Leistikow 2000
Leistikow 2000
Leistikow 2000
4: 1 -85 (2001)
Gesellschaft fr Biologische Systematik
http://www.senckenberg.uni-frankfurt.de/odes/01-04.htm
Abstract
South America is diverse in climatic and thus vegetational zonation, and even the uniformly looking tropical rain forests are a mosaic of
different habitats depending on the soils, the regional climate and also the geological history. An important part of the nutrient webs of the
rain forests is formed by the terrestrial Isopoda, or Oniscidea, the only truly terrestrial taxon within the Crustacea. They are important,
because they participate in soil formation by breaking up leaf litter when foraging on the fungi and bacteria growing on them.
After a century of research on this interesting taxon, a revision of the terrestrial isopod taxa from South America and some of the Antillean
Islands, which are traditionally placed in the family Philosciidae, was performed in the last years to establish monophyletic genera. Within
this study, the phylogenetic relationships of these genera are elucidated in the light of phylogenetic systematics. Several new taxa are
recognized, which are partially neotropical, partially also found on other continents, particularly the old Gondwanian fragments. The
monophyla are checked for their distributional patterns which are compared with those patterns from other taxa from South America and
some correspondence was found. The distributional patterns are analysed with respect to the evolution of the Oniscidea and also with
respect to the geological history of their habitats.
Species that were probably introduced are marked with an asterisk (*), nomina dubia are marked with an accent (^).
Key words: Crustacea, Isopoda, Oniscidea, Crinocheta, phylogenetical systematics, phylogeny, biogeography, Neotropis, South America
Contents
1
2
3
4
4.1
4.2
4.2.1
4.2.2
4.2.3
4.3
5
5.1
5.2
1
Introduction ..................................................
History ..........................................................
Material and Methods ...................................
Taxonomic Section .......................................
Phylogenetically Informative Characters .....
Generic Account ...........................................
The South American Genera ........................
Nomina dubia ...............................................
Genera used as Outgroups ............................
Key to the Genera of "Philosciidae"
of the New World ........................................
Phylogeny of South American
"Philosciidae" ..............................................
Evolution of Lung Structures .......................
Phylogenetic Reconstruction ........................
2
3
5
8
8
23
23
51
51
6
6.1
6.2
6.3
6.4
6.4.1
6.4.2
6.5
6.6
6.7
52
53
53
54
7
8
9
63
63
67
68
69
71
72
73
74
74
75
76
76
1 Introduction
The taxon Oniscidea plays an important role in the
decomposition of organic material within the terrestrial
biota. They can occur in high densities, comprising the
bulk of terrestrial detritivores with numbers of up to
500 individuals per square meter (Davis 1984). In
South America, they are found in tropical ecosystems
where they are both terrestrially and arboreally distributed (Picardo 1913, Paoletti 1989, Nadkarni and
Longini 1990). Their contribution to the consumption
of decaying plant material may be up to 12% (Grnwald 1988), thus they play an important role in soil
formation. Although they are ubiquitous, little is
known about their life history and the phylogeny of
this interesting suborder of the Isopoda. The Oniscidea
is the only truly terrestrial taxon of the Crustacea.
The oniscidean fauna of Europe is well documented; and an identification key for the approximately
1230 nominal species is available (Schmlzer 1965),
although new species and even genera are still being
discovered (e.g. Schmalfuss 1998a). A survey of African Oniscidea listed 575 species (Ferrara and Taiti
1979). For the Americas, a number of 254 species of
terrestrial Isopoda was given by van Name (1936) and
several more were listed in subsequent publications
(van Name 1940 and 1942). A recent survey led to an
increase to up to 521 described nominal species in
1997 (Leistikow and Wgele 1999). But this is only
the tip of the iceberg since every expedition to the
tropical rain forests reveals new species and even
genera.
Up to now, about 4000 species of Oniscidea are
known, half of all known isopods (Gruner 1993).
Apart from some smaller taxa, the suborder is
subdivided into the monophyletic adelphotaxa Synocheta and Crinocheta, the latter comprising the bulk
of the species. A summary of the attempts made in the
last years to establish a phylogenetic system of the
higher taxa of Oniscidea was given by Erhard (1998).
Until recently, the phylogeny within the Crinocheta
had been poorly studied, with almost every known
family forming a paraphyletic assembly rather than reflecting the natural relationships within this taxon (H.
Schmalfuss, pers. comm.). New insight into the phylogeny of this group was provided by Schmidt (1999).
Among the Crinocheta, about one hundred genera
are placed in the so-called family "Philosciidae", the
phylogeny of it has not been discussed sufficiently yet.
This basal group is, besides the Armadillidae, one of
the most important groups of Oniscidea in tropical
habitats (Schmalfuss 1984a: 12), in South America
about one fifth of the nominal species belong to this
In order to make the determination of South American philosciids possible for non-specialists, a key to
the genera is included, keys on species level were
published for Androdeloscia (Leistikow 1999c), Erophiloscia (Leistikow 2001a), Ischioscia (Leistikow in
press) and Benthana (Leistikow and Arajo, in press).
They should facilitate the recognition of the species
and aid in the discrimination of the representatives of
this taxon in ecological studies where they are mostly
referred to just as "Isopods".
2 History
In this chapter, the history of the exploration of the
diversity of South American philosciids is presented in
a review of the enhancement of our knowledge from
both a taxonomic and faunistic approach over the last
100 years. At the begining of the century, the genus
Philoscia Latreille, 1804 was a "melting pot" for species which at least have the narrow pleon in common.
Budde-Lund (1904, 1908) made some efforts to resolve this genus into more natural groups which led to
the establishment of several subgenera, later elevated
to generic rank by subsequent authors, e.g. Aphiloscia
Budde-Lund, 1908, Benthana Budde-Lund, 1908, and
Pseudophiloscia Budde-Lund, 1904. Verhoeff (1908a,
b) contributed to the systematics on supra-specific
level, he mainly focussed on European representatives
of Philoscia. From the Americas, the first "philosciid"
species were mentioned in the middle of the last
century. Dana (1853) described some species from
Chile, as did Nicolet (1849); Miers (1877) focussed on
the Crustacea of Central America and described Philougria nitida Miers, 1877 from Nicaragua, the first
species of a taxon now known as Ischioscia Verhoeff,
1928. Budde-Lund (1885, 1893) and Dollfus (1890,
1893) added further species of "Philoscia" from Venezuela and Chile. Van Name (1936) adopted the subgeneric division of Budde-Lund (1908) but hesitated in
raising them to the rank of genera. New species were
still described as belonging to Philoscia although he
recognised the differences in these taxa. It was the
merit of Verhoeff (1928, 1939, 1941, 1951) to establish the first new genera of Philosciidae for the neotropical fauna. The works of Vandel on the faunas of
Venezuela (1952), Ecuador (1968) and Colombia
(1972) enhanced our knowledge on the philosciids of
South America, although several genera are described
superficially. But in many cases it is not conforming to
the rules of the ICZN or simply lacking the phylogenetic background. An example is the establishment
of the genus Ecuadoroniscus Vandel, 1968 with the
single species E. orientalis Vandel, 1968 based on
America. For the reconstruction of phylogeny, additional species of Oniscidea were examined for outgroup
comparison. A list of the species examined is given
below (table 1).
