarsus, bulb uniform, embolus simple and slen-
der, embolus short, 0.25-0.55 x bulb length. Sper-
mathecae (Figs. 35A-E, 40A-D, 45A-C) paired, sim-
ple, unbranched, straight to distally curved, length
0.67—1.2 X distance between bases, with pores
throughout, unsclerotized, with distinct head and stalk,
head 1 .8-7.5 x stalk width.
Namral History. — ^The biology of only a few
species is known. All Paramigas adults for which col-
lection data are available have been arboreal but some
adult females have been taken from pitfall traps and
some juveniles have also been sifted from leaf litter.
Pocock (1895:190, figs, la, lb) described two trap
door nests received with the specimens of Paramigas
subrufus. These are oval with a single wafer type door,
woven of silk, and covered on three sides and the door
with moss, lichen and pieces of bark (Figs. 21A— B).
Several nests off! oracle were collected from depres-
sions on tree trunks and vines at Ranomafana. These
nests are oval with length 2 x the spider's total length,
with a single wafer type door at the upper end. The ex-
posed surfaces are covered with moss, lichen and
pieces of bark, effecting a superb camouflage. All nests
are made of fragments of bark and lichen woven to-
gether with silk (Figs. 21C-D; Pocock 1895, figs, la,
lb).
Composition. — Eleven species.
Distribution. — Probably all of Madagascar (Fig.
68).
Paramigas alluaudi (Simon)
(Figs. 28A-E, 45C, 65, 68)
Myrtale alluaudi Simon 1903: 133 (type female, Madagas-
car. Fort Dauphin. MNHN. examined).
Paramigas alluaudi, Roewer 1942: 193. Bonnet 1958: 3329.
Plamick200I.
Diagnosis. — Females are distinguished from other
Paramigas that lack a dense vestiture of long silky
hairs beneath legs I and II by the dense lateral network
of striae on the caput (Fig. 28A), longitudinal banding
pattern on legs (Fig. 28B), and the thinly distributed
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
21
cuspules on the pedipalpal coxae (<20) and labium
(<8); male unknown.
Description. — Female (holotype): Total length
12.7. Carapace, chelicerae, and legs light brown, ven-
ter, coxae, and trochanters yellow-brown, lateral mar-
gins of caput yellow-brown with light brown band ex-
tending from PME to thoracic fovea; dense lateral net-
work of light brown striae on caput and extending from
behind ocular area to thoracic fovea, also radiating
from thoracic fovea to carapace margin (Fig. 28A);
carapace with dark rebordered margin; ocular area
light brown, black surrounding AME, behind ALE and
between PLE and PME; clypeus dark; femora, patellae
and tibiae with narrow dorsal and broad pro-and retro-
lateral maculate bands, these united at base of patellae
and bases and apices of femora and tibia, area between
bands may have reticulate markings, tarsi yellow-
white, unmarked; abdomen dark purple-brown includ-
ing basal segment of spinnerets, distally segments yel-
low-brown.
Carapace 2.9 long, 2.55 wide, height at thoracic
fovea 0.39 >< carapace width; smooth. Caput inchned,
height 1.4 x that at thoracic fovea, width 0.76 x cara-
pace width; median ocular seta present with a pair of
setae anterior to this, prefoveal setae minute, visible at
greater than 80x magnification; clypeus height 0.4
times length OAL, margin straight. Thoracic fovea re-
curved, tripartite with weak posterior depression, width
0.23 X that of carapace, 6.0 x wider than long.
Ocular area width 0.41 x caput, 2.13 x wider than
long; AER 0.8 wide, 1.0 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 0.53: 1.0: 0.60: 0.80, diame-
ter AME 0.1; AME separated by 1.25 x their diameter,
PME by 3.25 x their diameter. Ocular quadrangle 0.40
X wider than long, posterior width 1.53 x anterior.
Sternum 2.35 long, 1.67 wide, widest behind coxa
II and narrowed anteriorly, sparsely setose; sigilla ir-
regularly oval, adjacent to coxae II and III, width 0.25
X width sternum, distance between 0.71 x distance
from margin. Labium with 6 and pedipalpal coxae with
16—18 cuspules; labium 0.5 long, 0.6 wide, pedipalpal
coxae 0.72 long, 0.55 wide, apex produced to a blunt
point. Chelicerae 0.45 long, fangs broad, flaring pro-
laterally before apex, promargin of fang furrow with 3
teeth, retromargin with 4 teeth (Fig. 28D).
Femur I (Fig. 28B) 0.80, tibia I 0.44, femur IV
0.76, and tibia IV 0.41 x width carapace. Spination:
pedipalpus: tibia pi -0-0, tarsus p 1-1-1, r 1-0-0; leg I:
tibia p 1-1-1-1-1, r 1-1-1-0, metatarsus p 1-3-2, r 1-3-
2; tarsus p 1-1, r 1-1; leg II: tibia p 1-0-2, r 0-1-1-1,
metatarsus p 3-1-1, r 1-0-2, tarsus p 0-2, r 1-0; leg III:
patella with approximately 16, tibia with approximate-
ly 20 spinules, metatarsus: vO-O-la, tarsus with 4—5
proapical spinules; leg IV: patella with approximately
50 slender spinules, metatarsus v 0-0- la. Femur II with
proximal ventral row of 5— 6 stout setae, retrodorsum of
tibiae I and II with 7—8 apical, metatarsi I and II with
3—6 basal stout procumbant setae. STC teeth (pro-
retro): I, II (2-2), III, IV (1-1); ITC simple, pedipalp
claw with 1 tooth (Fig. 28E). Leg measurements
(Femur + Patella + Tibia + Metatarsus + Tarsus =
[Total]): I: 2.05 + 1.17 + 1,12 + 0.87 + 0.70 = [5.91];
II: 1.82 + 1.05 + 0.87 + 0.75 + 0.72 = [5.21]; III: 1.5 +
0.9 + 0.8 + 0.77 + 0.75 = [4.72]; IV: 1 .95 + 1 . 1 2 + 1 .05
+ 0.87 + 0.87 = [5.86]; pedipalpus: 1.25 + 0.67 + 0.72
+ (absent) + 0.47 = [3.11].
Abdomen 3.0 long, 2.25 wide, sparsely covered
with short setae. Spermathecae with broad head and
short, narrow stalk, length spermathecae equal to dis-
tance between and to their head diameter, diameter
head 3.7 x diameter stalk, head length 3.6 x length
stalk (Figs. 28C, 45C).
Material Examined. — Madagascar: Toliara: Fort
Dauphin, 8 November 1901, Ch. Alluaud (Myrtale al-
luaudi female, type, MNHN 26248, AR4123).
Distribution. — Known only from the type locality
at the southern tip of Madagascar (Fig. 68).
Paramigas andasibe Raven, new species
(Figs. 29, 30A-G, 31A-H, 32A-C, 65, 68)
Types. — Holotype male from An' Ala, 9 km E An-
dasibe, 840 m, 15 February 1995, primary rainforest, F.
Glaw, deposited in ZFMK.
Note. — Robert Raven provided us with the speci-
men and descriptive notes. We attribute authorship of
the new species to him.
Etymology. — ^The specific name is a noun in ap-
position from the type locality.
Diagnosis. — Distinguished from other Paramigas
males by having pallid, swollen, weakly sclerotized
areas on the retrolateral surface of metatarsus I (Figs.
30B, 31D-E) and AME smaller than ALE (Fig. 29).
Description. — Male (holotype): Total length 17.1.
Carapace dark red-brown with a dark brown band mar-
ginal surrounding; caput with light red-brown areas
along lateral margin; ocular area dark red-brown, black
behind ALE and between PME and PLE; chelicerae
dark red-brown; sternum, coxae, and trochanters or-
ange-brown except stemo-labial junction red-brown;
dorsal surface of legs I-IV red-brown gradually fading
to yellow-brown on tips of tarsi and on ventral sur-
faces, femora with dorsal and retrodorsal longitudinal
dusky bands; prolateral surface of metatarsus I fading
from red-brown to yellow-brown, retrolateral surface
22
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
yellow-white, bulging, tarsi I and II yellow-brown;
pedipalpi yellow-brown; abdomen dark purple-brown;
spinnerets and book lung covers yellow-brown.
Carapace 8.0 long, 7.9 wide, height at thoracic
fovea 0.24 x carapace width; strongly rugose. Caput
low, height 1.15 x that at thoracic fovea, width 0.51 x
carapace width; with minute pair of prefoveal setae;
median ocular seta present; pair of setae situated ante-
riad of this; clypeus height 0.34 x length OAL, trans-
versely striate, margin straight. Thoracic fovea tripar-
tite (Fig. 29), weakly recurved, width 0.15 x that of
carapace, 1.71 x wider than long.
Ocular area width 0.59 x caput, 2.13 x wider than
long; AER 2.45 wide, 1.08 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 1.3: 0.6: 0.5, diameter
AME 0.5; AME separated by 0.2 x their diameter,
PME by 3.33 x their diameter. Ocular quadrangle 1.39
X wider than long, posterior width 1.6 x anterior.
Sternum 6.4 long, 4.2 wide, widest behind coxa 11
and narrowed anteriorly, sparsely setose laterally and
on surface; sigilla irregularly shaped, adjacent to coxae
II, 0.33 X width sternum, distance between 0.1 x dis-
tance from margin. Labium and pedipalpal coxae lack-
ing cuspules; labium 1.5 long, 1.4 wide, pedipalpal
coxae 3.3 long, 1.6 wide, apex produced to a blunt
point. Chelicerae 1.6 long, promargin of fang flirrow
with 4—5 teeth and retromargin with 1 basal tooth and
4-5 distal denticles (Fig. 31G)
Femur 1 1.15, tibia 1 0.83, femur IV 1.08, and tibia
IV 0.89 X width carapace. Scopulae beneath distal Vi of
tarsus I and beneath entire tarsi II— IV (Figs. 3 IE, F).
Spination: leg I: patella p 0-2-2, v 0-0-4, tibia p 1-0-0-
0, r 2-3-1- la, metatarsus p 2-2-0-1, r 1-0-1-1, tarsus p
1-1-1, r 1-1-1; leg 11: patella r 0-0-1, tibia p 0-0-1-1, r
2-3-0-2, tibia p 2-2-2- 1 , r 2-0-0-2, tarsus p 1 - 1 - 1 - 1 , r 1 -
1-0-0; leg III: patellae with approximately 41 prodorsal
and 18 retrodorsal spinules, tibia with approximately
50 apical spinules, metatarsus v 0-0-0-4; leg IV: patel-
la with approximately 150 prodorsal spinules, metatar-
sus v 0-0-2-3. STC teeth (pro, retro): 1, 11 (1-2), III, IV
(l-I) (Fig. 31H). Leg measurements (Femur + Patella
+ Tibia + Metatarsus + Tarsus = [Total]): 1: 9. 1 + 4.5 +
6.6 + 6.1 + 2.0 = [28.3]; II: 8.0 + 4.0 + 5.6 + 5.3 + 2.2
= [25.1]; III: 6.1 + 3.2 + 4.2 + 4.1 + 2.7 = [20.3]; IV:
8.6 + 3.9 + 7.1 + 5.8 + 3.5 = [28.9]; pedipalpus: 4.9 +
2.2 + 3.7 + (absent) + 1.5 = [12.3].
Pedipalpus (Figs. 31A— C) with femur 0.62, tibia
0.46 X carapace width; femur 3.26, tibia 2.46 x length
tarsus; tibia widest basally, height 0.48 x length; tarsus
with approximately 10 slender apical spinules; bulb
width 1.06 X tarsus length; embolus length 0.34 x bulb
width. Abdomen 7.5 long, 5.3 wide, sparsely covered
with short setae.
Variation (N=4).— Total length 8.15-17.1. The
size difference between the largest and smallest speci-
mens is dramatic but specimens of intermediate size
exhibit intermediate development of most distinctive
characters. Prosoma yellow-brown to dark red-brown,
legs orange-brown to dark red-brown. Caput 0.52-0.6
X carapace width, height 1.28-1.6 x that at fovea; tho-
racic fovea width 0.28-0.29 x caput width, straight to
slightly recurved; fang furrow with or without denti-
cles (Figs. 30C, 31G). Spines of leg I small and incon-
spicuous in smallest individual (Fig. 30B) to stout in
largest (Figs. 31D0150E). STC II with 2-3 teeth (Fig.
30G). Femur 1 0.48—1.15 x carapace width; pedipalpal
femur 0.55-0.62 x, tibia 0.35-0.47 x carapace width;
embolus length 0.25-0.37 x bulb (Figs. 30D-F,
31A-C, 32A-C).
Natural History. — The Ambohitantely specimen
was collected in pitfall traps at 1450m in disturbed
transitional montane mossy forest; other specimens are
recorded from forest.
Additional Material Examined. — Madagascar:
Toamasina: Manakambahiny near Vavatene, foret,
February 1995, A. Pauly (2 males, MRAC). Antana-
narivo: 1450m in R.S. d' Ambohitantely, 24 km NE
Ankazobe (18°10.1'S, 47°16.6'E), in pitfall traps, 7-12
December 1997, S. Goodman (1 female, FMNH).
Distribution. — East central Madagascar in An-
tananarivo and Toamasina provinces (Fig. 68).
Paramigas goodmani, new species
(Figs. 33, 34A-E, 35D, 65, 68)
Type. — Holotype female collected at 80m eleva-
tion in Foret de Vohibasia, 59 km NE Sakaraha
(22°27.5'N, 44°50.5'E), Toliara Province, Madagascar,
10-16 January 1996 by S. Goodman, deposited in
FMNH.
Etymology. — The specific name honors Steve
Goodman, whose extraordinary collecting efforts in
Madagascar discovered the type and numerous other
new species.
Diagnosis. — Distinguished from all other Parami-
gas that have a dense vestiture of long, silky hairs be-
neath patellae-metatarsi I and II by having dorsal
thorns at the apices of metatarsi I and II (Fig. 33).
Description. — Female (holotype): Total length
20.5. Carapace (Fig. 33) dark red-brown with a dark
brown band surrounding its margin; caput light red-
brown along lateral margin; ocular area dark red-
brown, dusky between AME and black mesad of ALE
and PLE; chelicerae dark red-brown; sternum, coxae,
and trochanters orange-brown (Fig. 34B); legs and
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
23
pedipalpi dark red-brown, except orange-brown tarsi,
with orange-brown setal bases giving a mottled pat-
tern; abdomen dark purple-brown including book lungs
and epigastric furrow; spinnerets dark red-brown (Fig.
34B).
Carapace 8.9 long, 8.2 wide, height at thoracic
fovea 0.31 x carapace width; smooth. Caput inclined
(Fig. 34A), height 1 .30 x that at thoracic fovea, width
0.73 X carapace width; median ocular seta situated an-
teriad of AME and twelve setae positioned anteriad to
this; clypeus 0.30 x length OAL, margin straight. Tho-
racic fovea recurved and tripartite, width 0.21 x that of
carapace, 1.8 x wider than long; with pair of prefoveal
setae (Fig. 33).
Ocular area width 0.50 x caput, 2.30 x wider than
long; AER 3.0 wide, 1.09 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 0.66: 0.66: 0.5, diameter
AME 0.6; AME separated by 0.33 x their diameter,
PME by 2.25 x their diameter Ocular quadrangle 1 .48
X wider than long, posterior width 1 .68 x anterior
Sternum 8.0 long, 5.3 wide, widest behind coxa II
and narrowed anteriorly, setose along lateral margin
and sparsely setose on surface; sigilla indistinct, shal-
low, irregularly oval, adjacent to coxa II, 0.13 x width
sternum, distance between 0.57 x distance from margin
(Fig. 34B). Labium with 41, pedipalpal coxae with
46-48 cuspules; labium 1.3 long, 1.65 wide, pedipalpal
coxae 3.0 long, 1.65 wide, apex produced to a blunt
point. Chelicerae 2.0 long, promargin of fang flirrow
with 4 teeth, retromargin with 6 teeth, pro- and retro-
margin interspersed with 1—3 denticles (Fig. 34D).
Femur I 0.84, tibia I 0.53, femur IV 0.85, and tibia
IV 0.54 X carapace width. Ventral surfaces of patellae,
tibiae, metatarsi, and tarsi of legs I and II densely cov-
ered by long filiform setae (Fig. 33). Spination: pedi-
palpus: tibia p 1 -0-0, tarsus p 1 - 1 - 1 , r 1 -0- 1 ; leg I: tibia
p 1-2-3-3, r 2-3-3-6, d 12 thorns, metatarsus p 2-6-2, r
3-3-3-3, d 26 thorns, tarsus p 1-1-1, r 1-1-0; leg II: tibia
p 2-3-3, r 1-3-4-2-3, d 15 thorns, metatarsus p 3-3-3-2-
la, r 3-3-Ia, d 28 thorns, tarsus p 2-1-1-1, r 1-1-0; leg
III: patella with approximately 69, tibia with approxi-
mately 75, metatarsus with approximately 40, and tar-
sus with approximately 12 spinules; leg IV: patella
with approximately 80 slender spinules. STC teeth
(pro-retro): I (3-2), II (3-3), III, IV (1-1); ITC simple,
pedipalp claw with 1 tooth (Fig. 34E). Leg measure-
ments (Femur + Patella + Tibia + Metatarsus -i- Tarsus
= [Total]); I: 6.9 + 4.0 + 4.4 + 3.0 + 2.0 = [20.3]; II: 6.2
+ 3.8 + 4.0 + 2.9 + 1.9 = [18.8]; III: 5.0 + 3.1 + 3.2 +
2.6 + 2.1 = [16.0]; IV: 7.0 + 4.0 + 4.5 + 3.9 + 2.5 =
[21.9]; pedipalpus: 4.0 + 2.1 + 2.6 + (absent) + 2.2 =
[10.9].
Abdomen 11.6 long, 8.7 wide, sparsely covered
with short setae. Spermathecae with narrow head and
long stalk, length spermathecae 0.78 x distance be-
tween them and 1 .85 x head diameter, diameter head
1.81 X diameter stalk, head length 0.90 x length stalk
(Figs. 34C, 35D).
Material Examined. — Only the type.
Distribution. — Known only from the type locality,
an isolated montane forest in south-western Madagas-
car (Fig. 68).
Paramigas macrops, new species
(Figs. 36, 37A-H, 65, 68)
Type. — Holotype male collected at elev. 440 m in
Reserve Naturelle Integrale d' Andohahela, parcel 1,
12.5 km NW Eminiminy (24°37.6'S, 46°45.9'E), To-
liara Province, Madagascar, 19-28 October 1995 by S.
Goodman, deposited in FMNH.
Etymology. — ^The specific name refers to the large
anterior median eyes.
Diagnosis. — Distinguished from other Paramigas
males except P. manakambus by having AME larger
than ALE (Fig. 36), the thoracic fovea recurved and
metatarsus I cylindrical, and from P. manakambus by
having a retroventral row of 4 spines on tibia I (Fig.
37C) and tarsi III and IV cylindrical (Fig. 36).
Description. — Male (holotype): Total length 6.15.
Prosoma (Figs. 36, 37A) light yellow-brown with dark
yellow-brown areas extending along lateral margins of
caput and ocular area darkening to black at edge of
clypeus; chelicerae dark yellow-brown; ocular area
dark except between PME; sternum, coxae, and
trochanters light yellow-brown (Fig. 37B); dorsal sur-
face of legs I-IV dark yellow-brown fading to light yel-
low-brown on tips of tarsi and on ventral surfaces;
pedipalpi light yellow-brown; abdomen pale purple-
brown; spinnerets and book lung covers light yellow-
brown.
Carapace 2.6 long and wide, height at thoracic
fovea 0.19 X carapace width; weakly rugose. Caput in-
clined (Fig. 37A), height 1.5 x that at thoracic fovea.
width 0.55 X carapace width; ocular area with single
setae situated anteriad of AME and a pair of setae situ-
ated anteriad of this; clypeus height 0.35 x length
OAL, margin recurved. Thoracic fovea tripartite and
recurved, width 0. 17 x that of carapace, 1.8 x wider
than long, prefoveal setae minute, nearly invisible ex-
cept at high (80x) power.
Ocular area width 0.70 x caput, 1 .78 x wider than
long; AER 1.02 wide, 1.0 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 0.63: 0.33: 0.67. diame-
ter AME 0.27; AME separated by 0.55 x their diame-
24
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
ter, PME by 2.6 x their diameter. Ocular quadrangle
1.0 X wider than long, posterior width 1.21 x anterior.
Sternum 1.82 long, 1.4 wide, widest behind coxa
II and narrowed anteriorly, sparsely setose laterally and
on surface; sigilla 0. 10 x width sternum, oval, adjacent
to coxa II, distance between 3.82 x distance from mar-
gin. Labium 0.35 long, 0.47 wide, pedipalpal coxae
0.75 long, 0.45 wide, apex rounded (Fig. 37B). Che-
licerae 0.6 long, promargin of fang furrow with 3 teeth,
retromargin with 1 large basal and 3 distal teeth (Fig.
37D).
Femur I 1.0, tibia I 0.73, femur IV 0.96, and tibia
IV 0.71 X width carapace. With sparse patches of scop-
ular hairs beneath apices of tarsi. Tibia III with weak
dorsobasal excavation before protuberance at mid-seg-
ment (Fig. 36). Tarsi III and IV elongate and nearly
straight dorsally. Spination: leg I: tibia r 0-1-1-1-1,
metatarsus r 0-0-0-1; leg III: patella with approximate-
ly 1 5 and tibia with approximately 9 minute lateroapi-
cal spinules. STC teeth (pro. retro): I. II (3-3), III (3-2),
IV (2-2) (Fig. 37H). Leg measurements (Femur -i-
Patella + Tibia -i- Metatarsus + Tarsus = [Total]): I: 2.6
H- 1.3 H- 1.9 -I- 1.75 + 0.65 = [8.2]; II: 2.5 -H.H- 1.65 -H
1.5 + 0.6 = [7.35]; III: 2.0 -^ 0.9 -H.25 -i- 1.15 + 0.65 =
[5.95]; IV: 2.5 -i- 1.25 + 1.85 + 1.6 + 0.8 = [8.0]; pedi-
palpus: 1.15-1-0.6 + 0.95 + (absent) + 0.45 = [3.15].
Pedipalpus (Figs. 37E-G) with femur 0.44, tibia
0.36 X carapace width; femur 2.5, tibia 2.11 x length
tarsus; tibia broad proximally and narrowed distally.
height 0.57 x length; tarsus with 2—3 apical spinules;
bulb width 0.93 x tarsus length; embolus length 0.52 x
bulb width. Abdomen 2.95 long, 2.2 wide, sparsely
covered with short setae.
Natural Histor>'. — ^The specimen was collected in
a pitfall trap.
Material Examined. — Only the type.
Distribution. — ^Known only from the type locality
in far southern Madagascar (Fig. 68).
Paramigas manakambus. new species
(Figs. 38A-H. 65. 68)
Type. — HolotN'pe male collected at Foret bac
jaune, Manakambahing Am., Madagascar, 1—17 Janu-
ary 1991 by A. Pauly. deposited in MRAC (#174.501).
