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HARVARD UNIVERSITY

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SPIXIANA °» Band 24 ® Heft1 » 1-96 *° München, 01. März 2001 » ISSN 0341-8391

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Umbellula monocephalus Pasternak, 1964, eine seltene Pennatularia
aus dem südlichen Westeuropäischen Becken

(Anthozoa, Octocorallia, Pennatularia)
Ludwig Tiefenbacher

Tiefenbacher, L. (2001): Umbellula monocephalus Pasternak, 1964, a rare Pennatu-
laria from the southern West European Basin (Anthozoa, Octocorallia, Pennatu-
laria). —- Spixiana 24/1: 1-4

During cruise 198 of F.R.V.“Walther Herwig III” in August/ September 1998 in
the southern part of the West Europe Basin two specimens of Umbellula monocepha-
lus Pasternak, 1964, this very rare pennatulacean octocoral of the abyssal, were
collected. Only eight specimens of this deep-sea species are known till now.

Dr. Ludwig Tiefenbacher, Zoologische Staatssammlung, Münchhausenstr. 21,
D-81247 München.

Einleitung

Die Pennatularia, zu deutsch Seefedern, sind hochspezialisierte sessile Anthozoa des Benthos, die über
alle Ozeane vom Flachwasser bis zum Abyssal verbreitet sind. Sie sind angepaßt an das Leben auf
Sediment wie Sanden und Schlick, in das sie mit ihrem Stiel teilweise eingebettet sind. Die Pennatularia
bilden üblicherweise Tierstöcke mit mehreren bis vielen Polypen. Innerhalb der Familie Umbellulidae
findet sich jedoch mit Umbellula monocephalus Pasternak, 1964, eine ‘Ausnahme’. Umbellula monocepha-
lus besitzt auf dünnem, hohen Stiel anscheinend nur einen einzigen großen Polypen.

Umbellula monocephalus Pasternak, 1964

Das der Originalbeschreibung von Pasternak zugrunde liegende Exemplar von Umbellula monocephalus
wurde von F.S. “Vitjaz” auf der Expedition 1959-1961 im Indischen Ozean südlich von Sri Lanka
(“Station 4911; 01°55'S/83°05'0”) aus einer Tiefe von “4809-4794 m” gefangen. Von der gleichen Expe-
dition, Station 4360 (“03°10'N/67°00'0; Tiefe 3490 m”), westlich der Malediven, führt Pasternak ein
weiteres Exemplar an.

Broch beschreibt 1957 eine Umbellula durissima Kölliker, 1880, nördlich von NO-Brasilien, die von
der Swedish Deep-Sea Expedition (“Station 357 (Haul No 6), 26.-27.07. 1948: N 02°26'/W 39°26' —
N 02°24'/W 39°12'. Depth 4474-4430 m”) eingebracht wurde. Er vermerkt für dieses Exemplar: “ Only
one specimen of this very characteristic species was brought home by the Swedish Atlantic expedition,
and the specimen is defective, the lower part wanting, and, with its single autozooid it seems a little
aberrant.” Er erkannte also nicht, daß er eine noch unbekannte Art vor sich hatte, obwohl ihm das
‘einzelne’ Autozooid aufgefallen war.

Grasshoff (1972) erhielt von der “Meteor”- Expedition 19 unter anderen 3 Exemplare von Seefedern
(“Ostatlantik, Horseshoe-Ebene; 33°46'N, 15°33'W; 3910-4002 m. “Meteor” - St. 19 - 191, Agassiz-Irawl
125, Hj. Thiel leg. 7.11.1970”), die ebenfalls nur ein ‘einzelnes’ Autozooid aufwiesen und die er als neue

Art Umbellula thieli beschrieb. Dieser Art fügte er das oben genannte, von Broch (1957) als Umbellula
durissima Kölliker, 1880, beschriebene Exemplar als “Paratypoid” zu. Grasshoff kannte wohl den Titel
der Arbeit von Pasternak (1964), doch war die Publikation für ihn damals nicht beschaffbar und aus
dem Titel allein war die Beschreibung einer neuen Art nicht zu erschließen.

Anläßlich der Veröffentlichung der Ergebnisse der französischen BIOGAS-Expeditionen u.a. je-
doch korrigiert Grasshoff (1981a,b) seinen Irrtum von 1972 und ordnet Umbellula thieli als Synonym
Umbellula monocephalus zu und ebenso das von Broch als Umbellula durissima bezeichnete Exemplar.
Aus dem Material der BIOGAS-Expedition IV kann Grasshof zwei weitere Exemplare von Umbellula
monocephalus isolieren, von denen er das eine “mit ca. 25cm Polypenlänge” als “das größte bisher
gefundene” (“CP 17°) bezeichnet und von dem zweiten (“CP 15”) angibt, daß es nur wenig kleiner ist
und sein Stiel mit ca. 52 cm Länge weitgehend erhalten ist. [“CP 15; Stn.4; 46°34'N, 10°26'W; 4715 m.
CP 17, Stn.; 46°31'N / 10°19'W; 4706 m”. Die Koordinaten wurden von mir aus der “Karte 17” (Grasshoff,
1981b) herausgemessen.].

Während der Expedition 198 mit F.F.S. "Walther Herwig III” konnten nun auf Station 50 (Hol 8) mit
einem Agassiz-Trawl nach über 20 Jahren seit den letzten Nachweisen erstmals wieder zwei Exemplare
dieser bisher außerordentlich selten gefundenen und offensichtlich ausschließlich auf das Abyssal als
Lebensraum beschränkten Art erbeutet werden.

Material

Fangdaten: 08.09.1998; Fangzeit (Agassiz-Trawl am Boden!) 00.10-04.00 UTC; Koordinaten: Anfang:
46°02.69'N / 16°44.95'W — 45°55.75'N / 16°41.40'W; 4635,75 m — 4704.25 m. Bodenbeschaffenheit: Etwas
verfestigter, blaugrauer Tiefseeton.

Die beiden Exemplare (Abb. 1A,B) wurden an Bord in 4%igem Formalin fixiert und an Land in 70 %
Alkohol übergeführt. Die folgenden Maße wurden an den fixierten Tieren abgenommen. Die Stiele
sind offensichtlich knapp über der Basis abgebrochen, da die vorhandenen Enden der Stiele einen
etwas größeren Durchmesser aufweisen.

Exemplar A: Abbruchstelle des Stiels ® 2 mm; Mitte Stiel © 1,5 mm; Stiel unter Polypenbasis
&4mm; Stiellänge von Bruchstelle bis Polypenbasis 153 mm; Länge der Rhachis (Polypenbasis bis
Ende der Achse) 43 mm; Ende der Rhachis bis Tentakelspitze 40 mm; Übergang Rhachis - Tentakelkro-
ne © 16 mm.

Exemplar B: Abbruchstelle des Stiels & 3 mm; Mitte Stiel © 1,5 mm; Stiel unter Polypenbasis
&6 mm; Stiellänge von Bruchstelle bis Polypenbasis 182 mm; Länge der Rhachis 64 mm; Ende der
Rhachis bis Tentakelspitze 60 mm. Übergang Rhachis - Tentakelkrone © 21 mm.

Die beiden Exemplare werden in der Zoologischen Staatssammlung, München, aufbewahrt.

Diskussion

Umbellula monocephalus gehört zu der Gruppe der Umbellulidae, die sich durch Sklerite in der Außen-
wand des Stiels, der Polypen und ihrer Tentakeln auszeichnen. Bei Umbellula monocephalus fällt dabei
die Masse und die dichte Packung der Sklerite auf. Im Querschnitt sind die Sklerite rund. Die großen
Sklerite der Tentakeln sind an einem, teilweise auch an beiden Enden etwas keulenförmig verdickt. Die
ebenfalls langen, jedoch im Vergleich zu den genannten etwa um die Hälfte bis ein Drittel schlankeren
Sklerite der Polypenwand sind spindelförmig. In gleicher Weise geformt, aber nur '/s bis '/ı so lang
sind die Sklerite an Stiel und Rhachis. Die Oberfläche der Sklerite erscheint bei stärkerer Vergrößerung
genoppt. Pasternak (1964) und Grasshoff (1972) geben hierzu detaillierte Abbildungen.

Grasshoff (1972) sieht als Vorläufer von Umbellula monocephalus (= Synonym von Umbellula thieli)
eine Art, die Umbellula durissima Kölliker, 1880 nahegestanden haben muß. “Die Größe der Polypen,”
... “die runde Hornachse und die großen, runden Sklerite in den Wänden und Tentakeln” sprechen
wohl dafür. “Unter Beibehaltung bzw. Verstärkung der genannten Eigenheiten kam es zu einer
Vergrößerung” des Primärpolypen im Laufe der Evolution “und zur Reduzierung der übrigen
Polypen.” Grasshoff (1972) stellt für U. thieli die Frage: “Tierkolonie — Solitärer Organismus?” und
vermerkt: “diese Seefeder erscheint als solitäres Wesen.” Er erklärt jedoch weiter, daß der Primärpolyp,
der “äußerlich nicht als solcher erkennbar”, den Stiel, die Hornachse, den Autozooiden “und die

Abb. 1. Die Exemplare A und B von Umbellula monocephalus Pasternak, 1964 aus dem südlichen Westeuropäis-
chen Becken. phot. M. Müller, Zoologische Staatssammlung, München.

unzähligen Siphonozooide” ... “in der Außernwand des Stiels und des großen Autozooiden produ-
ziert. Die Siphonozooide sind so stark reduziert, daß sie “nachgerade als Körperorgane des einen
großen Polypen” erscheinen und damit “mit der typischen Gestalt eines Oktokorallen - Polypen kaum
noch etwas gemein haben.” Es ist “die Lebensform eines solitären Organismus” entstanden, aber eben
nur das ‘Erscheinungsbild‘ dessen.

Im gleichen Fang, aus dem die beiden Exemplare von Umbellula monocephalus stammen, fanden sich
neben mehreren Holothuroidea und Asteroidea, die nicht näher bestimmt wurden, folgende Crusta-
cea, Decapoda (Tiefenbacher, 2001): 1 Carapax von Willemoesia leptodactyla (Willemoes-Suhm, 1873),
1 Männchen von Parapagurus abyssorum (Filhol, 1885), 1 Männchen und 2 Weibchen von Munidopsis
crassa Smith, 1885, das erste bekannt gewordene, eiertragende Weibchen von Munidopsis parfaiti
(A. Milne-Edwards & Bouvier, 1894), 1 Männchen und 2 Weibchen von Benthesicymus iridescens Bate,
1881 und 1 Weibchen von Plesiopenaeus armatus (Bate, 1881). Die überwiegende Anzahl dieser Orga-
nismen sind Sedimentfresser. Das Gleiche ist auch wahrscheinlich für Umbellula monocephalus. Grass-
hoff (1972) ist ebenfalls dieser Ansicht.

Danksagung

Herrn Dr. M. Grasshoff, Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt a.M., danke ich ganz
herzlich für seine freundliche Hilfe bei der Beschaffung eines Teils der angegebenen Literatur und für das
Entleihen seines Typenmaterials von U. thieli. In gleicher Weise gilt mein Dank Herrn Dr. Ted von Proschwitz,
Naturhistoriska Museet, Göteborg, Schweden, für das Entleihen des von Broch (1957) als U. durissima beschrie-
benen Exemplars.

Literatur

Broch, H. 1957. Pennatularians (Umbellula). - Rep. Swed. Deep-Sea Exp. 1947-48, Göteborg 2 (3): 347-364

Pasternak, Th. A. 1964. The deep-sea Pennatularians and Antipatharians obtained by R/S “Vitjaz” in the Indian
Ocean and the resemblance between the faunas of the Pennatularians of the Indian Ocean and the Pacific.
— Trudy Inst. Okeanol., Moskau 69: 183-215 (russ.)

Grasshoff, M. 1972. Eine Seefeder mit einem einzigen Polypen: Umbellula thieli n. sp. Die von F.S. “Meteor” 1967-
1970 im östlichen Nordatlantik gedredschten Pennatularia (Cnidaria: Anthozoa). - “Meteor” Forsch.-
Ergebnisse, Reihe D, No. 12: 1-11

-- 1981la (1982). Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der Biskaya (Cnidaria,
Anthozoa). Ergebnisse der französischen Expeditionen Biogas, Polygas, Geomanche, Incal, Noratlante und
Fahrten der “Thalassa”. I. Allgemeiner Teil. - Bull. Mus. natn. Hist. nat., Paris, 4° ser., 3, 1981, sect. A,n° 3:
731-766

-- 1981b (1982). Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der Biskaya (Cnidaria,
Anthozoa). Ergebnisse der französischen Expeditionen Biogas, Polygas, Geomanche, Incal, Noratlante und
Fahrten der “Thalassa”. II. Taxonomischer Teil. - Bull. Mus. natn. Hist. nat., Paris, 4° ser., 3, 1981, sect. A,
n° 4: 941-978

Tiefenbacher, L. 2001. Recent samples of mainly rare decapod Crustacea taken from the deep-sea floor of the
southern West Europe Basin. - Proceedings of the 7" Colloquium Crustacea Decapoda Mediterranea,
Hydrobiologia: (im Druck)

SPIXIANA München, 01. März 2001 ISSN 0341-8391

Eine neue Heteronemertine von der Küste Japans

(Nemertini)
Wolfgang Senz

Senz, W. (2001): Eine neue Heteronemertine (Nemertini) von der Küste Ja-
pans. — Spixiana 24/1: 5-13

Lineus nipponensis, spec. nov. from the coast of Japan is described and illustrated.
Characters of special interest are: the mouth opening lies a short distance behind
the brain; the preseptal outer longitudinal muscle layer includes a well-developed
circular musculature; brain lies distal to the circular muscle layer of the body wall.

Dr. Wolfgang Senz, Zoologisches Institut, Universität Wien, Althanstraße 14,
A-1090 Wien.

Einleitung

Die Nemertinenfauna der Küste Japans ist vor allem von U. Takakura, T. Yamaoka und F. Iwata
erforscht worden. Wie für beinahe alle Meeresgebiete, so gilt auch für die Küste Japans, daß die
Nemertinenfauna erst teilweise bekannt ist. In vorliegendem Aufsatz wird eine für die Wissenschaft
neue Nemertinenart dieses Küstengebietes beschrieben.