The material was examined mainly by means of a
Wild stereoscope, for the appendages an Olympus
light microscope was used. The animals were dissected
and the appendages were mounted in glycerine jelly.
Line drawings were prepared using a camera lucida.
As far as museum material is concerned, microscopic
slides of the type material were examined. In cases
where it was not possible to elucidate details of the
morphology, the structures were examined via scanning electron microscopy (SEM Hitachi S 450). The
material was critical point dried with a Balzers Union
Critical Point Dryer and then coated with a 30 nm coat
of gold with an Anatech Hummer VII sputter.
Lung structures, which can be found in some
species, were investigated in detail: in addition to the
SEM analysis, the pleon was mounted in unicryl resin
and cut with a Leica microtome in 0.25 mm slides
which were stained with bromine-toluidine blue and
then examined with a light microscope. Ink drawings
of some representative slides were prepared.
For the analysis of phylogenetic relationships, the
Hennigian principles of outgroup comparison, determination of character homology and polarity, and tree
construction using the maximum parsimony method
were used. For the analysis of the phylogeny of the
taxa examined, groundpatterns were reconstructed for
the taxa recognized. Characters that are commonly
found within a taxon and particularly in their basal
representatives were adopted for the groundpattern,
both autapomorphies of the taxon and plesiomorphic
characters. In terminal taxa, in this work mainly genera, attention has to be paid to apomorphic characters
which evolved within a taxon. Such characters can not
belong to the groundpattern. Subsequently, groundpatterns for higher taxonomic hierarchies are reconstructed by comparing the groundpatterns of the subordinate taxa. The groundpatterns were used for the
reconstruction of the cladogram presented in chapter
5.2 (fig. 13a-d); the sistertaxon relationships were
evidenced by means of synapomorphies, the polarity
of character states was fixed by outgroup comparison.
All taxa, which do not belong to the taxon under
consideration were used as outgroups.
New genera were established to accommodate some
species which formerly were included in Philoscia
Latreille, 1804, but do not share the autapomorphies of
this genus. For the definition of new genera, the characters of the species were compared to those present
in existing genera. After a check monophyly of the
Table 1. List of the species examined. Species that were probably introduced are marked with an asterisk (*), nomina dubia are marked
with an accent (^).
Ligiidae:
Ligia baudiniana Milne Edwards, 1840,
Costa Rica, own collection
Detonidae:
Deto echinata Gurin, 1836, South Africa, NMHU
Philosciidae:
4 Taxonomic Section
4.1 Phylogenetically Informative Characters
This chapter deals with the phylogenetically informative characters of the morphology of the oniscidean
taxa examined with particular respect to the putative
groundpattern of Oniscidea and the groundpattern of
the Crinocheta recently worked out by Schmidt (1999).
Although Racovitza (1923a, b) proposed an orientation and nomenclature for the isopod body and its
appendages, some modifications are made in this study. The appendages are all orientated towards their
plesiomorphic position with respect to the groundpattern of Mandibulata hanging down ventrally regardless their functional orientation in the animal. Hence,
the terms "medial", "lateral", "rostral" and "caudal" are
used for describing structures on the appendages: the
rostral side of both the maxillula or pleopod 1 is the
side facing towards the cephalothorax, the lateral side
towards the lateral margin.
Org. Divers. Evol. 1, Electr. Suppl. 4 (2001)
10
11
Ha4-1
Ha4-2
Ha4-3
Ha4-4
The antennula or first antenna (fig. 2) is threearticulate in all examined species. This is the condition
in the groundpattern of the Oniscidea. In basal species
like Ligia baudiniana and Halophiloscia couchii
(Kinahan, 1875), the articles are elongate and cylindrical [A1-1], whereas in other species a derived
condition is found: the proximal article is bulky, the
distal articles - particularly the medial one - being
much shorter [A1-2], the members of the genus
Ischioscia (fig. 2B) have the distal margin of the
proximal article drawn out to form a shield-like
protrusion [A1-3]. On the distal articles, several sets of
aesthetascs are inserted, in all cases a set of two aesthetascs apically, the more medio-proximally aesthetascs are inserted in pairs in stepped order (fig. 2B/C)
which is certainly the plesiomorphic character state in
the Oniscoidea [Ae1-1], because it reflects the aesthetasc pattern of formerly unfused flagellar articles as
seen in other isopod taxa. These structures may be
modified in several ways: the proximal article can be
further shortened or enlarged, the sets of medial aesthetascs can be arranged in different positions. In the
genus Prosekia (fig. 2E), the lateral aesthetascs form a
tuft [Ae1-3]; in Philoscia Latreille, 1804 (fig. 2D), all
the aesthetascs are gathered in an apical group [Ae12]. A further characteristic arrangement is found in the
genera Caraiboscia Vandel, 1968, Colombophiloscia
Vandel, 1968 and Nesophiloscia Vandel, 1968. Here,
the aesthetascs are not attached to the article, they are
erected and form a cockscomb-like row [Ae1-4].
12
13
Lateral maxillular article with 5+7 teeth, plumose slender stalk, inner set ctenate (plesiomorphic)
Lateral maxillular article with 5+6 teeth, teeth
of inner set simple
14
15
Ml1-3
Ml1-4
Ml1-5
Ml1-6
Ml1-7
Ml1-8
Ml2-1
Ml2-2
Ml2-3
16
Fig. 6. Aspects of the morphology of the maxilliped in the Crinocheta, arrows indicating [Mp2-6].