Etymology. — The specific name is an arbitrarv'
combination of letters.
Diagnosis. — Distinguished from other Paramigas
males except P. macrops by having .A.ME larger than
ALE (Fig. 38A), the thoracic fovea recur\ed and
metatarsus I not swollen retrolaterally (Fig. 38C), and
from P. macrops by having only an apical megaspine
on tibia I, metatarsus I flattened dorsally (Fig. 38C)
and tarsi III and IV cur\'ed and swollen (Fig. 38D).
Description. — Male (holotype): Total length 7.9.
Carapace (Fig. 3 8 A) dark red-brown with a dark band
around margin, dusky on clypeus and along anterior
margins of caput; ocular area black except between
PME; sternum, coxae, and trochanters yellow-brown;
dorsal surfaces of legs I through IV dark red-brown
gradually fading to yellow-brown on ventral surfaces
and on tarsi; pedipalpi dark yellow-brown; abdomen
dark purple-brown except spinnerets and anteriad of
epigastric fiirrow yellow-brown.
Carapace 3.35 long, 3.4 wide, height at thoracic
fovea 0.26 x carapace width, rugose. Caput low, height
1.38 X that at thoracic fovea, width 0.57 x carapace
width; ocular area with single seta situated between
AME and a pair of setae positioned anteriad to this;
clypeus length 0.34 x length OAL, slightly procurved.
Thoracic fovea tripartite, recurved, width 0. 1 6 x that of
carapace, 2.3 x wider than long, with minute pair of
prefoveal setae.
Ocular area width 0.55 x caput, 1.86 x wider than
long; AER 1.05 wide, 1 .07 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 0.6: 0.4: 0.5, diameter
AME 0.25; AME separated by 0.4 x their diameter,
PME by 3.6 x their diameter. Ocular quadrangle 1 .22 x
wider than long, posterior width 1.03 x anterior.
Sternum 1.9 long, 1.75 wide, widest behind coxa
II and narrowed anteriorly; sigilla 0.2 x width sternum,
irregular, adjacent to coxa II, distance between 1.25 x
distance from margin. Labium and pedipalpal coxae
lacking cuspules; labium 0.6 long, 0.65 wide, pedipal-
pal coxae 1.1 long, 0.6 wide, apex produced to a blunt
point. Chelicerae 0.9 long, promargin of fang furrow
with 3 teeth, retromargin with single basal tooth and
6-7 distal denticles (Fig. 38B).
Femur I 0.94, tibia I 0.63, femur IV 0.92, and tibia
IV 0.69 X width carapace. Scopulae restricted to tarsi
I— IV, I very weak. Metatarsus I (Fig. 38C) slightly flat-
tened dorsally, tibia III with weakly defined dorsobasal
excavation and tarsi III and IV elongate, dorsally con-
cave and swollen ventrally (Fig. 38D). Spination: leg I:
tibia r 0-0-0- la. d 5 minute apical spinules, metatarsus
d 17 minute spinules at tibia-metatarsus joint; leg II:
tibia d 8 minute apical spinules. metatarsus d 13 minute
spinules at tibia-metatarsus joint; leg III: patella with
approximately 3 1 spinules, tibia with approximately 14
minute dorsoapical spinules; leg TV: patella with ap-
proximately 50 spinules. STC teeth (pro, retro): I-IV
(l-I). Leg measurements (Femur + Patella -i- Tibia +
Metatarsus + Tarsus = [Total]): I: 3.2 + 1.5 + 2.15 +
2.15 + 0.9 = [9.9]; II; 2.95 +\A5+ 1.85 + 1.9 -i- 1.05
= [9.2]; III: 2.1 + 1.0 + 1.4 -i- 1.5 + 1.2 = [7.2]; IV: 3.15
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
25
+ 1.5 + 2.35 + 2.05 + 1.55 = [10.6]; pedipalpus: 1.4 +
0.75 + 0.95 + (absent) + 0.6 = [3.7].
Pedipalpus (Figs. 38E-G) with femur 0.72, tibia
0.49 X carapace width; femur 2.33, tibia 1.58 x length
tarsus; tibia ventrally concave (Fig. 38G) with approx-
imately 10 elongate setae on ventral surface, height
0.52 X length; approximately 4 spines at apex of tarsus;
bulb width 1.16 x tarsus length; embolus length 0.53 x
bulb width. Abdomen 3.65 long, 2.7 wide, sparsely
covered with short setae.
Material Examined. — Only the type.
Distribution. — ^Known only from the type locality
in eastern Madagascar (Fig. 68).
Paramigas milloti, new species
(Figs. 1, 35E, 39A-E, 65, 68)
Type. — Holotype female collected at 600 m eleva-
tion at "1 Av. Antongoniviksika" in November 1947 by
J. Millot, deposited in MNHN.
Etymology. — The specific name honors Jacques
Millot, collector of the type and many other interesting
Malagasy arthropods.
Note. — ^The hand-copied label may have a tran-
scription error. The type locality is probably Antongo-
nivitsika, which is a mountain north of the col d' Am-
batondradama. This latter mountain is 35-40 km NE of
Maroantsetra on the path between Maroantsetra and
Antalaha (Viette 1991).
Diagnosis. — Distinguished from other Paramigas
that lack a dense vestiture of long silky hairs beneath
legs I and II by having these legs densely spinose, tibia
I with more than 40 ventrolateral spines (Fig. 1), pre-
foveal setae large, and the spermathecae long and nar-
row, length greater than 1.63 x maximum diameter,
head diameter less than 1.81 x diameter stalk (Figs.
35E, 39C).
Description. — Female (holotype): Total length
20.1. Carapace and ocular area orange-red (Figs. 1,
39A) with faint longitudinal striations extending from
PME to thoracic fovea and radiating from thoracic
fovea to carapace margin, black surrounding AME and
mesad of ALE and PLE; sternum, coxae, and
trochanters light yellow-brown (Fig. 39B); legs and
pedipalpi orange-brown gradually fading to light yel-
low-brown on tarsi, with dark lateral longitudinal mac-
ulations; abdomen dark purple-brown; spinnerets and
book lung covers yellow-white.
Carapace 8.5 long, 8.2 wide, height at thoracic
fovea 0.26 x carapace width; smooth. Caput width 5.5
X carapace width, inclined (Fig. 39A), height 1 .4 x that
at thoracic fovea, with weak lateral dimples; ocular
area with single seta situated between AME and a pair
of setae situated anteriad to this; clypeus length 0.4 x
length OAL, margin straight, with 5—6 setae along
margin in center. Thoracic fovea recurved and tripar-
tite, width 0.21 X that of carapace, 1.8 x wider than
long, with pair of long prefoveal setae (Fig. 1).
Ocular area width 0.5 x caput, 2.2 x wider than
long; AER 2.75 wide, 1.12 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 1.45: 1.09: 0.72, diame-
ter AME 0.275; AME separated by 1.27 x their diam-
eter, PME by 4. 1 6 x their diameter. Ocular quadrangle
1.48 X wider than long, posterior width 2.05 x anterior.
Sternum 6.3 long, 5.2 wide, widest behind coxa II
and narrowed anteriorly (Fig. 39B), setose along mar-
gin and on surface; sigilla shallow, irregularly oval, ad-
jacent to coxa II, width 0.28 x width sternum, distance
between 0.086 x distance from margin. Labium with
24 and pedipalpal coxae with 25-42 cuspules; labium
1.75 long, 1.7 wide, pedipalpal coxae 3.5 long, 1.8
wide, apex produced to a sharp point. Chelicerae 2.0
long, promargin of fang furrow with 5 teeth, retromar-
gin with large basal tooth, distad of this 5 teeth and 4
denticles (Fig. 39D).
Femur I 0.80, tibia I 0.51, femur IV 0.82, and tibia
IV 0.54 X width carapace. Spination: pedipalpus: tibia
p 1-0-0, tarsus pl-1-1, rl-1-1; leg I: tibia promargin
with 24, retromargin with 28 spines, metatarsus pro-
margin with 20, retromargin with 23 spines, tarsus p 1-
1-1-1, r 3-2; leg II: tibia promargin with 16, retromar-
gin with 19 spines, metatarsus promargin with 15,
retromargin with 9 spines, tarsus p 3- 1 - 1 - 1 , r 1 - 1 - 1 ; leg
III: patella with prodorsal and dorsal bands totaling 70
spinules, tibia with prodorsal and dorsal bands totaling
55 spinules, v 0-0- la, metatarsus with prodorsal and
dorsal bands totaling 60 spinules, v 0-0-2a, tarsus with
more than 20 dorsal and more than 15 proapical spin-
ules; leg IV: patella with prodorsal and dorsal bands of
more than 120 spinules, tibia with prodorsal and dorsal
bands of 55 spinules, metatarsus v 0-1 -2a. Femur II
with rows of 20-25 stout proximal ventral setae,
retrodorsum of legs I and II with sparse patches of
stout, procumbant setae: 20-25 apical on tibiae and
20-25 basal on metatarsi. STC teeth (pro-retro): I (2-
2), II (2-1), III, IV (1-1); ITC simple, pedipalp claw
with 1 tooth (Fig. 39E). Leg measurements (Femur +
Patella + Tibia + Metatarsus -i- Tarsus = [Total]): 1: 6.6
+ 4.1 + 4.2 + 3.2 + 1.8 = [19.9]; II: 6.0 + 3.9 + 3.7 H-
3.0 + 1.9 = [18.5]; III: 4.6 + 2.9 + 3.0 -i- 2.7 -i- 2.2 =
[15.4]; IV: 6.8 + 4.0 + 4.5 + 3.6 + 2.7 = [21.6]; pedi-
palpus: 3.6 H- 1.9 -I- 2.2 -I- (absent) -i- 2.0 = [9.7].
Abdomen 9.6 long, 7.6 wide, sparsely covered
with short setae. Spermathecae with broad head and
narrow stalk, length spermathecae 0.77 x distance be-
26
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
tween them and L63 x head diameter, diameter head
2.44 X diameter stalk, head length 2.30 x length stalk
(Figs. 35E, 39C).
Material Examined. — Only the type.
Distribution. — ^Known only from the type locality
in northern Madagascar (Fig. 68).
Paramigas oracle, new species
(Figs. 3C-D, 21C-D, 25C, 27A-D, 40A-B, 41A-G, 42A-D,
43A-D, 65, 68)
Type. — Holotype female collected in primary for-
est at ca. 1200 m at Vatoharanana (21°16.7'S,
47°26.rE), Pare Nationale Ranomafana, Fianarantsoa
Province, Madagascar, on 29 April 1998 by C. E. Gris-
wold, deposited in CASC. Paratype female, also from
R N. R. at Talatakely (21°14.9'S, 47°25.6'E) ca. 900 m
elev., collected on 27 April 1998 by D. Ubick, deposit-
ed in CASC.
Etymology. — ^The specific name thanks the Oracle
Foundation, whose support made possible the 1998
California Academy of Sciences' expeditions to Mada-
gascar.
Diagnosis. — Distinguished from all other Parami-
gas that have a dense vestiture of long, silky hairs be-
neath patellae-metatarsi 1 and II (Fig. 41 B) by (contra
P. goodmani) having slender setae ra*her than thorns at
the apices of metatarsi 1 and II (Figs. 27A-C, 4 IB) and
(contra P. perroti) by having the spermathecae short
with large heads, length less than 1.67 x head diameter
and head diameter greater than 2.14 x diameter stalk
(Figs. 40A-B, 41C-D).
Description. — Female (holotype): Total length
11.0. Carapace dark yellow-brown with yellow-brown
striae extending along lateral margins of caput and a
dark band rebordering carapace (Fig. 41 A); ocular area
and clypeus dark, black surrounding AME and extend-
ing between these and ALE, PLE and PME; sternum,
coxae, and trochanters light yellow-brown gradually
darkening to orange-brown on pedipalpal coxae and
labium; legs and pedipalpi dark yellow-brown fading
to orange-brown on tarsi; patella IV yellow-white dor-
sally; abdomen dark purple-brown, venter and spin-
nerets paler.
Carapace 4.5 long, 4.1 wide, height at thoracic
fovea 0.29 x carapace width; smooth. Caput inclined,
height 1.6 x that at thoracic fovea, width 0.70 x cara-
pace width; anteromedian ocular seta present with a
pair of setae situated anteriad to this; clypeus length
0.38 X length OAL, margin straight. Thoracic fovea re-
curved and tripartite, width 0. 1 8 x that of carapace, 1 .6
X wider than long, prefoveal setae absent (Fig. 41 A).
Ocular area width 0.52 x caput, 1.96 x wider than
long; AER 1.5 wide, 1.07 x width PER. Ratio of eyes:
AME: ALE; PME: PLE: 1.0: 0.76: 0.84: 0.53, diame-
ter AME 0.32; AME separated by 0.38 x their diame-
ter, PME by 1.81 X their diameter. Ocular quadrangle
1 .96 X wider than long, posterior width 1 . 1 7 x anterior.
Stemum 3.0 long, 1.32 wide, widest behind coxa
II and narrowed anteriorly, setose along margin and
sparsely setose on surface; sigilla shallow, oval, adja-
cent to coxa II, width 0. 15 x width stemum, distance
between 0.61 x distance from margin. Labium 0.42
long and wide; labium with 25 and pedipalpal coxae
with 36-37 cuspules; pedipalpal coxae 0.8 long, 0.5
wide, apex produced to a blunt point. Chelicerae 1.2
long, fang with prolateral flange, promargin of fang
furrow with 3 teeth, retromargin with 4 teeth inter-
spersed with a denticle (Fig. 4 IF).
Femur I 0.82, tibia I 0.47, femur IV 0.81, and tibia
IV 0.48 X width carapace. Ventral surfaces of patellae,
tibiae, metatarsi, and tarsi of legs I and II densely cov-
ered by long filiform setae (Fig. 4 IB). Spination: pedi-
palpus: tibia p 1 -0-0, tarsus p 1 - 1 - 1 , r 0- 1 - 1 ; leg I: tibia
p 1-1-1-la, r 3-2-3a; metatarsus p l-l-2-l-l-3a, r 2-2-
2- la; tarsus p 2-1-1, r 2-0-0; leg II: tibia p 2-2-1 -2a, r
3-2-3a, metatarsus p 2-2-2- la, r 2-2-2- la, tarsus p 1-1-
1-1, r 1-1-0; leg III: patella with approximately 45,
tibia with approximately 59, metatarsus with approxi-
mately 20, and tarsus with 5 spinules; leg IV patella
with approximately 70 slender spinules. Femur II with
row of 5— 6 stout proximal ventral setae, retrodorsum of
legs 1 and II with dense patches of stout, smooth,
procumbant setae: 13—18 apical on tibiae, 20-24 basal
on metatarsi. STC teeth (pro-retro): I, II (2-2), III, IV
(1-1); ITC simple, pedipalp claw with 1 mukidentate
tooth (Figs. 3C-D, 41G). Leg measurements (Femur +
Patella -i- Tibia -i- Metatarsus + Tarsus = [Total]): I: 3.4
+ 1.9 + 1.95 + 1.5 + 0.95 = [9.7]; II; 3.0 + 1.85 + 1.55
+ 1.4 -Hl.l = [8.9]; III: 2.55 + 1.5 + 1.55 + 1.25 + 1.15
= [8.0]; IV: 3.35 + 1.95 + 2.0 + 1.7 + 1.35 = [10.35];
pedipalpus: 1.95 + 1.10 -i- 1.15 + (absent) + 1.15 =
[5.95].
Abdomen 5.3 long, 4.3 wide, sparsely covered
with short setae. Spermathecae with broad head and
short stalk, length spermathecae 0.67 x distance be-
tween them and 1.37 x head diameter, diameter head
3.0 X diameter stalk, head length 1.28 x length stalk
(Figs. 40B, 41C).
Variation (N=3).— Total length 8.8-1 1.0; height at
fovea 0.29-0.34 x carapace width. Caput 0.70-0.77 x
carapace width, height 1.2-1.6 x that at fovea; diame-
ter ALE 0.61-1.0 X AME, PLE 0.72-1.0 x PME;
clypeus length 0.33-0.44 x OAL; thoracic fovea width
1 .25-1 .6 X length, pre-foveal setae absent or reduced
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
27
to a single seta. Sternal sigilla width 0.071-0.15 x ster-
num width; labium with 20-42 and pedipalpal coxae
with 16-40 cuspules (Figs. 43C-D); retromargin of
fang furrow with 4-5 teeth (Figs. 41E-F). Metatarsus I
with 5—7 retrolateral spines; tibia II with 5-7 and
metatarsus II with 5-7 prolateral spines, pedipalp claw
with 1—3 teeth. Spermathecae length 0.67-1 .05 x dis-
tance between them and 1.33-1.67 x head diameter, di-
ameter head 2.14—3.0 x diameter stalk, head length
1.0-1.28 X length stalk.
Natural History. — Specimens from Vatoharanana
and Talatakely were taken from nests on the trunks of
small trees or large vines in forest understory. Three
nests observed have the same form: they are oval, ori-
ented vertically on the substrate, and with a single
wafer type door at the upper end. The nest of the holo-
type is 29 mm long, 13 mm wide, and 9 mm deep; the
door is 10 mm long and 12 mm wide (Figs. 21C— D).
All nests are made of fragments of bark and lichen
woven together with silk.
Distribution. — Central to south-central Madagas-
car (Fig. 68).
Additional Material Examined. — Madagascar:
Fianarantsoa: Pare Nationale Ranomafana: Talatakely,
21°14.9'S, 47°25.6'E, 5-18 April 1998, C. E. Griswold
& D. H. Kavanaugh (1 fragmentary female, CASC), 30
October-21 November 1998, V. Lee & K. Ribardo (1
female, CASC). Antananarivo: 3 km 41° NE Andra-
nomay, 11.5 km 147° SSE Anjozorobe, 18°28.4'S,
47°56.6'E, elev. 1300 m, montane rainforest, 5-13 De-
cember 2000, CAS/PBZT Spider- Ant Class ( 1 female,
CASC).
Paramigas pauliani (Dresco & Canard)
(Figs. 40C-D, 44A-I, 65, 68)
Legendrella pauliani Dresco & Canard, 1975:783-788 (fe-
male holotype, 2 female paratypes, and 3 additional speci-
mens [Dresco & Canard reported that all 3 are juveniles but
2 are adult females], Fort Dauphin, Madagascar, collected
by Paulian, 1961, in MNHN, examined), Brignoli 1983:
119.
Paramigas pauliani. Raven, 1985:155. Platnick 1989:73.
Platnick2001.
Diagnosis. — ^Females are distinguished from other
Paramigas that lack long silky hairs beneath legs I and
II (Fig. 44C) by having tarsus III with more than 15
prolateral spinules (Figs. 44D, I) and by the form of the
spermathecae with the head diameter greater than 7 x
that of the very narrow stalk (Figs. 40C-D, 44G-H),
and from P. rothorum by having a pair of long pre-
foveal setae (Fig. 44A); male unknown.
Description. — Female (holotype): Total length
19.0. Carapace orange-brown with faint brown striae
posteriad of PME and extending to thoracic fovea (Fig.
44A); ocular area and clypeus orange-brown, black
surrounding AME and mesad of ALE and PLE; che-
licerae orange-brown, sternum, coxae, and trochanters
light orange-brown (Fig. 44B); legs and pedipalpi or-
ange-brown; patellae, especially IV, orange-brown dor-
sally; abdomen dark brown; spinnerets and book lung
covers pale yellow-white.
Carapace 7.7 long, 7.0 wide, height at thoracic
fovea 0.25 x carapace width; smooth. Caput inclined,
height 1.4 x that at thoracic fovea, width 0.71 x cara-
pace width, with shallow dimples just anteriad of tho-
racic fovea; median ocular seta present with two pairs
of setae situated anteriad to this; clypeus length 0.63 x
length OAL, margin weakly procurved, with one setae
near margin. Thoracic fovea recurved and tripartite,
width 0.18 X that of carapace, 1.6 x wider than long,
with long prefoveal setae (Fig. 44A).
Ocular area width 0.51 x caput, 1.62 x wider than
long; AER2.55 wide, 1.15 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 2.1: 1.3: 1.1, diameter
AME 0.25; AME separated by 1.4 x their diameter,
PME by 2.71 x their diameter. Ocular quadrangle 1.42
X wider than long, posterior width 1.87 x anterior.
Sternum 6.4 long, 4.3 wide, widest behind coxa II
and narrowed anteriorly, setose along margin and
sparsely setose on surface; sigilla shallow, oval, adja-
cent to coxa II, width 0.162 x width sternum, distance
between 0.4 x distance from margin (Fig. 44B). Labi-
um with 24 and pedipalpal coxae with 41-43 cuspules;
labium 1.2 long, 1.5 wide, pedipalpal coxae 2.9 long,
1.4 wide, apex produced to a sharp point. Chelicerae
1.8 long, promargin of fang furrow with 4 teeth, retro-
margin with 1 large basal tooth, distad of this row of 7
small teeth (Fig. 44E).
Femur I 5.0, tibia I 3.2, femur IV 5.5, and tibia IV
3.6 X width carapace. Spination: pedipalpus: tibia p 1-
0-0, tarsus p 1-1-1-1, r 1-1-1; leg I (Fig. 44C): tibia p
1-3-5-3, r 3-3-3-8; metatarsus p 4-4-3, r 8-4-4; tarsus p
1-1-1, r 1-1-1-1; leg II: tibia p 4-2-3, r 4-2-3-1,
metatarsus p 3-2-3-4, r 4-2-4-1, tarsus p 1-1-1, r l-I-l;
leg III (Fig. 44D): patella with approximately 46, tibia
with approximately 52, metatarsus with approximately
56, and tarsus with 15 proapical spinules; leg IV patel-
la with approximately 90 slender spinules. Femur II
with row of 5—6 stout proximal ventral setae, retrodor-
sum of legs I and II with sparse groups of stout, serrate,
procumbant setae: 4—6 apical on tibiae, 4—6 basal on
metatarsi. STC teeth (pro-retro): I, III, IV (1-1), II (2-
2); ITC simple, pedipalp claw with 1 simple tooth (Fig.
441). Leg measurements (Femur + Patella + Tibia +
28
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Metatarsus + Tarsus = [Total]): I: 5.0 + 3.0 + 3.2 + 2.5
+ 1.4 = [15.1]; IL 4.4 + 2.9 + 2.7 + 2.3 + 1.5 = [13.8];
IE: 3.7 + 2.4 + 2.1 + 2.0 + 1.8 = [12.0]; TV: 5.5 + 3.1 +
3.6 + 2.8 + 2.5 = [17.5]; pedipalpus: 3.0 + 1.7 + 1.9 +
(absent) + 1.8 = [8.4].
Abdomen 9.5 long, 6.7 wide, sparsely covered
with short setae. Spermathecae (of paratype) with
broad head and short, very narrow stalk; although the
heads are nearly contiguous the length of the sper-
mathecae is 0.86 X distance between them and 1.05 x
head diameter, diameter head 7.33 x diameter stalk,
head length 5.47 x length stalk.