Material und Methoden

Der Untersuchung liegen zwei Individuen zugrunde. Von dem Körpervorderende und Teilen des
Mitteldarmbereichs jedes Tieres sind histologische Schnittserien hergestellt worden. Die Schnittserien
(Einbettung in Paraplast, Schnittdicke: 10 um) wurden in Haematoxylin-Eosin gefärbt. Das Material ist
in der Naturhistorisches Museum Wien — Evertebrata Varia Sammlung (NHMW-EV) aufbewahrt. Es
ist an der Küste Japans von dem Arzt v. Roretz gesammelt worden; Acquisitionsjahr (Naturhistorisches
Museum Wien): 1881.

Lineus nipponensis spec. nov.
Abb. 1-9

Typen. Holotypus: NHMW-EV 17026/3990. — Paratypus: NHMW-EV 17027/3991.
Etymologie. Die Art ist nach dem Fundgebiet benannt.

Diagnose. Laterale Kopfspalten flach und nicht unmittelbar bis zu den Cerebralorganen reichend;
äußere Längsmuskelschicht ohne Bindegewebeschicht; starke Ringmuskulatur in der äußeren Längs-
muskelschicht des Preseptalbereiches; Ringmuskelschicht der Körperwand stark entwickelt; Mundöff-
nung signifikant hinter dem Gehirn liegend; Gehirn vollständig distal der Ringmuskelschicht der
Körperwand positioniert; äußeres Neurilemma des Gehirns sehr gering entwickelt; Cerebralorgane
lediglich mit ihren Hinterenden gegen die Seitengefäße vordringend; Ocellen fehlen; Rhynchodaeum

in zwei Abschnitte unterteilt; Rüssel aus vorderem Abschnitt und Hauptabschnitt bestehend; Rüssel-
epithel ohne rhabditoide Strukturen; Rüssel mit zwei Muskelkreuzen.

Beschreibung

Äußere Erscheinung. Die Körperlänge beträgt knapp 10cm. Der Körper ist in der Vorderdarm- und
der vordersten Mitteldarmregion weitgehend zylindrisch (Durchmesser: 2,8mm); dahinter tritt eine
deutliche Abflachung auf (Breite: 3,6 mm, Höhe: 1,4mm). Seitliche Kanten des Körpers fehlen. Die
Mundöffnung ist relativ groß. Ein Paar lateraler Kopfspalten ist ausgebildet. Sie sind flach und reichen
von der Kopfspitze bis deutlich vor die Mundöffnung. Ein Caudalecirrus fehlt. Die fixierten Tiere sind
von einheitlich gelblich-weißer Farbe.

Körperwand. Die Epidermis weist keine Besonderheiten auf. Die distale Basalmembran der äußeren
Längsmuskelschicht ist an den histologischen Schnitten kaum erkennbar; die proximale Basalmembran
ist geringfügig besser ausgebildet. Im Preseptalbereich weist die äußere Längsmuskelschicht eine
kräftige Ringmuskulatur auf (Abb. 1, 3). Diese besteht in der Kopfspitze aus locker angeordneten, mehr
oder weniger dorsoventral orientierten Fasern (Abb. 1), seitlich des Rhynchodaeums. Nach hinten zu
entwickelt sich hieraus ein kompakter Muskelzylinder. Dieser liegt im distalen Drittel der äußeren
Längsmuskelschicht. Im Bereich der proximalen Kopfspaltenden fächert die Ringmuskulatur teilweise
auf, so daß einige Fasern ober- bzw. unterhalb der Kopfspalten distad ziehen (Abb. 1). Die Ringmus-
kulatur reicht bis in die Gehirnregion zurück. In ihrem Bereich befinden sich die meisten der insgesamt
wenigen Dermaldrüsenzellen.

Im Vorderdarm- und vorderen Mitteldarmbereich ist die äußere Längsmuskelschicht knapp dop-
pelt so dick wie die sehr gut entwickelte Ring- und die Längsmuskelschicht der Körperwand zusam-
men. In diesem Bereich dominieren im muskulären Teil der äußeren Längsmuskelschicht Längs- und
Radiärmuskelfasern, wie auch radiär angeordnete Bindegewebestränge. Letztere sind in geringem
Ausmaß miteinander verflochten. Im distalen Bereich der äußeren Längsmuskulatur tritt ein Komplex
aus Bindegewebeelementen, Ringmuskelfasern und einigen Dermaldrüsen auf. Im Paratypus sind die
Dermaldrüsen stellenweise signifikant zahlreicher ausgebildet als im Holotypus. Eine Bindegewebe-
schicht zwischen dem muskulären Teil der äußeren Längsmuskulatur und den Dermaldrüsen fehlt.
Dorsoventrale Muskel- und Bindegewebefasern, wie sie zum Beispiel bei Cerebratulus Renier, 1804
auftreten, fehlen. Dies gilt für die gesamte äußere Längsmuskelschicht

Hinter dem vorderen Mitteldarmbereich wird die äußere Längsmuskulatur zusehends dünner,
wobei sie vor allem ihre Radiärmuskulatur verliert, wie auch der proximale Gewebemantel stark
reduziert wird.

Etwa auf halber Höhe des Preseptalbereiches, das Rhynchodaeum erfährt hier eine abrupte Einen-
gung, tritt das Vorderende der Dorsalkommissur des Gefäßsystems auf. Zudem liegt hier das Vorder-
ende des Zentralzylinders, in Form seines dorsalen Bogens (Abb. 1d). Dieser umwächst nach hinten zu
die Dorsalkommissur und das Rhynchodaeum ventrad. Die Ringmuskulatur des Zentralzylinders ist
in ihrem Ursprungsgebiet schwach entwickelt, dahinter stärker, wobei sie tangential in die äußere
Längsmuskelschicht ausstrahlt. Die Längsmuskelschicht des Zentralzylinders bildet zunächst einen
einheitlichen Mantel um Rhynchodaeum und Dorsalkommissur. Nach hinten zu wird dieser weitge-
hend in ein Paar dorsolaterale Pakete und ventrolaterale Teile differenziert (Abb. 3). Letztere sind nicht
überall eindeutig gegenüber der Längsmuskulatur des Rhynchodaeums abgrenzbar. Die dorsomedia-
ne Verdrängung der Längsmuskulatur ist durch eine Erweiterung der Dorsalkommissur des Gefäßsy-
stems bedingt. Im hinteren Drittel des Preseptalbereiches steigt das Rhynchodaeum innerhalb des
Zentralzylinders in eine zentrale Position auf (Abb. 1f), die Dorsalkommissur beendend. Seitlich des
Rhynchodaeums liegen somit die großen Seitengefäße. Der dieserart gegebene Komplex wird von der
nun dünnen Längsmuskulatur des Zentralzylinders umgrenzt. Mediodorsal und -ventral fließt sie mit
der Längsmuskulatur des Rhynchodaeums zusammen.

Die Längs- und die Ringmuskelschicht der Körperwand sind im Gehirnbereich relativ schwach
entwickelt (Abb. 5). Dahinter, aber noch vor der Mundöffnung, gewinnen sie, abgesehen von ihren
ventromedianen Teilen, an Dicke (Abb. 4). Mit dem Auftreten der Mundbucht wird der ventromediane
Teil der Ringmuskelschicht zum Muskelbalken differenziert, wird also von der übrigen Muskelschicht
abgetrennt. Deren blinde ventrolateralen Enden - wie auch jene der Längsmuskelschicht - umwachsen
die Mundbucht ventrad, so daß hinter der Mundöffnung wieder ein geschlossener Muskelzylinder
auftritt. Der Muskelbalken selbst ist median mit der ventralen Längsmuskelplatte verflochten und

Abb. 1. Lineus nipponensis, spec. nov.: Darstellung einiger Organe des Preseptalbereiches (nach Zeichnungen
mit dem Zeichenspiegel, schematisiert). a = 12. Schnitt (Stern: Rhynchodealöffnung; Pfeilspitzen: Kopfspalten),
b = 20. (Stern = Rhynchodaeum,; Pfeilspitzen: Kopfspalten), ce = 35. (Pfeilspitze: mögliches Frontalorgan an der
Hinterwand des vorderen Rhynchodaeum-Abschnittes), d=40. (Stern = Rhynchodaeum; Pfeilspitze: Vorder-
ende der Dorsalkommissur des Gefäßsystems), e=46. (Stern = Rhynchodaeum; Pfeilspitze: ventraler Zusam-
menschluß des Zentralzylinders) (vgl. Abb. 3), £f=90. (Pfeilspitzen: Seitengefäße); Maßstab: 1 mm; Abkür-
zungen: dm = dorsoventral orientierte Muskelfasern (Ursprung der Ringmuskulatur der äußeren Längsmuskel-
schicht), rm = Ringmuskulatur der äußeren Längsmuskelschicht, rs = Rhynchodealsphinkter, vzz = Vorderende
des Zentralzylinders (dorsaler Bogen).

kann bis in den Vorderdarmbereich zurück verfolgt werden. Seitlich bildet der Balken Muskelfahnen
aus, das Vorderende der Vorderdarm-Radiärmuskulatur.

Von der inneren Ringmuskelschicht sind in geringem Umfang Horizontal- und Dorsoventralmus-
keln des Gehirnbereichs, sowie Dorsoventralmuskeln des Mitteldarmbereichs (Abb. 8, 9) ausgebildet.
Letztere formen zwischen den Seitentaschen des Mitteldarms teilweise gut entwickelte Muskelblätter
aus.

Die ventrale Längsmuskelplatte nimmt ihren Ursprung von jenen Längsmuskelfasern der Körper-
wand-Längsmuskelschicht, die zwischen den Seitengefäßen und dem Vorderende des Rhynchocoels
liegen (Abb. 4). Im Vorderdarmbereich gehen die Seitenränder der Muskelplatte kontinuierlich in die
Vorderdarm-Längsmuskulatur über. Im Vorderdarm- und vorderen Mitteldarmbereich ist die Mus-
kelplatte gut entwickelt. Dahinter sind ihre Seitenteile stark reduziert.

Zentralraum und Mesenchym. Im Vorderdarmbereich treten aufgrund des Vorderdarm-Gefäßnetzes
einfache Leisten auf (Abb. 7). Im Mitteldarmbereich sind in Zusammenhang mit den Seitentaschen des
Mitteldarms gut entwickelte Leisten ausgebildet (Abb. 8). Trotz der offenen Zentralraum-Organisation
fehlt Mesenchym in nennenswerten Kontingenten weitestgehend.

Darmtrakt. Die Mundöffnung liegt deutlich hinter dem Gehirn. Die Distanz zum Gehirn ist aber
wesentlich kürzer als jene vom Gehirn zum Körpervorderende. Die Mundöffnung ist relativ groß. Die
Wand der Mundbucht ist drüsenreich und geringfügig in Falten gelegt. Der Mundbucht schließt der
gerade nach hinten ziehende Vorderdarm an. Dessen Wand, sie weist kaum Faltenbildung auf, besitzt
subepitheliale Drüsen (Abb. 7). Die Vorderdarm-Muskulatur besteht aus Ring-, Längs- und Radiär-
muskulatur (Abb. 7). Die Radiärmuskeln sind zumeist kräftig. Der Vorderdarmwand liegt die Ring-
muskulatur an, der die Längsmuskulatur folgt. Letztere bildet eine fast vollständige Ummantelung der
ventralen und lateralen Vorderdarmwand. Da diese Muskulatur beinahe kontinuierlich in die Seiten-
ränder der ventralen Längsmuskelplatte übergeht, ist der Vorderdarm allseits von Längsmuskulatur
umgeben.

Der Übergang in den Mitteldarm erfolgt kontinuierlich. Abgesehen von seinem vordersten Bereich
weist der Mitteldarm relativ tiefe, geringfügig in Falten gelegte Seitentaschen auf (Abb. 8, 9). Sie sind
tiefer, als das Zentralrohr des Mitteldarms breit ist.

Rüsselapparat. An der Kopfspitze liegt eine große Öffnung (Abb. 1a), an die ein depresser, waagerecht
nach hinten ziehender Kanal anschließt (Abb. 1b). Dieser besitzt ein relativ dünnes Epithel. Etwa in der
Mitte des Preseptalbereiches erfährt der Kanal eine abrupte Einengung auf seinen ventralen Bereich
(Abb. Ic). Die hierdurch gegebene Rückwand des Kanales bildet Falten aus, die möglicherweise ein
Frontalorgan beinhalten (Abb. 1c). Die Fortsetzung des Kanals ist zunächst ein wesentlich dünneres
Rohr (Abb. 1d, e), das sich caudad dorsad erweitert (Abb. If). Letztlich reicht es (im Querschnitt) bis
zum mediodorsalen Teil des Zentralzylinders. Dieserart erstreckt es sich bis zum Septum. Der gesamte
Kanal - von der Körperspitze bis zum Septum - ist das Rhynchodaeum (vgl. unten). Das Epithel des
vorderen Rhynchodaeum-Abschnittes — also jenem vor dem Zentralzylinder - ist deutlich dünner als
jenes des hinteren Abschnittes. Zudem ist der hintere Rhynchodaeum-Abschnitt im Querschnitt nicht
depress geformt. Beide Übergänge erfolgen aber kontinuierlich. Dem vorderen Rhynchodaeum-Ab-
schnitt liegen Längsmuskelfasern an. Der dorsale Teil dieser Muskulatur fließt in das Vorderende der
Längsmuskelschicht des Zentralzylinders ein. Dieser befindet sich dorsal des Vorderendes der Dorsal-
kommissur des Gefäßsystems. Eine kontinuierliche Verbindung mit der Längsmuskulatur des hinteren
Rhynchodaeum-Abschnittes ist nicht feststellbar. Ventral liegen dem Vorderende der Dorsalkommis-

Abb. 2-9. Lineus nipponensis, spec. nov. 2. Querschnitt durch den Hauptabschnitt des Rüssels; Maßstab: 0,4 mm;
Pfeilspitzen: Arme des stärker entwickelten Muskelkreuzes. 3. Querschnitt auf Höhe des Vorderendes des
hinteren Rhynchodaeum-Abschnittes (entspricht in etwa Abb. le); Maßstab: 0,3 mm; Pfeilspitzen: Ringmusku-
latur der äußeren Längsmuskelschicht. 4. Querschnitt durch den Bereich zwischen Gehirn und Mundbucht;
Maßstab: 0,4 mm; obere Pfeilspitze: Ursprung der ventralen Längsmuskelplatte; untere Pfeilspitze: Vorderdarm-
nerv. 5. Querschnitt durch den Cerebralorgankanal-Bereich; Maßstab: 0,4 mm; senkrechte Pfeilspitze: Cerebral-
organkanal; waagerechte Pfeilspitze: Körperwandmuskulatur proximal des dorsalen Nervenpols. 6. Quer-
schnitt durch den Mundbuchtbereich; Maßstab: 0,4 mm; obere Pfeilspitze: Rüsselsektor mit den grob granulierte
Sekretgranula enthaltenden Drüsen; untere Pfeilspitze: Rüsselsektor mit den blau färbbares Sekret enthaltenden
Drüsenzellen. 7. Querschnitt durch den Vorderdarmbereich; Maßstab: 0,4 mm; Pfeilspitzen: Radiärmuskeln der
Vorderdarm-Muskulatur. 8. Querschnitt durch die Mitteldarmregion; Maßstab: 0,5 mm; Pfeilspitze: Dorsoven-
tralmuskulatur. 9. Längsschnitt durch die Mitteldarmregion; Maßstab: 0,5 mm; Pfeilspitzen: Dorsoventral-
muskulatur; Sterne: Gonaden. Abkürzungen: alm = äußere Längsmuskelschicht der Körperwand, da = Dorsalast
des posterioren dorsalen Nervenpoles, dk = Dorsalkommissur des Gefäßsystems, go = Gonade, Im = Längsmus-
kelschicht der Körperwand, In = Längsnervenstrang, Inw = Längsnervenstrangwurzel, mdt = Mitteldarmtasche,
osg = oberster Ast des Vorderdarmgefäßnetzes, rc = Rhynchocoel, rd = Rhynchodaeum, rm = Ringmuskelschicht
der Körperwand, sg = Seitengefäß, svd = subepitheliale Drüsen des Vorderdarms, vdl = Vorderdarmlumen.