A: Deto echinata Gurin, 1836 [Mp1-1/Mp2-2/Mp3-1]; B: Ischioscia guamae Leistikow, 2001 [Mp1-3/Mp2-3/Mp3-2]; C: Philoscia muscorum
(Scopoli, 1793) [Mp1-2/Mp2-4/Mp3-4]; D: Androdeloscia formosa (Mulaik, 1960) [Mp2-4/Mp3-3]; E: Caraiboscia christiani Leistikow, 2001
[Mp2-5/Mp3-3]
17
Fig. 7. Some examples of the structure of the pereopod 1 carpus and propus in the Crinocheta. A: Deto echinata Gurin, 1836; B:
Benthana taeniata Arajo and Buckup, 1994; C: Androdeloscia formosa (Mulaik, 1960); D: Philoscia muscorum (Scopoli, 1793); E:
Plataoniscus borellii (Dollfus, 1897)
18
19
PE2-1
PE8-1
PE9-1
20
21
22
The male pleopod 1 and 2 endopodites are transformed to copulatory organs which interact with the
genital papilla. Thus, several interlocking structures
have evolved, as recently demonstrated by Erhard
(1997) and first described by Legrand (1946). The
endopodite 1 has a caudal furrow, its medial margin
bears a row of small spines [PL4-2], which may be
reduced in several taxa [PL4-1]. The pointed shape of
the endopodite is deviated several times and may be
specifically complex in some neotropic "philosciids".
The elongate endopodite 2 can reach a length of more
than twice the length of the exopodite [PL5-2]. This
faint structure is then retained by the pleopod 5
exopodite, which is medio-proximally protruding and
may show a caudal furrow for holding the endopodite
2 [PL6-3]. Such a "guide slot" certainly evolved several times convergently in taxa with extremely prolonged pleopod 2 endopodites. For Porcellio dilatatus
Koch, 1846, Legrand (1946) described the structure in
detail. He also stated that this guide slot which he
called "gouttire marginale" is present in several Synocheta, all higher Crinocheta, but is lacking in the
Scyphacidae primarily, and in the Trachelipodidae and
Hemilepistus Auduoin, 1825 due to the short pleopodite 2 endopodite. Verhoeff (1920) misinterpreted this
guide slot as part of a water conducting system. He
referred to the rim - the "Innenrandlngsrinne" - as a
structure to divert a surplus of water from the pleoventral area.
Pleopodal lungs [PL1-1] are evolved several times
in the Oniscoidea; among the taxa of philosciid facies,
only few have pleopodal lungs [PL1-2]. The members
Balloniscidae (pl. 150, E/F) both bear semi-covered
[PL2-1] and covered lungs [PL2-2], semi-covered
lungs are present in Atlantoscia rubromarginata
Arajo and Leistikow, 1999, too.
PL2-1 Pleopods with semi-covered lungs on exopodites 1 to 5, bearing transverse clasps (plesiomorphic)
PL2-2 Pleopods with monospiracular covered lungs
on exopodites 1 to 5
PL3-1 Male pleopod 1 exopodite pointed (plesiomorphic)
PL3-2 Male pleopod 1 exopodite rounded
PL3-3 Like PL3-1, but with lateral incision
PL3-4 Male pleopod 1 exopodite almost subquadrangular
PL3-5 Male pleopod 1 exopodite drop-shaped
The shape of the uropod is uniform within the examined taxa, most have rather short endo- and exopodites which are inserted on a subtriangular, laterally
grooved protopodite. The insertion may be at the same
level [UR2-1], as in some species of Benthana; or the
endopodite is inserted more proximally than the
exopodite as in the bulk of taxa [UR1-2]. The insertion
at the same level is found in Ligia, thus it is thought to
represent the plesiomorphic character state; similarly,
the protopodites surpassing the pleotelson caudally
[UR3-1] are found in Ligia and are referred to as
Org. Divers. Evol. 1, Electr. Suppl. 4 (2001)
23
24
25
26
27
28
29
30
31
32
33
34
broader. Therefore, the teeth are standing in a semicircle laterally, similar to Ligia Fabricius, 1798, representing the position found in the groundpattern of
Oniscidea. This character can not be used to define the
adelphotaxon of Benthana nor the close relationship of
the forms bearing ctenate teeth.
The generic independence of Benthana and Benthanoides has to be doubted, it seems as if the species of
Benthanoides are simply the more basal species which
lack the characteristic dentiform protrusion of the pleopod 1 exopodite. This character was used by Lemos de
Castro (1958b) for separation of Benthana pauper
Jackson, 1928 in the genus Benthanoides. For a formal
synonymization of the two genera, a detailed phylogenetic study of all the species hitherto known has to
be undertaken. It is likely that the genus Benthanoides
represents the basal representatives of the genus Benthana in its original extent (Benthana s.l.), with the
autapomorphies of Benthana s.str.:
n Cordiform shape of pleopod 1 exopodite with a lateral dentiform protrusion [PL3-3] o exopodite pointed
without protrusion [PL3-2]
n Ornamental sensory spine of carpus 1 with hand-like
apex, slightly merged with cuticular stalk [7-2] o
ornamental sensory spine with serrate double-fringe
[PE7-1]
The second character is at first sight similar to that
seen in Ischioscia Verhoeff, 1928, but details indicate
that the similarity is a convergency: In Ischioscia and
its allies, the stalk is clearly separated from the handlike apex, whereas it is intervening in the present genus. It thus is postulated to have evolved independently.
More information on the phylogeny and a description
of the hitherto unknown male of B. convexa Lemos de
Castro, 1958 were given in Arajo and Leistikow
(1999), the status of Benthanoscia Lemos de Castro,
1958 was discussed in Leistikow and Arajo (2000b).
and the short noduli laterales. The structure of the cephalothorax is quite similar to that in the genera Erophiloscia Vandel, 1972, Androdeloscia Leistikow,
1999 or Xiphoniscus Vandel, 1968. The shape of the
antennula and the presence of lateral lobes on the
cephalothorax are plesiomorphies within the Oniscoidea. The apomorphic characters of this genus are:
n Sensory spines of carpus and merus with up to five
subapical tips, sensillum laterally o sensory spines
with up to two subapical tips, sensillum apically
n Reduction of the tip on the medial endite of the
maxillula [Mm1-3] o medial endite with apical tip
[Mm1-2]
n Lateral lobe of the maxilla apically truncate o
lateral lobe apically rounded
These apomorphies have to be evaluated with respect
to the closest relative of the genus Atlantoscia, the
genus Benthana, with which it shares the shape of the
maxilliped. They are mainly confined to the mouthparts and may be susceptible to convergence. At least
the first character, the shape of the sensory spines is
quite unique among the philosciids and supports
monophyly of the genus.
35
lacking, compound eyes composed of about 15 ommatidia. Antennula three-articulate, slender, antenna long
with three-articulate flagellum, apical organ longer
than distal article.