Variation (N=5). — ^Total length 16.8-19.9; height
at fovea 0.21-0.25 x carapace width. Caput height
1.3-1.6 X that at fovea; diameter ALE 1.8-2.1 x AME;
clypeus length 0.6-0.7 x OAL, margin straight to
weakly procurved; thoracic fovea width 1.6—2.0 x
length. Sternal sigilla width 0.16-0.20 x sternum
width; labium with 19-26 and pedipalpal coxae with
32^5 cuspules; retromargin of fang furrow with large
basal tooth and 3—7 small teeth (Figs. 44E— F). Metatar-
sus I with 13—18 retrolateral spines; tibia II with 9-13
and metatarsus II with 11—12 prolateral spines. STC
with 2-3 teeth. Spermathecae (N=2) length 0.78-0.86
X distance between them and 0.89-1 .05 x head diame-
ter, diameter head 7.33—7.40 x diameter stalk, head
length 5.00-5.47 x length stalk (Figs. 40C-D, 44G-H).
Material Examined. — Madagascar; Toliara: Fort
Dauphin, 1961, R. Paulian (Legendrella pauliani ho\o-
type female, 2 paratype females, 2 females, 1 juvenile,
MNHN)
Distribution. — Known only from the type locality
at the southern tip of Madagascar (Fig. 68).
Paramigas pectinatus, new species
(Figs. 45A, 46A-E, 65, 68)
Type. — Holotype female collected 20 March 1994
by Alaine Pauly at the Piscine Naturelle at Isalo, Fia-
narantsoa Province, Madagascar, deposited in MRAC
(#201.283).
Etymology. — The specific name refers to the
comb or rake of procumbant setae on the anterior legs.
Diagnosis.— Females are distinguished from other
Paramigas that lack a dense vestiture of long silky
hairs beneath legs I and II by the presence of dense
patches of conspicuously serrate, procumbant setae at
apices of tibiae and bases of metatarsi 1 and II (Fig.
46B), carapace with small prefoveal setae and dimples
(Fig. 46A); spermathecae short, with base narrow, head
diameter greater than 3.7 x stalk diameter (Figs. 45A,
46C); male unknown.
Description. — Female (holotype): Total length
10.3. Carapace yellow-brown with faint longitudinal
striae extending from AME and PLE on caput to tho-
racic fovea (Fig. 46A) and faint dark striae along later-
al margins of caput; ocular area dark, black surround-
ing AME and extending between all eyes; sternum,
coxae, and trochanters light yellow-brown; legs and
pedipalpi yellow-brown gradually fading to light yel-
low-brown on tarsi; abdomen dark purple-brown, paler
ventrally; spinnerets light yellow-brown.
Carapace 4.4 long, 3.75 wide, height at thoracic
fovea 0.26 x carapace width; smooth. Caput inclined,
height 1.8 x that at thoracic fovea, width 0.76 x cara-
pace width, with weak lateral dimples; ocular area with
a single seta positioned between AME, 2 behind AME,
and a pair of setae anteriad to median ocular seta;
clypeus length 0.59 x length OAL, procurved, with
several setae. Thoracic fovea recurved and tripartite,
width 0.16 X that of carapace, 1.3 x wider than long,
with a pair of short prefoveal setae (Fig. 46A).
Ocular area width 0.44 x caput, 1.8 x wider than
long; AER 1.27 wide, 1.06 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 1.17: 0.70: 0.88, diame-
ter AME 0.17; AME separated by 1.0 x their diameter,
PME by 3.8 x their diameter. Ocular quadrangle 1.5 x
wider than long, posterior width 1.25 x anterior.
Sternum 2.85 long, 2.15 wide, widest behind coxa
II and narrowed anteriorly, setose along margin and
sparsely setose on surface; sigilla shallow, irregularly
oval, adjacent to coxa II, contiguous medially, width
0.30 X width sternum, distance from sternal margin
0.77 X diameter Labium with 24, pedipalpal coxae
with 15—21 cuspules; labium 0.32 long and wide, pedi-
palpal coxae 1 .2 long, 0.85 wide, apex produced to a
blunt point. Chelicerae 1.0 long, fang with prolateral
flange, promargin and retromargins of fang fiirrow
with 3 teeth (Fig. 46D).
Femur I 0.86, tibia I 0.46, femur IV 0.74, and tibia
IV 0.4 X width carapace. Spination: pedipalpus: tarsus
p 1-1-1, r 1-1-1; leg I: tibia p 2-2-2, r 2-2-4, metatar-
sus p 1 -2-2- 1 a, r 2-2-2, tarsus p 0- 1 , r 1 - 1 -0; leg II : tibia
p 2-2-1, r 2-2-2, metatarsus p 2-2- la, r 1-1-1-1, tarsus
p 1-1-0, r 1-0-0; leg III: patella with approximately 38,
tibia with approximately 56, and metatarsus with ap-
proximately 17 spinules, tarsus with 4—5 proapical
spinules; leg IV patella with approximately 80 slender
spinules, metatarsus v 0-0-2a. Femur II with row of
5—6 stout proximal ventral setae, retrodorsum of legs I
(Fig. 46B) and II with dense patches of stout, serrate
procumbant setae: 20-25 apical on tibiae, 25-27 basal
on metatarsi. STC teeth (pro-retro): I, II (2-2), III, IV
(1-1), ITC simple, pedipalp claw with 1 simple tooth
(Fig. 46E). Leg measurements (Femur + Patella +
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
29
Tibia + Metatarsus + Tarsus = [Total]): I: 3.25 + 1.8 +
1.75 + 1.5 + 0.85 = [9.15]; II: 2.75+ 1.6+ 1.35+ 1.25
+ 0.75 = [7.7]; III: 2.1 + 1.3 + 1.25 + 1.05 + 1.05 =
[6.75]; IV: 2.8 + 1.6 + 1.5 + 1.15 + 1.05 = [8.1]; pedi-
palpus: 1.5 + 0.95 + I.O + (absent) + 0.95 = [4.4].
Abdomen 4.9 long, 3.65 wide, sparsely covered by
short setae. Spermathecae with broad head and short
stalk, length spermathecae 1.125 x distance between
them and 0.90 ^ head diameter, diameter head 3.75 x
diameter stalk, head length 3.25 x length stalk (Figs.
45A, 46C).
Natural History. — The label with the type states
"Piscine naturelle, massif rocheux." This is an area of
large rocks within Isalo National Park, which is sur-
rounded by arid grassland.
Material Examined. — Only the type.
Distribution. — Known only from the type locality
in south-central Madagascar (Fig. 68).
Paramigas perroti (Simon)
(Figs. 21A-B, 35A-C, 47A-I, 65, 68)
Myrtale perroti Simon, 1891:300 (lectotype female from
Tamatave, Madagascar, in MNHN, examined). Simon
1892:84, 1903:883.
Kolosvarya perroti. Strand 1934:272.
Paramigas perroti, Roewer 1942:193. Bonnet 1958:3329.
Dresco & Canard 1975:785. Platnick 2001.
Paramigas subrufus Pocock, 1895:189 (lectotype female,
here designated in order to ensure the name's proper and
consistent application, and paralectotype female, from Sen-
brendrana, Madagascar, in BMNH, examined). Simon
1895:1066. Roewer 1942:193. Bonnet 1958:3329. Platnick
2001. New Synonymy
Myrtale subrufus, Simon 1903:883.
Synonymy. — In 1895, Pocock described a new
migid, Paramigas subrufus, with some reservation as
his specimens shared many characteristics with Myr-
tale perroti Simon (Pocock, 1895). However, using the
illustration provided in Simon (1892), Pocock believed
that his specimens were sufficiently different to war-
rant new status. Indeed, Simon's illustration of the
cephalothorax of Myrtale reflects a very distinct look-
ing spider (Simon 1892, fig. 84). However, upon ex-
amination of the holotypes, we have found Simon's il-
lustration to be an inaccurate representation of Myrtale
perroti specimens that differ from Paramigas subrufus
only by color.
Diagnosis. — Distinguished from other Paramigas
that have a dense vestiture of long, silky hairs beneath
patellae-^netatarsi I and II (Fig. 47C) by (contra P.
goodmani) lacking thorns at the apices of metatarsi I
and II and (contra P. oracle) having the spermathecae
long with narrow heads, length greater than 1.67 x
head diameter and head diameter greater than 2.0 x di-
ameter stalk (Figs. 35A-C, 47G-H).
Description. — Female (lectotype): Total length
20.7. Carapace orange-brown, caput with faint longitu-
dinal striations behind ocular area (Fig. 47A); ocular
area and clypeus orange-brown, black surrounding
AME and mesad of ALE and PLE; chelicerae, ster-
num, coxae, and trochanters light orange-brown (Fig.
47B); legs and pedipalpi orange-brown; abdomen light
brown; spinnerets and book lung covers pale yellow-
brown.
Carapace 8.6 long, 6.7 wide, height at thoracic
fovea 0.28 x carapace width; smooth. Caput inclined,
height 1.47 x that at thoracic fovea, width 0.71 x cara-
pace width; median ocular seta present with two pairs
of setae situated anteriad to this; clypeus length 0.7 x
length OAL, margin weakly procurved. Thoracic fovea
recurved and tripartite, width 0.5 x that of carapace,
1.57 X wider than long, with pair of small prefoveal
setae.
Ocular area width 0.5 x caput, 2.4 x wider than
long; AER 2.35 wide, 1 .02 x width PER. Ratio of eyes;
AME: ALE: PME: PLE: 1.14: 1.0: 1.0: 0.71, diameter
AME 0.4; AME separated by 0.5 x their diameter,
PME by 2.7 1 x their diameter. Ocular quadrangle 1 .5 x
wider than long, posterior width 1.36 x anterior.
Sternum 6.4 long, 4.2 wide, widest behind coxa II
and narrowed anteriorly, setose along margin and
sparsely setose on surface; sigilla shallow, oval, adja-
cent to coxa II, width 0. 1 1 x width sternum, distance
between 0.45 x distance from margin (Fig. 47B). Labi-
um with 36 and pedipalpal coxae with 39-47 cuspules;
labium 1.3 long, 1.4 wide, pedipalpal coxae 2.8 long,
1.5 wide, apex produced to a sharp point. Chelicerae
2.3 long, promargin of fang furrow with 5 teeth, retro-
margin with 1 large basal tooth and distal row of 4—6
small teeth.
Femur I 0.86, tibia I 0.52, femur IV 0.82, and tibia
IV 0.50 X width carapace. Spination: pedipalpus: tibia
p 1-0-0, tarsus p 1-1-1, r 1-1-1-1; leg I: tibia p 2-1-4, r
3-3-3-3; metatarsus p 2-2-3-2, r 2-2-3-3; tarsus p 1-1-
1, r 1-2; leg II: tibia p 3-2-3, r 2-4-2-2, metatarsus p 2-
3-2-2, r 2-1-1-1-1, tarsus p 1-1-1, r 1-0-0; leg III:
metatarsus vO-0-0-2, patella with approximately 50
prolateral and retroapical, tibia with approximately 47
pro- and retrolateral, metatarsus with approximately
21, and tarsus with 13 prolateral spinules (Figs. 47D,
I); leg IV patella with approximately 90 slender prolat-
eral spinules. Femur II with proximal ventral row of
5—6 stout setae, retrodorsum of legs I (Fig. 47C) and II
with dense patches of stout, smooth procumbant setae:
13—20 apical on tibiae, 30—35 basal on metatarsi; with
a dense vestiture of long, silky hairs beneath patellae-
30
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
metatarsi I and II. STC teeth (pro-retro): I (2-3), II (2-
2), III (1-2), IV (I- 1); ITC simple, pedipalp claw with
1 tooth having small denticle (Fig. 471). Leg measure-
ments (Femur + Patella -i- Tibia + Metatarsus -i- Tarsus
= [Total]): I: 5.8 + 3.2 -i- 3.5 + 2.4 + 1.5 = [16.4]; IL 5.0
+ 2.8 + 3.1 + 2.3 + 1.5 = [14.7]; III: 4.1 -t- 2.3 + 2.5 +
2.1 + 1.7 = [12.7]; IV: 5.5 + 3.2 -i- 3.4 + 2.7 + 1.8 =
[16.6]; pedipalpus: 3.0 + 1.5 + 1.7 + (absent) + 2.0 ^
[8.2].
Abdomen 9.8 long, 7.8 wide, sparsely covered
with short setae. Spermathecae of lectotype not dis-
sected: for description see variation below.
Variation (N=5). — Prosoma orange-brown to dark
red-brown with light red-brown areas along lateral
margins of ocular area and extending to edge of
clypeus; chelicerae yellow-brown to dark red-brown;
sternum, coxae, and trochanters yellow-brown to light
red-brown; legs and pedipalpi unicolorous or with dor-
sal surface red-brown gradually fading to light red-or-
ange on apices of metatarsi and tarsi; abdomen brown
to purple-gray. Total length 14.0-20.7; height at fovea
0.23-0.33 X carapace width. Caput 0.66-0.75 x cara-
pace width, height 1.38—1.55 x that at fovea; diameter
ALE 0.57-0.87 x AME, PLE 0.66-0.71 x PME;
clypeus length 0.41-0.8 x OAL, margin straight to
weakly procurved; thoracic fovea width 1.22—2.0 x
length. Sternal sigilla width 0.11-0.16 x sternum
width; labium with 23^5 and pedipalpal coxae with
45—107 cuspules; retromargin of fang furrow with
basal tooth and 3—5 small teeth, with or without 1-3
denticles between tooth rows (Figs. 47E— F). Tibia I
with 10-14 and metatarsus I with 8—12 retrolateral
spines; tibia II and metatarsus II each with 8—12 pro-
lateral spines. STC with 2—3 teeth. Spermathecal
length 0.78-0.90 x distance between, 1.67-2.05 x head
diameter, head diameter 2.00-2.33 x stalk diameter,
head length 0.89-1.0 x stalk length (Figs. 35A-C,
47G-H).
Natural History. — ^The nest of the large Paramigas
subrufus syntype is oval, 37 mm long, 20 mm wide,
and 18 mm deep with a single wafer type door at one
end that is 13 mm long and 17 mm wide (Figs. 21 A— B;
Pocock 1895, figs, la, lb). The nest is made of frag-
ments of bark and lichen woven together with silk.
Pocock (1895) reports that he received two nests with
the syntypes and that they appear to have been taken
from the trunk of a tree.
Material Examined. — ^M'\dagascar: no locality,
label states 'comp with type' (2 females, MRAC
122.888). Toamasina: Tamatave, Perrot "revu par
Dresco 1974,"(Mr/a/e perroti Simon, lectotype fe-
male, designated by PLC Benoit, ES #9915, AR 4134,
and syntype female, MHNH); Senbrendrana, 1891,
"purchase of Gerrard, Mons. Magestre, BMNH
1891.7.1.4-5" (2 female syntypes of Paramigas subru-
fus Pocock, BMNH).
Distribution. — East central Madagascar in Toa-
masina province (Fig. 68).
Paramigas rothorum, new species
(Figs. 45B, 48, 49A-E, 65, 68)
Type. — Holotype female collected at Montague
d'Ambre (12°30'57"S, 49°11'04"E) in Antsiranana
Province, Madagascar, on 12 August 1992 by V. and B.
Roth, deposited in CASC.
Etymology. — The specific name honors Barbara
and the late Vincent Roth, collectors of the type and
many other new and interesting spiders from Madagas-
car.
Diagnosis. — ^Females are distinguished from other
Paramigas that lack a dense vestiture of long silky
hairs beneath legs I and II by the pattern of longitudi-
nal striae extending from the PER to the thoracic fovea
(Fig. 48), by having legs I and II with sparse dorsal
patches of procumbant setae (<15) at apices of tibiae
and bases of metatarsi, the spermathecal base narrow,
having head maximum diameter greater than 4.0 x
stalk diameter (Figs. 45B, 49C), and from P. pauliani
by lacking prefoveal setae (Figs. 48, 49A); male un-
known.
Description. — Female (holotype): Total length
10.65. Carapace yellow-brown with dark longitudinal
striae extending from AME and PME to thoracic fovea
(Fig. 48) and with faint dark striae along lateral mar-
gins of caput; ocular area dark, black surrounding
AME, behind ALE, and extending between PME and
PLE; sternum, coxae, and trochanters light yellow-
brown (Fig. 49B); legs and pedipalpi yellow-brown
gradually fading to light yellow-brown on tarsi; ab-
domen dark purple-brown; spinnerets and book lung
covers light yellow-brown.
Carapace 3.85 long, 3.2 wide, height at thoracic
fovea 0.39 x carapace width; smooth. Caput inclined
(Fig. 49A), height 1.2 x that at thoracic fovea, width
0.75 X carapace width; ocular area with a single seta
positioned between AME and a pair of setae anteriad to
this; clypeus length 0.47 x length OAL, procurved,
with 2 setae. Thoracic fovea tripartite, recurved, width
0.19 X that of carapace, 2.48 x wider than long, pre-
foveal setae absent (Fig. 48).
Ocular area width 0.5 x caput, 2.28 x wider than
long; AER 1.17 wide, 1.04 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 1.13: 0.54: 0.56, diame-
ter AME 0.22; AME separated by 0.77 x their diame-
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
31
ter, PME by 4.0 x their diameter. Ocular quadrangle
1.5 X wider than long, posterior width 1.3 x anterior.
Sternum 2.3 long, 2.05 wide, widest behind coxa
II and narrowed anteriorly, setose along margin and
sparsely setose on surface (Fig. 49B); sigilla shallow,
oval, adjacent to coxa II-III, width 0.21 x width ster-
num, distance between 0.55 x width, distance from
sternal margin 0.89 x width. Labium with 19, pedipal-
pal coxae with 22—24 cuspules; labium 0.6 long, 0.7
wide, pedipalpal coxae 1.25 long, 0.8 wide, apex pro-
duced to a blunt point. Chelicerae 1.2 long, pro-and
retromargins of fang furrow with 4 teeth (Fig. 49D).
Femur I 0.8 1, tibia I 0.46, femur IV 0.82, and tibia
IV 0.5 X width carapace. Spination: pedipalpus:
metatarsus p 1-0-0, tarsus p 1-1-1, r 0-1-1; leg I: tibia p
1-1-2-2, r 1-1-2-1-2, metatarsus p 1-1-2-1-1-1, r 2-2-3,
tarsus p 1-1-1, r 1-1-0; leg II: tibia p 2-2-2, r 0-2-2-2,
metatarsus p 2-2-2, r 0-2-1, tarsus p 1-1-1, r 1-1-0; leg
III: patella with approximately 18, tibia with approxi-
mately 37, and metatarsus with approximately 31 spin-
ules, tarsus with 8—9 proapical spinules; leg IV: patella
with approximately 34 spinules. Femur II with row of
5—6 stout proximal ventral setae, retrodorsum of legs I
and II with sparse groups of stout, procumbant setae:
4—6 apical on tibiae, 5—7 basal on metatarsi. STC teeth
(pro-retro): I, II, III (2-1), IV (1-1); ITC simple, pedi-
palp claw with 1 tooth (Fig. 49E). Leg measurements
(Femur -i- Patella -i- Tibia -i- Metatarsus -i- Tarsus =
[Total]): I: 2.6 + 1.5+ 1.5 -^ 1.15 + 0.85 = [7.6]; II: 2.3
+ 1.45 + 1.35 + 1.0 + 0.85 = [6.95]; III: 1.95 + 1.15 +
1.05 + 1.0 + 0.85 = [6.0]; IV: 2.65+ 1.4+1.6+1.35 +
1.15 = [5.5]; pedipalpus: 1.6 + 0.85+ 1.05 + (absent) +
0.9 = [4.4].
Abdomen 5.9 long, 4.2 wide, sparsely covered by
short setae. Spermathecae close together with broad
head and short, narrow stalk, length spermathecae 0.74
X distance between them and 1.33 x head diameter, di-
ameter head 4.20 x diameter stalk, head length 3.67 x
length stalk (Figs. 45B, 49C).
Material Examined. — Only the type.
Distribution. — ^Known only from the type locality
in an isolated montane forest in far northern Madagas-
car (Fig. 68).
Genus Thyropoeus Pocock 1895
Thyropoeus Pocock 1895:191—192 (type species, by mono-
typy, Thyropoeus mirandus Pocock 1895). Simon
1903:883. Roewer 1942:193. Bonnet 1959:4607. Dresco &
Canard 1975:784. Raven 1985:144. Platnick 1989:71. Dip-
penaar-Schoeman & Jocque 1997:77. Platnick 2001.
Heteromigella Strand 1908:454 (type species, by monotypy,
Heteromigella malagasa Strand 1908). Roewer 1942:193.
Bonnet 1957:2184. First synonymized by Raven 1985:144.
Diagnosis. — Distinguished from all other migid
genera by having deep, lunate shaped sternal sigilla
(Figs. 5 IB, 53B, 58B), clavate spermathecae that grad-
ually taper from narrow base to broad apex (Figs.
56A— D), prolaterally expanded male tibia I (Figs. 52,
55A) and a large tooth on STC III (Figs. 5 IE, 58G).
Note. — The diagnostic characters (above) are pos-
sible synapomorphies for the genus.
Description. — Medium to large sized, 13.0-44.1
in length. Carapace smooth in female, rugose in male,
length 1.00-1.08 x width, posteriorly narrowed in fe-
male, rounded in male; caput raised (Figs. 51 A, 58A),
with median ocular seta, prefoveal setae present (7^ mi-
randus) or absent (T. malagasus). Ocular area
0.41-0.65 X width caput; AER straight to slightly re-
curved, ALE diameter equal to or less than AME; PER
straight to slightly recurved, PLE diameter equal to or
less than PME; ocular quadrangle narrowed anteriorly.
Thoracic fovea tripartite, straight or weakly recurved,
0.21-0.26 X width carapace. Sternum length 1.09-1.69
X width, margin sinuate; setose along margin, sparsely
setose on surface; sigilla lunate, deep, length 0.02—0. 1 1
X sternum width. Thorns absent from coxae. Leg for-
mula 4123 (female) or 1423 (male); femur III with
ventral membrane extending to base, tibia III with deep
dorsobasal excavation (Figs. 50, 57), anterior ridge
present or absent. Male with tibia I expanded prolater-
ally (Figs. 52, 55A), with retrolateral tibia I megaspine
(Figs. 53C, 55B), bulb unifomi (Figs. 53E-G, 54A-C),
embolus simple and slender, length 0.55—0.61 x bulb
length. Spermathecae gradually tapering from narrow
base to broad apex, straight to distally curved, with
pores throughout, unsclerotized (Figs. 51C, 56 A— D,
58C, D).
Natural History. — Collection data with 77n'-
ropoeus malagasus suggest that they may be terrestri-
al.
Composition. — ^Two species.
Distribution. — Southern and western Madagascar
(Fig. 68).
Thyropoeus malagasus (Strand) 1908
(Figs. 50, 51A-E, 52, 53A-H, 54A-C, 55A-D, 56D, 65. 68)
Heteromigella malagasa Strand 1908:454 (holotype female
from St. Marie de Marovoay, Madagascar, in NHR Stock-
holm, examined).