=
EN

DIET

sur des Gefäßsystemes aber Längsmuskelfasern an, wie sie auch weiter hinten in dieser Lage ausge-
bildet sind. Hier reicht das Rhynchodaeum aufgrund seiner posterioren Erweiterung bis nahe an die
Dorsalkommissur heran und besagte Längsmuskelfasern erweisen sich als solche des Rhyncho-
daeums. Insofern besteht eine gewisse Verbindung zwischen der Längsmuskulatur beider Rhyncho-
daeum-Abschnitte. Der hintere Abschnitt des Rhynchodaeums besitzt zudem Ringmuskulatur, die vor
dem Septum einen dicken Sphinkter ausbildet.

Das Septum ist beinahe geschlossen. Das Rhynchocoel weist weder Divertikel noch Diskontinui-
täten auf und reicht bis in den hinteren Körperbereich. Seine Wand besitzt eine Ring- und eine
Längsmuskelschicht (Abb. 4-6). Sie zeigen keine Verflechtung mit der Körperwandmuskulatur.

Der Rüssel ist nicht gespalten. Über seine gesamte Länge besteht seine Wand aus einem dicken
Epithel, einer anschließenden Ring- und Längsmuskelschicht, sowie dem Endothel. Ein relativ kurzer
vorderer Rüsselabschnitt - Durchmesser: 0,5 mm - (Abb. 6) und der Hauptabschnitt - Durchmesser bis
zu 1,0 mm - (Abb. 2) sind zu unterscheiden. Im vorderen Abschnitt bildet das Epithel vier Pakete aus,
von denen zwei - einander gegenüberliegende - Pakete wesentlich größer sind als die übrigen beiden.
In einem der beiden großen Pakete treten Drüsenzellen auf, mit im proximalen Epithelbereich liegen-
dem blau färbbarem Sekret. In den übrigen Paketen dominieren Drüsenzellen mit bräunlich färbbarem,
grob granuliertem Drüsensekret. In geringem Ausmaß können in diesen Paketen auch Zellen mit blau
färbbarem Sekret auftreten. Zumeist ist die Muskulatur jenes Rüsselsektors, der die Drüsenzellen mit
blau färbbarem Sekret enthält, deutlich schwächer entwickelt als in den übrigen Sektoren. Im vorderen
Rüsselabschnitt sind die Rüsselnerven kaum identifizierbar, wie auch die beiden Muskelkreuze
schwach entwickelt sind. Zumeist aber ist das Muskelkreuz in dem Sektor mit den grob granuliertes
Sekret enthaltenden Drüsenzellen kräftiger ausgebildet als das diesem gegenüberliegende Muskel-
kreuz.

Im Hauptabschnitt des Rüssels sind die Rüsselnerven und die Muskelkreuze deutlich besser
entwickelt, wobei das Nerven- und das Muskelkreuz-Paar im rechten Winkel zueinander angeordnet
sind. Die beiden Muskelkreuze sind fortgesetzt unterschiedlich stark ausgebildet. Die Paketbildung im
Bereich des Rüsselepithels geht verloren, da die grob granulierten Drüsenzellen nun in weitaus
geringerer Anzahl auftreten, wohingegen die Zellen mit blau färbbarem Sekret über das gesamte
Epithel verstreut vorliegen. Die Rüsselwand ist nun deutlich stärker als davor in Falten gelegt.

Ein Retraktormuskel ist ausgebildet.

Nervensystem. Im Bereich des anterioren Nervenpols entspricht jede Gehirnhälfte (Terminologie zur
Gehirnanatomie gemäß Senz & Tröstl 1997) im Querschnitt einem vergleichsweise großen Längsner-
venstrang. In diesem Gehirnbereich befindet sich die relativ dicke Dorsalkommissur. Die Faserkerne
der Gehirnhälften weisen erst hinter dem Vorderende des Mittelteils des Gehirns, also in etwa hinter
dem Vorderende der Ventralkommissur, eine nennenswerte Kompartimentierung auf. Die Ventral-
kommissur besitzt ventral ihres Faserkernes Ganglienzellkörper. Am Hinterende des Mittelteils des
Gehirns tritt eine Aufspaltung in den dorsalen und ventralen posterioren Nervenpol auf. Im Ur-
sprungsbereich ist der dorsale Pol im Querschnitt annähernd um die Hälfte größer als der ventrale Pol.
Der dorsale Pol spaltet terminal auf. In ihrem Ursprungsbereich sind seine beiden Äste ungefähr gleich
groß. Die Faserkerne beider Äste sind von einem einheitlichen Mantel aus Ganglienzellkörpern umge-
ben. Der dorsale Ast reicht bis knapp über das Vorderende des Cerebralorgans zurück. An seinem
Hinterende fließt er in die Nervenschicht der Körperwand ein. Dies ist im Paratypus umfangreicher
ausgebildet als im Holotypus. Der ventrale Ast geht in das Cerebralorgan über. Das Gehirn - somit
auch die Äste des dorsalen posterioren Nervenpoles - liegt vollständig distal der Ringmuskelschicht
der Körperwand (Abb. 5). Das innere Neurilemma des Gehirns ist zumeist gut entwickelt. Ein distink-
tes äußeres Neurilemma fehlt. Trotzdem ist eine relativ scharfe Grenze des Gehirns gegenüber der
äußeren Längsmuskelschicht gegeben.

Die Längsnervenstränge zweigen kontinuierlich aus dem ventralen posterioren Nervenpol des
Gehirns ab. Die Neurilemmata der Längsnervenstränge sind wie jene des Gehirns ausgebildet. Zwi-
schen dem Faserkern und den Ganglienzellpaketen der Längsnervenstränge treten Radiärmuskelfa-
sern auf. Neurochorde, Seitenstamm-Muskelfasern, accessorische Faserstränge und andere Besonder-
heiten fehlen. Der Dorsalnerv ist unmittelbar hinter dem Gehirn gut entwickelt, ansonst nur an
einzelnen Schnitten gegenüber der Nervenschicht der Körperwand histologisch abgrenzbar. Aus den
Innenwänden der Längsnervenstrangwurzeln zweigen die Vorderdarmnerven ab. Diese sind vor der
Mundöffnung miteinander verbunden. Ab der Mundbucht konnten die Vorderdarmnerven nicht mehr
erkannt werden. Das Ursprungsgebiet der Rüsselnerven konnte nicht eruiert werden.

Kopfdrüse und Sinnesorgane. Ein typisches Frontalorgan fehlt, doch mag das Hinterende des vorde-
ren Rhynchodaeum-Abschnittes ein Frontalorgan beinhalten (vgl. oben). Die Kopfdrüse ist auf den
Preseptalbereich beschränkt. Sie weist keine Besonderheiten auf.

Ein Paar lateraler Kopfspalten ist ausgebildet (Abb. 1). Sie reichen von der Kopfspitze bis zu den
Öffnungen der Cerebralorgankanäle. Sie sind auffallend flach. Ihr Epithel ist arm an Drüsenzellen.
Ganglienzellpolster bzw. Sinneszellpolster sind nicht mit den Kopfspalten assoziiert. Als einzige
Differenzierung in Zusammenhang mit den Kopfspalten tritt jene der Ringmuskulatur der äußeren
Längsmuskelschicht auf (vgl. oben). Knapp, aber signifikant vor den Cerebralorganen (teilweise
kontraktionsabhängig) enden die Kopfspalten in jeweils einer einfachen kolbenförmigen Erweiterung.
Diese erreichen das Gehirn nicht. Von jeder Erweiterung zweigt daher ein vergleichsweise langer
Cerebralorgankanal ab, der zwischen den dorsalen und ventralen posterioren Nervenpol des Gehirns
vordringt (Abb. 5). Auf gleicher Höhe spaltet der ventrale posteriore Nervenpol auf, wobei der ventrale
Ast zusammen mit dem Cerebralorgankanal nach hinten zu das eigentliche Cerebralorgan ausbildet.
Die Cerebralorgane sind gut entwickelt, liegen zunächst aber der Ringmuskelschicht der Körperwand
distal an. Diese - wie auch die Längsmuskelschicht der Körperwand - wird an den Berührungsstellen
mit den Cerebralorganen caudad dünner, wobei distal der Cerebralorgane Muskelfasern auftreten, die
der Ringmuskelschicht zuzuordnen sind. Teilweise stellen sie laterale Verlängerungen des dorsome-
dianen Bogens der Ringmuskelschicht der Körperwand dar. Im hinteren Bereich der Cerebralorgane
werden aus diesen Fasern dorsolaterale Bögen der Ringmuskulatur, die caudad jene proximal der
Cerebralorgane ersetzen. Dieserart gelangen die Hinterenden der Cerebralorgane proximad der Kör-
perwand bzw. in Kontakt mit den Seitengefäßen. Die Abgrenzung der Cerebralorgane gegenüber der
äußeren Längsmuskelschicht der Körperwand entspricht jener des Gehirns. Die Abgrenzung jedes
Cerebralorganes gegenüber dem dorsalen Ast des dorsalen posterioren Nervenpols ist erst im Bereich
von dessen Hinterende gegeben.

Ocellen und weitere Sinnesorgane sind nicht ausgebildet.

Gefäßsystem. Am Vorderende der Einengung des Rhynchodaeums tritt jenes der Dorsalkommissur
des Gefäßsystems auf (Abb. 1d). Die Dorsalkommissur nimmt jenen Platz ein, der durch die Veren-
gung des Rhynchodaeums frei wird. Caudad gewinnt das Rhynchodaeum wieder an Durchmesser,
womit die Spaltung der Dorsalkommissur in die beiden Seitengefäße einhergeht (Abb. 1f). Diese ziehen
seitlich des Rhynchodaeums bis zum Septum. Hinter diesem erweitern sie sich, womit die Ventralkom-
missur des Gefäßsystems zur Ausbildung gelangt. Von diesem zweigt das Dorsalgefäßs ab. Es steigt
sogleich gegen das Rhynchocoel auf (Abb. 6). Diese Lage verläßt es knapp hinter dem Vorderende des
Mitteldarms. Der Komplex aus den Seitengefäßen und der Ventralkommissur wird von den Seitentei-
len des Muskelbalkens in geringem Ausmaß zerklüftet. Hieraus erwächst nach hinten zu das Vorder-
darm-Gefäßnetz, sowie die Radiärmuskulatur. Das Gefäßnetz ist gut entwickelt (Abb. 7) und reicht bis
zum Vorderende des Mitteldarms. Im Mitteldarmbereich sind somit das Dorsalgefäß und die Seiten-
gefäße gegeben, die über seriale Kommissuren miteinander verbunden sind. Die Seitengefäße weisen
kaum eigene Muskulatur auf. Um das Dorsalgefäß ist die ventrale Längsmuskelplatte auch in jenem
Körperbereich gut entwickelt, in dem sie ansonst weitgehend reduziert ist.

Exkretionsapparat. Der Exkretionsapparat ist auf den hinteren Vorderdarmbereich beschränkt. Die
Kanäle des Exkretionsapparates liegen vor allem lateral im Körper, in direktem Kontakt mit dem
Gefäßnetz des Vorderdarms. Im hinteren Bereich des Exkretionsapparates besitzt dieser in jenen
beiden Gefäßen des Gefäßnetzes die unmittelbar neben dem Rhynchocoel liegen - also in den beiden
größten Gefäßen - je einen Hauptsammelkanal. Diese sind bis zu 0,25 mm dick und ziehen knapp über
das Hinterende des Vorderdarms hinaus. Soweit es den Schnittserien zu entnehmen ist, weist jede
Körperseite einen Ausführgang des Exkretionsapparates auf.

Fortpflanzungsapparat. In beiden untersuchten Individuen liegen zwischen den Seitentaschen des
Mitteldarms große, endothelial umkleidete Hohlräume, die Gonaden (Abb. 8, 9). In den Gonadenwän-
den treten einige undifferenzierte Geschlechtszellen auf. Weitere Geschlechtszellen fehlen. Gonoducte
konnten an den Schnittserien nicht festgestellt werden.

all

Diskussion

Grundlage der folgenden Diskussion ist, daß gegenwärtig keine Systematik des Taxons Nemertini
bzw. eines Nemertinentaxons geboten werden kann, so daß es bei einer klassifikatorischen Einteilung
belassen werden muß (vgl. Senz 2000 und hierin zitierte Literatur).

Aufgrund der äußeren Längsmuskelschicht der Körperwand, des Baus und der Lage der Cerebral-
organe sowie des Gefäßsystems ist das Material den Heteronemertinen zuzurechnen und innerhalb
dieser aufgrund der Rüsselmuskulatur in die Gruppe Lineidae sensu Gibson (1985) einzureihen.
Innerhalb der Lineidae ist es die Gattung Lineus Sowerby, 1806, deren Anatomie — gemäß der Gattungs-
definition von Gibson (1985, 1990a) - mit jener des untersuchten Materials vollständige Übereinstim-
mung aufweist.