Mandible with molar penicil consisting of about ten
free branches, maxillula with lateral endite bearing 4
and 6 teeth, inner set with five teeth cleft, slender stalk
present, maxilla with both lobes subequal in breadth,
maxilliped with endite setose, bearing prominent penicil rostrally, palp three-articulate, with three prominent
setal tufts.
Pereopods slender, ornamental sensory spine of carpus
1 with serrate double-fringe, dactylus with interungual
seta long, more conspicuous than dactylar seta, inner
claw short, coxal plates without gland pores, sulcus
marginalis and nodulus lateralis present, all noduli
inserted at same distance from lateral margin. Male
pereopod 1 propus and carpus inflated, bearing more
or less prominent setal brushes.
Pleopods without respiratory areas, exopodites with
about five sensory spines laterally, endopodites bilobate. Uropod with protopodite laterally grooved, endopodite inserting more proximally than exopodite.
Genital papilla with short ventral shield, orifices on
small to conspicuously elongate protrusions of the
terminal spatula.
Type species: Philoscia richardsonae Holmes and
Gay, 1904
Number of nominal species: 23, four reported from the
Neotropics
Distribution: Interestingly, the genera Halophiloscia
and Littorophiloscia vicariate each other. As stated
above, the genus Halophiloscia is distributed in the
Mediterranean area and on the Atlantic coasts of
Europe. On the other hand, Littorophiloscia is found in
the temperate Pacific, the western Atlantic coasts and
the subtropical to tropical regions of the world.
Remarks: The genus was instituted by Hatch (1947)
for the eastern pacific L. richardsonae. Lemos de
Castro (1968) gave a synopsis of the genus and
presented a generic definition without indicating the
apomorphies of the genus. A close relationship of
Littorophiloscia, Halophiloscia, and Alloniscus Dana,
1853 was postulated based on the similarities of the
maxillipedal palp and the presence of a dactylar seta in
the three genera. As stated above, the former character
is a symplesiomorphy, and this is true for the latter
character. Moreover, dactylar setae are present in most
of the Crinocheta. Another character was interpreted
erroneously as an ancestral state: the bifurcate genital
papilla of Halophiloscia was thought to be intermediate between the two separate genital papillae of Ligia
(plesiomorphic state) and the fused papilla of Crino-
36
37
38
39
40
41
42
female.
Distribution: Confined to Ecuador.
Remarks: Unfortunately, the antennae, maxillae, pereopods, pleopods and uropods are missing in the single
known specimen. Since the specimen is a female and
due to the incompleteness of the preserved material,
the genus Oreades is merely a nomen dubium similar
to Ecuadoroniscus. It is described here with respect to
the phylogeny of the South American Oniscidea of
philosciid facies.
As an autapomorphy of the genus one character can be
mentioned:
n Mandible with simple molar penicil [Md2-4] o molar penicil composed of about seven branches [Md2-2]
There may be more and better characters of this kind
in this form, but they remain unknown due to the loss
of the rest of the material, i.e. the pereopods and
pleopods of this species.
Oreades lativentris Vandel, 1968 (fig. 18)
Material: microscopic slides of female (body length
4 mm): Ecuador, Oriente, tropical forest at Puyo, alt.
800 m a.s.l., humus, leg. IV.1965, N. and J. Leleup,
VC 5237.1-7
Colour: Vandel (1968) wrote: "Violace. Le vertex et
la partie mdiane sont couverts de linoles blanches,
correspondant aux insertions musculaires. Les pleurpimres sont entirement colors. Le plon est entirement pigment l'exception des deux premiers segments qui presentent, dans la rgion antrieure, une
zone dpigmente."
Cephalothorax: Linea frontalis and lamina frontalis
lacking, faint linea supra-antennalis present, compound
eyes composed of about 14 ommatidia arranged in four
rows.
Pereon: Tegument smooth, only few tricorn-like setae,
coxal plates with sulcus marginalis, lacking gland
pores and noduli laterales (fig. 18, Cx3).
Pleon: Neopleurae of pleonite 3 to 5 continuing body
outline of pereon, pleotelson with concave lateral margins and semicircular pit ventrally.
Appendages:
Antennula: Three-articulate with distal article pointed,
bearing some aesthetascs apically, single aesthetasc
more proximally (fig. 18, An1).
Antenna: Lacking in the only specimen.
Mandible: Only left mandible preserved, molar penicil
simple, slender, pars intermedia bearing few coniform
setae and two penicils, additional plumose seta more
proximally (fig. 18, Mdl).
Maxillula: Only left lateral endite preserved, apically
with 4+6 teeth, five of inner set deeply cleft, pointed
triangular lobe rostrally, lateral fringe of trichiform
Org. Divers. Evol. 1, Electr. Suppl. 4 (2001)
43
44
45
plex differentiation of the male pleopods. The monophyly of this group is supported by the autapomorphies
listed below:
n Antennula with the medial aesthetascs gathered in a
tuft, directed more or less medio-distally, not attached
to article 3 [Ae1-3] o antennula with aesthetascs
paired in a medial line [Ae1-1]
n Transverse fold between aesthetasc tuft and distal
pair of aesthetascs o distal article coniform
n Male pleopod 1 with hyaline lamellae near apex
[PL4-3] o no such lamellae on pleopod 1 exopodite
[PL4-2]
In contrast to Vandel's statement (1952), no flagelliform noduli laterales could be observed in Prosekia
rutilans (Vandel, 1952). The noduli are slightly longer
than the tricorn-like setae but not considerably different to those of the genus Chaetophiloscia Verhoeff,
1908 in which the species was originally included. In
Metaprosekia Leistikow, 2000, the noduli are not much
longer than in Chaetophiloscia, too. Long Noduli
laterales are confined to the more derived taxa of the
Prosekiini. If flagelliform noduli were present in the
groundpattern of the Prosekiini, an independent reduction of the length of the noduli in Prosekia and Metaprosekia has to be postulated. There is no evidence for
such a reduction of the size, therefore, flagelliform
noduli laterales are considered not to be a character of
the groundpattern of this group.
46
47
48
49
50
51
52
13
14
17
18
25
34
31
53
Andenoniscus
32
A detailed analysis of the lung morphology was performed by Hoese (1982 and 1983). His excellent work
led to some misinterpretations of the observed structures, and a simple morphological row was constructed
and found its way even in treatises on zoology (e.g.