Heteromigella malagassa, Petrunkevitch 1928:69 (unjusti-
fied emendation). Roewer 1942:193. Bonnet 1957:2184.
Thyropoeus malagassa. Raven 1985:145. Platnick 1989:71.
Thyropoeus malagasus, Platnick 2001.
Synonymy. — Raven (1985) first proposed the syn-
onymy of Heteromigella Strand under Thyropoeus
Pocock. He noted the similarity in the deep, lunate
32
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
shaped sternal sigilla and the dorsobasal excavation on
tibia III shared by Heteromigella malagasa Strand and
Thyropoeus mirandus Pocock. Although the dor-
sobasal excavation on tibia III is shared by other Mala-
gasy Migidae, the lunate sternal sigilla are unique to
Thyropoeus and Heteromigella. The types of Het-
eromigella malagasa and Thyropoeus mirandus have
been examined and found to share these features, con-
firming Raven's decision.
Notes. — Petrunkevitch (1928:69), without com-
ment, used malagassa instead of Strand's name mala-
gasa. This change was followed as an emendation by
Roewer (1954) and Bonnet (1957), the latter stating "II
vaut mieux ecrire malagassa" (Bonnet 1957:2184).
This emendation is unjustified under the rule of
nomenclature (ICZN 1999, Article 33.2.3).
The male is described here for the first time. Al-
though the type locality of//, malagasa in Mahajanga
Province is far from Vohimena in Toliara Province, the
great similarity of females from each locality indicate
that they are conspecific.
Diagnosis. — Distinguished from Thyropoeus mi-
randus by its smaller size (length 12.9-14.8), thoracic
fovea that is straight to weakly procurved medially
(Fig. 50) in females, and the basal tooth on the fang
(Figs. 51B, 53B).
Female. — (Holotype): Total length 14.8. Speci-
men faded. Carapace pale yellow-brown with faint
dark areas along lateral margins of caput; ocular area,
chelicerae, sternum, labium, coxae, pedipalpi and legs
pale yellow-brown; abdomen yellow-white including
spinnerets.
Carapace 5.8 long, 5.4 wide, height at thoracic
fovea 0.22 x carapace width; smooth. Caput highly
arched, height 2.5 x that at thoracic fovea, width 0.79
X carapace width; median ocular seta present with a
pair and single seta anterior to this, one pair of setae on
clypeus; clypeus 0.42 x length OAL, margin straight.
Thoracic fovea weakly procurved medially and re-
curved at sides, width 0.25 x that of carapace, 7.0 x
wider than long; prefoveal setae absent.
Ocular area width 0.46 x caput, 2.85 x wider than
long; AER 2.0 wide, 1.08 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 1.6: 1.6: 1.0, diameter
AME 0.13; AME separated by 1.3 x their diameter,
PME by 3.0 x their diameter. Ocular quadrangle 0.92 x
wider than long, posterior width 2.08 x anterior.
Sternum 3.6 long, 3.2 wide, widest behind coxa II
and narrowed anteriorly, sparsely setose along margin
and on surface; sigilla adjacent to coxa II, 0.03 x width
sternum, distance between 0.77 x distance fi^om mar-
gin. Labium with 17 and pedipalpal coxae with 32—33
cuspules; labium 1.0 long, 1.3 wide, pedipalpal coxae
2.0 long, 1.4 wide, apex produced to a blunt point.
Chelicerae 2.5 long, promargin of fang furrow with 3
teeth, retromargin with 6 teeth, pro-and retromargins
interspersed with 8 denticles.
Femur I 0.59, tibia I 0.37, femur IV 0.68, and tibia
IV 0.39 X width of carapace. Spination: pedipalpus:
tibia p 1-2, r or 0-0-1, tarsus p 2-2-1-1, r 1-1-1-1; leg
I: tibia p 3-2-2, r 2-4-2, metatarsus p 2-4-4, r 3-3-3, tar-
sus r 2; leg II: tibia p 3-3-2, r 0-2-2, metatarsus p 2-3-
3, r 3-2-2, tarsus r 2-0; leg III: patella with approxi-
mately 55, tibia with approximately 75, metatarsus
with approximately 54, and tarsus with approximately
20 spinules; leg IV: patella with approximately 70 and
tibia with approximately 30 spinules, metatarsus vO-0-
2a. STC (pro-retro) I, II, and IV (l-I), III (1-2), ITC
simple, pedipalpal claw with 1 simple tooth (Fig. 5 IE).
Leg measurements (Femur -i- Patella + Tibia -i- Metatar-
sus + Tarsus = [Total]): I: 3.2 -(- 2.2 + 2.0 +1.55 + 1.0
= [9.95]; II: 2.95 + 2.2 + 1.9 + 1.65 + 1.2 = [9.9]; III:
2.65 + 2.05 + 1.5 + 1.65 + 1.4 = [9.25]; IV: 3.7 + 2.35
+ 2.15 + 2.05 + 1 .7 = [ 1 1 .95]; pedipalpus: 2.4 + 1 .35 +
1.5 + (absent) + 1.5 = [6.75].
Abdomen 6.5 long, 5.5 wide, sparsely covered
with short setae. Spermathecae not dissected but
through cuticle they appear to be like those of FMNH
female (see below).
Variation (N=2). — Total length 13.0-14.8. Mark-
ings of a fresh female specimen from Vohimena are
pale yellow-brown on prosoma except dusky on mid-
dle of and along margin of caput, with black surround-
ing each AME and mesad of each ALE, PME and PLE;
abdomen yellow-white, unmarked (Figs. 50, 51A— B).
Height at fovea 0.16-0.22 x carapace width. Caput
0.79-0.84 X carapace width, height 2.5-^3.62 x height
at thoracic fovea; width ocular area 0.41—0.46 x caput
width, diameter ALE 1.6-1.7 x AME, PLE 1.4-1.6 x
PME; clypeus length 0.83-1.15 x OAL, 3^ setae an-
terior of ocular area; thoracic fovea width 6.2-7.0 x
length. Teeth of fang furrow interspersed with 6-9 den-
ticles (Fig. 5 ID). Sternal sigilla width 0.03-0.1 1 x ster-
num width; labium with 16-17, pedipalpal coxae with
33-40 cuspules. Tibia I with 8, metatarsus I with 9-10
retroventral spines, tibia II with 6-8 proventral spines.
STC may have small denticle distad to tooth. Sper-
matheca (of FMNH specimen from Vohimena) length
3.0 X maximum diameter, length 0.88 x base width,
maximum diameter 1.45 x minimum, head and stalk
not distinguishable (Figs. 51C, 56D).
Male. — (Vohimena): Total length 11.7. Carapace
red-brown with faint dark areas along median and mar-
gins of caput and around thoracic fovea; ocular area
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
33
and chelicerae red-brown, black between AME and
mesad of ALE, PME and PLE (Figs. 52, 53A); ster-
num, coxae, and trochanters pale yellow-brown (Fig.
53B); pedipalpi and legs 11 through IV pale yellow-
brown with dorsal surface of femora distinctly darker
than ventral surface; leg I red-brown fading to pale yel-
low-brown ventral ly and on tarsus and apex of metatar-
sus (Figs. 52, 53C); abdomen yellow-white including
spinnerets.
Carapace 5.0 long, 4.7 wide, height at thoracic
fovea 0.25 x carapace width; rugose, margin rebor-
dered. Caput inclined (Fig. 53 A), height 1.5 x that at
thoracic fovea, width 0.65 =< carapace width; median
ocular seta present with a pair of setae anterior to this;
clypeus 0.66 x length OAL, rugose, margin straight.
Thoracic fovea T-shaped, tripartite, with deep posteri-
or part, width 0.21 x that of carapace, 1.6 x wider than
long, prefoveal setae absent (Fig. 52).
Ocular area width 0.41 x caput, 2.16 x wider than
long; AER 1.35 wide, 1.02 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 0.63: 1.57: 0.36, diame-
ter AME 0.27; AME separated by 0.45 x their diame-
ter, PME by 3.14 x their diameter Ocular quadrangle
1.5 X wider than long, posterior width 1.5 x anterior.
Sternum 2.95 long, 2.45 wide, widest behind coxa
II and narrowed anteriorly; sigilla adjacent to coxa II,
0.06 X width sternum, distance between 1.18 x dis-
tance from margin (Fig. 53B). Labium and pedipalpal
coxae lacking cuspules; labium 1.0 long, 0.95 wide,
pedipalpal coxae 2.0 long, 1 .05 wide, apex produced to
a blunt point. Chelicerae 1.7 long, fang long and slen-
der, with longitudinal basal tooth, promargin of fang
fitrrow with 4 teeth, retromargin with 8 teeth, pro-and
retromargins interspersed with 8 denticles (Fig. 53D).
Tibia 1 prolaterally swollen and with retrolateral
megaspine (Figs. 52, 53C, 55A— B). Femur 1 1.0, tibia 1
0.78, femur IV 0.89, and tibia IV 0.63 x width cara-
pace. Scopulae weak beneath tarsi 1 and II, dense and
entire beneath tarsi 111 and IV and apically on 1/3 of
metatarsus IV. Spination: leg 1: patella p 0-0-1-1, v 0-
0-0-3, r 0-0-1-0, tibia p 0-2-2-2-2a, v 0-1-1-0, metatar-
sus p 0-1-1-1-la, r 0-1-1-1-la, tarsus r 2-1-0; leg 11:
patella p 0-0-1, v 0-0-1, tibia p 1-1-1-1, v 1-2-1-2-la,
metatarsus p 0-1-1-I-la, r 1-2-1-1-la, tarsus r 1-1-0;
leg III: patella with approximately 45, tibia with ap-
proximately 50, and metatarsus with approximately 30
spinules; leg IV: patella with approximately 50 and
tibia with approximately 25 spinules. STC (pro, retro)
I (1-1), II (1-2), III, IV (1-1), ITC simple (Fig. 53H).
Leg measurements (Femur + Patella + Tibia + Metatar-
sus + Tarsus = [Total]): I: 4.7 + 2.5 + 3.7 + 2.8 + 1.2 =
[14.9]; II: 4.0 + 2.0 + 2.7 -i- 2.6 + 1.4 = [12.7]; III: 3.4
+ 2.0 + 2.0 + 2.1 + 1.9 = [11.4]; IV: 4.2 + 2.2 + 3.0 +
2.6 + 2.3 = [14.3]; pedipalpus: 3.5 + 1.5 + 2.9 + (ab-
sent) + 0.8 = [8.7].
Pedipalpus with femur 0.74, tibia 0.62 x carapace
width; femur 4.37, tibia 3.62 x length tarsus; tibia
stout, prolaterally swollen (Figs. 53E— G, 54A), height
0.34 X length; tarsus without apical spinules; bulb
width 1.75 X tarsus length; embolus length 1.25 x bulb
width (Fig. 53F). Abdomen 5.0 long, 3.8 wide, sparse-
ly covered with spiniform setae (Fig. 52).
Variation (N=5).— Total length 8.8-12.9; height at
fovea 0.14—0.25 x carapace width. Caput inclined,
caput height 1.33—2.5 x that at thoracic fovea, width
0.31—0.38 X carapace width; width ocular area
0.41-0.53 X caput width, diameter AME 0.20-0.32,
PME 0.10-0.20, PME interdistances 3.14-6.0 x their
diameter; clypeus length 0.57—0.88 x OAL; thoracic
fovea width 1.6-4.0 x length. Sternal sigilla width
0.06-0.115 X width sternum; retromargin of fang fur-
row with 4—6 teeth and 2-4 denticles. Metatarsus II
with 4—5 proventral spines. STC with 1—3 teeth. Bulb
length 2.13-2.93 x width.
Natural History. — A female and many male speci-
mens were collected in pitfall traps during January
1996. Whereas males were probably wandering, the
presence of a female in these terrestrial traps suggests
that this species may be terrestrial.
Distribution. — ^Western Madagascar (Fig. 68).
Material examined. — Madagascar: Mahajanga:
St. Marie de Marovoay, 22 November 1906 (holotype
female of Heteromigella malagasa Strand 1908,
NHR). Toliara: Foret de Vohimena, 35 km SE Sakara-
ha, 22°41.0'S, 44°49.8'E, elev. 780m, 17-24 January
1996, S. Goodman, (2 males CAS; 1 female, 23 males,
FMNH).
Thyropoeus mirandus Pocock (Figs. 56A— C, 57,
58A-G, 65, 68)
Thyropoeus mirandus Pocock 1895:192 (holotype female
from S. Central Madagascar, in BMNH, examined). Simon
1903:84. Roewer 1942:192. Bonnet 1959:4607. Dresco &
Canard 1975:784. Raven 1985:145. Platnick 2001.
Diagnosis. — Distinguished from Thyropoeus
malagasus by its exceptionally large size (>30 mm
body length), the absence of a basal tooth on the fang
(Fig. 58B), and the broad and short ocular area that is
more than 3 x wider than long (Fig. 57).
Description. — Female (holotype): Total length
44. 1 . Carapace dark yellow-brown with faint dark lon-
gitudinal reticulate markings extending along margins
of caput and medially from PME to thoracic fovea
(Figs. 57, 58A). Chelicerae, pedipalpi, and legs dark
34
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
yellow-brown, femora with dorsal longitudinal bands,
integument darker beneath dorsal and lateral setal
bands on patellae-tibiae. Sternum, labium, and coxae
dark orange-brown except yellow-brown transverse
band at stemo-labial junction (Fig. 58B). Abdomen
dark brown including spinnerets. Abdomen split open
and stuffed with cotton, connected to prosoma by a pin.
Carapace 18.5 long, 17.0 wide, height at thoracic
fovea 0.26 x carapace width; smooth. Caput highly
arched (Fig. 58A), height 2.1 x that at thoracic fovea,
length 0.52 x carapace width; median ocular seta pres-
ent with 13 setae positioned anteriorly; clypeus length
0.03 X length OAL, margin straight. Thoracic fovea
deep, T-shaped, width 0.25 x that of carapace, 8.6 x
wider than long (Fig. 57). Prefoveal setae present.
Ocular area width 0.65 x caput, 3.5 x wider than
long; AER 8.8 wide, 1.1 x width PER. Ratio of eyes:
AME: ALE: PME: PLE: 1.0: 0.3: 0.6: 0.5, diameter
AME 0.6; AME separated by their diameter, PME by
6.27 X their diameter. Ocular quadrangle 2.25 x wider
than long, posterior width 2.51 x anterior.
Sternum 16.1 long, 9.5 wide, widest behind coxae
II and narrowed anteriorly; sigilla adjacent to coxa II,
0.06 X width sternum, distance between 1.16 x dis-
tance from margin (Fig. 58B). Labium with 28 and
pedipalpal coxae with 39-43 cuspules; labium 3.7
long, 4.7 wide, pedipalpal coxae 7.2 long, 4.3 wide,
apex produced to a blunt point. Chelicerae 5.0 long,
promargin of fang furrow with 4 teeth, retromargin
with 3 teeth, pro and retromargin interspersed with 16
denticles (Fig. 58E).
Legs I and II with dense ventral distribution of
long, slender setae. Femur I 0.64, tibia I 0.44. femur IV
0.61, and tibia IV 0.41 x width carapace. Spination:
pedipalpus: tibia p I-I, r 0-2, tarsus p 0-3, r 2-2; leg I:
tibia p 2-3-3-4. r with 29 spines in two longimdinal
rows, metatarsus p 2-4-3-3-2-2-4-3, r 3-4-3-2-3-3-4-8,
tarsus p 1-1, r 3-0; leg II: tibia p 0-1 -1-2-2 -2 -2-1, r 2-
3-5-5-3-3-2, metatarsus p 3-3-5-8-2, r 4-2-2-2-5-4, tar-
sus p 1-1, r 2-0; leg III: patella with approximately 90,
tibia with approximately 140, and metatarsus with ap-
proximately 90 spinules; metatarsus with approximate-
ly 1 80 and tarsus with approximately 80 slender spin-
ules; leg IV: patella with 60 spinules, metatarsus v 0-0-
0-3a. STC (pro. retro) I (2-1), II (2-2), III. IV (1-2),
ITC simple, pedipalp claw with a single large tooth.
Leg measurements (Femur + Patella + Tibia + Metatar-
sus + Tarsus = [Total]): I: 10.9 + 6.0 + 7.6 + 5.6 + 1.6
= [31.7]; II: 9.6 + 6.0 + 6.5 + 5.5 + 1.9 = [29.5]; III: 8.7
+ 5.0 + 5.1 + 5.2 + 3.2 = [27.7]; IV: 10.5 + 8.0 + 7.0 +
7.1 + 3.6 = [36.2]; pedipalpus: 7.7 + 3.2 + 5.1 + (ab-
sent) +4.1 = [20.1].
Abdomen 23.0 long, 21.1 wide, sparsely covered
with short setae. One spermathecae dissected out, with
very narrow base and broad apex, length 1.79 x maxi-
mum diameter, maximum diameter 4.75 x minimum
(Figs. 56B, 58C).
Variation (N=3). — ^Total length 32.0-44.1; height
at fovea 0.21-0.26 x carapace width. Caput 0.76-0.84
X carapace width, 2.0-3.0 x height at thoracic fovea;
width ocular area 0.56-0.69 x caput width, OAW
3 .5-4.0 X OAL, diameter ALE 1 . 1 4-1 .28 x AME, PLE
= PME; clypeus length 1.12-1.54 x OAL, margin
weakly procurved to straight; thoracic fovea width
1.92—3.53 X length. Fang ftirrow interspersed with
14—16 denticles (Figs. 58E-F). Sternal sigilla width
0.03-0.07 X stemimi width, distance between 0.6-2.0
X distance from sternal margin; labium with 25-40,
pedipalpal coxae with 35-50 cuspules. Tibia I with
12-29, metatarsus I with 30-32 retroventral spines,
tibia II with 11-18, metatarsus II with 20-22 proven-
tral spines. Legs I and II may have sparse to dense
vestiture of fine ventral setae. STC I and II have 1—3
teeth (Fig. 58G). Spermathecal length 1.61-1.79 x
maximum diameter, maximum diameter 4.75—5.14 x
minimum (Figs. 56A-C, 58C— D).
Distribution. — Probably southern Madagascar: the
type locality is vague but the Paris specimens are
recorded from Ft. Dauphin at the southern tip of the is-
land (Fig. 68).
Material Examined. — Madagascar: "S. Central
Madagascar," 1894, J. Last, BM 1894.2.27.2 (holotype
female of Thyropoeus mirandus Pocock, BMNH). To-
liara: Fort Dauphin (3 females, MNHN AR4135).
PHYLOGENETICS
Data
A data matrix was assembled (Appendix) com-
prising 45 characters scored for all 14 Malagasy
species, three exemplars of the outgroup families Id-
iopidae, Ctenizidae and Actinopodidae, and 11 exem-
plars of the 6 non-Malagasy migid genera Calathotar-
sus, Heteromigas, Mallecomigas, Migas, Moggridgea
and Poecilomigas. This matrix was analyzed under a
variety of parameters to obtain trees of minimum
length and of maximum fit {sensu Goloboff 1993c).
Exemplars for the non-Malagasy migids and out-
groups are listed in Table 2. When possible the type
species of each genus was chosen, i.e., Calathotarsus
cownatus, Heteromigas dovei, Mallecomigas schlin-
geri. and Poecilomigas abrahami. Poecilomigas
basilleupi was also included because it differed from P.
abrahami in several characters of potential phyloge-
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
35
netic importance (e.g., male tibia I megaspine, dense
setal vestiture beneath legs 1 and II of female). The
type species of Moggridgea (M. dyeri) was inappropri-
ate because it is known only from females. Instead,
Moggridgea species were chosen to reflect the phylo-
genetic diversity of the genus as proposed in Griswold
(1987a): M. peringneyi from the peringueyi group, M.
intermedia from the quercina group, and M rupicola
from the crudeni group. The Australian Moggridgea
tingle was also included to test the monophyly of the
genus. Exemplar selection for Migas was perhaps the
most difficult problem. The species that we examined
include the type species and comprise species from the
full geographic range of the genus: Migas afftnis from
New Caledonia, M. distinctus. M. gatenbyi, M. giveni.
M. paradoxus, M. taierii and a male of an undescribed
species from New Zealand, and M. nitens and M. vari-
apalpus from Australia. In our dataset the exemplar
Migas comprises data from female M. gatenbyi and
male M. taierii: M. vellardi from Chile is included as a
separate taxon to test its placement within the Migidae.
All Malagasy taxa are coded from the specimens listed
in the monograph.
Outgroups
The Migoidea, comprising Migidae and Actinopo-
didae, has been well supported in the studies of Plat-
nick and Shadab (1976), Raven (1985) and Goloboff
(1993a). In order to polarize characters within the
Migoidea additional outgroups are necessary. Raven
proposed Ctenizidae and Idiopidae as successive out-
groups to the Migoidea and Goloboff depicted the
same families as forming a trichotomy with the
Migoidea. Accordingly, we selected representatives of
the Actinopodidae (Actinopus), Idiopidae (Idiops) and
Ctenizidae (Bothriocyrtum) as outgroups to the Migi-
dae. Exemplars were chosen on the basis of available
material of both sexes at CASC; in addition, the exem-
plars of the Actinopodidae and Idiopidae are the type
genera of their families.
Analysis
We analyzed our data with Hennig86 (Farris 1988)
and NONA 1.8 (Goloboff 1993b) to find minimum
length trees and Pee- Wee 2.6 (Goloboff 1997) to obtain
trees that maximize implied weights across all charac-
ters ("fittest" sensu Goloboff 1993c), with all charac-
ters equally weighted (wt = 1) and unordered. We dis-
cuss the results with reference to our preferred clado-
gram (Fig. 65). Hennig86 analysis using the options t;
bb*; gave 42 trees of 96 steps with c.i (consistency
index) = 0.52 and r.i. (retention index) = 0.76. The
strict consensus of these trees is like Figure 65 except
for lacking clades G through K. The results of these
analyses were subjected to several rounds of succes-
sive weighting using the routine xs; w; and the trees
obtained were compared with the weights reset to 1.
Successive weighting gave 36 trees of 97 steps (1 step
longer than the minimum). The consensus of these
trees is like our preferred tree (Fig. 65) except for no
resolution among the Paramigas species (i.e., nodes
G — K were absent) and in lacking clade C, i.e. the
placement of Heteromigas is ambiguous. We imple-
mented Nona shuffling the input order of taxa 50 times
(command mult*50). Trees were calculated accepting
(amb =) and rejecting (amb -) ambiguous branch sup-
port. Accepting ambiguous branch support gave 45
trees, rejecting ambiguous branch support gave only
four trees: all were of 96 steps. In each case the con-
sensus gave no resolution among the Paramigas
species (i.e., nodes G — K were absent). We imple-
mented Pee- Wee shuffling the input order of taxa 50
times (command mult*50), rejecting ambiguous sup-
port (default) and using the concavity values 1 to 6.