“Amongst the 90 or more species which have been included in the genus Lineus, a comparatively
few have been adequately described and a secure diagnosis for the taxon is still not possible” (Gibson
1990b: 123). Dies erschwert die Identifikation des Materials auf Artniveau. Es zeichnet sich aber durch
eine Reihe von Merkmalen aus, die es von allen bisher beschriebenen Arten eindeutig unterscheiden
läßt. Diese Merkmale sind: Laterale Kopfspalten flach und nicht bis direkt zu den Cerebralorganen
reichend; äußere Längsmuskelschicht mit Ringmuskulatur; Rhynchodaeum in zwei Abschnitte unter-
gliedert; Mundöffnung signifikant hinter dem Gehirn positioniert; Ringmuskelschicht der Körper-
wand stark entwickelt; Gehirn vollständig distal der Ringmuskelschicht der Körperwand liegend;
Cerebralorgane nur mit ihren Hinterenden gegen die Seitengefäße vordringend. Für das Material wird
daher die Art Lineus nipponensis, spec. nov. eingerichtet.

Insbesondere die Anatomie der Kopfspalten, die Lage der Cerebralorgane und der Mundöffnung
sowie die Stärke der Ringmuskelschicht der Körperwand in L. nipponensis stimmen mit den Verhält-
nissen in Lineus molochinus Bürger, 1892 überein (vgl. Bürger 1895 für L. molochinus). Diesen Überein-
stimmungen stehen aber mehrere Unterschiede gegenüber (vgl. Tab. 1). In den angeführten Merkma-
len stimmt L. nipponensis zudem mit Lineus insignis Senz, 1993 überein (vgl. Senz 1993 für L. insignis).
Sichere Unterscheidungsmerkmale zwischen beiden Arten ergeben sich unter anderem aus der Rüssel-
Anatomie und jener der Kopfspalten im Gehirnbereich (vgl. zudem Tab. 1).

Zum Rhynchodaeum von L. nipponensis ist anzumerken: Alternativ zu der hier getroffenen Aussa-
ge, daß das Rhynchodaeum in zwei Abschnitte differenziert ist, kann begründet überlegt werden, ob
der hier so genannte vordere Rhynchodaeum-Abschnitt als Atrium zu bezeichnen ist, das Rhynchodae-

um also auf den Bereich ab dem Vorderende des Zentralzylinders beschränkt ist. Hierfür könnte |

angeführt werden, daß die Längsmuskulatur des vorderen Kanalabschnittes kontinuierlich in jene des
Zentralzylinders übergeht, nicht aber in jene des hinteren Kanalabschnittes (vgl. oben). Von dieser
Interpretation ist Abstand genommen worden, vor allem, da die Epithelien der einzelnen Kanalab-
schnitte keine wesentlichen histologischen Unterschiede aufweisen bzw. die auftretenden Unterschie-
de kontinuierlich ineinander übergehen. Die angeführte fehlende kontinuierliche Verbindung der
Längsmuskulatur der beiden Rhynchodaeum-Abschnitte ist demzufolge in Zusammenhang mit der
generell schwachen Entwicklung der Muskulatur proximal der äußeren Längsmuskelschicht im Be-

Tab.1. Vergleich von Lineus molochinus Bürger, 1892, Lineus insignis Senz, 1993 und Lineus nipponensis, spec. nov.

1; 2% 3. 4. 5. 6. v% 8. 9, 10. 11. 12.
L. molochinus + + + + + + + _ ? = 1 a!
L. insignis = _ + + + —_ + + + — 0 b
L. nipponensis + + + + + _ — + = + 2

1. = Kopfspalten flach und nicht bis zu den Cerebralorganen reichend; 2. = Ringmuskelschicht der Körperwand |

dick; 3. = Mundöffnung nicht unmittelbar hinter dem Gehirn; 4. = Cerebralorgane nur mit den Hinterenden
gegen die Seitengefäße vordringend; 5. = Ocellen fehlen; 6. = Gehirn mit Ventralverlagerung; 7. = dorsaler Ast
des posterioren dorsalen Nervenpoles auffallend groß; 8. = Ringmuskulatur in der preseptalen äußseren Längs-

muskelschicht vorhanden; 9. = Rhynchodaeum mit Untergliederung; 10. = äußeres Neurilemma des Gehirns |
stark entwickelt; 11. = Anzahl der Muskelkreuze des Rüssels; 12. = berichtetes Verbreitungsgebiet, a: Golf von |

Neapel, b: Küste Rovinjs, c: Küste Japans.

' “Isler’s report (1902: 278) of this species occuring in Chile is of doubtful validity” (Gibson 1995: 399).
? Die beiden Äste des dorsalen posterioren Nervenpols sind im Ursprungsgebiet aber gleich groß.

reich der Kanalverengung zu sehen. Es bleibt aber zu beachten, daß der Begriff “Atrium” in der
Nemertinenforschung erst vage definierbar ist. Gegenwärtig kann auch nicht angegeben werden,
welche funktionellen Aspekte mit der Untergliederung des preseptalen Kanals - sowie mit der hiermit
zusammenhängenden Konzentration des Zentralzylinders auf den Bereich des hinteren Rhynchodae-
um-Abschnittes — verbunden sind. Die Analyse hiervon müßte zudem auf mögliche funktionelle
Zusammenhänge hiervon mit der Ringmuskulatur der äußeren Längsmuskelschicht Bedacht nehmen,
eine weitere Struktur, die für Heteronemertinen untypisch ist.

Danksagung

Der Autor möchte sich bei Dr. Helmut Sattmann (Naturhistorisches Museum in Wien) für die Bereitstellung des
Materials bedanken.

Literatur

Bürger, ©. 1895. Die Nemertinen des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. - Fauna
Flora Golf. Neapel 22: 1-743

Gibson, R. 1985. The need for a standard approach to taxonomic descriptions of nemerteans. - Amer. Zool. 25:
5-14

-- 1990a. The macrobenthic nemertean fauna of the Albany region, Western Australia. In: Wells F. E., Walker
D. I., Kirkman H. and Lethbridge R. (Eds.): Proc. Third Int. Mar.-Biol. Workshop: The Marine Flora and
Fauna of Albany, Western Australia Perth. - Western Australian Museum 1: 89-194

-- 1990b. The macrobenthic nemertean fauna of Hong Kong. In: Morton B. (Ed.): Proc. Sec. Int. Mar.-Biol.
Workshop: The Marine Flora and Fauna of Hong Kong and Southern China; Hong Kong. - Hong Kong
University Press 1: 33-212.

-- 1995. Nemertean genera and species of the world: an annotated checklist of original names and description
citations, synonyms, current taxonomic status, habitats and recorded zoogeographic distribution. - J. Nat.
Hist. 29: 271-562

Isler, E. 1902. Die Nemertinen der Sammlung Plate. — Zool. Jb., Suppl. 5: 273-280

Senz, W. 1993. Nemertinen europäischer Küstenbereiche (nebst ergänzenden Angaben zur Anatomie von
Apatronemertes albimaculosa Wilfert & Gibson, 1974). - Ann. Naturhist. Mus. Wien 94/95B: 47-145

-- 2000. Neue Nemertinen aus dem Golf von Arabien. 1. Palaeonemertini. - Ann. Naturhist. Mus. Wien 102B:
321-373

-- &R.A. Tröstl 1997: Überlegungen zur Struktur des Gehirns und Orthogons der Nemertinen. - Sitzungsber.
Akad. Wiss., Wien, Naturwiss.-Math. Kl. 204: 63-78

Buchbesprechungen

1. Kornacker, P.M.: Checklist and key to the snakes of Venezuela - Lista sistemätica y clave para las serpientes |

de Venezuela. - PaKo-Verlag, Rheinbach, 1999. 270 S., 90 Farbfotos, 69 Textfig. u. Zeichn., ISBN 3-9806240-0-5.

Nach den “Schlangen von Venezuela” (Lancini & Kornacker 1989, Caracas [Armitano Editores], 381 S.) liegtnun

die zweite Bearbeitung dieser Tiergruppe aus der Feder Paul Kornackers vor. Geboten wird eine zweisprachig
(englisch/spanisch) verfaßte Liste der Schlangen des Landes im Paperback-Format; die für jedes Taxon (Gat-

tung, Art, Unterart) gegebenen Informationen umfassen dessen vollständige Synonymie (inklusive der
Cresonyme), sowie bei Arten und Unterarten die Angabe der terra typica und eine Beschreibung des Gesamt- |

areals nach Ländern. In einem zentralen Bildteil sind Farbphotos etwa der Hälfte der aufgeführten Taxa
zusammengestellt. Bestimmungsschlüssel ermöglichen eine Determination bis auf Unterartniveau. Schließlich

finden sich Listen und Graphiken zu den endemischen Formen des Landes, eine Liste der englischen und

spanischen Trivialnamen, sowie eine tabellarische Übersicht der Verbreitung der venezuelanischen Schlangen
in allen weiteren Staaten Lateinamerikas. Ein Index sowie ein über 600 Zitate umfassendes Literaturverzeichnis
runden das Buch ab.

Gegenüber der teils eher populärwissenschaftlich gehaltenen Darstellung von Lancini & Kornacker (1989)
werden mittlerweile 28 weitere, inzwischen für das Land neu nachgewiesene oder neu beschriebene Taxa
berücksichtigt (182 aktuell vs. 154). Weiterhin sind in der 1989’er Bearbeitung lediglich Bestimmungsschlüssel

enthalten, die eine Gattungs-Determination ermöglichen. Schließlich hätten wohl die jetzt präsentierten umfan-
greichen Synonymielisten sowie das detaillierte Literaturverzeichnis den Rahmen der älteren Venezuela-Mono-

graphie gesprengt bzw. einen Teil des seinerzeit anvisierten, breiten Leserkreises abgeschreckt. Damit wird klar,
daß die aktuelle Checkliste keineswegs als gleichrangige Konkurrenz zur ersten Monographie von 1989 zu
verstehen, sondern eher als eine Ergänzung dazu gedacht ist.

Die Farbphotos des zentralen Bildteils sind bis auf wenige Ausnahmen gut und lockern (wie auch die als
Textabbildungen eingegliederten, überaus genau gezeichneten Portraits) den ansonsten naturgemäß eher trock-
enen Stoff etwas auf. Dabei hätten die Unterschriften der Photos, die nur aus dem wissenschaftlichen Namen
der Art bestehen, aber teils etwas ausführlicher sein können: z.B. fehlen bei den Abb. 43-45 Verweise auf Jugend-
bzw. Adultfärbung bei Mastigodryas boddaerti und bei Abb. 73 ein Verweis auf den dort dargestellten Geburts-
vorgang bei Bothriechis schlegelii. Eine schöne Ergänzung des Buches wäre auch eine tabellarische Übersicht über
das Vorkommen der einzelnen Arten in den naturräumlichen Einheiten Venzuelas gewesen. Dies sind letztend-
lich aber nur randliche, kleinere Defizite, die den insgesamt sehr guten Gesamteindruck kaum schmälern
können.

Kornackers Checkliste ist aufgrund ihres relativ geringen Umfangs als Bestimmungswerk auf Exkursionen |

gut mitzuführen. Darüber hinaus machen die verschiedenen Listen das Werk auch zu einer Datengrundlage für

umfassendere Analysen zur Biodiversität. Schließlich ermöglicht das ausführliche Literaturverzeichnis auch
wenig erfahrenen Personen einen leichten Einstieg in die Schlangenfauna von Venezuela. Rundum also eine
schöne, professionelle (und aus wissenschaftlicher Sicht notwendige) Ergänzung der früheren Darstellung, die
in keiner Literatursammlung zur südamerikanischen Reptilienfauna fehlen darf. Es ist zu hoffen, daß besonders

durch die nun zur Verfügung stehenden, umfassenden Bestimmungshilfen die Erforschung der Schlangenfauna
des Landes weiterhin katalysiert wird, ein Effekt der sich zuvor schon bei vielen ähnlichen Werken eingestellt
hat. M. Franzen

SPIXIANA 15-18 München, 01. März 2001 ISSN 0341-8391

Taxonomic notes on chitons. 1.
Trochodochiton de Rochebrune, 1884 -
a genus which was fallen into oblivion

(Mollusca, Polyplacophora, Mopaliidae)
Enrico Schwabe

Schwabe, E. (2001): Taxonomic notes on chitons. 1. Trochodochiton de Roche-
brune, 1884 — a genus which was fallen into oblivion (Mollusca, Polyplacophora,
Mopaliidae). — Spixiana 24/1: 15-18

The taxonomic position of de Rochebrune’s genus Trochodochiton is discussed.
The genus is characterized by the presence of precephalic tentacles. Investigations
of the syntypes of Chiton coronatus Fischer MS, Locard, 1898 have shown that this
species refers to de Rochebrune’s description and, therefore, the species herein is
designated as the type species of the genus Trochodochiton. As C. coronatus is a junior
synonym of Placiphorella atlantica (Verrill & S. 1. Smith, 1882), Trochodochiton falls
into the synonymy of Placiphorella Carpenter MS, Dall, 1879. A lectotype and
paralectotype were selected from the syntypes of Chiton coronatus.

Enrico Schwabe, Zoologische Staatssammlung München, Münchhausenstrasse
21, D-81247 München. E-mail: Enrico.Schwabe@zsm.mwn.de

Introduction

The taxonomy of de Rochebrune’s genus Trochodochiton is doubtfull, as its description was not related
to a certain species. The material on which the description of the new genus was based, was collected
during the Talisman — expedition from off West Sudan and is still available in the Museum National
d’Histoire Naturelle in Paris (MNHN).

An investigation of the syntypes shows, that Chiton coronatus Fischer MS, Locard, 1898 in all
respects agrees with de Rochebrune’s description, and it is herein designated to be the type species of
Trochodochiton.

Because the description of the new taxon is available in the sense of the International Code of
Zoological Nomenclature and a type species is now known for it, the genus should no longer be
ignored by recent authors.

Abbreviations used in text

nom. null. - misspelling of a taxon
OD - orginal designation

SD - subsequent designation

M - designation by monotypy

MS - manuscript name

1:5

Systematic part

Polyplacophora Gray, 1821
Mopaliidae Dall, 1889

Trochodochiton de Rochebrune, 1884

Trochodochiton, Van Belle 1983: 115 (marked with a “?” in synonym of Placophoropsis Pilsbry, 1893)
Trochochiton, Kaas & Van Belle 1994: 318 [nom. null.]