Gruner 1993). Further research revealed the fact that
the lungs, which until then on the basis of available
data were supposed to have evolved only once in the
Oniscoidea and then been modified in the course of
further evolution must have evolved several times,
starting with folded lateral areas of the exopodites and
forming respiratory organs of different complexity, the
most specialized being covered lungs, which have contact to the surrounding air solely by a spiracle. For example, this morphological row can be traced in the Eubelidae (Ferrara et al. 1990). Furthermore, the thorough examination of the genus Aphiloscia BuddeLund, 1908 ended in the discovery of a convergent
evolution of covered lungs within this genus (Ferrara
et al. 1994). This contrasts with the opinion that within
the family "Philosciidae" lungs never occur [cf. Vandel's (1962) definition of the family]. For this study,
the respiratory organs of the genera Balloniscus, Benthana and Atlantoscia were examined and a converOrg. Divers. Evol. 1, Electr. Suppl. 4 (2001)
54
55
56
57
58
59
60
61
62
medial and the lateral margins of the pleopod 1, another possible autapomorphy.
The genus Pulmoniscus has pleopods which resemble those of Agnara Budde-Lund, 1908: The position
of the lungs, the structure of the cuticle of the perispiracular area and the outline of the pleopod exopodites
are similar. If Pulmoniscus belonged to to a monophyletic group with the Agnaridae, the mandible
would be simplified: it lacks the multiple intermediate
penicils which are present in Agnaridae. Moreover, the
antennal flagellum of Pulmoniscus is three-articulate,
whereas the Agnaridae have a two-articulate flagellum.
In the case of a close relationship of Pulmoniscus and
the Agnaridae, the similarity in the mandible of the
Agnaridae and the Porcellionidae, Cylisticidae and
Porcellionidae, the multiplication of the intermedial
penicils, and the two-articulate flagellum must be convergencies. For the moment, the phylogenetic relationships of Pulmoniscus remain obscure.
The genus Oniscophiloscia, too, is part of this monophyletic group characterized by the presence of Yshaped tricorns [Ha3-3], but similar to Balloniscus, the
pleotelson is different from the pleotelson with triangular distal part found in Philoscia, the Oniscidae, Trachelipodidae and Porcellionidae, among others. Anyway, the ascription of Oniscophiloscia to the Oniscidae (Strouhal 1960) is not justified by any character. It
is more likely an early descendant of the unnamed
taxon 6 of Schmidt (1999), which is quite isolated
within this taxon.
Finally, the discussion of the phylogeny of South
American Crinocheta of philosciid facies again has to
focus on the noduli laterales. These "philosciids" belong to the recently established by Schmidt (1999).
Thus, noduli laterales should be present in all the taxa
dealt with. Interestingly, in some of the Scleropactidae
and in the Roraimoscia genus-group, such receptors
have not been found. Since they are present in the
basal genera of the Halophilosciidae and in both Atlantoscia and Benthana, they must have been reduced in
these taxa. Within the Scleropactidae, one of the tricorn-like dorsal receptors differentiated to a "nodulus
lateralis". These noduli differ in their position from
those known from other taxa. No ultrastructural work
has been done on these noduli, so the question is still
unanswered if these receptors were all homologous or
if they evolved several times. The noduli laterales of
the Prosekiini differ remarkably in size from those
found in Benthana or the Porcellionidae, and they
must not necessarily be homologues if there was a selective pressure to evolve specialized dorsal receptors.
To clarify this question, more ultrastructural research
on the dorsal receptors is necessary. Some first results
6 Biogeographic Section
6.1 Geological and Geographical Background
South America is part of the former supercontinent
Gondwana. Until the late Triassic, Gondwana comprised Australia, Antarctica, Africa, India and South
America. Beginning at about 200 mio. years ago in the
Mozambique basin off East Africa (Mukasa and
Dalziel 1996), continental drift caused the split of
Gondwana into the continents of our days (fig. 20A).
The largest fragment was the South America-Africa
land mass which stayed unfragmented until the early
Cretaceous (fig. 20B). With the spread of the Atlantic
Ocean, the separation of Africa and South America
began. The oldest sea floor in the southern Atlantic is
about 130 mio. years old, it was formed in the Valangian stage of the Jurassic (Tarling 1980). Other authors
date the split off to 150 mio. years in the south and due
to the relative rotation of Africa around an Euler pole
in Western Africa in the region of the Niger delta, to
110 to 95 mio. years in the north (Pindell and Dewey
1982, Pitman et al. 1993). In the late Cretaceous, both
continents were separated by at least 200 km distance
(fig. 20C). The shallow water might not have effectively inhibited a faunal exchange, but the young ocean
was a great barrier for smaller organisms. An effective
barrier is postulated for a period of the last 100 mio.
years (Pielou 1979) although there was a long-lasting
stepping stone route between northeastern Brazil and
eastern West Africa due to the position of the Euler
pole in West Africa: presumably a swampy area with
small islets was filling the gap between the two continental crusts (Briggs 1987). For a period from 100 to
84 mio. years B.P. the pole of relative rotation shifted
to a position in the central Atlantic (Pitman et al.
1993), with persisting anticlockwise relative rotation
of Africa.
While South America and Africa have been separated for at least 80 mio. years, the connection of South
America and Antarctica existed for a much longer time
(fig. 20D). Although there is evidence for rocks in the
Weddell Sea as old as 160 mio. years (Musaka and
Dalziel 1996), the opening of the Drake passage began
35 mio. years ago, and the present position was reached
63
64
Fig. 20 A. The face of the earth in the Triassic (240-200 million years ago)
White: land masses, Light Blue: epicontinental water, Blue: ocean; after Cox (1981)
FIG. 20 B. The face of the earth in the upper Jurassic (Kimmeridgean stage: 145 million years ago)
White: land masses, Light Blue: epicontinental water, Blue: ocean; after Cox (1981)
65
Fig. 20 C. The face of the earth in the lower Cretaceous (Hautevivian stage: 125 million years ago)
White: land masses, Light Blue: epicontinental water, Blue: ocean; after Cox (1981)
Fig. 20 D. The face of the earth in the upper Cretaceous (Senonian stage: 90-80 million years ago):
White: land masses, Light Blue: epicontinental water, Blue: ocean; after Cox (1981)
66
Fig. 20 E. The face of the earth in the Eocene (45 million years ago):
White: land masses, Light Blue: epicontinental water, Blue: ocean; after Cox (1981)
Today South America spreads from about 12N to
45S and has the longest north-south extension of all
modern continents. It is quite clear that this situation
leads to interesting climatic conditions, the southern
areas are dominated by warm-temperate climates (Cfa,
Csa and Cw of the Kppen system) north to the Tropic
of Capricorn and in the higher mountain ranges of the
Andes just crossing the equator. Dry climates of the
BWh type are found in coastal Venezuela, BWk and
BSk are confined to most parts of Argentina and the
Atacama desert at the western slopes of the Andes.