Concavity 1 gave 16 trees of fit 309.1. The strict con-
sensus of these trees groups Heteromigas with clade S
(Calathotarsus, Goloboffia vellardi, and Mallecomi-
gas) and provides no resolution within Paramigas. The
other concavity values each gave the same four trees:
concavity 2 fit = 340.2, concavity 3 fit = 359.9, con-
cavity 4 fit = 372.4, concavity 5 fit = 380.7 and con-
cavity 6 fit = 387.2. In each case the strict consensus
was like our preferred tree except for no resolution
within Paramigas.
The least restrictive assumption (accepting am-
biguous branch support) allows numerous trees that
suggest no resolution with Paramigas. More restrictive
assumptions (i. e., rejecting ambiguously supported
branches or preferring fittest trees derived with most
concavity values) gave a smaller set of possibilities: 4
trees. Although the consensus of these also suggests no
resolution within Paramigas, examination the 4 trees
reveals a simple problem: resolution is lost in the con-
sensus because P. manakambus and P. macrops may
trade positions. Each in turn may be the sister group of
P. oracle or belong to clade H. In one tree P. macrops
is the sister group of P. oracle and P. manakambus be-
longs to clade H, and in three others P. manakambus is
the sister group of P. oracle and P. macrops belongs to
clade H. The first tree is also most fully resolved.
Preferred tree
We believe that the most efficient character opti-
mization and most realistic discussion of evolution
36
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
should be done on the shortest, best resolved tree pos-
sible. A tree is a hypothesis, and maximum resolution
and minimum length make the maximum number of
predictions that can be tested. We prefer the tree that is
of minimum length and most completely resolved
given only unambiguous node support (Fig. 65). This
tree requires 96 steps and has a consistency index of
0.52 and a retention index of 0.76. Readers should keep
in mind that equally short but less resolved solutions
for Paramigas occur, and that their consensus offers no
resolution within Paramigas.
Character optimization and
branch support
We used MacClade 3.0 (Maddison and Maddison
1992) and Clados 1.2 (Nixon 1992) to optimize char-
acters on the tree. If optimizations were ambiguous, we
usually resolved them using the ACCTRAN option
(Farris optimization), which favors secondary loss over
convergence to explain homoplasy and therefore max-
imizes homology. Character optimizations and evolu-
tion are discussed below. Branch support indices (Bre-
mer 1994) were calculated with Nona for the clado-
gram depicted in Figure 65 using the options
hold25000 bsupportS. The "Bremer Support"
("Decay Index") for a given node in the shortest un-
constrained tree is the number of additional steps re-
quired in the shortest trees for which that node collaps-
es. Due to lengthy calculation times the search for
branch support was truncated at values of 5, therefore
the Bremer support values reported range between
and 5 or greater. Bremer support (decay indices) for the
nodes in Figure 65 are A (2), B (3), C ( 1), D (4), E (5),
F (1), G— K (0), L (4), M (2), N (>5), O (4), P (4), Q
(3),R(l),S(2)andT(l).
CHARACTER DESCRIPTIONS AND
INTERPRETATIONS
We describe the characters and their states and
also discuss their evolution as implied by our analyses.
Character evolution is referred to our preferred clado-
gram (Fig. 65).
1. OAW/ Caput width: (0) less than 0.41, (1)
greater than 0.45.
A traditional character used to support the mono-
phyly of the Migoidea has been the broad distribution
of eyes on the caput (Platnick & Shadab 1976; Raven
1985; Goloboff 1991). This feature optimizes at node
A as a synapomorphy for the Migoidea.
2. Ocular area width: (0) normal for Migoidea, (1)
extra wide.
The ocular areas of Calathotarsus coronatus (Fig.
4), C. simoni (but not C. pihuychen) and Goloboffia
vellardi are especially wide, greater than 0.60 x cara-
pace width (Fig. 14 A). Most migids and their out-
groups have the ocular area with less than 0.40 x cara-
pace width, and even the wide ocular areas of Actino-
pus and Thyropoeus mirandus (Fig. 57) are less than
0.55 X carapace width. The extra wide ocular area op-
timizes as a synapomorphy at node T for Calathotarsus
plus Goloboffia vellardi with presumed reversal in C.
pihuychen.
3. Prefoveal setae: (0) absent, (1) present, small,
length < 1 X interdistances, (2) present, enlarged, length
>lx interdistances.
A pair of setae is usually present on the caput near
the anterolateral margins of the thoracic fovea (Figs. 1,
18). These setae may be small in or absent from males.
Bothriocyrtum and Idiops have at least three bands of
setae extending from the PME to the thoracic fovea. In
Idiops an enlarged pair of setae, which we code as pre-
foveals, is closer to the ocular area than to the thoracic
fovea. In Actinopus females there are setae near the an-
terior margin of the thoracic fovea, which the highly-
arched caput renders nearly invisible when viewed
from above. In some Migidae the pre-foveal setae are
absent, in others conspicuously enlarged. Our clado-
gram implies a complex evolutionary history for these
setae. Small prefoveals (state 1) is the most likely ple-
siomorphic state (Fig. 46A). Also coded as state 1 are
the infinitesimal prefoveals of Paramigas alluaudi
(Fig. 28A) and some Paramigas males. Enlarged pre-
foveals (state 2) have evolved in parallel five times: at
node Q as a synapomorphy for Poecilomigas plus
Migas (Figs. I3A— B, 18), and in Idiops, Moggridgea
tingle, Paramigas pauliani (Fig. 44A), and P. milloti
(Fig. 1). Prefoveals have been lost in parallel (state 0)
four times: in Moggridgea intermedia (Figs. 15A— B),
Thyropoeus malagasus (Figs. 50, 5 1 A), and in Parami-
gas oracle (Fig. 41 A) and P. rothorum (Fig.48).
4. Caput setation: (0). Extensive, with four to
many setae posteriad of ocular area. (1) reduced, with
prefoveal setae only or lacking setae altogether poste-
riad of ocular area.
Idiopids, ctenizids and actinopodids have the
caput with two to several rows of setae extending back
from the ocular area. Most Migidae also have setae on
the caput. Migas (Figs. 13A— B), Heteromigas (Fig. 8),
Mallecomigas (Fig. 12 A) and Poecilomigas (Fig. 18)
may have three strong longitudinal rows of setae; most
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
37
Moggridgea (Figs. 15 A, B), Calathotarsus , and
Golobofjia vellardi have two to three weak rows; Mog-
gridgea tingle, other Moggridgea and Migas affinis
have at least a row of 2—3 setae as well as prefoveal
setae. The Malagasy Migidae lack setae on the caput
posteriad of the ocular area with the exception of pre-
foveal setae (Figs. 2A, 23A, 50). This loss is a synapo-
morphy at node D.
5. Female caput height/ caput length: (0) highly
arched (height > 0.72 x length), (1) low (height < 0.7 x
length).
When viewed from a lateral perspective, the fe-
male carapace of Actinopodidae, Idiopidae, Ct-
enizidae, and some Migidae (i.e., Calathotarsus [Fig.
5 A], Thyropoeus [Figs. 51 A, 58A], Heteromigas [Fig.
9A]) is strongly arched. This character is most dramat-
ic in the Actinopodidae. The arch may be absent or re-
duced in males. Most migids have the caput only
slightly higher than or equal to the height at the tho-
racic fovea (Figs. 13 A, 15 A, 23 A, 34A). A ratio com-
paring the height of the caput to its length (from tho-
racic fovea to clypeus) quantifies this character. The
arched caput is plesiomorphic with a low caput derived
in parallel three times: as synapomorphies at nodes E
and N, and in Goloboffia vellardi.
6. Fovea: (0) procurved, (1) recurved.
The thoracic foveae of the outgroups Actinopodi-
dae, Idiopidae, Ctenizidae are strongly procurved. All
Migidae have the fovea recurved: even Thyropoeus
malagasus, which has the median part of the fovea
slightly procurved, has the comers recurved (Fig. 50).
The recurved fovea (Fig. 2A) is a synapomorphy at
node B for the Migidae.
7. Fovea shape: (0) simple, (1) tripartite.
Some migids have a longitudinal, posterior groove
connecting with the transverse foveal groove (Figs.
2A, 22, 41A). Pocock (1895) was the first to recognize
this tripartite fovea in Paramigas subrufus and used it
to diagnose the genus Paramigas. Both males and fe-
males of Heteromigas and all Malagasy migids except
Thyropoeus malagasus have a conspicuous posterior
groove, though the groove in Heteromigas is shallow
(Figs. 8, 10) whereas that of the Malagasy migids is
deep. As noted by Griswold (1987a, figs. 106—116,
149—150) the character may be variable at least within
some Moggridgea. Some Calathotarsus individuals
may have this weakly developed and it is faintly visi-
ble in Mallecomigas (Fig. 12A). We consider the
groove reduced in these taxa, and code their foveae as
simple. Interestingly, all Malagasy Paramigas share
this fovea shape including very small immature indi-
viduals (Fig. 2A). The tripartite condition optimizes as
plesiomorphic for those taxa with recurved foveae with
the simple fovea arising twice: as a synapomorphy at
node M and in Thyropoeus malagasus.
8. Cheliceral tooth row number: (0) 2 rows only,
(1) with denticles scattered between the rows of large
teeth.
Some Migidae and all outgroup taxa except Idiops
show complex dentition patterns on the chelicerae in-
cluding large teeth and much smaller denticles, which
are scattered on the fang furrow between the tooth
rows and beneath the fang when it is closed. Idiops has
a promarginal row of large teeth and a retrobasal row
of small teeth, which we have not coded as denticles.
Denticles occur in Calathotarsus (Fig.5D), Heteromi-
gas (Fig. 9C), Thyropoeus (Figs. 5 ID, 58E-F) and Mi-
cromesomma (Fig. 23C). Goloboffia vellardi has 2—3
denticles (Fig. 14E), as do some Paramigas (Fig. 39D).
Other migids have only 2 rows of teeth without denti-
cles (Figs. 13D, 15E, 41E-F). The plesiomorphic state
is equivocal at the base of our cladogram, but denticles
optimize unambiguously as plesiomorphic for the
Migoidea. Loss of denticles occurs in Idiops, at nodes
H and K within Paramigas, and possibly at node M
with their reappearance at node T as a synapomorphy
for Calathotarsus plus Goloboffia vellardi. Alterna-
tively their presence in Calathotarsus and Goloboffia
vellardi may be a plesiomorphic retention: this requires
parallel loss at node N and in Mallecomigas.
9. Intercheliceral basal swellings: (0) absent; (1)
present.
This is a small unsclerotized swelling located
basally on the cheliceral promargin proximad of the
teeth (Figs. 24E, 43B) in Actinopus, Migas, Poecilomi-
gas, Paramigas, Micromesomma and some Mog-
gridgea. This feature appears to have arisen three
times: in Actinopus and as synapomorphies at nodes E
and N (with loss of this feature in Moggridgea
peringueyi).
10. Male intercheliceral tumescence: (0) absent;
(1) present.
Intercheliceral tumescence refers to pale, unscle-
rotized areas presumed to be of glandular origin on the
interior surfaces of the chelicerae in some male myga-
lomorphs. This feature has been recorded by Raven
(1985) and scored and used cladistically by Goloboff
(1993a). On our cladogram (Fig. 65) intercheliceral
tumescence arises as a synapomorphy at node M with
loss within Moggridgea at node R
11. Rastellum: (0) absent, (1) present.
A traditional character used to unite the Migidae
38
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
has been the absence of a rastellum at the apex of the
chelicerae (Raven 1985, Goloboff 1991). This charac-
ter is corroborated by our analysis as a synapomorphy
at node B for the Migidae.
12. Fang shape: (0) round, (1) quadrate and
keeled.
A traditional character used to support the mono-
phyly of the Migidae has been the presence of strong
keels on the fang (Goloboff 1991. Raven 1985). The
keeled fangs of Migidae are also quadrate in shape
with minute serrations basally on the pro-and retrolat-
eral margins (Figs. 2C, 43A). This character is corrob-
orated by our analysis as a synapomorphy at node B for
the Migidae.
13. Fang basal tooth: (0) absent, (1) present.
The monophyly of the Miginae (Migas and Poe-
cilomigas) has been suggested by the presence of a
small basal tooth on the fang (Fig. 59D) (Griswold
1987a, 1987b; Raven 1985). Goloboff and Platnick
(1987) were first to recognize potential problems with
this character as a similar tooth is also present in
Actinopus. Thyropoeus malagasus has a basal tooth
(Fig. 5 IB) although its sister species, T. mimndus, does
not (Fig. 58B). Our analysis suggests that the basal
tooth has arisen in parallel 4 times: as a synapomoiphy
at node Q for Migas plus Poecilomigas, and as autapo-
morphies in Actinopus, Goloboffia vellardi, and Thy-
ropoeus malagasus.
14. Fang orientation: (0) vertical, (1) diagonal.
The diagonal fang orientation found in the
Actinopodidae and Migidae (Figs. 2C, 12B) was used
by Raven (1985) in support of the Migoidea. The char-
acter is further corroborated as support at node A for
the Migoidea in our analysis.
15. Pedipalpal coxa cuspule distribution: (0)
across coxa to apex, (1) proximally near labium.
Females of all Chilean Migidae (including
Goloboffia vellardi) share a cuspule distribution that is
concentrated at the proximal edge of the pedipalpal
coxa near the labium (Figs. 5B, 12, 14B), which is a
synapomorphy at node S. Bothriocyrtum also has this
state, but this is most parsimoniously interpreted as a
parallelism.
16. Pedipalpal coxa cuspule anterior distribution:
(0) broadly across coxa to apex, (1) only proximally
near labium.
Goloboff and Plamick (1987) noted that Heteromi-
gas and Mallecomigas have the cuspules on the pedi-
palpal coxae extending to the anterior face of the seg-
ment (rather than confined to the ventral surface) and
suggested that this feature is primitive and might indi-
cate that either of these genera, or both together, might
represent the sister group of all remaining migids. In
fact cuspules extend onto the anterior surface of the
pedipalpal coxae in all taxa studied. In Actinopus and
Heteromigas cuspules extend broadly onto the anterior
surface, whereas in all others the cuspules extend only
onto the anteromedian comer. We have optimized
broad anterior extension as a synapomorphy for
Migoidea at node A with reversal to anterior extension
only at the comer as a synapomorphy at node C. Alter-
natively broad extension may have arisen in parallel in
Actinopus and Heteromigas.
17. Thorns on coxae II-III: (0) absent, (1) present.
Thorns are stout, shortened setae that have a
length 5 X or less than their basal diameter (Figs. 23B,
25D). They differ from cuspules in gradually tapering
to a point rather than first expanding from the base
(Fig. 25D). Pocock (1895) used the distribufion of
thorns on the coxae as support for his new genus Mi-
cromesomma . Thorns occur in both females and males
of some Moggridgea, though reduced in size and num-
ber in the latter (the male of Micromesomma is un-
known). Griswold (1987a) also used the distribution of
thorns on the coxae in his analysis of Moggridgea and
suggested that these features indicate a sister-group re-
lationship between Micromesomma and Moggridgea.
Our cladogram refutes this, suggesting that thorns
arose in parallel in Micromesomma and Moggridgea.
18. Anterior sternal sigilla: (0) present, (1) absent.
Migidae lack the anterior stemal sigilla, which are
present in their outgroups. Goloboff (1993) used this
character (his character 69: "posterior" is a typo
[Goloboff pers. commun.]). Loss of anterior stemal
sigilla is a synapomorphy at node B for the Migidae.
19. Deeply excavate, lunate shaped sternal sigilla:
(0) absent, (1) present.
Pocock (1895) recognized the deeply excavated,
lunate shaped stemal sigilla as diagnostic for Thy-
ropoeus (Figs. 5 IB, 58B). Raven (1985) recognized
the significance of similar sigilla in Heteromigella
malagasa Strand and synonymized Heteromigella with
Thyropoeus. Our analysis corroborates this morpholo-
gy as a synapomorphy at node L for Thyropoeus.
20. Female tarsi I and II spines: (0) present; (1) ab-
sent.
Most migids and their outgroups have spines on
female tarsi 1 and 11 (Figs. 1, 14C). Migas (Fig. 13C),
Poecilomigas (Fig. 18), and Moggridgea (Figs. 15C,
1 6B) lack these spines, a synapomorphy at node N.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
39
21. Female leg I and II shape: (0) rounded; (1)
dorsoventrally flattened.
When viewed from a lateral perspective, the tibi-
ae, metatarsi, and tarsi of legs I and II appear dorsoven-
trally flattened in all Migidae although they may be
less flattened in males. We quantified this character by
measuring the cross section of the tibia I apex at the
tibia-metatarsus joint. Idiops and Actinopus have near-
ly round tibiae I and II, with the ratio of breadth to
height less than 1.1, whereas Bothriocyrtum and all
Migidae have the ratio of breadth to height greater than
1.2. Golobofifand Platnick (1987) suggested that Mal-
lecomigas had these segments more rounded than other
migids, but we find the cross section of Mallecomigas
tibia I to be little different from that of Migas. This
character cannot be polarized with our data set, but
comparison to other mygalomorphs suggests that flat-
tened legs (shared by Bothriocyrtum and the Migoidea)
are derived. We optimize flattened legs as a synapo-
morphy at node B with parallelism in Bothriocyrtum.
22. Patellar ventral lamellate setae: (0) absent, (1)
present.
Recognized by Cambridge (1875) and Hewitt
(1913c), these erect, distally expanded setae (Fig. 61D)
beneath at least patellae I, II and IV were postulated as
a synapomorphy for Moggridgea by Griswold (1987a).
Main (1991) discovered these setae in two new species
from Australia, which she placed in Moggridgea. The
setae occur in both males and females of the African
species but only in females of the Australian M. tingle.
We have found these setae beneath patellae I, II and IV
in a female of the Chilean migid Goloboffia vellardi
Zapfe (Fig. 14C). Although erect setae may occur in a
similar position beneath the patellae oi Migas gatenbyi
these do not have the characteristic lamellate shape.
Our analysis suggests that lamellate setae are a synapo-
morphy at node O for Moggridgea (Figs. 15C, 16B)
with parallel evolution in Goloboffia vellardi.
23. Leg I and II long, fine, curved ventral setae: (0)
absent to weak, (1) dense.
The densely distributed, long, fine, curved ventral
setae that extend beyond the spine tips of legs I and 11
of females (Figs. 27A, 41B) were first recognized by
Pocock (1895) in Paramigas subrufus and mentioned
as a diagnostic character for Paramigas by Raven
(1985). Griswold (1987b) recorded such setae in Poe-
cilomigas abrahami. These setae are very densely dis-
tributed. In Migas gatenbyi more than 1 5 setae can be
counted on a transverse line between the spine rows on
tibia I (Fig. 13C); in Poecilomigas abrahami, and
Paramigas oracle (Figs. 27A, 4 1 B), fl goodmani (Fig.
33) and P. perroti (Fig. 47C) there are more than 20.
Some specimens of Thyropoeus mirandus have long,
curved setae beneath legs I and II: although these are in
longitudinal rows rather than being evenly distributed
we code this state as present. Coding T. mirandus either
way makes no difference to the resulting trees. These
setae do not occur in males. Our analysis suggests that
the dense setal vestiture has arisen in parallel within
Paramigas at node J, in Thyropoeus mirandus, in Poe-
cilomigas abrahami and in Migas. It could be a
synapomorphy at node Q for Migas plus Poecilomigas,
but, given the variability in Poecilomigas and Migas,
we prefer to minimize its importance until Migas is re-
vised.
24. Dorsal setae at apex of tibia I and II: (0) erect,
(1) procumbant and stout.
A conspicuous retrodorsal patch of thickened
procumbant setae is present on the apices of tibiae 1
and II near the tibia-metatarsus joint of Paramigas
(Figs. 27A-C). These setae may be dense (e.g.,
Paramigas perroti, P. pectinatus) or sparse (e.g., P.
pauliani, P. alluaudi). Micromesomma has thorns in
the same position (Figs. 22, 24A-B). In most other
specimens examined the setae in this region are evenly
distributed and erect (Figs. 13C, 14C). Thyropoeus mi-
randus has extensive longitudinal dorsal bands of
procumbant setae on legs I and II (Fig. 57): whereas
there are retrodorsal procumbant setae at the tibia-
metatarsus joints such setae also occur dorsally and ex-
tend to mid-segment. We have not coded the condition
in T mirandus as homologous to that in Micromesom-
ma and Paramigas. These setae do not occur in males.
Our analysis suggests that these setae are a synapo-
morphy at node E for Micromesomma plus Paramigas.
25. Dorsal setae at base of metatarsus I and II: (0)
erect, (1) procumbant and stout.
A conspicuous retrobasal patch of thickened
procumbant setae is present on the bases of metatarsi I
and II near the tibia-metatarsus joint of Paramigas
(Figs. 27A— B). Like the previous character these setae
may be dense or sparse. Micromesomma lack procum-
bant setae in this position (Figs. 24A, B). These setae
do not occur in males. Our analysis suggests that these
setae are synapomorphic at node F for Paramigas.
26. Femur III ventral membrane: (0) short. (1)
elongate.
The length of the ventral membrane of the femur-
patella joint on femur III may be restricted to 1/4 (Fig.
59B) to 1/3 (Figs. 15D, 61C) the length of the femur
(most migids and their outgroups), or extend at least
4/5 the length of the femur (Malagasy migids including
40
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Micromesomma, Paramigas and Thyropoeus) (Figs.
42C, 44D). This character appears to be independent of
the tibia III dorsobasal excavation (see below) found in
Malagasy migids: in other taxa with a tibia III dor-
sobasal excavation (e.g., Moggridgea; Ctenizidae Um-
midia) the membrane is short (Figs. 15D, 61C). This
feature does not occur in males. Our analysis suggests
that this morphology is a synapomorphy at node D for
the Malagasy Migidae.
27. Tibia III dorsobasal excavation: (0) absent, (1)
present.
Most Migidae and their outgroups have tibiae III
that are cylindrical (Figs. 4, 8, 14D, 59A). The pres-
ence of a dorsobasal excavation on tibia III was used
by Raven (1985) to support inclusion oi Moggridgea in
the Paramiginae. The dorsobasal excavation is a de-
pressed, glabrous, usually shiny dark area at the base of
tibia III (Figs. 1, 25A-C). In most Moggridgea (in-
cluding M. tingle from Australia) it is small and shal-
low (Fig. 15D, 61A— B), although in some species it
may be absent (e.g., M. crudeni, Griswold 1987a fig.
18). In Moggridgea breyeri the dorsobasal excavation
is deep and equals nearly one quarter the length of the
segment (Griswold 1987a, fig. 86). In all Malagasy
Migidae the dorsobasal excavation is deep and equals
nearly one third of the segment length. All dorsobasal
excavations, whether shallow or deep, are coded as ho-
mologues. A vestige of the dorsobasal excavation may
or may not occur in males: it is visible, at least as a
darkened area, in the males of Malagasy Paramigas
(Figs. 29, 36). Our analysis suggests that the dor-
sobasal excavation has arisen in parallel in Mog-
gridgea (node O) and the Malagasy Migidae (node D).
When this character was weighted until forcing the
monophyly oi Moggridgea plus the Malagasy Migidae
the resulting trees were 4 steps longer than minimum
length trees.