De Rochebrune (1884: 754) has described his new genus as follows:

“Le Talisman a recueilli quelques especes de ce groupe tres rare dans les grandes profondeurs;
quelques Echantillons d’un type remarquable meritent d’attirer l’attention; par leur aspect general on
les prendrait a premiere vue pour des specimens de Lepidopleurus de taille ordinaire, mais ce quiles
distingue tout particulierement, c’est la presence autour de la region cephalique d’appendices quad-
rangulaires regulierement espaces et donnant a la partie qu’ils occupent la forme d’une roue dentelee;
ce caractere special et que l’on ne retrouve chez aucune autre espece de cette classe suffit pour autoriser
la creation d’un genre, qui des lors devrait &tre inscrit sous le nom de Trochodochiton (de Rochebrune).”

[The Talisman has collected some species of this group which are very seldom in great depths.
Several specimens of a remarkable type deserve special attention. From their general appearance one
would rather associate them with the species of Lepidopleurus. But what makes them especially different
are rectangular appendages of regular intervals in the cephalic region which gives the part they occupy
the shape of a toothed wheel; this feature and the fact that it is unknown from other species of this class
brings about the necessity to relate them to a new genus which from now on should be termed as
Trochodochiton (de Rochebrune).]

[Die Talisman hat einige Arten dieser Gruppe gesammelt, die in großen Tiefen sehr selten sind;
einige Proben eines bemerkenswerten Typus verdienen besondere Aufmerksamkeit; von ihrem Äuße-
ren würde man sie auf den ersten Blick als Arten von Lepidopleurus von gewöhnlicher Größe halten,
was sie aber ganz speziell unterscheidet, sind viereckige Anhängsel von regelmäßigem Abstand im
Kopfbereich, was der von ihnen besetzten Partie die Form eines gezähnten Rades gibt; dieses spezielle
Merkmal und die Tatsache, daß man es bei keiner anderen Art dieser Klasse findet, ist Anlaß genug,
sie einer neuen Gattung zuzuordnen, die von nun an mit dem Namen Trochodochiton (de Rochebrune)
bezeichnet werden soll.]

The species of Polyplacophora collected during the Talisman — expedition (Locard 1898) are Chiton
coronatus Fischer MS, Locard, 1898 and Acanthochites [= Acanthochitona] fascicularis (Linnaeus, 1767).
The latter species can be excluded, as it doesn't show the features described by de Rochebrune: it lacks
the appendages in the head area and the valve surface is in no way Lepidopleurus — like, and the
tegmentum is strongly granulated, divided into pleural and jugal areas, which is not the case in
Lepidopleurus. Beside this features the girdle in Acanthochitona bears usually 18 hair tufts, whereas in
Lepidopleurus the girdle is clothed with scales interspaced with hyaline spicules.

The syntypes of Chiton coronatus (collected at stations 78 & 71 in a depth of 640-698 m from off West
Sudan) fits in all respects with de Rochebrune’s description, i.e. regarding the presence of precephalic
tentacles (Fig. 1A). Therefore Chiton coronatus is here designated to be the type species of Trochodochiton.

From the type material (MNHN), the author designates herewith the complete specimen (by
Locard 1898 as fig. 25 on plate IV) as lectotype (Figs 1B-1C) and the other partly disarticulated specimen
(it lacks the first three plates) as paralectotype (figured as 23 & 26, by Locard).

The lectotype measures ca. 11.4x7mm and bears 9 gills on the right and 8 gills on the left side of
the foot. The paralectotype (ca. 16.9x 9.8 mm) has 14 gills on the right and 16 on the left side of the foot.
Both specimens are strongly curled and the girdle is bent inwards. It appears that due to contraction
of the specimens Locard just estimated the sizes (21x 16 mm by Locard); the external features figured
by Locard (Figs 23-26) agree with the material examined herein.

Thiele (1909: 9, 31) has correctly shown, that Chiton coronatus is identical with Placiphorella atlantica

(Verrill & S. I. Smith, 1882), which makes Trochodochiton a junior subjective synonym of Placiphorella

Carpenter MS, Dall, 1879.

Fig. 1. Chiton coronatus Fischer MS, Locard, 1898. A. Paralectotype in ventral view, showing the precephalic
tentacles. B. Lectotype, whole animal in dorsal view. C. Lectotype, detail anterior part, showing the girdle
expansion.

Based on the preceeding recognitions the synonymy of Placiphorella should read as follows:

Placiphorella Carpenter MS, Dall, 1879 (pp. 298, 303) (Placiphorella velata Carpenter MS, Dall, 1879), OD

Euplacophora Verrill & S. I. Smith, 1882 (p. 365 [footnote]) (Placophora (Euplacophora) atlantica Verrill &
S. I. Smith, 1882), M

Trochodochiton de Rochebrune,1884 (p. 754) (Chiton coronatus Fischer MS, Locard, 1898), SD, herein

Placophorella Fischer, 1885 [nom. null.] (p. 882)

Placophoropsis Pilsbry, 1893 (p. 313) (Placophora (Euplacophora) atlantica Verrill & S. I. Smith, 1882), M

Plaxiphorella Pelseneer, 1898 [nom. null.] (p. 14)*

Langfordiella Dall, 1925 (p. 96) (Langfordiella japonica Dall, 1925), OD

Euplaciphora Kaas & Van Belle, 1994 [nom. null.] (p. 318) non Carpenter in Dall, 1879 (= nom. null. pro
Euplaxiphora Shuttleworth, 1853 = Plaxiphora Gray, 1847)

Trochochiton Kaas & Van Belle, 1994 [nom. null.] (p. 318)

Praciphorella Matsukuma & Tsubaki, 1995 [nom. null.] (p. 92)

* only Plaxiphorella tentaculifera Pelseneer was listed, without a description of this species.

Zusammenfassung

Die Gattung Trochodochiton, von de Rochebrune für eine Tiefwasserart von W-Sudan aufgestellt, die während
der Talisman — Expedition gesammelt wurde, ist charakterisiert durch das Vorhandensein von Tentakeln im
Kopfbereich.

Da das Talisman - Material im Pariser Nationalmuseum archiviert ist, war eine Untersuchung der Art
möglich, die der Beschreibung von de Rochebrunes neuer Gattung zugrunde lag. Es handelt sich um Chiton
coronatus Fischer MS, Locard, 1898, eine Art die tatsächlich Tentakeln im Kopfbereich aufweist und deshalb
hiermit als Typart für Trochodochiton festgelegt wird. Da aber C. coronatus ein jüngeres Synonym von Placiphorella
atlantica (Verrill & S. I. Smith, 1882) ist, kann de Rochebrunes Gattung nicht als valides Taxon benutzt werden,
sondern muß in die Synonymie von Placiphorella Carpenter MS, Dall, 1879 transferiert werden.

Für die Syntypen von C. coronatus werden der Lectotyp und der Paralectotyp festgelegt.

Acknowledgements

Dr. Philippe Bouchet and Dr. Virginie Heros (Museum National d’Histoire Naturelle, Paris) are thanked for the
loan of the syntypes of Chiton coronatus such as for the important hints for literature. Thanks are also due to Eva
Lodde and Michael Schrödl (Zoologische Staatssammlung München) for correcting the english parts. Special
thanks are due to Mrs Albrecht (Zoologische Staatssammlung München) who kindly translated the french text.

References

Dall, W. H. 1879. Report on the limpets and chitons of the Alaskan and Arctic regions, with descriptions of genera
and subspecies believed to be new. — Proc. U. S. nat. Mus. 1: 281-344, figs A-E, pls 1-5

-- 1925. New shells from Japan. — Nautilus 38: 95-97

de Rochebrune, A. T. 1884. Les Vers, les Mollusques, les Echinodermes, les Zoophytes, les Protozoaires et les
animaux de grandes profondeurs. A. E. Brehm. edition francaise par le Dr A. T. de Rochebrune, Paris, J. B.
Bailliere et fils [1884]: VI + 780 pp. Merveilles de la nature: l’homme et les animaux 7

Fischer, P. 1880-1887. Manuel de conchyliologie et de paleontologie conchyliologique, pp. 1-1369. — Paris

Gray, J. E. 1847. On the genera of the family Chitonidae. — Proc. zool. Soc. Lond. 15: 63-70

Kaas, P. & R. A. Van Belle 1994. Monograph of living chitons (Mollusca: Polyplacophora) 5. Suborder Ischno-
chitonina: Ischnochitonidae: Ischnochitoninae (concluded) Callistoplacinae; Mopaliidae; Additions to Vol-
umes 1-4: 1-402, figs 1-141, maps 1-57. - E. ]J. Brill/ W. Backhuys, Leiden

Locard, A. 1898. Mollusques testaces. Expedition scientifiques du “Travailleur” et du “Talisman” pendant les
annees 1880,1881,1882,1883: 1-515, pls 1-18. — Paris

Matsukuma, A. & Y. Tsubaki 1995. Radular morphology and feeding tracks of Liolophura japonica (Mollusca.
Polyplacophora). - Sci. Rep. Dep. Earth Planet. Sci. Kyushu Univ. 19(1): 81-92, figs 1-13, tab. 1-2

Pelseneer, P. 1898. Recherches morphologiques et phylogenetiques sur les Mollusques archaiques. - Mem. cour.
Acad. Roy. Belg. 57: 1-113, pls 1-24

Pilsbry, H. A. 1892-1894. Monograph of the Polyplacophora. In: G. W. Tryon, Manual of Conchology 14: 1-128,
pls 1-30 (1892); i-xxxiv, 129-350, pls 31-68; 15: 1-64, pls 1-10 (1893); 65-133, pls 11-17 (1894). - Academy of
Natural Sciences, Philadelphia

Shuttleworth, R. J. 1853. Diagnosen neuer Mollusken. 4. Ueber den Bau der Chitoniden, mit Aufzählung der die
Antillen und die Canarischen Inseln bewohnenden Arten. — Mitt. naturf. Ges. Bern 286-291: 169-207

Thiele, J. 1909. Revision des Systems der Chitonen. I. Teil. - Zoologica Stuttg. 22: 1-70, pls 1-6

Van Belle, R. A. 1983. The systematic classification of the chitons (Mollusca: Polyplacophora). — Inf. Soc. belg.
Malac. 11(1-3): 1-178, pls 1-13

Verrill, A. E. 1882. Notice of the remarkable marine fauna occupying the outer banks off the southern coast of
New England, No. 7, and of some additions to the fauna of Vineyard Sound. — Amer. J. Sci. (3) 24: 360-371

SPIXIANA 24 19-27 München, 01. März 2001 ISSN 0341-8391

Description of Allocyclops montenegrinus, spec. nov. and a revision
of the genus Allocyclops Kiefer, 1932

(Crustacea, Copepoda, Cyclopoida)
Tomislav Karanovic

Karanovic, T. (2001): Description of Allocyclops montenegrinus, spec. nov. and a
revision of the genus Allocyclops Kiefer, 1932 (Crustacea, Copepoda, Cyclopoida).
- Spixiana 24/1: 19-27

A new species of the genus Allocyclops Kiefer, 1932 is described on the basis of
a single female collected from subterranean waters in Montenegro (SE Europe).
This genus is revised and divided into three subgenera: Allocyclops s. str., Psammo-
cyclops Kiefer, 1955, and Stolonicyclops Reid & Spooner, 1998. Also, two species from
the genus Speocyclops (S. transsaharicus and S. orcinus) are transferred to the genus
Allocyclops. The species Allocyclops ritae Dumont & Lamoot, 1978 is found as a
synonym of Psammocyclops excellens Kiefer, 1955. With the addition of the new
species, and after this revision, the genus Allocyclops now includes twelve species
throughout the world. At the end of this paper a key for their determination is
given.

Tomislav Karanovic, via Brescia 3, 84092 Bellizzi (SA), Italy

Introduction

The genus Allocyclops was established by Kiefer (1932) to accommodate a new species, A. chappuisi,
which he redescribed very soon after that description (Kiefer 1933), also from Ivory Coast. At the same
time Kiefer (1933a) described one new species from Macedonia, Cyclops (Diacyclops) minutissimus, but
in some further papers he doubted about its taxonomic position (Kiefer 1937, 1937a). Redescription of
this species by Petkovski (1971), and its allocation to the genus Allocyclops, finally resolved its taxonom-
ic status.

However, another taxonomic problem caused more confusion which maintained until today. It is
related to the description of a new species from France, Speocyclops orcinus, which is based only on a
single male (Kiefer 1937b). The isolated position of this species within the genus was noticed already
by Petkovski (1954). Dussart (1967) even refused to place it in the genus Speocyclops. However, a lot of
copepodologists left this species in the genus Speocyclops, without particular comments (Rylov 1948,
Lescher-Moutoue 1967, 1973, 1986, Kiefer 1978, Dussart & Defaye 1985). Although the description was
very poor in detail, there is no doubt that this species belongs to the genus Allocyclops. The appearance
of the endopodite of the fourth swimming leg, as well as the fifth leg (although it was hardly visible
at that time and especially difficult for the verbal description) confirm this claim. Similar situation is
with Metacyclops arenicolus, which was described from Lake Nyasa (Fryer, 1956) and until now consid-
ered as a member of the genus Bryocyclops (see Dussart & Defaye, 1985). Chappuis (1951) described
Allocyclops cavicola from a cave in Zaire, and Petkovski (1971) described Allocyclops kieferi from intersti-
tial waters in Macedonia. Later Dumont & Lamoot (1978) described Allocyclops ritae from Ivory Coast,
but strange enough as the second species in the genus (even though four species were known by that
time).

Even more strange was that the same authors three years later (Lamoot et al. 1981) described a very
similar species from Ivory Coast in the genus Speocyclops (S. transsaharicus). This species also belongs
to the genus Allocyclops, although in the meantime it was uncritically accepted by few authors as the
first representative of the genus Speocyclops in tropical Africa (Dussart & Defaye 1985, Lescher-
Moutoue 1986). Plesa (1981) described Allocyclops botosaneanui from a cave in Cuba. Dussart (1984)
described Allocyclops neotropicalis from Venezuela only after the male which Reid (1988) correctly
transferred to a new genus, Yansacyclops. Rocha & Bjorberg (1988) described Allocyclops silvaticus from
Brazil and remarked: “Allocyclops badly needs revision”. Reid & Spooner (1998) described Stolonicyclops
heggiensis as a new genus and new species from the USA, but there are not enough differential
characteristics between this genus and the genus Allocyclops. They did not compare it with Allocyclops
ritae, which also has endopodite of the fourth swimming leg slightly fused. Partial oligomerization of
that appendage was known as a specific variability (Monchenko 1974) in some species in the genus
Metacyclops. We think that Stolonicyclops heggiens also belongs to the genus Allocyclops.