The remainder of the continent is dominated by
tropical climates of the Af and the Aw type. The
climatic situation has changed remarkably since the
ice-ages. Van der Hammen (1982) reported of about
20 glacial cycles. During the glacials, the Andean
forest for example was found at an altitude of max.
2000 m whereas nowadays its upper limit is at 3200 to
3500 m (van der Hammen 1974). Since the climate in
the pleniglacial of the last ice-age at about 30.000 to
10.000 B.P. was much drier (although it is difficult to
say to what extent), the ecological zones were not only
lowered, they were also vertically compressed due to
the increase of the temperature gradient: The temperature was about 3 to 6C lower than today in the higher
Andes, for the lowlands a temperature 2 to 3C lower
67
The high diversity of many taxa is of special interest. Haffer (1969) correlated species diversity and endemism with paleogeographic data and hence laid the
cornerstone for applying the refuge theory on tropical
ecosystems. The refuge theory explains diversity on a
subordinate taxonomic level, whereas for higher taxonomic levels, vicariance theories taking continental
drift in consideration are more appropriate (Haffer
1982). Evolution rates in the Neotropics seem rather
high and e. g. in the vertebrate taxa, most species may
be younger than 1.8 to 2 mio. years (Haffer 1979).
Hence, for the species and subspecies level the 'refuge
theory' as explained by Haffer in several publications
(1969, 1974, 1990) seems to give an appropriate explanation for the patterns observed. The refuge theory
does not exclude vicariance models, "refuge biogeography is but a recent chapter of vicariance biogeography on a sub-continental scale" (Whitmore and
Prance 1987: 188).
But it is not only changes in precipitation postulated
by the refuge theory (Haffer 1969) that caused a
drastic change in vegetation. The fluctuations of the
sea level are of importance, too. Inundation cylces, for
example, caused the submergence of vast parts of
central Amazonia three times in the last 100.000 years;
this may be a cause for the separation of the Caribbean
Amazonian and southern Amazonian region (Erwin
and Adis 1982). The changing course of river beds and
fluctuating water levels over millions of years contributed to the diversity of the Neotropis. For taxa with
low vagility and even for some vertebrate taxa of
Central Amazonia like forest birds (Haffer 1974) and
primates (Hershkovitz 1977) Erwin and Adis (1982)
postulated a strong influence of microbarriers like
climatic differences or small rivers. Sometimes, even
more motile taxa seem to be incapable of overcoming
water bodies, e.g. 21 bird species of tropical Australia
failed to cross the Torres Strait from Cape York peninsula to New Guinea despite the presence of stepping
stones (Garnett 1991). Moreover, in South America,
areas of endemism evidenced by bird distribution are
greatly congruent to those obtained from data on butterfly distribution (Cracraft 1985). Hence, there are no
general constraints to a direct comparison of distributional patterns of birds and terrestrial isopods in spite
of differences in motility.
Biogeographic analyses were performed for many
of the taxa mentioned above. For South America, a
synopsis of the biogeography of the well-studied tree
flora, the avifauna and the distributional patterns of
several Lepidoptera revealed areas of high species
diversity and endemism (Whitmore and Prance 1987).
Several other authors found similar patterns, e.g.
Spassky et al. (1971) described five regions for Drosophila paulistorum semispecies: the Centroamerican,
Orinocan, Amazonian, Interior, and Andean-Brazilian
(coast) region.
Within the Neotropics, the easterns slopes of the
Andes intervening into the wide Amazon lowland,
which is lower than 500 m, are especially rich in
species of terrestrial isopods of philosciid facies. These
lowlands of the western part of Amazonia in eastern
Ecuador and northeastern Peru do not only harbour a
rich Oniscidean fauna, they are also among the most
diverse regions in relation to birds (Haffer 1990), reptiles (Dixon 1979), amphibians (Duellman 1988), butterflies, mammals and plants (cf. Gentry 1988). Interestingly, high diversity is not evidenced for the Gastropoda, which are quite abundant in other parts of South
America. Fittkau (1981) related this discrepancy to the
lack of calcium in the soils and to the nutrient cycles
which give place for many destruents (like Oniscidea)
but only little for phytophagous animals (like Gastropoda).
The climatic history of Central America seems to be
similar to that of Colombia, thus cylces of harsher
climates during the ice ages (cooler/drier) may have
led to the retraction of the tropical rain forests, e.g in
the Petn Itza and Izabal lake region of Guatemala
(Toledo 1982). The spread of rain forest persisted until
recently as can be evidenced from xeric floral elements
within the tropical rain forests (Prance 1982). Nowadays the tropical forests spread northwards to the 22nd
parallel. It is characterized by a low rate of endemism
and species diversity (Toledo 1982).
In a summary of the distribution of Andean Arthropoda, Morrone (1992) published a map detailing the
biogeographic subregions of South America.
In this work, the main focus is directed to the diverse Oniscidean fauna of the Neotropis, however
some considerations on the distributional patterns
found in southern South America and their consequences for the relationships to other continents are
discussed. Some of the findings on the biogeography
of South American Oniscidea of philosciid facies
coincide with those made in various taxa of both plants
and animals from the Neotropics and are discussed
below, starting in the north in Central America and
going south to the temperate climate of Argentina. The
most interesting fact is, that the phylogeny is reflected
by the biogeography of the taxa: while the temperate
South America is devoid of members of the taxon B,
members of Balloniscidae and Benthana are exclusively found in this region. Conversely, the tropical taxon
B is widespread in the Neotropis. The distributional
patterns of e.g. Prosekiini and Ischiosciini coincide
68
Interestingly, both species of Androdeloscia, A. formosa and A. valdezi Leistikow, 2000 belong to different
subtaxa within the genus. A possible explanation may
be found in the distant position of nuclear Central
America and Yucatn in Triassic times (Pindell and
Dewey 1982). The presence of a member of the group
bearing three hooks on the merus 7 (i.e. A. formosa)
and of the hookless group (i.e. A. valdezi) in Guatemala was due to the colonization by an ancestor of the
hook-bearing group of Proto-Yucatn from Venezuela.
An ancestor of the hookless group must have colonized the "Protoguatemalan" portion (Chortis block)
from the Ecuadorian region in the Triassic before a
northward drift of the Chortis block led to a separation
from South America. Alternatively, Central America
might have been colonized twice after the Isthmus of
Panama was formed.