28. Tibia III anterior surface: (0) convex, (1) with
diagonal ridge.
The anterior surface of tibia III of most migids and
their outgroups is convex (Figs. 14D, 59A, 61 A). The
Malagasy migids (except Thyropoeus malagasus) have
a weak to strong diagonal ridge extending fi^om the
midpoint of the dorsobasal excavation to the middle or
nearly to the apex of the segment (Figs. 25A, 42A,
47D). No vestige of this ridge is found in males. We
optimize this as a synapomorphy at node D for the
Malagasy Migidae with loss in Thyropoeus malagasus.
Alternatives are parallel evolution in Thyropoeus mi-
randus and clade E.
29. Spines on female patella and tibia III: (0) stout,
(1) slender.
The spines on the anterior surfaces of patella and
tibia III of most taxa studied are short and stout, with a
length that is less than 5 fimes base width (Figs. 1, 25).
Heteromigas (Fig. 8), Calathotarsus (Fig. 4), Malle-
comigas (Fig. 12D), and Goloboffia vellardi (Fig. 14D)
resemble one another in having slender spinules with
length greater than 10 times base width. This state op-
timizes as a synapomorphy at node S with parallel evo-
lution in Heteromigas.
30. Dark dorsal and lateral maculations forming
bands on leg tibia: (0) absent, (1) present.
Many migids have dorsal and lateral maculations
on the leg tibiae and metatarsi (e.g.. Fig. 28B). In Poe-
cilomigas these marks are overlain by diffuse dark
bands that completely surround the segment forming
conspicuous basomedian leg bands most prominent on
the tibiae (Fig. 18). This banding was commemorated
by Simon's naming the genus Poecilomigas (Simon
1903) and was proposed as a synapomorphy for this
genus by Griswold (1987b). Our analysis corroborates
this character as a synapomorphy for Poecilomigas at
node R.
3 1 . Preening comb at apex of metatarsus IV: (0)
absent, (1) mixed and/or separate at base (calathotar-
sine type), (2) identical and on common base {Mog-
gridgea type).
We define preening combs as rows or clusters of
apical setae that differ conspicuously from their sur-
roundings and that may be regularly arranged (see also
Raven 1984:381; Griswold 1987a; Goloboff & Plat-
nick 1987). In most migids and their outgroups, setae
at the metatarsal apex do not form combs. Calathotar-
sus, Mallecomigas, Goloboffia vellardi, most Migas
(but not M. affiinis) and most Moggridgea have unam-
biguous preening combs at the apex of metatarsus IV.
The nature of these setae may differ among taxa. In
most Moggridgea the comb setae are similar in length
and thickness and arise juxtaposed from a common
base (Figs. 62C-D). In Calathotarsus (Figs. 62A— B)
and many Migas long setae are widely spaced and may
alternate with short setae. In Goloboffia vellardi and
Mallecomigas (Goloboff & Platnick 1987:3, 9) and
other Migas (Fig. 59C) the setae are uniformly long
and separated at the base by distances greater than their
diameter. Vestiges of these combs occur in males, e.g.,
Calathotarsus males have well developed combs on
metatarsi III and IV, and in Moggridgea males there is
a comb on metatarsus IV. Most migids may be scored
unambiguously as having or not having combs. Het-
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
41
eromigas dovei has 3-5 slender setae regularly
arranged at the retroapex on metatarsus IV whereas
other Heteromigas species have nothing resembling a
comb (Raven 1984, pers. commun.)- We score the
comb as absent in Heteromigas. A comb of widely-
spaced setae optimizes as a synapomorphy at node M
with loss of the comb occurring at node R for Poe-
cilomigas. The Moggridgea type of comb optimizes as
a synapomorphy at node P for the African Moggridgea.
32. Preening comb at apex of metatarsus III: (0)
absent, (1) present.
Calathotarsus (Fig. 4), Mallecomigas, and
Goloboffia vellardi (Fig. 14D) have combs at the apex
of metatarsus III that may be similar to those at the
apex of metatarsus IV. The metatarsus III comb of
Goloboffia vellardi alternates long and short setae, dif-
fering from the comb of uniform setae on metatarsus
rV. This comb is a synapomorphy at node S uniting Ca-
lathotarsus, Mallecomigas and Goloboffia vellardi.
33. Preening comb extent: (0) narrow, (1) extend-
ing around more than one fifth of segment circumfer-
ence.
Whereas the preening combs of Moggridgea and
Migas consist of a few closely-spaced setae (Figs. 59C,
62C— D), the combs of Calathotarsus. Mallecomigas,
and Goloboffia vellardi consist of many setae that ex-
tend part way around the ventral apex of the segment
(Figs. 4, 62A— B). These broad combs are most con-
spicuous when the segment is viewed end-on. Opti-
mization of this character is ambiguous. We prefer the
broad comb as a synapomorphy at node S uniting Ca-
lathotarsus, Mallecomigas and Goloboffia vellardi. Al-
ternatively the narrow comb could be a synapomorphy
at node N uniting Moggridgea, Migas, and Poecilomi-
gas (where it is lost).
34. STC III and FV: (0) claw larger than tooth, (1)
tooth larger than claw.
The superior tarsal claws of legs III and IV may
have one (e.g. Paramigas pauliani) to three teeth (e.g.,
Paramigas macrops) beneath the apex (Figs. 37H,
441). In most taxa the teeth are shorter than the apex of
the claw but in both Thyropoeus malagasus (Fig. 5 1 E)
and T. mirandus (Fig. 58G) the tooth is as long as or
longer than the apex of the claw. This is particularly
pronounced on STC III. In Idiops the tooth of STC III
is also large, though not as large as in Thyropoeus. This
enlarged tooth is a synapomorphy at node L uniting the
two species of Thyropoeus.
35. ITC III size: (0) large, greater than 1/3 length
of STC III, (1) small, less than 1/5 length of STC III.
Goloboff and Platnick (1987) noted that Malle-
comigas resembles the Actinopodidae in having a nor-
mal (rather than reduced) ITC III, implying that the
large inferior tarsal claw is a shared primitive feature.
We quantified ITC III size by comparing its length to
that of STC III. Idiops, Actinopus, Heteromigas dovei,
Mallecomigas schlingeri and Thyropoeus mirandus
have ITC III that are at least 1/3 the length of the STC
and all other taxa studied have reduced ITC. Large ITC
optimize as primitive with reduced ITC appearing in
Bothriocyrtum and at node C within the Migidae. The
large ITC of Mallecomigas and Thyropoeus mirandus
are reversals.
36. Male femur I venter: (0) convex, (1) carinate.
Male femora are typically cylindrical or laterally
flattened (Figs. 7D, IID, 30B). The ventral surface of
femur I (and femur II of many species) of African
Moggridgea is so strongly laterally flattened that it
forms a ventral carina or ridge (Fig. 60C; Griswold
1987a, fig. 13). The carinate femur I optimizes as a
synapomorphy at node P for the African Moggridgea.
37. Male pedipalpal tibia apex form: (0) Ectal lobe
blunt, not or only slightly longer than mesal, ( 1 ) ectal
lobe pointed, much longer than mesal.
The apex of the male pedipalpal tarsus has two
lobes that are of near equal length in most migids and
their outgroups (Figs. 7A-C, llA-C, 32C, 548). The
ectal lobe is much broader than the mesal, and in Mog-
gridgea intermedia, M. peringueyi and M. rupicola (as
well as most other African Moggridgea) this lobe is
pointed and extends far beyond the apex of the mesal
lobe (Figs. 16A, 32D). The morphology of unequal
apical lobes of the pedipalpal tarsus optimizes as a
synapomorphy at node P for the African Moggridgea.
38. Male pedipalp tarsus apical spinules: (0) ab-
sent, (1) present.
One or more elongate spines, conspicuously
stouter than the surrounding setae, may be present at
the apex of the male pedipalpal tarsus in some Migidae
(Poecilomigas [Figs. 20A— B], Paramigas [Figs. 32C,
37E-G], and Goloboffia vellardi) and Idiopidae. Our
analysis suggests that apical spinules evolved four
times: as synapomorphies at node F for Paramigas (the
male of Micromesomma is unknown) and node Q for
Migas plus Poecilomigas, and in Idiops and Goloboffia
vellardi.
39. Male retrolateral tibia I megaspine: (0) absent,
(1) present.
We define a megaspine as a spine that is conspic-
uously larger that others near it (Figs. 7D, 55B).
42
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
African Moggridgea lack enlarged spines at the
retroapex of femur I (Fig. 16B), as do Heteromigas
(Fig. IID) and Goloboffia vellardi. Poecilomigas
abrahami lacks such a spine but Poecilomigas basille-
upi has an enlarged retroapical spine (Griswold 1998a
fig. 7). Similar enlarged spines occur in Moggridgea
tingle. Migas variapalpus, and Calathotarsus (Fig.
7D). A single large apical spine is present on the retro-
lateral surface of tibia I on all Malagasy migids. This is
at the apex in Paramigas macrops (Fig. 37C) and P.
manakambus (Fig. 38C) and arises from a subapical
spur in Paramigas andasibe (Figs. 30B, 3 IE). Idiops
has two and Bothriocyrtum several enlarged retroapical
spines: we have coded these as homologous to the
migid megaspine. Coding these as non-homologous
makes no difference to the resulting tree. If we consid-
er the spines in Idiops and Bothriocyrtum as
megaspines this feature optimizes as plesiomorphic
with megaspines lost in parallel five times: in Actino-
pus, Heteromigas dovei. Goloboffia vellardi, Poe-
cilomigas abrahami, and as a synapomorphy at node P
for the Afi-ican Moggridgea. Alternatively loss of the
megaspine could be a synapomorphy at node A for the
Migoidea with regain at node C within the Migidae.
40. Male metatarsus I shape: (0) cylindrical, (1)
curved, with retrolateral pale swelling.
The males of Paramigas andasibe have metatarsi
I that are curved, with a retrolateral pale swelling (Figs.
29, 30B, 31D-E). Calathotarsus (Figs. 6, 7D) and
Bothriocyrtum have male metatarsi that are strongly
bent retrolaterad at mid-segment; we do not consider
this as homologous to the state discussed here. This
feature is an autapomorphy for Paramigas andasibe.
41. Male tarsus III shape: (0) cylindrical, (I)
curved, sausage shaped, strongly convex ventrally.
Most rastelloid males have tarsi III and IV cylin-
drical (Figs. 6, 10, 36). Males of the Malagasy migids
Paramigas andasibe (Fig. 3 IF) and P. manakambus
(Fig. 38D) ha\e tarsi III and IV weakly curved, slight-
ly swollen, and strongly convex ventrally. We optimize
this as a synapomorphy at node G, predicting its pres-
ence in the undiscovered males of P. alluaudi, P. pecti-
natus. P. pauliani, and P. rothorum.
42. PLS apical segment: (0) triangular, (1) domed.
As noted by Raven (1985), the apical segment of
some mygalomorph spiders is triangular in shape, most
easily observed in the Ctenizidae. All Migidae and
Actinopodidae (the Migoidea, clade A) share a domed
apical segment of the PLS (Figs. 3A— B).
43. PLS spigot distribution: (0) all articles, (1) me-
dian and apical articles only.
Goloboffand Platnick (1987) first recognized the
unique distribution of spigots on the PLS of Chilean
Migidae and suggested it as a potential synapomorphy
for the family. All migid taxa surveyed have their spig-
ots limited to the median and apical segments of the
PLS (Figs. 3A, B), corroborating this condition as a
migid synapomorphy at node B.
44, Distribution of spermathecae sclerotization:
(0) uniform, (1) a median band.
Most migids and their outgroups have the sper-
mathecae evenly sclerotized (Figs. 35A— E, 64A-C).
The degree of sclerotization may vary from light to
heavy. Uniquely, Moggridgea (including M. tingle
from Australia) have a narrow band of sclerotization
across the middle of the spermatheca (Figs. 17A— C).
This sclerotized band may even be visible through the
cuticle. This is a synapomorphy at node O for Mog-
gridgea.
45. Spermatheca shape: (0) short, cylindrical, HS
equal to or slightly wider than stalk, (1) head narrower
than long stalk, (2) long stalk with wider head, (3) head
much wider than short, narrow stalk, (4) clavate.
The spermathecae of Actinopus (Fig. 64B) and
Bothriocyrtum (Fig. 64C) are short, stout and have the
head only slightly wider than the stalk. Similar sper-
mathecae occur in Goloboffia vellardi (Figs. 63E— F)
and Calathotarsus (Fig. 63A). Heteromigas dovei (Fig.
63B) and Mallecomigas schlingeri (Figs. 63C, D) have
spermathecae that are long and have the head narrow-
er than the stalk. A slender stalk and broad head is
characteristic of most migids, i. e., Migas (Fig. 19C),
Moggridgea (Figs. 17A— C), Poecilomigas (Figs.
19A— B) and several Paramigas (Figs 35A— E, 40A— B).
Idiops (Fig. 64A) and the Malagasy migids Paramigas
rothorum (Fig. 45B), P. alluaudi (Fig. 45C), P. pecti-
natus (Fig. 45A), P. pauliani (Figs. 40C-D), and Mi-
cromesomma (Figs. 26A-C) have the head much wider
than short, narrow stalk. The stalk is especially narrow
in Paramigas rothorum. P. alluaudi, P. pectinatus, and
P. pauliani. Thyropoeus mirandus (Figs. 56A— C) and
T. malagasus (Fig. 56D) have clavate spermathecae
that taper gradually from the base to the head: that of T.
mirandus arises from a very narrow base. State
(short, cylindrical) optimizes as plesiomoiphic. State 1
(head narrower than long stalk) arises in parallel in
Heteromigas and Mallecomigas. State 3 (head much
wider than short, nairow stalk) arises in parallel in Id-
iops and as a synapomorphy at node E uniting Parami-
gas and Micromesomma, and state 2 (long stalk with
wider head) is a synapomorphy for clade N (Mog-
gridgea plus the Miginae) and at node I within
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
43
Paramigas. State 2 is predicted to occur in the undis-
covered female of Paramigas macrops.
Characters not included in data matrix
Goloboff and Platnick (1987:7) state that unlike
most migid genera, Heteromigas and Mallecomigas
have short spines on tibiae and metatarsi I and II and
imply that retention of this primitive feature might in-
dicate that either of these genera, or both together,
might represent the sister group of all remaining
migids. In fact Heteromigas and Mallecomigas spines
are not particularly short: the longest spines on tibiae
and metatarsi I and II of these genera have a
Ie^gth^ase width ratio equal to that of many species of
Migas, Moggridgea and Paramigas.
PHYLOGENY OF MIGID AE
Our preferred tree (Fig. 65) contains the maximum
resolution supported by the data. We discuss clades on
this tree with their synapomorphies listed in parenthe-
sis (Character: State).
Clade A (Migoidea) — Synapomorphies are the
wide ocular area, with OAW/ Caput width greater than
0.45 (1:1), diagonal fang orientation (14:1), and domed
PLS apical segment (42:1). A potential synapomorphy,
ambiguously optimized, is the cuspules on the pedipal-
pal coxae that extend broadly onto the anterior surface
(16:1).
Clade B (Migidae) — Synapomorphies are the re-
curved thoracic fovea (6:1), loss of the rastellum
(11:1), quadrate and keeled fang (12:1), loss of anteri-
or sternal sigilla (18:1), flattened female anterior legs
(21:1, ambiguous) and PLS spigots limited to median
and apical articles only (43: 1).
Clade C (all migids other than Heteromigas) —
The reduced ITC III is a synapomorphy (35:1). A po-
tential synapomorphy, ambiguously optimized, is the
cuspules on the pedipalpal coxae that are restricted to
the anterior median comer (16:0).
Clade D (Malagasy migids, Paramiginae sensu
Simon) — Synapomorphies are the reduced caput seta-
tion, with prefoveal setae only or lacking setae alto-
gether posteriad of ocular area (4: 1 ), femur III ventral
membrane that extends at least 4/5 the length of the
femur (26:1), and tibia III with a deep dorsobasal ex-
cavation (27:1). The diagonal ridge on tibia III (28:1)
is an ambiguous synapomorphy.
Clade E {Micromesomma plus Paramigas) —
Synapomorphies are the low female caput, with height
<0.7 X length (5:1), cheliceral basal swellings (9:1),
procumbant and stout dorsal setae at the apices of tibi-
ae I and II (24:1), and spermathecae with the head
much wider than the short, narrow stalk (45:3).
Clade F {Paramigas) — Synapomorphies are the
procumbant and stout dorsal setae at bases of metatar-
si I and II (25:1) and male pedipalp tarsus apical spin-
ules (38:1). The latter feature could instead be a
synapomorphy uniting Micromesomma and Parami-
gas, as the male of Micromesomma is unknown.
Clade G (six species of Paramigas) — The
synapomorphy is male tarsi III and IV that are weakly
curved, slightly swollen, and strongly convex ventrally
(41:1). This has been observed only in Paramigas an-
dasibe (Fig. 3 1 F) and P. manakambus (Fig. 38D) and is
predicted to occur in the species of clade G that are
known only from females.
Clade H (five species of Paramigas) — Loss of
denticles from between the cheliceral teeth is the
synapomorphy (8:0).
Clade I (five species of Paramigas) — Spermath-
ecae with a long stalk with wider head (45:2) is a
synapomorphy uniting these species.
Clade J (four species of Paramigas) — The dense-
ly distributed, long, fine, curved ventral setae that ex-
tend beyond the spine tips of legs I and II of females
are the synapomorphy (23:1).
Clade K (Paramigas macrops plus P. oracle) —
Loss of denticles from between the cheliceral teeth is
the synapomorphy (8:0).
Clade L (Thyropoeus) — Synapomorphies are the
deeply excavate, lunate shaped sternal sigilla (19:1),
ITC III and IV with the tooth longer than the claw
(34:1) and clavate spermathecae that taper gradually
from the base to the head (45:4).
Clade M (Calathotarsus, Goloboffia. Mallecomi-
gas, Migas, Moggridgea and Poecilomigas) — Synapo-
morphies are the simple, recurved fovea lacking poste-
rior extension (7:0), loss of denticles from between the
cheliceral tooth rows (8:0, ambiguous), presence of
male intercheliceral tumescence (10:1) and metatarsus
IV preening comb (31:1 ).
Clade N (Miginae sensu Simon) — Migas. Mog-
gridgea and Poecilomigas are united by the synapo-
morphies low female caput, with height < 0.7 x length
(5:1), basal cheliceral swellings (9:1), loss of spines
from female tarsi I and II (20:1) and by ha\ing the
spermathecae with a long stalk with wider head (45:2).
Clade O (Moggridgea) — Synapomorphies are the
patellar ventral lamellate setae (22:1), tibia III dor-
sobasal excavation (27: 1), and narrow band of scleroti-
zation across the middle of the spennatheca (44:1).
Clade P (African Moggridgea) — Synapomor-
phies are loss of male intercheliceral mmescence
44
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
(10:0), metatarsus IV preening comb setae that are sim-
ilar in length and thickness and arise juxtaposed from a
common base (31:2), carinate male femur I venter
(36:1), male pedipalpal tibia apex with the ectal lobe
pointed, much longer than mesal (37:1) and loss of the
male retrolateral tibia I megaspine (39:0).
Clade Q (Migas plus Poecilomigas) — Synapo-
morphies are the enlarged prefoveal setae (3:2, am-
biguous), basal tooth on the fang (13: 1), and male pedi-
palp tarsus apical spinules (38:1).
Clade R {Poecilomigas) — Synapomorphies are
the dark dorsal and lateral maculations forming annuli
on the leg tibiae (30:1) and loss of the metatarsus IV
preening comb (3 1 :0).
Clade S (South American Migidae) — This newly
proposed clade comprising Calathotarsus, Mallecomi-
gas and Goloboffia vellardi is united by the synapo-
morphies of pedipalpal coxa cuspule distribution that is
concentrated at the proximal edge near the labium
(15:1), slender spinules densely situated on patella-
tibia III of females (29:1), a preening comb at the apex
of metatarsus III (32:1) and preening combs that con-
sist of many setae that extend part way around the ven-
tral apex of the segment (33:1, ambiguous).
Clade T (Calathotarsus plus Goloboffia
vellardi) — Synapomorphies are the extra wide ocular
area, greater than 0.60 x carapace width (2:1) and den-
ticles scattered between the cheliceral tooth rows (8:1,
ambiguous).
CLASSIFICATION
This phylogenetic analysis corroborates the mono-
phyly of Migoidea (Actinopodidae plus Migidae)
(Platnick & Shadab 1976; Raven 1985; GolobofF
1993a) and Migidae. We reject the Calathotarsinae
(sensu Simon 1903) and note that it is paraphyletic
with respect to the Miginae and Paramiginae. On the
other hand, our results corroborate both the Miginae
(sensu Simon 1903) and Paramiginae (Petrunkevitch
1939, sensu Simon 1892, but not sensu Raven 1985).
Simon (1892, 1903) believed that the migids fell into
three groups. His Migeae (Miginae), comprising
Migas. Moggridgea and Poecilomigas, included taxa
that have simple thoracic foveae and third tibiae and
low caputs. His Calathotarsae (Calathotarsinae), com-
prising Calathotarsus and Heteromigas, included taxa
that have caputs raised as in Actinopus. His Myrtaleae
(Paramiginae Petrunkevitch 1939), comprising Mi-
cromesomma, Paramigas, and Thyropoeus. included
taxa that have tripartite foveae and dorsobasal excava-
tions on tibiae III. Raven (1985) reexamined migid
classification and modified the limits of Miginae and
Paramiginae by transferring Moggridgea from the
Miginae to Paramiginae. He noted that many Mog-
gridgea species have a dorsal excavation on tibia III.
He retained the Calathotarsinae although noting
(Raven 1985:57) that they appear to lack a synapomor-
phy. Griswold accepted Raven's relimited Paramiginae
(Griswold 1987a) and Miginae (Griswold 1987b).
GolobofF and Platnick (1987) described Mallecomigas
and the female of Migas vellardi. Migas vellardi was
tentatively placed in the Miginae based on the basal
tooth on the fang and low caput. Goloboff and Platnick
(1987) noted that Mallecomigas retained presumably
plesiomorphic features, e.g., rounded tibiae and
metatarsi I and II (shared with Heteromigas) and a nor-
mal (rather than reduced) ITC III, and suggested that
Mallecomigas, Heteromigas, or both together might
represent the sister group of the remaining migids. Our
conclusions differ from some of those of Raven (1985)
and of Goloboff and Platnick (1987). Although the dor-
sally excavate tibia III is included in the dataset (char-
acter 27) for both Moggridgea and the Paramiginae it
is most parsimonious to place Moggridgea in the Migi-
nae with Migas and Poecilomigas. When character 27
was weighted to force the monophyly of the Paramigi-
nae sensu Raven 1985 (including Moggridgea), result-
ing trees were 1 00 steps, 4 steps longer than the mini-
mum for unweighted data. Our results suggest that the
South American Migas vellardi, Mallecomigas and
Calathotarsus form a clade. Migas vellardi does not
belong in Migas, and we propose the new generic name
Goloboffia for it. Goloboffiia appears to be the sister
genus of Calathotarsus, but to place Migas vellardi in
Calathotarsus would render this genus heterogeneous
in that Migas vellardi has ventral lamellate setae on the
patellae but lacks the characteristic Calathotarsus
synapomorphies of modified female pedipalpal tarsus
bearing a ventral expansion and dorsal group of cus-
pules and bent and apically swollen male metatarsus I.