At the end we must mention one more curiosity. Kiefer (1955, 1956) twice described Psammocyclops
excellens, as a new genus and new species from Madagascar. His description of the fifth leg was very
provisional, although later it was uncritically accepted by many authors. From the drawings of that
appendage of the male it is clear that the fifth leg is not a distinct segment, but completely fused to the
somite. Even more, we think that this species is synonymous (of course the older one) with Allocyclops
ritae Dumont & Lamoot, 1978 which is mentioned before. So, until now eleven species were known in
the genus Allocyclops. During an investigation of the copepod fauna in Montenegro, a further new
species of that genus was identified. This new species is herein described as A. montenegrinus, spec. nov.
Also a revision of the genus Allocyclops is proposed.

Material and Methods

The sample was collected using the Karaman-Chappuis method from interstitial waters of a very small
and nameless stream in the village Vrela, near the town Cetinje, Montenegro, SE Europe (type locality),
on May 9, 1998. The material was preserved by adding several drops of 36 % formaldehyde. Copepods
were separated with a Wild-M5 stereomicroscope and moved to 70 % ethyl alcohol. Specimens were
dissected in a mixture of equal parts of distilled water and glycerol, with fine entomological needles
(mark 000). Dissected appendages were placed on a slide, in the same mixture of distilled water and
glycerol, and covered with a coverslip. For larger parts (abdomen, etc.) two human hairs were mounted
between slide and coverslip, so the parts could not be crushed. During the examination water slowly
evaporates, and after some time appendages remain in the pure glycerol. All drawings have been
prepared using a drawing attachment (tube) on a Leica-DMLS microscope, with C-PLAN achromatic
objectives. Dissected appendages were preserved in Faure’s medium. Non-dissected specimens, after
examination, were again preserved in 70 % ethyl alcohol. In that sample following species were found:

Diacyclops bicuspidatus (Claus, 1857) - 284, 62?

Diacyclops bisetosus (Rehberg, 1880) - 584, 52? (3 ovigerous)

Allocyclops montenegrinus, spec. nov. — 1? (holotype)

Bryocamptus (s. str.) minutus (Claus, 1863) — 6865, 132 (6 ovigerous), 2 copepodids
5. Bryocamptus (Rheocamptus) pygmaeus (Sars, 1863) - 583, 1722 (2 ovigerous)

HEN DE

All specimens are deposited in the author’s collection in Italy. The holotype of the new species
(Allocyclops montenegrinus) was completely dissected and mounted on a slide in Faure’s medium
(Number: 9/43/0606/e). In the description, diagnosis, keys and figure legends no abbreviations were
used.

Figs 1-3. Allocyclops montenegrinus spec. nov., holotype (female 0.563 mm). 1. Urosome, dorsal view. 2. Uro-
some, lateral view. 3. Urosome, ventral view. Scale = 0.1 mm.

Results

Allocyclops montenegrinus, spec. nov.
(Figs 1-18)

Holotype: 2, stream near Vrela, near Cetinje, Montenegro, SE Europe, May 9, 1998.

Description

Female. Body length, excluding furcal setae, 0.563 mm. Habitus compact, dorsoventrally compressed.
Prosome comprising cephalothorax, incorporating first pedigerous somite, and 3 free pedigerous
somites. Surface of dorsal shield covering cephalothorax without any ornamentation, as well as 3 free
pedigerous somites (Fig. 13). Body widest at prosomite first pedigerous somite. Urosome comprising
fifth pedigerous somite, genital double-somite (representing fused genital and first abdominal
somites), and 3 free abdominal somites. More or less sclerotized joint (as pseudosomite) present
between prosome and urosome, as well as between fifth pedigerous somite and genital double-somite
(Figs 2, 13). Body colourless, nauplius eye absent. Genital double-somite about 1.3 times broader than
long, trapeziform, with hind margin ventrally smooth and dorsally serrated (Figs 1, 3). Genital
apertures placed dorsolaterally at the first third, and covered by operculum derived from fused sixth
legs. Seminal receptacle with broad anterior and ovoid posterior expansions (Fig. 3). First and second
free abdominal somites with hind margins ventrally smooth and dorsally serrated. Anal somite
ornamented with pair of sensillae, and with row of spinules along posterior margin. Anal operculum
convex, not reaching beyond limit of anal somite. Anal sinus smooth (Fig. 1). Furcal rami slightly
divergent, close, without ornamentation, and about 2.7 times longer than wide (Fig. 3). Lateral seta
inserted dorsolaterally, just to midlength of ramus. Dorsal seta slightly longer than ramus. Outermost

8.9.10
yaı

Figs 4-11. Allocyclops montenegrinus, spec. nov., holotype (female 0.563 mm). 4. First svimming leg. 5. Anten-
nula. 6. Second swimming leg. 7. Antenna. 8. Fifth leg. 9. Sixth leg. 10. Labrum. 11. Maxillula. Scales =
0.1] mm.

apical seta very stout (spiniform), subterminal, and about twice longer than innermost apical seta. Two
middle apical setae broken (Fig. 1). Rostrum large, even wellrounded, but not reaching beyond end of
antennula first segment (Fig. 13). Antennula 11-segmented, shorter than cephalothorax, with short
aesthetasc on eighth segment and setal formula as follows: 7.3.7.1.2.2.3.2.2.2.8 (Fig. 5). Distal seta on
fifth segment very stout and short (maybe spiniform). Antenna 4-segmented, without seta representing
exopodite (Fig. 7). No ornamentation visible on surface of basipodite. This appendage makes right
angle with body ose (Fig. 13), and with setal formula as follows: 2.1.5.6 (Fig. 7). Labrum with strong
teeth on posterior margin, but without any other ornamentation (Fig. 10). Mandibula with strong teeth
on distal end of coxa (Fig. 18), and with palp represented by only 1 very thin and short seta (Fig. 17).
Maxillula comprised of elongated praecoxa and 1-segmented palp (Fig. 11). Praecoxa arithrite with 7
smooth setae and spines, while palp bears 2 apical (plumose) and 3 lateral (smooth) setae. Maxilla
5-segmented, comprising praecoxa, coxa, basis, and 2-segmented endopodite (Fig. 15). Praecoxa with
proximal endite bearing 2 setae (proximal one broken), and distal endite unarmed and very small. Coxa
with 2 endites; proximal with 1 smooth seta, distal endite highly mobile and bearing 1 plumose and
1 smooth setae. Basis drawn out into claw, with 5 teeth on inner margin and 2 setae. First endopodite
segment armed with 2, second with 3 setae (Fig. 15). Maxilliped 4-segmented, with setal formula as
follows: 2.1.1.2 (Fig. 16). All swimming legs with smooth coxae, and 1 plumose seta on their inner-distal
corner (Fig. 4, 6, 12 and 14). Couplers (intercoxal sclerites) without surface ornamentation. Basis of each
swimming leg with epipodite seta on outer margin, especially well-developed on first leg (Fig. 4). Basis
of first leg also with short and stout spine on distomedial corner. That corner on other swimming legs
with small spinous process. All swimming legs with 2-segmented endopodites and exopodites. First
exopodite segment of all legs lacking seta, and bearing 1 outer spine. Second exopodite segments with
spine formula 3.4.4.3, and setal formula 5.4.4.4. First endopodite segment of all legs bearing 1 seta on
inner-distal corner. Second endopodite segment of first swimming leg with 3 inner setae, 1 smooth and

1.17.18

Figs 12-18. Allocyclops montenegrinus, spec. nov., holotype (female 0.563 mm). 12. Third swimming leg.
13. Habitus, lateral view. 14. Fourth swimming leg. 15. Maxilla. 16. Maxilliped. 17. Mandibula. 18. Mandibula.
Scales = 0.1 mm.

curved apical spine, and 1 outer seta (Fig. 4). Second endopodite segment of second swimming leg with
2 inner setae, 1 apical seta, 1 apical spine, and 1 outer seta (Fig. 6). The same segment of third leg with
1 more inner seta (Fig. 12). Second endopodite segment of fourth swimming leg with 3 inner setae, 2
apical spines (inner spine about 2.4 times longer than outer one), and 1 outer seta (Fig. 14). This segment
about 1.7 times longer than broad. Outer margins of endopodites, as well as inner margins of ex-
opodites, of all swimming legs (except first exopodite segment of first leg) ornamented with rows of
long pinnules. Also, all setae (except epipodite setae on second and third legs) are plumose. Fifth leg
inserted laterally and fused to somite (Fig. 2). Remnant of proximal segment only 1 plumose and short
(in comparison with somite) seta. Distal segment like small protrusion, and with inner short and stout
spine and even shorter outer seta (Fig. 8). Sixth leg inserted dorsolaterally, consisting of small plate
bearing 2 short and smooth setae (dorsal seta about 2.7 times longer) and 1 short and smooth spine,
completely fused to leg (Fig. 9). This leg also fused to somite, and covers genital aperture.

Male. Unknown.

Etymology. The species name montenegrinus is taken from the name of republic Montenegro where the material
was collected, i.e., as an adjective agreeing in gender with the (masculine) generic name.

Distribution. At present Allocyclops montenegrinus, spec. nov. is known only from type locality. We
suppose that it inhabits a wide area of south Dinaric Alps.

Revision of the genus Allocyclops Kiefer

Order Cyclopoida Sars, 1886
Family Cyclopidae Burmeister, 1834
Subfamily Cyclopinae Dana, 1853

Genus Allocyclops Kiefer, 1932

Diagnosis (emended). Small species, body length ranging from 0.41 to 0.8 mm. Genital double-somite
broader than long, with genital apertures placed at the first half. Anal operculum broad, convex or
quadrate, slightly shorter than anal somite, equal, or slightly longer. Furcal rami stout, from 1.5 to 3.5
times longer than wide, and with lateral seta inserted after the first third. Antennula 11-segmented,
shorter than cephalothorax. Maxilliped 4-segmented. All swimming legs with 2-segmented en-
dopodites and exopodites (that are almost equally long), without any sexual dimorphism. Sometimes
endopodite of fourth swimming leg slightly fused. Fifth leg inserted laterally and fused to somite.
Remnant of proximal segment only 1 seta. Distal segment like small protrusion, bearing two short setae
or (more frequently) inner spine and outer seta.

Type species: Allocyclops chappuisi Kiefer, 1932.

Subgenus Allocyclops Kiefer, 1932

Diagnosis. Antenna without seta representing exopodite. Maxillular palp 1-segmented. Coxae of all
swimming legs with seta on their inner-distal corner. Second endopodite segment of fourth swimming
leg with 3 inner setae, 2 apical spines (inner spine longer than outer one), and 1 outer seta. Seta remnant
proximal segment of fifth leg short in comparison with somite.

Type species: Allocyclops (s. str.) chappuisi Kiefer, 1932.

Additional species: Allocyclops (s. str.) cavicola Chappuis, 1951; A. (s. str.) botosaneanui Plesa, 1981;
A. (s. str.) orcinus (Kiefer, 1937) comb. nov.; A. (s. str.) montenegrinus, spec. nov.; A. (s. str.) kieferi
Petkovski, 1971; A. (s. str.) minutissimus (Kiefer, 1933); A. (s. str.) arenicolous (Fryer, 1956), comb. nov.

Key to the species of the subgenus Allocyclops

1. Innermost apical seta on furcal rami longer than outermost one .........useseseesesenensenenensensnsonenennensnnenenn 2:
- Innermost apical seta on furcal rami shorter than outermost one.........eueneenseseneesenensonensnnensnneneneenene 4.
25% Anallopereulumshortiand'quadratere.. n.eneenenesenanseeenereenan een sense RR 8:
= Anal’opereulumsdearlyseonverm re en rsenn A. (s. str.) chappuisi Kiefer, 1932
3. Furcal rami more than 3 times longer than wide............ A. (s. str.) cavicola Chappuis, 1951
- Furcal rami less than 3 times longer than wide .............n.. A. (s. str.) botosaneanui Plesa, 1981
4... Anall/opereulum.smooth..................u...:2e22078se22000200200sdstnsefonasns een onstarnsasunean sense nen ee RE S%
Ze AnalKopereulumktinelyzserrateder ern nn A. (s. str.) orcinus (Kiefer, 1937) comb. nov.
5.2 Basis.of first les with spine.an distomedialkeormer...............22222202Beusesenansenernennre cases en eennene 6.
- Basis of first leg without that spine ............n. A. (s. str.) arenicolous (Fryer, 1956) comb. nov.
6. Setal formula on second exopodite segments of swimming legs is 5.3.5.9 c...enenenenenenenenenenenenennnnnnn 7

- Setal formula on second exopodite segments of swimming legs is 5.4.4.4 ....essnnssesenenene
Manetseaskadnsnsätenehssnerssebrshehrsnsnulhtnegehiep er nesemueen nennen ers ennensenete rer trirete A. (s. str.) montenegrinus, spec. noV.

7. Furcal rami about 3 times longer than wide; innermost apical seta on ramus about 2 times shorter
thanfoutermostrone. a. on. A. (s. str.) kieferi Petkovski, 1971

- Furcal rami about 1.5 times longer than wide; innermost apical seta on ramus slightly shorter than
OULELINOSTIONE ee ee er A. (s. str.) minutissimus (Kiefer, 1933)

Subgenus Psammocyclops Kiefer, 1955

Diagnosis. Antenna with short seta representing exopodite. Maxillular palp 1-segmented. Coxae of all
swimming legs with seta on their inner-distal corner. Second endopodite segment of fourth swimming
leg with 2 or 3 inner setae, 1 apical spine, and 1 outer seta. Seta remnant proximal segment of fifth leg
long in comparison with somite, changing habitus of animal in dorsal view.

Type species: Allocyclops (Psammocyclops) excellens (Kiefer, 1955) comb. nov. [synonym: Allocyclops ritae Dumont
& Lamoot, 1978].

Additional species: Allocyclops (Psammocyclops) transsaharicus (Lamoot, Dumont & Pensaerat, 1981)
comb. nov.; A. (P.) silvaticus Rocha & Bjornberg, 1988.

Key to the species of the subgenus Psammocyclops

1. Second endopodite segment of fourth swimming leg with 3 inner setae .......nnen 2:

- Second endopodite segment of fourth swimming leg with 2 inner setae .......ennnen:
once RESTE HERREN A. (P.) excellens (Kiefer, 1955) comb. nov.

2. Spine formula on second exopodite segments of second, third and fourth swimming legs is 3.3.2
ee N N NE ER A. (P.) silvaticus Rocha & Bjornberg, 1988

- Spine formula on second exopodite segments of second, third and fourth swimming legs is 4.4.3
KR RER NEL U, A. (P.) transsaharicusus (Lamoot, Dumont & Pensaert, 1981) comb. nov.