From the data available in the literature (Brian
1929, Schultz 1977b and 1984), the genus Troglophiloscia Brian, 1929 is restricted to Yucatn peninsula and the adjacent areas, i. e. in recent times Belize
and Cuba. This genus may be close to Colombophiloscia from northern South America. A faunistic link
between the Greater Antilles and nuclear Central
America was also found for several poeciliid fishes in
the Caribbean, which occur only in fresh water (Rauschenberger 1988).
For Costa Rica, besides the genera Androdeloscia,
Ischioscia and Mirtana, records of Pentoniscus and
"Chaetophiloscia" were reported (Arcangeli 1930;
Richardson 1913; van Name 1936). Especially in the
genus Ischioscia, many species occur in the southern
part of the Central American land bridge: seven species were described in recent times(Leistikow 1997a
and 1999d). Southern Central America must have been
colonized by representatives of the taxon Ischiosciini
at least twice: a first colonization gave rise to Mirtana
costaricensis, subsequently, members of Ischioscia
colonized the land bridge. The very distinct species of
the genus Ischioscia present in Panama, I. zebricolor,
suggesting a correlation of the Central American faunas and those of trans-andean Amazonia even found in
Scopiones (Loureno 1984). A further colonization of
Central America must have taken place from Venezuela, the core area of the martinae-variegata subtaxon.
It are members of this subtaxon, which probably
reached the rain forest of northern Guatemala. A resolution of this question will be possible when the males
of these forms will be found (cf. Chapter 4.2).
The Greater Antilles have shifted from a more westerly to their present day position since the late Cretaceaous to early Tertiary (Briggs 1987, Pindell and
Dewey 1982, Pitman et al. 1993), so the ancestor of
69
70
Fig. 21. Projection of the dendrogram of the Ischiosciini on the map of South America:
Each square colour represents a genus
71
Fig. 22. Projection of the dendrogram of the Prosekiini on the map of South America:
Each square colour represents a genus
72
postulated pleistocene forest refuges which were evidenced from various biogeographical studies (Cracraft
1985, Prance 1982). Some bird taxa are endemic to the
Caracas and Paria cordilleras (Whitmore and Prance
1987: 145). Interestingly, these restricted-range species
partially occur sympatrically with I. variegata which is
found in almost all the areas where distinct regional
species were collected. These regional species are
found in locations with a high rate of endemism. I.
variegata might be an expansive species which colonized many regions of Venezuela relatively recently. It
lives synantropically in many areas (Leistikow 2001f).
The phenomenon of one widespread species which
lives in sympatry or parapatry with restricted-range
species is well-kown in many taxa as diverse as marine
Peracarida (C.O. Coleman, pers. comm.), birds
(several monographs on neotropical bird taxa) or
plants: the genus Tecoma of the family Bignoniaceae,
for example, is composed of one widespread species
which can be found from the southern U.S.A. to
Argentina, two widespread South African species and
a set of about ten species endemic to certain Andean
ridges (Gentry 1992). Nonetheless, some of the records
of I. variegata from northern South America and Central America have to be reconsidered (Richardson
1914, Arcangeli 1930). With respect to the records
from the Caribbean islands, a different species might
be involved.
Some elements of the Guyanan and southern Venezuelan fauna may be derived from the evolutionary
line leading to the last common ancestor of the Ischiosciini, i.e. Pentoniscus, Formicascia and Roraimoscia. They may also be close to the fauna of the Caribbean islands. The avifauna of the Pantepu region is
related both to the Venezuelan cordilleras and the Andes (50 per cent of species) and to lowland Amazonia
(30 per cent) (Whitmore and Prance 1987).
73
tion on one of the Windward Islands north of Venezuela in the Caribbean is biogeographic evidence for the
paraphyly of Balloniscidae, with the remaining species
found in southeastern and southern Brazil and Argentina.
Lastly, the genera Benthana and Benthanoides are
true austral faunal elements, which penetrated the Neotropis at its southeastern edge. The apparently disjunct
distribution of Benthana and Benthanoscia in southern
South America - if not a collecting artifact - may have
been caused by the lowering of the vegetational zones
during the glacials. The common ancestor of this stock
may have occurred in the intervening areas but is now
extint due to the drier climate in the cerrado areas of
Brazil, where no representatives of Benthana have been
found. Such a disjunct distribution has been found for
several taxa (Simpson 1973). Bigarella and de Andrade-Lima (1982) reported of a disjunct distribution of
the Araucaria-forests in the Andes and Brazil. They
found a colonization of Brazil by Andean genera in the
Oligocene with the pre-Andes as a ressource for this
colonization. This would coincide with a phylogenetic
scenario of Benthanoides being the less derived and
Benthana the more derived taxon. Benthana/Benthanoides is thought to be the sole strong link of the South
American and African fauna with Benthanops from
South Africa the closest recent relative (H. Schmalfuss, pers.comm.). Similarly, despite considerable coincidence the herpetofauna of South America and
Africa are distinct as regards the vicariance of the two
continents (Laurent 1979).
74
7 Summary
The analysis of South American Crinocheta of philosciid facies revealed the fact, that the family "Philos-
75
76
8 Acknowledgements
For the possibility to work on terrestrial isopods and their
phylogeny I would like to thank Prof. Dr. J.W. Wgele
(Ruhr-Universitt Bochum), who let me work in his department and use the facilities of the institute. He always
was a good supervisor of this work and I could freely
discuss special and common questions of phylogeny with
him. Similarly, I am indepted to Prof. Dr. T. Bartolomaeus (Universitt Bielefeld), Dr. H. Schmalfuss (SMNS),
Dr. S. Taiti (University of Florence), P.B. de Arajo
(Universidade Federal de Rio Grande do Sul) for fruitious
discussions on the manuscript. My special thanks are for
C.O. Coleman (MNHU), A. Brandt and G. Wegener
(Universitt Hamburg), H. Dalens (University of Toulouse), D. Defaye (MNHN), Y.-R. Kim and M. Lawrence
(AMNH), M. Lowe (BMNH), M. Schotte (USNM), L.
Tiefenbacher (ZSM) and P. Young (NMRJ) for their uncomplicated aid in the loan of material. For the financial
and ideological support I would like to thank the Studienstiftung des deutschen Volkes, especially making possible my collecting trip to Guatemala.
9 References
Absy, M.L. (1982). Quaternary palynological studies
in the Amazon basin. pp. 67-73 in: Biological diversification in the Tropics. Edited by G.T. Prance.
New York.