Mallecomigas is not a relatively primitive migid. We
include the shape of tibiae and metatarsi I and II in the
dataset (character 2 1 ) but we find the cross section of
Mallecomigas tibia I to be flattened and little different
from that of Migas. The large ITC of Mallecomigas
(character 35) appears to be an independent reversal to
the plesiomorphic form. Our results do agree with
those of Goloboff and Platnick (1987) in suggesting
that Heteromigas is a primitive migid: it is probably the
sister group to the rest of the family.
BIOGEOGRAPHY
Due to their strikingly disjunct southern distribu-
tion (Fig. 66) the migids have long drawn the attention
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
45
of biogeographers. Pocock (1903: 352) first offered an
explanation for their distribution suggesting that they
had arisen in the Afro-Mascarene area (i.e., Africa plus
Madagascar) and subsequently migrated to Australia-
New Zealand and later to South America. Legendre
(1979) next commented on their distribution, writing
after the widespread acceptance of continental drift as
an influence on biogeography. Legendre accepted
migid distribution as strong evidence for Gondwana-
land, considering them "couvrent incontestablement
I'ancienne Gondwanie et ont ete souvent invoquees
comme preuve de I'existence du continent de Gond-
wana" (Legendre 1979:45-46). Platnick was the first
to examine the migids in the context of cladistics and
vicariance biogeography (Platnick 1981; Nelson &
Platnick 1984). Lacking a cladistic treatment of the
family, Platnick accepted the then current classification
as the best available phylogenetic hypothesis (Platnick
1981:90, fig. 7) and used the then conventional hy-
pothesis of Gondwanaland breakup (i.e., [[Africa,
Madagascar] [New Zealand [Australia, South Ameri-
ca]]]). Recognizing that changes in migid classification
and geological interpretation might alter the basic data,
he nonetheless found that two components in migid
classification were consistent with vicariance theory
and explained by Gondwanaland breakup (Platnick
1981:89, fig. 6, components 4 [Australia plus South
America] and 5 [New Zealand plus Australia plus
South America]). Since Platnick's work our under-
standing of Gondwanaland breakup has changed some-
what (Smith et al. 1994) and with our new cladogram
several components of migid classification have
changed. Main (1991) described new Mogghdgea
fi-om southern and western Australia, suggested that
the genus may have originated in the Jurassic (140 ma),
and compared the distribution of Moggridgea to pseu-
doscorpions, scarab beetles, and midge flies (see
below). Griswold (1991a) examined migids as part of
the Afromontane biota (Griswold 1991a; White 1978,
1983). Using a parsimony method to combine cladistic
biogeographic information from the spider families
Microstigmatidae (Microstigmata), Migidae (the Mog-
gridgea quercinca group) and Phyxelididae (the Vi-
doleini and Phyxelidini, then considered Amaurobi-
idae), he proposed a cladogram for areas of temperate
and montane forest in Africa and Madagascar (Gris-
wold 1991a, fig. 5) that contained a sister-area rela-
tionship between Madagascar and the mountains of
East Africa and predicted that members of the Mog-
gridgea quercinca group should be discovered in
Madagascar.
Dispersal cannot be ruled out a-priori in the case
of migid disjunctions although mygalomorphs like
migids have long been considered poor in dispersal
abilities and therefore excellent subjects for studies of
historical biogeography. Most mygalomorphs are not
thought to disperse by ballooning (becoming airborne
and floating on silken threads) though spiderlings of
some mygalomorphs exhibit stereotyped ballooning
behavior and float for short distances on silken threads
(Coyle 1983, 1985). Rafting on floating vegetation also
cannot be ruled out for migids. Many migids are tree
dwellers with tightly-sealed trap door nests. As Raven
(1980) pointed out, a gravid female in a sealed trap
door nest on a floating tree trunk might succeed in dis-
persing over water. At least the ancestors of Mog-
gridgea nesiota may have arrived on the oceanic is-
lands that make up the Comoros in this way. To favor
a vicariance explanation for the disjunctions in migid
distribution these patterns must correlate with some
potential vicariance event in earth history and/or be
general, that is, not characteristic of the migids alone.
As shown below, migid disjunctions are both correlat-
ed with events in earth history and similar to distribu-
tion patterns found in other organisms.
Our new cladogram bears upon previous hypothe-
ses and suggests several new insights. The Migidae can
reasonably be considered a Gondwanan family, occur-
ring in several parts of that former southern supercon-
tinent (Africa, Australia, Madagascar, New Caledonia,
New Zealand, and South America) and nowhere else
(Fig. 66). An area cladogram (Fig. 67) derived from
our taxon cladogram (Fig. 65) suggests several inter-
continental relationships. Australia appears three times
on the cladogram: related to Africa {Moggridgea), re-
lated to New Zealand and New Caledonia together and
to eastern and southern Africa (clade Q: Migas plus
Poecilomigas). and related to the areas Madagascar,
South America, Africa and Australia, New Zealand
and New Caledonia together (this relationship speci-
fied by Heteromigas). The complex area relationships
shown by Australia suggest that this continent might be
subdivided for biogeographic studies. For example,
Moggridgea suggests a relationship of western and
southern Australia to Africa, the Miginae suggest a re-
lationship of eastern Australia to Africa, New Zealand
and New Caledonia, and Heteromigas suggests a rela-
tionship of eastern Australia to all the other parts of
Gondwanaland occupied by migids. Africa appears
twice, once related to Australia {Moggridgea) and once
to Australia, New Zealand and New Caledonia (clade
Q: Poecilomigas plus Migas). South America (repre-
sented by clade S, comprising Calathotarsus, Malle-
comigas and Goloboffia vellardi) is related to Africa,
46
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Australia, New Zealand and New Caledonia together
(represented by clade N, comprising Moggridgea,
Migas and Poecilomigas). Madagascar arises near the
base of the cladogram (clade D: Paramiginae) and is
the sister area of all other Gondwanan components in-
habited by Migidae, i.e. there is a fundamental split be-
tween Madagascar and the remainder of Gondwana-
land. This is compatible with a recent, widely accepted
hypothesis of Gondwanaland breakup. Smith, Smith
and Funnell (1994) map a water gap between the east
coast of Africa and west coast of Madagascar that may
date back to the Sinemurian (200 ma); these areas re-
mained in contact via Antarctica to the south. Mada-
gascar (plus India) had separated from the remainder of
Gondwanaland by the Aptian (120 ma). Other Gond-
wanan fragments remained in contact at this time. The
fiindamental division between the Malagasy Paramigi-
nae and migids of the rest of Gondwanaland suggests
that the family existed at least 120 million years ago
and that the ancestors of Paramiginae may have been
stranded on the Madagascar-India plate during the Apt-
ian. The fact that Heteromigas (from eastern Australia)
is plesiomorphic to even this potentially ancient split
suggests that the Migidae may have geographically dif-
ferentiated before the breakup of Gondwanaland.
The distribution of the remaining migids cannot be
simply explained by continental drift and Gondwana-
land breakup. The simplified hypothesis of Gondwana-
land fragmentation suggested by modem works (e.g.,
Smith et al. 1994) is ((Madagascar, India) (Africa
(New Zealand (South America, Australia)))). This is of
course an oversimplification. Gondwanaland was not
homogeneous, nor was or is any of its fragments. Nu-
merous tropical west Africa-tropical South America
disjunctions on the one hand and Australia-New
Zealand-Chile disjunctions on the other suggest the
fragmentation of tropical and temperate biotas that
were already distinct before the breakup of Gondwana-
land. Migidae exhibit at least two cases of trans-Indian
Ocean disjunction: within Moggridgea (Africa-Aus-
tralia) and clade Q (Poecilomigas [Africa] and Migas
[Australia, New Caledonia and New Zealand]). Trans-
Indian Ocean disjunctions occur in several other
arthropod groups. The millipede families Harpago-
phoridae and Sphaerotheriidae occur in Africa, Mada-
gascar, south Asia and Australia (Kraus 1978). The
millipede distributions include taxa in Madagascar and
south Asia, a situation not known in the trans-Indian
Ocean Migidae. Trans-Indian Ocean disjunctions (for
taxa that do not also occur in South America) have
been called "East gondwanwan" by Kraus (1978). This
conception of "East Gondwana" differs from others
(e.g., Truswell 1977; Pielou 1979) in which East Gond-
wana included India, Antarctica, Australia, New
Zealand and New Caledonia, but excluded Madagas-
car, which was considered part of "West Gondwana-
land" along with Africa and South America. The spider
family Cyatholipidae describes another trans-Indian
Ocean disjunction, occurring in Africa, Madagascar,
Australia and New Zealand but probably absent from
South America (Griswold 2001). Harvey (1996a,
1996b) records several trans-Indian Ocean disjunctions
in pseudoscorpions that he attributes to fragmentation
of former ranges that encompassed parts of Gond-
wanaland. Cranston, Edward and Colless (1987) record
an Africa-Australia disjunction in the chironomid
midge fly genus Archaeochlus, and Mathews (1976)
and Howden (1981) report Africa- Australia disjunc-
tions in various groups of scarab beetles. These dis-
junct distributions may result from vicariance of a
biota that was already differentiated in Gondwanaland.
Given the Aptian age (120 ma) for Migidae implied by
a basal split in the family it is likely that continental
drift has influenced the distribution of the non-
Paramiginae, probably acting on formerly restricted
distributions and shaped by extinctions. The solution to
migid biogeography requires a better understanding of
the environmental history of Gondwanaland as well as
a more complete picture of migid phylogeny and dis-
tribution.
Our study refutes the prediction of Griswold
(1991a) that Madagascar should be home to close rela-
tives of migids from the East African mountains. No
members oi Moggridgea have been found in Madagas-
car, although the species Moggridgea nesiota lives on
the nearby Comoro islands (Griswold 1987a). The East
Africa-Madagascar sister area relationship proposed by
Griswold (1991a) does appear to be a general pattern
for spiders, though. It has been replicated by new data
for two groups of cyatholipid spiders (Griswold 2000,
2001 ) and a pair of zorocratid spider genera (Griswold
1993, 2000), and parallels vicariance patterns suggest-
ed for lemurs (Yoder et al. 1996) and tenrecs (Asher
1997).
What can be said of the distribution of migids
within Madagascar? Unfortunately the sampling is un-
even and the cladogram is poorly resolved. Indeed, the
consensus of equally short cladograms (but not neces-
sarily equally preferable: see discussion above under
"Preferred tree") offers no resolution among Madagas-
car migids. On the preferred cladogram (Fig. 65) some
distributions of interest do emerge. With the exception
of Paraniigas goodmani from the southwestern part of
the country, all clade J species (Paramigas perroti, P.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
47
macrops and P. oracle) occur in the central-eastern part
of the country. The sister group of clade G, Paramigas
milloti, occurs in the north. Clade H comprises one
species from the eastern escarpment {Paramigas man-
akambus), three species from the south or southwest of
the country (Paramigas alluaudi, P. pauliani, and P.
pectinatus) and one {Paramigas rothorum) from the far
north. It is unfortunate that relationships cannot be fur-
ther resolved among the members of this eastern,
northern and western clade. A special relationship be-
tween northern and southwestern species has been sug-
gested by Raxworthy and Nussbaum (1997, fig. 5.5),
who performed a parsimony analysis of endemicity
(PAE) on the distributions of 200 Malagasy reptile
species. Unfortunately, understanding the biogeogra-
phy of migids within Madagascar will require far
denser sampling than has been done to date.
CONCLUSIONS
Migid trap door spiders are an ancient group
whose complex distribution has been influenced by
continental drift but defies simple explanation. They
may have been geographically differentiated even be-
fore the breakup of Gondwanaland. It appears that the
Paramiginae evolved from ancestors stranded by conti-
nental drift on Madagascar, probably while this was
still connected to India. This suggests that we need to
look for Migidae in south Asia, particularly Sri Lanka
and southern India. No migids have been found in
these areas, but their highly cryptic habits may have led
to them being overlooked. Much remains to be learned,
especially about the Malagasy fauna. We need to asso-
ciate more males and females, observe the biology of
Thyropoeus and Micromesomma, and document the
distribution of Micromesomma. The monograph and
phylogeny presented herein is only a preliminary look
at this ancient and fascinating group of Malagasy spi-
ders.
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FIGURES
1-68
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
53
5.0mm
Figure 1. Paramigas milloti, new species, holotype female, dorsal. Illustration by JS.
54
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 2. Paratmgas spp., juveniles. A. B. Resene Speciale Ivohibe. Madagascar. C. D. Ambanizana. Masoala, Madagas-
car, A. Cephalothora.x. dorsal. B. Cephalothorax, anterior. C. Ape.x of sternum, labium, and mouthparts, ventral. D. Ocular
area, dorsal. Scale bars; A = 1000 nm, B = 430 ^m, C = 500 (im. D =250 urn.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
55
Figure 3. Morphology of Paramigas. A, B. Juvenile from Ambanizana, Masoala, Madagascar C, D. Paramigas oracle, new
species, female from Talatakely. Madagascar A. Spinnerets, ventral. B. Left spinnerets, ventral. C. Pedipalpal tarsus claw. D.
Tarsus I claws. Scale bars: A = 250 \xm. B = 120 (xin, C = 100 |am, D = 200 |.un.
56
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
3.0mm
Figure 4. Calalbolarsus coronatiis. female from Cerro La Campana. Chile, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
57
1.0mm
2.0mm
0.5mm
0.3mm
Figure 5. Calathotarsus female morphology. A, B. Calathotarsiis sinioni from Cerro Negro. Argentina. C-E. Calathotarsiis
coronatus from Cerro La Campana. Chile. A. Cephalothora.x. lateral. B. Cephalothorax, ventral. C. Right pedipalp, prolater-
al. D. Dentition of right chelicera. E. Right tarsal claws: pedipalp, lea 1 retrolateral, leg II retrolateral, les III prolateral. lea
IV prolateral. Illustrations A-C by JS, D by JL, E by CG. "
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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
3.0mm
Figure 6. Calatholarsus coronatiis, male from Cerro La Campana, Chile, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
59
O.Smm
B
• ••
0.3mm
O.Smm
Figure 7. Calathotarsus coronaius, male from Cerro La Campana, Chile. A-C. Left pedipalp tibia and tarsus. A. Prolateral.
B. Ventral. C. Retrolateral. D. Leg I, retrolateral. E. Dentition of right chelicera. F. Left tarsal claws: leg 1 retrolateral. leg II
retrolateral, leg III prolateral, leg IV prolateral. Illustrations A-D by JS, E by JL, F by CG.
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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
3.0mm
Figure 8. Heteromi^as dovei. female from from Paiersonia. Tasmania, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
61
3.0mm
00 ••
B
3.0mm
0.5mm
0.5mm
"yr^
>^
^
Figure 9. Hetewmigas dovei, female from from Patersonia, Tasmania. A. Cephalothorax. lateral. B. Cephalothorax, ventral.
C. Dentition of right chelicera. D. Left tarsal claws: pedipalp, leg I retrolateral, leg II retrolateral, leg 111 prolateral. leg IV
prolateral. lllustra'tions A-B by JS, C by JL, D by CG.
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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 10. Hetemmigas dovei, male from from Patersonia, Tasmania, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
63
0.3mm
'a
'^
\^
'7=^
0.3mm
Figure U. Hetewmigas dovei, male from from Patersonia, Tasmania. A-C. Left pedipalp tarsus. A. Retrolateral. B. Ventral.
C. Prolateral. D. Left leg L retrolateral. E. Dentition of right chelicera. F. Left tarsal claws: leg I retrolateral, leg II retrolat-
eral, leg III prolateral, leg IV prolateral. Illustrations A-D by JS, E by JL, F by CG.
64
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 12. Mallecomigas schlingeri, holotype female. A. Cephalothorax, dorsal. B. Cephalothorax. ventral. C. Left leg I,
retrolateral. D. Right leg III. prodorsal. E. Dentition of right chelicera. F. Tarsal claws: right pedipaip, left leg I retrolateral,
left leg II retrolateral, right leg 111 prolaterai, right leg IV prolateral. Illustrations A-D by JS, E by JL, F by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
65
Figure 13. Migas gatenbyi, female from Wellington, New Zealand. A. Cephalothorax. lateral. B. Cephalothorax, dorsal. C.
Right leg 1, retrolateral. D. Dentition of right chelicera. E. Left tarsal claws: pedipalp, leg I retrolateral, leg II retrolateral, leg
III prolateral, leg IV prolateral. Illustrations A-C by VK, D by JL, E by CG.
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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
0.2mm
Figure 14. Goloboffia vellardi, female from Guanaqueros. Chile. A. Cephalothora.x. dorsal. B. Cephalothorax, ventral. C.
Right leg I. retrolateral. D. Left leg III, prolateral. E. Dentition of right cheiicera. F. Left tarsal claws: pedipalp, leg I retro-
lateral, leg II retrolateral. leg III retrolateral. leg IV prolateral. Illustrations A-D by JS. E by JL, F by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
67
Figure 15. Moggridgea intermedia, female from Diepwalle. South Africa. A. Cephalothorax, lateral. B. Cephalothorax, dor-
sal. C. Right leg 1, retrolateral. D. Right leg 111, prolateral. E. Dentition of right chelicera. F. Left tarsal claws: pedipalp, leg
I retrolateral, leg II retrolateral, leg III prolateral, leg IV prolateral. Illustrations A-D by VK. E by JL, F by CG.
68
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
B
Figure 16. Moggridgea peringiieyi, male from Oudtshoom, South Africa. A. Left pedipalp tibia and tarsus, retrolateral. B.
Right leg I, retrolateral. Illustrations by VK.
Figure 17. Spermalhecae of Moggridgea spp., dorsal. A, B. M. liir^ , ^.■ (WAM 90/1 112). Valle\ ol ihc (jiants, Aus-
tralia. C. M. intermedia. Diepwalle. South .'\frica. Arrows to bands ot bclcrotization. Scale bars for A, C = U.2 mm, for B =
O.I mm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
69
Figure 18. Poecilomigas basilleiipi, female from Mazumbai, Tanzania, dorsal. Illustration by JS.
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Figure 19. Spermathecae of Migidae. dorsal. A. Poecilomigas basilleupi, Mazunibai, Tanzania. B. Poecilomigas abrahami,
Sordwana Bay, South Africa. C. Migas gateiibvi, Wellington, New Zealand. HS - speimathecai head, SS - spennathecal stalk.
Scale bars: = 0.2 mm.
1.0mm
Figure 20. Poecilomigas basilleupi. male from Mazumbai, Tanzania. A, B. Left pedipalp patella-tarsus. A. Ventral. B. Retro-
lateral. C. Dentition of right chelicera. Illustrations A, B by JS. C by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
71
Figure 2 1 . Nests of Paramigas. A, B. i^ perroti (lectotype oiP. subrufus). C, D. P. oracle, new species, holotype. A, C. Outer.
B. D. Lateral. Scale bars: = 1 cm.
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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 22. Micmmesomma cowani, female from MRAC, dorsal. Illustration by JS.
5.0mm
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
73
• ••
• •
c JL
0.3mm
B
O^mm
Figure 23. Micromesomma cowani. A-C, E. Female from MRAC. D. Female from MNHN. F. Lectotype female from
BMNH. A. Lateral. B. Ventral. C. Dentition of right chelicera of two individuals. D, E. Spermathecae, dorsal. F. Tarsal claws;
pedipalp, left lea I retrolateral, left leg II retrolaterai, left leg III prolateral, right leg IV prolateral. Illustrations A, B by JS,
C-EbyJL. FbyCG.
74
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 24. Micwmesomma cowani. female from MNHN. A. Leg I. retrolateral. B. Tibia-metatarsus I junction, dorsal, show-
ing thorns. C. Tarsus IV trichobothrial base. D. Tarsal organ IV. E. Right chelicera. Arrow to intercheliceral basal swelling.
Scale bars: A = 1000 ).im. B = 231 |im, C. D = 38 urn. E = 600 nm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
75
Figure 25. A, B, D. Microinesomma cowani, female from MNHN. C. Paramigas oracle, female from Talatakely. Madagas-
car. A-C. Tibia III. A. Prolateral. B. Retrolateral. C. Dorsal. D. Coxa III showing thorns. Arrow to prolateral ridge of tibia
III. Scale bars: A, B = 750 [im, C = 600 |im, D = 300 )im.
76
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
^HHI
IP
v*e'
-«*r
ase^
»?
"^_^^^^
^^.-.^M^MW^&^rili^u
■-.^
^
1^
i
^^^
SIK^W
mi
Ea?
Figure 26. Spemiathecae oi Micwmesomma cowanL dorsal. A. C. MNHN females. B. MRAC female. Scale bars: = 0.2 mm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
77
Figure 27. Pammigas oracle, new species, female from Falatakely, Madagascar. A. Fibia-metatarsus L retrolateral. B. Retro-
lateral view of tibia-metatarsus 1 junction showing serrate, procumbant setae. C. Dorsal view of seiTate, procumbant setae at
apex of tibia I. D. Farsus IH, dorsal, showing trichobothrial distribution and tarsal organ (an'ow). Scale bars: A = 1000 \xm..
B = 430 urn, C = 75 nm. D = 270 iim.
78
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
1.0mm
B
0.1mm
1.0mm
C
0.2mm
0.2mm
Figure 28. Paramigas alluaiidi. female holoUpe of Myrlle albiaiidi. A. Cephalothora.x. dorsal. B. Right leg 1. retrolateral. C.
Spermathecae. dorsal. D. Dentition of right chelicera. E. Left tarsal claws: pedipalp. leg 1 retrolateral, leg II retrolateral, leg
111 prolateral, leg IV prolateral. Illustrations A, B by JS, C, D by JL, F by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
79
2.0mm
Figure 29. Paramigas andasibe Raven, new species, male from Ambohitantely, Madagascar, dorsal. Illustration by JS.
80
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 30. Paramigas andasibe Raven, new species, male from Ambohitantely, Madagascar. A. Cephalothorax, ventral. B.
Right leg I, retrolateral. C. Dentition of right chelicera. D-F. Left pedipalp tibia-tarsus. D. Prolateral. E. Ventral. F. Retrolat-
eral. G. Left tarsal claws: lea I retrolateral, leg 11 retrolateral, leg 111 prolateral, lea IV prolateral. Illustrations A, B, D-F by
JS, C by JL, G by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
81
■5^\|' I
/'
0.5mm
H
0.5mm
3.0mm
Figure 31 Paramigas cmdasibe Raven, new species, holotype male. A-C. Left pedipalp patella-tarsus. A. Prolateral. B. Ven-
tral. C. Retrolateral. D. Right patella-tarsus I, dorsal. E. Right patella-tarsus I, retrolateral. R Left tibia-tarsus 111, retrolater-
al. G. Dentition of right chelicera. H. Tarsal claws: right leg 1, retrolateral, left leg II retrolateral, left leg 111 and right IV, pro-
lateral. Illustrations A-F by JS, G by JL, H by CG.