Subgenus Stolonicyclops Reid & Spooner, 1998

Diagnosis (emended). Antenna without seta representing exopodite. Maxillular palp 2-segmented.
Coxae of second, third and fourth swimming legs without seta on their inner-distal corner. Second
endopodite segment of fourth swimming leg with 3 inner setae, 1 apical spine, and 1 outer seta. Seta
remnant proximal segment of fifth leg relatively long in comparison with somite, but not changing
habitus of animal in dorsal view.

Type and single species: Allocyclops (Stolonicyclops) heggiensis (Reid & Spponer, 1998) comb. nov.

Discussion

The systematics of the genera included in the subfamily Cyclopinae recently was discussed by many
authors (Dussart & Defaye 1985, Reid 1993, 1999, Pesce 1996, Ferrari 1998, Rocha et al. 1998, Reid et al.
1999). Now, like fifty years ago, the most important systematic character at generic level is the
morphology of the fifth leg. Until now, only seven genera are known with fifth leg completely fused
to somite: Austriocyclops Kiefer, 1964; Bacillocyclops Lindberg, 1956; Bryocyclops Kiefer, 1927; Allocyclops
Kiefer, 1932; Haplocyclops Kiefer, 1952; Palaeocyclops Monchenko, 1972; and Yansacyclops Reid, 1988. The
genera Austriocyclops and Bacillocyclops have the fifth leg reduced to a single seta or spine. Five other
genera have the fifth leg completely fused to the somite, but all three setae remain (two from distal,
and one from proximal segment).

The genus Bryocyclops differs from the genus Allocyclops by the following features: sexual dimor-
phism in swimming legs; endopodite of the fourth swimming leg always considerably shorter than
exopodite (even when it is 2-segmented); distal segment of the fifth leg does not remain as a small
protrusion; and anal operculum always produced posteriorly. This genus indeed has confused system-
atics (Reid 1999), and also needs revision.

The genus Haplocyclops is revalidated by Rocha et al. (1998), and differs from Allocyclops as follows:
genital apertures placed at the second half of the genital double-somite; lateral seta on furcal rami
inserted in the first third; antennula without seta on the inner margin of the ultimate segment; distal
segment of the fifth leg does not remain as a small protrusion; and endopodite of the fourth swimming
leg 1-segmented.

The genus Palaeocyclops is monospecific, known from the Kisilkum Desert (Monchenko 1972). It
differs from Allocyclops by the very long anal operculum, as well as by presence of sexual dimorphism

in swimming legs, and absence of any spine on endopodite of the fourth swimming leg.

The genus Yansacyclops is also monospecific, known from Brazil (Reid 1988). It differs from the
genus Allocyclops by the following features: genital double-somite longer than broad; anal operculum
very short, placed in the first half of anal somite; antennula 10-segmented; antenna with very long seta
representing exopodite; and second endopodite segment of the fourth swimming leg with one apical
spine and one apical seta. This genus is, in our opinion, closest related to the genus Allocyclops. Many
other genera from the subfamily Cyclopinae have similar segmentation of the swimming legs and
antennula (Speocyclops Kiefer, 1937; Muscocyclops Kiefer, 1937; Fimbricyclops Reid, 1993; etc.), but the
fusion of their fifth leg to somite is of completely different nature, and it is never total.

We divided the genus Allocyclops into 3 subgenera: Allocyclops s. str., Psammocyclops Kiefer, 1955,
and Stolonicyclops Reid & Spooner, 1998. Their differential diagnoses, in our opinion, are not sufficient
for giveng them generic status. This especially refers to the monospecific subgenus Stolonicyclops,
which can be separated from the subgenus Psammocyclops only by the absence of the coxal setae. All
other characteristics are at specific, not at generic level. We already pointed out that Kiefer (1955; 1956)
very provisionally described Psammocyclops excellens as a new genus and new species. Even Pesce
(1996) accepted this, without any particular comments. But if we believe in Kiefer’s drawing of the
female’s fifth leg (which should be one separated article with three setae), how we can explain the
drawing of the male’s fifth leg (which is completely fused to the somite, and quite exact as the fifth leg
in the genus Allocyclops)? It seems that Kiefer made an error and drew some cuticular suture or curve
as a distinctive membrane of the fifth leg in the female. If we accept this, and after comparison of
Psammocyclops excellens with Allocyclops ritıe Dumont & Lamoot, 1978, we see that there are no
differential characteristics between these two species. In their description Dumont & Lamoot (1978)
wrote that A. ritae has the spine formula on exopodites of the swimming legs as 3.3.3.2. But reexam-
ination of the type material (Rocha & Bjornberg 1988) showed that this formula is 2.3.3.2, i.e. the same
formula as in Psammocyclops excallens. Shape of the fourth swimming leg, as well as dorsal view of the
urosome, and other details are the same in both species. Only Dumont & Lamoot (1978) noted that
separation of the segments of the fourth leg endopodite is hardly visible, but we already said that this
characteristic is known as highly variable in many species. Therefore we consider Allocyclops ritae as a
synonym of Psammocyclops excellens. It is probably trogloxen, or maybe a troglophilous species also in
Madagascar, but Kiefer (1955, 1956) there collected only interstitial fauna.

Allocyclops montenegrinus, spec. nov. clearly belongs to the subgenus Allocyclops. In fact, in that
subgenus there are two well distinguished groups of species. The first group contains three tropical
species, having the innermost apical seta of the furcal rami longer than the outermost one. The other
group to which A. montenegrinus belongs has the innermost apical seta shorter than the outermost one.
All species of that group inhabit subterranean waters of Europe, except A. (s. str.) arenicolous which was
known from the interstitial of Lake Nyasa (Africa). From all species in the subgenus Allocyclops the new
species is easily distinguishable by the shape of furcal rami, anal operculum, and spine and setal
formula on swimming legs. Very strong differential characteristics between species, as well as geo-
graphical distribution of the genus Allocyclops, tell us that this genus had the highest diversity during
Tertiary or even before. The genus Speocyclops, on the other hand, has very weak differential charac-
teristics between species, and it seems that this genus now has its highest diversity in subterranean
waters of Europe. After transferring Speocyclops transsaharicus and Speocyclops orcinus from that genus
to the genus Allocyclops (in this paper), many taxonomical and zoogeographical problems in that genus
are resolved. The observations of some populations of Speocyclops demetiensis from surface waters in
Norway by Hessen & Stene (1991) demonstrate that probably we do not need to search ancestors of
that genus in African surface waters. Maybe they inhabited mosses and other surface waters on
European high mountains before Quaternary major climatic oscillations.

References

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Buchbesprechungen

2. Bosch, D. T., Dance, S. P., Moolenbeek, R. G. & P. G. Oliver: Seashells of Eastern Arabia. - Motivate
Publishing, London, Abu Dhabi, 1995. 296 S., zahlreiche Farbabb. ISBN 1-873544-64-2.

In diesem optisch sehr ansprechenden Bildband werden 1273 (!) Arten und Unterarten schalentragender
Weichtiere aus dem Arabischen Golf und dem östlichen Arabischen Meer vorgestellt. Das Hauptgewicht liegt
natürlich bei den artenreichen Gastropoden und Muscheln, es werden aber auch einige relevante Scaphopoden,
Cephalopoden und Polyplacophoren behandelt. Zu (fast) jeder Art findet sich ein qualitativ hochwertiges
Farbfoto, z.T. auch als prächtige Großaufnahme, ein kurzer und prägnanter Text zur Schalenmorphologie sowie
Information zu Habitat und Verbreitung im Telegrammstil. Kleine Schalen werden meist als Zeichnungen oder,
sehr anschaulich, als raster-elektronenmikroskopische Aufnahmen präsentiert. Sammlern und faunistisch Inter-
essierten bietet sich so ein umfassendes Bestimmungs- und Nachschlagewerk arabischer Meeresweichtiere, in
dem auch einige Klein- und Tiefwasserformen enthalten sind. Anspruch auf Vollständigkeit besteht verständ-
licherweise jedoch nicht.

Für die Zuverlässigkeit der Angaben und Artbestimmungen stehen vier namhafte Autoren mit langjähriger
wissenschaftlicher Erfahrung und Leidenschaft. Ihre Recherchen schlossen aufwendige Vergleichsstudien an
internationalen Museen sowie Expertisen zahlreicher Spezialisten mit ein. Taxonomische Anmerkungen oder
Hinweise zur Synonymie sind jedoch bewußt knapp gehalten und nur in Kurzform an die Beschreibungen
gefügt, auf Fachliteratur wird nur in Einzelfällen hingewiesen. Dies werden wissenschaftlich Interessierte
sicherlich mehr bedauern als das Festhalten an einer traditionellen Großgruppeneinteilung.

Ihr selbst gestecktes Ziel “to provide a well-illustrated guide to most of the species of shell-bearing Mollusca
living around the coasts of Oman and the Arabian Gulf ...” haben die Autoren jedenfalls bestens erfüllt und ein
Standardwerk zu einem akzeptablen Preis geschaffen. M. Schrödl

3. Wallace, A.: The Origin of Animal Body Plan: A Study in Evolutionary Developmental Biology. - Cambridge
University Press, Cambridge. 1997. xii + 338 pp. ISBN 0-521-55014-9 hbk.

In order to fulfill the promise of its title, the author has compiled data from various sources and disciplines
including comparative developmental genetics, selection theory, population genetics, ecology, and phylogenet-
ics. Indeed, the strength of this book lies in the comparative and interactive consideration of many different
biological disciplines providing a nice example of what is called “reciprocal illumination”.

However, the morphologist and phylogenetist feels bad in recognizing a general confusion of “body plan”
(i.e. a compilation of characters shared by the majority of species of a taxon and thus a logic but purely idealistic
subject) and “stem species” (i.e. the hypothetical, probabilistic reconstruction of a historical reality). Only the
latter can and have to be explained, the former exists but in our minds alone. The same is true for ideas on the
origin and evolution of life cycles, which are based on “characteristic” instead of ancestral larval types or life-
cycle modes in the various taxa. Indeed, according to the given survey echinoderms, crustacean and molluscs
alone show various larval types, and practically not a single, truly ancestral life-cycle of any higher taxon is
presented.

Despite these shortcomings, which unfortunately can be found in many current contributions on develop-
mental evolution, the book is recommended to everyone who is interested in the evolution of development and
its underlying processes. G. Haszprunar

4. Flindt, R.: Biologie in Zahlen, eine Datensammlung in Tabellen mit über 10.000 Einzelwerten — Spektrum
Akademischer Verlag, Heidelberg, 5. Auflage. 2000. 285 S. ISBN 3-8274-0914-4.

Ein unentbehrliches Nachschlagewerk, dessen Erfolg mit der in kurzer Zeit notwendig gewordenen 5. Auflage
deutlich wird. Hier kann man schnell und einfach vergleichende Zahlen zu allen möglichen Fachgebieten der
Biologie nachschlagen. Die Tabellen informieren unter anderm über zoologische, botanische, physiologische,
genetische und zellbiologische Themen. Bei allen Tabellen sind die Originalquellen angegeben, über die man
dann bei Bedarf genauere Information finden kann. Ein Nachschlagewerk, das in keiner guten Bibliothek fehlen
sollte. K. Schönitzer

SPIXIANA 29-51 München, 01. März 2001 ISSN 0341-8391 ”

The genus Erophiloscia Vandel, 1972 <y
its phylogeny and biogeography,
with description of three new species

(Crustacea, Isopoda, Oniscidea)
Andreas Leistikow

Leistikow, A. (2001): The genus Erophiloscia Vandel, 1972 - its phylogeny and
biogeography, with description of three new species (Crustacea, Isopoda, Onisci-
dea). — Spixiana 24/1: 29-51

A reexamination of the type material of Erophiloscia longistyla Vandel, 1972
revealed the presence of a second species in Colombia within the type series.
E. waegelei, spec. nov. is quite similar to the preceding species, but differs in the
presence of a caudal row of spines onn the male pleopod 1 endopodite and a set of
teeth in endopodite 2. Furthermore, there are two new species from a collection
performed by Dr. W. Hanagarth at Panguana station in Peru in 1975/76: E. recur-
vata, spec. nov. which is characterized by the laterally bent male pleopods 1 and 2
and E. acanthifera, spec. nov. with some specific ornamentations on the male pleo-
pod endopodite 1. The new records throw new light on the phylogenetic relation-
ships of the species of this genus: The presence of a linea frontalis on the cephalo-
thorax is considered a plesiomorphy of E. longistyla and E. waegelei, whereas the
reduction of the caudomedial row of spines on the male pleopod 1 endopodite is
a synapomorphy of E. longistyla, E. recurvata, and E. acanthifera. The Peruvian
species are adelphotaxa due to the lack of a linea frontalis. The genus is found in
the valleys of the northern Andes, which have a tropical climate. It is close to several
genera forming the monophyletic Prosekia-group. A key for the four species is
presented.

Andreas Leistikow, Universität Bielefeld, Abteilung für Zoomorphologie und
Systematik, Morgenbreede 45, D-33615 Bielefeld.
e-mail: Leiste@Biologie.Uni-Bielefeld.de

Introduction

The genus Erophiloscia Vandel, 1972 was instituted for one new species of terrestrial isopods from
Colombia (Vandel 1972), namely Erophiloscia longistyla Vandel, 1972. In addition, two species from
Ecuador, first ascribed to the genus Andenoniscus Verhoeff, 1941 (Vandel 1968) were included. These
two species were superficially described solely on the basis of female specimens, and a recognition of
the males of these species is not warranted. Particularly since the descriptions were superficial and
focused on characters commonly found in other species, they were regarded as nomina dubia by
Leistikow (1998a) until new material, particularly males, are found.

Nonetheless, this genus is rather distinct from other South American genera of “philosciid” Onis-
cidea, especially the shape of pleopod 5 is unique within the Crinochaeta of this continent. A similarly
shaped pleopod 5 can be found among others in Chaetophiloscia hastata Verhoeff, 1929. At the first
glance, one should consider these forms congeneric. For elucidating the phylogenetic relationships of

Erophiloscia, the type material was re-examined and within this material, a new species could be
located. Other material was found in the collection which Dr. W. Hanagarth made in the 1970’s in Peru.
The first material described was those of the genus Ischioscia Verhoeff, 1928 (Schmalfuss 1980). Now
some more information on the interesting isopod fauna of the Biological station “Panguana” shall be
given: the examination of the collection revealed the presence of two new species of the genus
Erophiloscia.