Acosta, J. & E. Upuchi (1996). Transtensional tectonocs along the south Scotia Ridge, Antarctica. Tectonophysics 267: 31-56.
Arcangeli, A. (1930). Contributo alla conoscenza del
'Microgenton' de Costa Rica. Bull. Lab. Zool. gen.
agr. Fac. Agr. Portici 25: 1-29.
Arcangeli, A. (1932). Isopodi terrestri raccolti dal Prof.
Silvestri nel Nord-America. Boll. Lab. Zool. gen.
Agr. Portici 26: 121-140.
Arcangeli, A. (1954). I rappresentanti della famiglia
dei Porcellionidae (Crostacei isopodi terrestri) hanno siempre il flagello delle antenne esterne composto di due articoli. Boll. Ist. Mus. Zool. Univ. Torino
4(7): 119-112.
77
Budde-Lund, G. (1912). Terrestrial Isopoda, particularly considered in relation to the distribution of the
southern Indopacific species. Trans. Linnean Soc.
(2)15: 367-397.
Cardoso da Silva, J.M. & D.C. Oren (1996). Application of parsimony analysis of endemicity in Amazonian biogeography: an example with primates. Biol.
J. Linnean Soc. 59: 427-437.
Carpenter, J.M. (1992). Random cladistics. Cladistics
8: 147-153.
Carpenter, J.M. (1996). Uninformative bootstrapping.
Cladistics 12: 177-181.
Cocks, L.R.M. (1981). The evolving earth. Cambridge.
Colinveaux, P.A. (1998). A new vicariance model for
Amazon endemics. Global Ecol. Biogeogr. Letters
7: 95-96.
Cracraft, J. (1973). Continental drift, paleoclimatology, and the evolution and biogeography of birds.
J. Zool. Lond.169: 455-545.
Cracraft, J. (1985). Historical biogeography and patterns of differentiation within the South American
avifauna: areas of endemism. Ornithol. Monogr. 36:
49-84.
Crisci, J.V, M.M. Cigliano, J.J. Morrone and S. RoigJuent (1991). Historical biogeography of southern
South America. Syst. Zool. 48(2): 152-171.
Dalziel, I.W. (1982). West Antarctica: Problem child
of Gondwanaland. Tectonics 1: 3-19.
Dana, J.D. (1852-53). Isopoda, Part 2. pp. 696-805 and
plates XLVI-LIII in: United States exploring expedition during the years 1838, 1839, 1840, 1841,
1842, under the command of Charles Wilkes, U.S.N.
vol. 13, Crustacea.
Darlington, P.J. (1957). Zoogeography: the geographic
distribution of animals. New York.
Darlington, P.J. (1965). Biogeography of the southern
end of the world. New Haven.
Davis, R.C. (1984). Effects of weather and habitat
structure on the population dynamics of isopods in a
dune grassland. Oikos 42: 387-395.
de Wit, M.J. (1977). The evolution of the Scotia Arc as
a key to the reconstruction of southwestern Gondwanaland. Tectonophysics 37: 53-81.
Dixon, J. (1979). Origin and distribution of reptiles in
lowland tropical rain forests in South America.
Mus. Nat. Hist., Univ. Kansas, Monogr. 7: 217-240.
Dollfus, A. (1890). Isopodes terrestres du "Challenger". Soc. etudes sci. Paris 12: 1-8.
Dollfus, A. (1893). Vogage de M. E. Simon au Venezuela. Ann. Soc. Entomol. France 62: 339-346.
Dollfus, A. (1894). Viaggio del Dott. Alfredo Borelli
nella Repubblica Argentina e nel Paraguay. VI. Iso-
78
79
Holthuis, L.B. (1949). On a small collection of Isopoda Crustacea from the greenhouses of the Royal
Botanic Gardens, Kew. Ann. Mag. Nat. Hist., Ser.
11, 8: 122-137.
Jackson, H.G. (1927). Terrestrial Isopoda in: British
Museum of Natural History (ed.). Insects of Samoa,
part 8.1: 1-11.
Jackson, H.G. (1928). The morphology of the isopod
head. Part II. The terrestrial isopods. Proc. zool
Soc. London 1928: 561-595.
Kamp, P.J.J. (1980). Pacifica and New Zealand: proposed eastern elements in Gondwanaland's history.
Nature 288: 659-664.
Kellogg, J.N. & W.E. Bonini (1982). Subduction of
the Caribbean plate and the basement uplifts in the
overriding South American plate. Tectonics 1(3):
251-276.
Kesselyk, A. (1930). Halophilosciinae n. subfam. fr
die Gattungen Halophiloscia und Stenophiloscia.
Zool. Anz. 92: 282-286.
Laurent, R.F. (1979). Herpetofaunal relationships between Africa and South America in: The South
American Herpetofauna: Its origin, evolution, and
dispersal. pp. 55-71. Edited by W.E. Duellman.
Mon. Mus. Nat. Hist. Univ. Kansas 7.
Legrand, J.J. (1946). Les coaptations sexuelles des
Oniscoidea. Bull. biol. France Belg. 80:241-388.
Leistikow, A. (1997a). Terrestrial Isopods from Costa
Rica and redescription of Ischioscia variegata
(Dollfus, 1893); Canad. J. Zool. 75: 1415-64.
Leistikow, A. (1997b). Description of the terrestrial
isopod Mirtana costaricensis gen. et sp. n. from
Central America. Stud. neotrop. Fauna Environm.
32(2): 118-127.
Leistikow, A. (1997c). A new species of the genus
Reductoniscus Kesselyk, 1930 from Sabah, North
Borneo, Malaysia (Isopoda: Oniscidea: Armadillidae). Rev. suisse Zool. 104(4): 785-793.
Leistikow, A. (1998a). Redescriptions of Terrestrial
Isopoda (Crustacea: Oniscidea) from Chile and
Peru. Spixiana 21(3): 215-225.
Leistikow, A. (1998b). Considerations about the genus
Pentoniscus Richardson, 1913 (Crustacea: Isopoda:
Oniscidea) with description of a new species. J. Nat.
Hist. 32: 1339-1355.
Leistikow, A. (1998c). The genus Pseudophiloscia
Budde-Lund, 1912 in South America. Mitt. Mus.
Nat.kde. Berlin, zool. Reihe 74(2): 233-241.
Leistikow, A. (1999a). A new generic placement for
Philsocia richmondi Richardson, 1901 from Puerto
Rico (Crustacea: Isopoda: Oniscidea). J. Nat. Hist.
33: 1451-1460.
80
81
82
83
84
85