82
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 32. Pedipalpi ol'male Migidae. A-C. Paramigas andasibe Raven, new species, from Manakambahiny, Madagascar,
right pedipalp. A. Patella-tarsus, retroiateral. B. Patella-tarsus, prolateral. C. Tarsus, dorsal. D. Moggridgea pseiidocrudeni
from .Alicedale. South .Africa, apex of pedipalpal tarsus. Arrows to ectal lobe of tarsus. Scale bars: A. B = 1000 |im, C, D =
380 |jm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
83
5.0mm
Figure 33. Paramigas goodmani, new species, holotype female, dorsal. Illustration by JS.
84
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
I* ••••"
^
Figure 34. Paramigas goodmani, new species, holotype female. A. Lateral. B. Ventral. C. Spermathecae, dorsal. D. Denti-
tion of right chelicera. E. Left tarsal claws: pedipalp, leg 1 retrolateral, leg II retrolateral, leg 111 prolateral, leg IV prolateral.
Illustrations A, B by JS, C, D by JL, E by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
85
Figure 35. Spermathecae of Paramigas spp., dorsal. A. P. peiroti (large female, MRAC). B. P. perroli (small female,
MRAC). C. P. peiToti (syntype of Paramigas siihriifns). D. P. goodmani, holotype. E. P. milloti, holotype. HS - sperraathe-
cal head. SS - spemiathecal stalk. Scale bars; = 0.4 mm.
86
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
2.0mm
Figure 36. Paramigas macrops, new species, holotype male, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
87
0.5mm
H
<^
^v_
'O-^'^W
0.3mm
Figure 37. Pwaniigas macrops, new species, holotype male. A. Lateral. B. Cephalothora.x, ventral. C. Right leg 1, retrolat-
eral. D. Dentition of right chelicera. E-G. Left pedipalp patella-tarsus. E. Prolateral. F. Ventral. G. Retrolateral. H. Left tarsal
claws: leg I retrolateral, leg II retrolateral, leg III prolateral, leg IV prolateral. Illustrations A-C, E-G by JS, D by JL. H bv
CG.
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
2.0mm
0.3mm
0.3mm
Figure 38. Paramigas manakambus. new species, holotype male. A. Cephalothorax, dorsal. B. Dentition of right chelicera.
C. Right leg 1. relrolateral. D. Right leg IV. prolateral. E-G. Left pedipalp patella-tarsus. E. Prolateral. F. Ventral. G. Retro-
lateral. H. Left tarsal claw s: lea I retrolateral, lea II retrolateral. lea HI prolateral, leg IV prolateral. Illustrations A, C-G by
JS. B bv .IL. H bv CG. ^
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
89
Figure 39. Paramigas milloti, new species, holotype female. A. Lateral. B. Ventral. C. Spemiathecae, dorsal. D. Dentition
of right chelicera. E. Left tarsal claws: pedipalp, leg I retrolateral, leg II retrolateral, leg III prolateral, leg IV prolateral. Il-
lustrations A, B by JS, C, D by JL, E by CG.
90
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 40. Spermathecae of Paraniigas spp.. dorsal. A. P. oracle, Talatakely, Madagascar. B. P. oracle, holotype (SS fore-
shortened in this view). C. P. pauliani (small paratype of Legendrella pauliani). D. P. paidiani (large paratype of Legendrel-
la paultani). HS - spermathecal head, SS - spermathecal stalk. Scale bars; = 0.3 mm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
91
0.3mm
^'
</
-^
0.3mm
Figure 41. Paramigas oracle, new species, female. A-C, F, G holotype. D, E. Talatakeley. Madagascar. A. Cephalothorax,
dorsal. B. Right leg I, retrolateral. C, D. Spennathecae, dorsal. E, F. Dentition of right chelicera. G. Left tarsal claws: pedi-
palp, leg 1 retrolateral, leg II retrolateral, leg 111 prolateral, leg IV prolateral. Illustrations A, B by JS, C-F by JL, G by CG.
92
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 42. Paramigas oracle, new species, female from Talatakely, Madagascar. A. Tibia III, prolateral. B. Junction of patel-
la-tibia III. C. Venter of femur III showing membrane. D. Tarsal organ and trichobothrium on tarsus III. Arrow to prolateral
ridge of tibia 111. Scale bars: A = 600 nm.^B = 250 urn, C = 750 \xm, D = 38 urn.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
93
p
m
i
■
If
■^
■M
b':':' Jfl|^fl
m
1
■■■■■
^^9
Figure 43. Paramigas oracle, new species, female from Talatakely, Madagascar. A. Fang. B. Dentition of fang furrow (arrow
to basal swelling on chelicera). C. Right pedipalpal coxa. D. Cuspules on right pedipalpal coxa. Scale bars: A= 500 |jm. B
= 430 urn, C = 600 ixm, D = 120 urn."
94
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
• :
E • • :\.
— •
O.Smro _
^-
Figure 44. Paramigas pauliani. female. A-E. Holotvpe of Legendrella pauliani. F-I. Paratypes oi Legendrella pauliani. A.
Cephalothora.\, dorsal. B. Cephalothora.\. ventral. C. Right leg I. retrolateral. D. Left leg 111. prolateral. E, F. Dentition of
riaht chelicera. G, H. Spermathecae. dorsal. I. Tarsal claws: pedipalp, left lee 1 retrolateral, right leg II retrolateral, right leg
IlT prolateral, left leg IV prolateral. Illustrations A-D by JS, E-H by JL, 1 by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
95
fl^^sLx.
Figure 45. Spermathecae of Paramigas spp., dorsal. A. P. pectinatiis. holotype (narrow stalks of spemiathecae partially hid-
den). B. P. rothorwn, holotype. C. P. alluaudi (holotype of Myrtale alluaudi). Scale bars: = 0.2 mm.
96
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
^^^^^«
1.0mm
1.0mm
D C
0.3mm
0.2mm
Figure 46. Pammigas pectinaws. new species, holotype female. A. Cephalothora.x. dorsal. B. Right leg I, retrolateral. C.
Spermathecae, dorsal. D. Dentition of right chelicera. E. Left tarsal claws: pedipalp, leg 1 retrolateral, leg II retrolateral, leg
III prolateral, leg IV prolateral. Illustrations A. B by JS. C, D by JL. E by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
97
Figure 47. Paramigas perroti, female. A-E, I. Lectotype of Myrtale permti. H. Paralectotype of Paramigas subnifus. F, G.
MRAC specimens. A. Cephalothorax, dorsal. B. Cephalothorax, ventral. C. Right leg I, retrolateral. D. Left leg III, prolater-
al. E, F. Dentition of right chelicera. G, H. Spermathecae, dorsal. 1. Left tarsal claws; pedipalp, leg I retrolateral, leg II retro-
lateral, leg III prolateral, leg IV prolateral. Illustrations A-D by JS, E-H by JL, 1 by CG.
98
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
2.0mm
Figure 48. Paramigas roihonim, new species, holotype female, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
99
Figure 49. Paramigas rothonim, new species, holotype female. A. Lateral. B. Ventral. C. Spermathecae. dorsal. D. Denti-
tion of right chelicera. E. Left tarsal claws: pedipalp, leg 1 retrolateral, leg II retrolateral, leg III prolateral, leg IV prolateral.
Illustrations A, B by JS, C. D by JL. E by CG.
100
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
2.0mm
Figure 50. Thympoeus malagasus, female from Vohimena, Madagascar, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
101
2.0mm
0.2mm
B
2.0mm
0.5mm
0.3mm
Figure 51. Thyropoeus malagasiis. A-D. Female from Vohimena, Madagascar. E. Holotype female of Heteivmigella mala-
gasa. A. Lateral. B. ventral. C. Spermathecae, dorsal. D. Dentition of right chelicera. E. Left tarsal claws: pedipalp, leg I
retrolateral, leg II retrolateral, leg III prolateral, leg IV prolateral. Illustrations A, B by JS, C, D by JL, E by CG.
102
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
^ 2.0mm
Figure 52. Thyropoeus malagasus, male from Vohimena, Madagascar, dorsal. Illustration by JS.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
103
2.0mm
2.0mm
D {
0.5mm
H
c
0.3mm
0.5mm
S4
w
Figure 53. Thyropoeus malagasus, male from Vohimena, Madagascar. A, Lateral. B. Cephalothorax. ventral. C. Left leg \,
retrolateral. D. Dentition of right chelicera. E-G. Left pedipalp patella-tarsus. E. Retrolateral. F. Ventral. G. Prolateral. H.
Left tarsal claws: leg I retrolateral, leg II retrolateral, leg III prolateral, leg IV prolateral. Illustrations A-C, E-G by .IS, D bv
JL, H by CG.
104
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 54. Thyropoens malagasus, male from Vohimena, Madagascar, right pedipalpus. A. Tibia-tarsus, prolateral. B. Tar-
sus, dorsal. C. Tarsus, retrolateral. Scale bars: A = 1000 urn, B = 380 |im, C = 600 |im.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
105
Figure 55. Thyropoeus malagasus, male from Vohimena, Madagascar. A. Right tibia I. dorsal. B. Apex of right tibia I. retro-
lateral. C. Trichobothrial base, metatarsus I. D. Tarsal organ 1. Scale bars: A = 1200 )im, B = 500 |.im, C, D = 30 |im.
106
OCCASIONAL PAPERS OF THE CALIFORNIA AC ADEMY OF SCIENCES, NO. 151
^
HI
/j^B
'
<v
/^^^MJH^I
i ^
a
L.
' ^yrniijMBl^HfflB
/'":^Si-^.-'^
- ;■;- .^-^jU-^
■<^-:^
i^^^S
Figure 56. Spermathecae of Thyiopoeus spp., dorsal. A. T. mirandus, small female, Ft. Dauphin, Madagascar. B. T. miran-
diis, holotype, left. C. T. mirandus, large female. Ft. Dauphin, Madagascar. D. T. malagasus, Vohemena, Madagascar. Scale
bars: A-C = 0.4 mm, D = 0.2 mm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
107
10.0mm
Figure 57. Thyropoeus mirandus. female holotype, dorsal. Illustration by JS.
108
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
0.5mm
Figure 58. Thympoeus mirandus. female. A-C, E. Holotype female. D, F. G. Ft. Dauphin, Madagascar. A. Lateral. B.
Cephalothorax, ventral. C. Left spermatheca, dorsal. D. Spermathecae, dorsal. E, F. Dentition of right chelicera. G. Left tarsal
claws: pedipalp, leg 1 retrolateral, leg 11 retrolateral, leg III prolateral, lea IV prolateral. Illustrations A, B by JS, C-F by JL,
G by CG.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
109
Figure 59. Moqshology of Miginae. A-C. Migas gatenbyi, female from Wellington, New Zealand. D. Poecilomigas abm-
hami, female from Grahamstown, South Africa. A. Patella-tibia 111, prolateral. B. Leg 111 ventral, showing membrane, arrow
to proximal extent. C. Metatarsus-tarsus IV junction, showing widely spaced cornb setae (arrows). D. Fang base showina
tooth. Scale bars: A = 600 \i.m, B = 750 |im, C = 200 |im, D = 1 90 |im.
110
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 60 Morphology of male Migidae. A. Poecilomigas abrahami, from Town Bush, Pietermaritzburg, South Africa,
metatarsus-tarsus IV showing ventral scopula. B. Migas sp., from Tauranga. New Zealand, metatarsus-tarsus IV showing lack
of ventral scopula. C. Moggiidgea pseudocrudeni. from Alicedale, South Africa, femur 1 ventral showing carina. Scale bars:
= 500 nm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
111
1 '
'''1' ^ '' 4 /f .
^ J/i -^^
*
'■*/#/'' ■'•
"'<ji'' 11 //
i
dMi..^
Figure 61. Moggridgea intermedia, female from Diepwalle. South Africa. A. Patella-tibia 111, prolaterai. B. Patella-tibia III
junction. C. Leg III ventral showina membrane (arrow to pro.ximal e.xtent of membrane). D. Lamellate setae beneath patella
IV. Scale bars: A = 600 \im, B = 250 nm, C = 1000 |im, D = 150 |jm.
112
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 62. Metatarsus IV combs of Migidae. A. B. Calalhotarsus coronatiis, female from El Canelo. Chile, metatarsus IV
apical comh. retrox entral. C. Moggridgea cntdem, female from Alicedaie. South .Africa, metatarsus IV comb. D. Moggridgea
dveri, female from Uitenhage. South Afiica, metatarsus IV preening comb base. Scale bars: A = 200 \im, B = 100 \im, C =
200 nm, D = 25 ^m.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
113
Figure 63. Spermathecae of Migidae, dorsal. A. Calathotarsus simoni. Sierra de la Ventana. Argentina. B. Heteromigas
dovei, Patersonia, Tasmania. C, D. Mallecomigas schiingeri, hoiotype. D. Close up of left spermatheca. E. F. Goloboffia vel-
lardi, Guanaqueros, Chile. F. Close up of left spermatheca. HS - spermathecal head. SS - spennathecal stalk. Scale bars A,
B, D, E = 0.2 mm, C = 0.4 mm, F = 0.1 mm.
114
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Figure 64. Spermathecae of migid outgroups, dorsal. A. Idiops sp., Pietermaritzburg, South Africa. B. Actinopus sp., Lo-
malinda, Colombia. C. Bothhocyrtum califomicum. Eagle Rock, California, USA. Scale bars A, B = 0.2 mm, C = 0.4 mm.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
R-O
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116 OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
____ / ' tX fc
A Calathotarsus
• Heteromigas
n Mallecomigas
■ Micromesomma
O Migas
OGoloboffia
A Moggridgea
▼- Paramigas
•♦ Poecilomigas
V Thyropoeus
MIGIDAE
:^
Figure 66. Map showing world distribution of migid genera.
^^
<g^
<^ c#
<**
J>'
#* .^ ^*
/ /
/ / /
/ /
Figure 67. Area cladogram implied by preferred migid cladogram. E = eastern. NC = New Caledonia, NZ = New Zealand,
S = southern, W = western.
GRISWOLD AND LEDFORD: MADAGASCAR MIGIDS
117
A Thyropoeus malagasus
B Thyropoeus mirandus
C Paramigas alluaudi
D Paramigas andasibe
E Paramigas goodmani
F Paramigas macrops
G Paramigas manakambus
H Paramigas milloti
BCJ
I Paramigas oracle
J Paramigas pauliani
K Paramigas pectinatus
L Paramigas perroti
M Paramigas rothorum
• juvenile Paramigas
?Micromesomma cowani
Figure 68. Map of Madagascar showing records of Migidae. Locality data for juvenile Paramigas are listed in Table
APPENDIX
Character by taxon matrix. Rows represent characters. The first state Hsted is coded as "0", the second as "1", etc.,
"?" = unl<nown, "-" = non-applicable. Columns represent laxa. Character statistics are for our preferred tree (Fig 65.
which has length = 96 steps, consistency index = 0.52, retention index = 0.76, total fit [from PeeWee, concavity =3] =
359.9). The final five columns give the number of steps (St), the consistency index (CI), the retention index (RI). the
weight from successive weighting from Hennig86 (Wt) and the fit from Pee- Wee. Taxon abbreviations are (veaically):
Id = Idiops, Ac = Aclinopus, Bo = Bothriocyrium, Co= Calalholarsus spp.. Ma = Mallecomigas schlingeri. Ht =
Heteromigas dovei, Mg = Migas gatenbyi female + Migas taierii male, Mv = Goloboffia vel/ardi, Mp = Moggridgea
pehngueyi, Mr = Moggridgea rupicola. Mi = Moggridgea intermedia, Mt = Moggridgea tingle, Po = Poecilomigas
abrahami, Pb = Poecilomigas basilleupi, Ta = Thyropoeus malagasus, Tm = Thyropoeus mirandiis. Ms =
Micromesomma cowani, Av = Paramigas milloti. Is = Paramigas pectinatus, Pu = Paramigas alluaudi, Rm =
Paramigas oracle, Le = Paramigas pauliani, Vb = Paramigas goodmani, Md = Paramigas rothontm, Mb = Paramigas
manakambus. Ah = Paramigas macrops, Pp = Paramigas perroli. An = Paramigas andasibe. Character abbreviations
are (horizontally): abs = absent, ant. = anterior, apic = apical, aut = autapomorphy, Calath = calathotarsine type, cl =
clavate, cusp = cuspules, dist. = distribution, dorsoap. = dorsoapical, F = femur, H = height, H/L = height divided by
length, ITC = inferior tarsal claw, L = length, 1 = long stalk with head. Max. = pedipalpal coxa, med = median, Moggr
= Moggridgea type, mt = metatarsus, n = head narrower than stalk, ns = narrow stalk, OAW = ocular area width, Pat =
patella, palp = pedipalpus, PLS = posterior lateral spinnerets, procumb. = procumbant, prox. = proximal, prs = present,
s = short, cylindrical, scler. = sclerotization, segs = segments, SS = sternal sigilla, T = tibia, t = tarsus, vent = ventral, W
= width, < = less than, > = greater than, M = male, F = female.
Carapace
1. OAW/ Caput W: < 0.41; > 0.45;
2. Wide OAW: normal; excra wide;
3. Prefoveal setae: abs; small; enlarged;
4. Caput setation: extensive; reduced;
5. Caput H/ caput L: arched; low;
6. Fovea: procurved to straight; recurved;
7. Recurved Fovea: simple; tripartite;
Mouthparts
8. Cheliceral tooth row #: 2; plus denticles;
9. Intercheliceral basal swellings: abs; prs;
10. M intercheliceral tumescence: abs; prs;
11. Rastellum: prs; abs;
12. Fang shape: round; quadrate & keeled;
13. Fang basal tooth; abs; prs;
14. Fang orientation: vertical; diagonal;
15. F max cusp dist: across max; prox;
16. F max cusp ant dist: broad; corner only;
Legs and sternum
17. Thorns on coxae II- III: abs; prs;
18. Anterior sternal sigilla: prs; abs;
19. Deeply excavate SS : abs; prs;
20. F t I, II spines: prs; abs;
21. F ant. leg shape: round; flattened;
22. Pat vent lamellate setae: abs; prs;
23. F I-II fine, vent setae: abs; dense, long;
24. F T I,II dorsoap setae: erect; procumb;
25. F Mt I, II dorsoap setae: erect; procumb;
26. F F III vent membrane: short; elongate;
27 . F T III basal concavity: abs; prs;
28. F T III anterior ridge: abs; prs;
29. F P-T 3 spination: stout; slender;
30. T dark maculations: abs; prs;
31. Mt IV preening comb: abs; Calath; Moggr;
32. Mt III preening comb: abs; prs;
33. Preening comb form: narrow; encircling;
34. Claw 111,1V size relative to tooth: >; <;
35. ITC 3 size: large; small;
lABCMHMMMMMMPPTTMAIPRLVMMAPA
dcooatgvpritobamsvsumebdbhpn St
0101111111111111111111111111 1
-0-1000100000000000000000000 1
2111112111022201121102101111 9
000000000000001111111111--1- 1
0000001111111100111111111111 3
0001111111111111111111111111 1
— ooioooooooooiiiiimiiiii 2
lABCMHMMMMMMPPTTMAIPRLVMMAPA
dcooatgvpritobajnevsumebdbhpn St
0111010100000011110000100011 5
010Q0010071H100111111111111 4
0001-0110701110 00-0 2
0001111111111111111111111111 1
0001111111111111111111111111 1
0100001100001110000000000000 4
0101111111111111111111111111 1
001110010000000000000000--0- 2
010001000000000000000000--0- 2
lABCMHMMMMMMPPTTMAIPRLVMMAPA
dcooatgvpritobamsvsvunebdbhpa St
0000000001000000100000000000 2
0001111111111111111111111111 1
0000000000000011000000000000 1
000000101111110000000000--0- 1
001111111111111H1111111--1- 2
0000000111110000000000000000 2
000000100000100100001010--1- 4
000000000000000011111111--1- 1
000000000000000001111111--1- 1
000000000000001111111111--1- 1
0000000011110011111111111111 2
000000000000000111111111--1- 2
000111010000000000000000--0- 2
0000000000001100000000000000 1
0001101122210000000000000000 3
0001100100000000000000000000 1
11-010000 1
00
00
22
00
33
00
50
CI
. 20
0-25
1 .
1 .
,
1 .
50
00
00
25
00
. 50
. 50
CI
0.50
1 . 00
1 . 00
1 . 00
. 50
0000000000000011000000000000
0011001111111110111111111111
.00 10
,00 10
22
00 10
71 2
00 10
90 4
II wt
S3 1
62 1
.80 4
00 10
,00 10
40 1
00 10
66 3
.66
Fit
10.0
10 .
3 .
10.
6 .
10.
7 .
I Wt
00 10
00 10
00 10
75
40
00 10
00 10
1.00 10
1.00 10
0.83 5
1 . 00 10
1.00 10
1.00 10
0.25
Fit
4 .2
5 .0
7 . 5
10 .0
10 . 5
5.0
10.
7 . 5
7. 5
119
120
OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES, NO. 151
Male characters
36. M F I venter: convex; carinate;
37. M_ palp t form: ectal=mesal ; ectal>>me3al ;
38. M palp t apical spinules: abs; prs;
39. M Retrolateral T I megaspine : abs ; prs ;
40. M mt I shape: cylindrical,- pale swelling;
41. M t III shape: cylindrical; swollen;
Spinnerets
42. PLS apical segment: triangular; domed;
43. PLS spigot dist.: all segs . ; med & apic;
Female genitalia
44. Spermathecal scler.; uniform; median band;
45. Spermathecal shape: s; n; 1; ns; cl;
lABCMHMMMMMMPPTTMAIPRLVMHAFA
dcooatgvpritobamsvsumebdbhpn St
0000-0001110000 00-0 1
0000-0001110000 00-0 1
1000-0110000110 11-1 4
1011-0100001011 11-1 5
0000-0000000000 00-1 1
0000-0000000000 10-1 1
lABCMHMMMMMMPPTTMAIPRLVMMAPA
dcooatgvpri tobamavsiiinebdbhpn St
0101111111111111111111111111 1
0001111111111111111111111111 1
lABCMHMMMMMMPPTTMAIPRLVMMAPA
dcooatgvpritobamavsuinebdbhpn St
000000001111000000000000??0? 1
300011202222224432332323??2? 7
CI
RI Wt Fit
1
. 00
1.00
10
10
.0
1
.00
1. 00
10
10
.0
.25
0.57
1
5
.0
.20
0.33
4
.2
1
.00
aut
10
__.
--
1
. 00
1.00
10
10
.0
CI RI Wt Fit
1.00 1.00 10 10.0
1.00 1. 00 10 10.0
CI RI Wt Fit
1.00 1.00 10 10.0
. 57 .70 4 5.0
SMFTHSONIAN INSTFTUTION UBRARIES
3 9088 01302 6430