The new species are described in detail and their phylogenetic relationships are discussed. Type
specimens are deposited in Museum National d’Histoire Naturelle, Paris (MNHN) and in Staatliches
Museum für Naturkunde, Stuttgart (SMNS).

Systematic account
Genus Erophiloscia Vandel, 1972

Diagnosis. Cephalothorax with linea supra-antennalis, linea frontalis present or lacking, small lateral
lobes, profrons with two slight depressions above antennal sockets, eyes composed of about 5 to 10
ommatidia. Antennula three-articulate, distal joint with two divergent aesthetasc tufts, the distal one
directed more prominent than the vertical one, antenna with three-articulate flagellum, bearing a long
apical organ.

Mandibular penicil consisting of about 4-7 branches, medial endite of maxillula without apical tip,
lateral endite with 4+6 teeth, five of inner set cleft, outer lobe of maxilla up to two times broader than
inner lobe, maxilliped with palp bearing two setal tufts, endite with small penicil, lacking setation.

Pereopods with sparse setation, ornamental sensory spine of carpus 1 double-fringed serrate, coxal
plates with long nodulus lateralis, nodulus lateralis on coxal plate IV inserted more dorsally.

Pleopods without respiratory areas on exopodites, endopodites 1 and 2 of male elongate, especially
of pleopod 2, reaching or somewhat surpassing caudal tip of pleotelson. Pleopod 5 exopodites of males,
mediodistally drawn out, as for supporting pleopod 2 endopodites.

Uropod protopodite with lateral groove, endopodite inserting proximally of exopodite.

Type species. Erophiloscia longistyla Vandel, 1972 (by original designation)

Remark. This interesting genus is at once distinguished from other South American members of the
“Prosekia-group”, which have a synapomorphic morphology of the antennula as discussed elsewhere
(Leistikow 1998a), by the shape of the male pleopods, especially the drawn out pleopod 5 exopodites
and the extraordinarily long pleopod 2 endopodites already noted by Vandel (1972), a conspicuous set
of autapomorphies. The pleopod 5 exopodite bears no guide slot for the pleopod 2 endopodite, which
is holded simply by the medial margin of the exopodite 5. This character and the lack of a proximal
setal tuft on the palp of the maxilliped and of setation on the endite are synapomorpies of the species
of Erophiloscia. The closest relatives of Erophiloscia are the members of Prosekia Leistikow, 2000 and
Xiphoniscus Vandel, 1968 which have a synapomorphic structure of the antennula. Rather closely
related is Andenoniscus Verhoeff, 1941, with the noduli laterales extraordinarily long.

Erophiloscia waegelei, spec. nov.
Figs 1-6
Erophiloscia longistyla Vandel, 1972 (part).

Types. Holotype, 3, 2.5 mm, several paratypes: Colombia, montane forest near Tibabitä, under tree mosses,
2600 m-2800 m, leg. H. Sturm, 18.07.1969, MNHN Vandel Collection.

Note. Among the plentiful material of Erophiloscia longistyla Vandel, 1972 there was found a lot with
the males differing remarkably from the type. They are somewhat smaller and show differences in the
morphology of the male pleopods.

Description

Colour. Material somewhat faded.

Cephalothorax. Linea frontalis faint, most significant medially from compound eyes, continued by
lateral lobes, linea supra-antennalis present, faint lamina frontalis, compound eyes consisting of about
8 ommatidia (Fig. 1, Ctf).

Pereon. Tegument smooth with scattered tricorn-like setae, coxal plates (Fig. 6Cx3) with sulcus
marginalis and nodulus lateralis, nodulus on coxal plate IV inserted more distantly from the lateral
margin (Fig. 1, Cxp), gland pores lacking.

Pleon. Retracted from pereon, neopleurae of pleonites 3 to 5 small, pleotelson with almost straight
lateral margins, bearing scattered tricorn-like setae (Fig. 6, Tel).

Antennula. Three-articulate with prominent proximal article, distal joint bulbous, bearing two
distinct sets of aesthetascs (Fig. 1, Anl).

Antenna. Antennal peduncle composed of five articles with length ratio 1:2:2:3:4, densely cov-
ered with tricorn-like setae, flagellum composed of three articles, distal one bearing prominent apical
organ, as long as flagellar articles 1 and 2 together (Fig. 1, An2).

Mandible. Molar penicil composed of about seven branches, pars intermedia with two penicils on
left and one on right mandible, additional plumose seta more proximally (Fig. 2, Mdl/r).

Maxillula. Medial endite with two pointed penicils, no apical tip, lateral endite with apically 4+5
teeth, four of inner set cleft, laterally fringed (Fig. 2, Mx1).

Maxilla. Lateral lobe almost twice as broad as medial one, almost without setation, medial endite
bearing some cusps apically (Fig. 2, Mx2).

Maxilliped. Basipodite with sulcus lateralis, palp with two setal tufts, proximal one consisting of
three setae, endite without setation, bearing two teeth caudally; in examined specimen, rostral surface
with transverse scar at level where knob-like penicil inserted, probably broken (Fig. 2, Mxp).

Pereopods. Rather slender (Fig. 3, PE1-4, 4, PE5-7), carpus of pereopod 1 with small antenna-
grooming brush rostrally, ornamental sensory spine double-fringed serrate (Fig. 3, Scl), prominent
sensory spines with two subapical tips, dactylus with short inner claw, dactylar seta simple (Fig. 3,
Dac).

Pleopods. Shape of exopodites rhomboidal, laterally bearing two to six sensory spines, exopodite
5 with transverse row of long pectinate setae on caudal surface (Fig. 5, PL1-5).

Sexual dimorphism. Male ischium of pereopod 7 with fewer spines on medial margin than female
Male pleopod 1 exopodite circular, endopodite styliform, with longitudinal ridge on rostral surface,
subapically with slight transverse furrows rostrally, producing a striate appearance, mediocaudally
with longitudinal row of spines. Male pleopod 2 exopodite pointed, lateral margin sinuous with three
sensory spines subapically, endopodite conspicuously surpassing exopodite, flagelliform, distal quar-
ter with several hyaline, obtuse hooks, medioproximally directed. Even female endopodite mediodis-
tally pointed. Male pleopod 5 exopodite mediocaudally pointed, protrusion of one fifth the length of
medial margin.

Uropod. As described in generic diagnosis (Fig. 4, UR).

Genital papilla. Ventral shield coniform with almost parallel margins, mouths of ductus ejacula-
torii distinctly longer than ventral shield (Fig. 5, Gen).

Etymology. The new species is dedicated to Prof. Dr. J. W. Wägele for his merits to isopodology and molecular
phylogenetic systematics.

Remark. As described above, the new species was found among the samples of Erophiloscia longistyla,
which was collected at several sites in Colombia. Both species resemble each other, but they can be
determined by the male pleopods as follows: pleopod 1 endopodite of E. waegelei, spec. nov. less
pointed than in E. longistyla, with transverse furrows subapically and caudomedial row of spines
(Fig. 7; PL1-2). Both characters may represent plesiomorphies as regards the ground pattern of the
genus, they are lacking in E. longistyla, which has a mediocaudal row of hyaline lobes delimiting the
spermatic channel. In contrast to the relations in pleopod 1, the endopodite of pleopod 2 in E. waegelei
is the more derived: the obtuse hyaline hooks near the apex are not found in any other species and are
an apomorphic feature of this appendage. In pleopod 5 the protrusion is longer in E. longistyla. Thus,
it can be concluded that in this species the endopodite 2 is longer than in E. waegelei because it certainly
works as a supporting structure for the endopodite 2.

Fig. 1. Erophiloscia waegelei spec. nov. Holotype male. Anl: antennula; An2: antenna with detail of apical organ;
Ctf: cephalothorax in frontal view; Cxp: coxal plates with position of noduli laterales; Had: habitus in dorsal
view; Hal: habitus in lateral view.

Fig. 2. Erophiloscia waegelei, spec. nov. Holotype male. Mdl: left mandible, with detail of pars intermedia;
Mar: right mandible; Mxp: maxilliped with detail of endite in rostral view; Mx1: maxillula with detail of apex
of lateral endite in caudal view; Mx2: maxilla.

Fig. 3. Erophiloscia waegelei, spec. nov. Holotype male. Dac: dactylus 3 in rostral view; PE1-4: pereopods 1-4
(caudal view), with detail of carpus 1 in rostral view; Sb4: tricorn-like seta of basis 4; Sc1: ornamental sensory
spine of carpus 1; Sc2: longest sensory spine of carpus 2; Sp1: distal sensory spine of propus 1.

Fig. 4. Erophiloscia waegelei, spec. nov. Holotype male. PE5-7: pereopods 5-7 (caudal view); Sb5: tricorn-like seta
of basis 5; Sd6: dactylar seta of dactylus 6; Sm7: sensory spine of merus 7; UR: uropod.

PL1-5: pleopods

„

spec. nov. Holotype male. Gen: genital papilla, with detail of apex;

’

1-5, rostral view, with details of endopodite 1 in caudal view; endopodite 2 in rostral view.

Fig. 5. Erophiloscia waegelei

Bei

Cx3

BT On Tel \

50um

Fig. 6. Erophiloscia waegelei, spec. nov. Paratype female. Cx3: coxal plate 3 with detail of nodulus lateralis;
PE7: ischum of perepod 7; PL1-5: pleopod 1, 2, 5 in rostral view; Tel: pleotelson.

100um

PL2

200um

Fig. 7. Erophiloscia longistyla Vandel, 1972. Paratype male. PL1-2: pleopods 1-2, rostral view, with detail of
endopodite 1 in caudal and mediorostral view.

E. waegelei, spec. nov. is the most basal form of this genus in several aspects of cephalothorax
morphology: the linea frontalis is rather distinctive, especially medial from the compound eyes, and
a faint lamina frontalis is present, both characters belonging to the ground pattern of a taxon within
the crinochaete Oniscidea comprising most of the known species.

Erophiloscia longistyla Vandel, 1972
Fig 7

Vandel, 1972:

Types. Lectotype: microscopic slides of 8: Colombia, Bogotä, above calle 71, montane forest, leaf litter, 2750 m
a.s.l. Leg. H. Sturm, 14.02.1969, MNHN Vandel Collection

Remark. This species was described by Vandel (1972) and is well documented. For comparison with
E. waegelei, spec. nov., the male pleopods 1 and 2 are figured (Fig. 7, PL1-2). The male pleopod 1
endopodite lacks the caudomedial row of spines, it is pointed, bearing transverse lamellae at its distal
part. Male pleopod 2 endopodite elongate, more than three times longer than pointed exopodite which
bears about 4 sensory spines laterally near its apex.

Erophiloscia recurvata, spec. nov.
Figs 8-11

Types. Holotype: d, 3 mm, Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station “Pangua-
na” 9°37'S 74°56'W, altitude 250 m, Maniokfeld, 12.-27.XIIL 1975 leg. W. Hanagarth (SNMS coll. T439). — Para-
types: 286, 929, 2 juv, max. 3.5 mm, same data as holotype (SMNS coll. T440); 284, 32? max. 2.5 mm, Cocha,
X.1975-1.1976, leg. W. Hanagarth (SMNS coll. T441).

Description

Colour. Reddish brown with several white spots on the tergites, pleon without light markings,
cephalothorax dorsally with white spots.

Cephalothorax. Linea frontalis lacking, linea supra-antennalis and lamina frontalis present, small
lateral lobes, compound eyes consisiting of seven ommatidia (Fig. 8, Ctf).

Pereon. Tegument smooth and shiny, coxal plates lacking gland pores and sulcus marginalis,
noduli laterales present, long and flagelliform, on coxal plate IV more distantly from lateral margin
(Fig. 8, Cxp).

Pleon. Retracted from pereon, neopleurae visible, pleotelson with rounded distal margin, bearing
some tricorn-like setae (Fig. 8, Tel).

Antennula. Three-articulate, rather stout with distal article bearing prominent tuft of aesthetascs
medially and 2 aesthetascs apically (Fig. 8, Anl).

Antenna. Comparatively slender, length ratio of peduncular articles 1 to 5 1:2:2:4:5, flagellum
three-articulate with articles subequal in length, distal article slightly longer, as long as apical organ
(Fig. 8, An2).

Mandible. Pars molaris consisting of a four-branched molar penicil, pars intermedia bearing two
penicils on left and one on right mandible, intermedial penicil slender (Fig. 9, Mdl/r).

Maxillula. Medial endite bearing two penicils apically, no additional tip discernible. Lateral endite
with 4+4 teeth, inner set cleft (Fig. 9, Mx1).

Maxilla. Lateral lobe two times broader than medial one, bearing scattered trichiform setae and
pectinate scales, medial endite apically with 8 cusps (Fig. 9, Mx2).

Maxilliped. Basipodite with short sulcus lateralis, palp with one seta on proximal article, endite
with small knob-like penicil rostrally and two strong teeth caudally (Fig. 9, Mxp).

Pereopods. Rather similar to the preceding species (Fig. 10, PE1-7), dactylus with short inner claw
and flagelliform dactylar seta (Fig. 10, Dac).

Pleopods. Exopodites rather prominent, bearing laterally 1 to 4 sensory spines, endopodites with
two lobes (Fig. 11, PL1-5).

Sexual dimorphism. Male pereopod 7 ischium bearing only one sensory spine laterally instead of
two in the female. Male pleopod 1 exopodite small, rounded, endopodite long and slender, at
halflength bent laterally and apex turned caudally, row of small spines reduced, medial border of
spermatic furrow crenulate near apex (Fig. 11, PL1). Male pleopod 2 exopodite triangular with one
sensory spine laterally, endopodite slender, bent laterally (Fig. 11, PL2). Male pleopod 5 exopodite with
long protrusion of mediodistal edge, decurved laterally for holding the similarly shaped endopodite
(EN

Uropod. As in generic diagnosis.

Genital papilla. Ventral shield ovate, ductus ejaculatorii not surpassing apex of ventral shield
(Fig. 11, Gen).

Etymology. The species name “recurvata” is latin and means incurved, i.e. the shape of the male copulatory
devices bent laterally.

Remark. This species is more derived with respect to the lack of a linea frontalis and a well accentuated
lamina frontalis, yet Erophiloscia recurvata, spec. nov. is a typical member of its genus with elongate
pleopod 2 endopodite and prolonged tip of pleopod 5 exopodite in the male. The peculiar form of the
male endopodite 1 and the laterally bent distal half of all the copulatory devices are unique among its
congeners. Similar to the other species of Erophiloscia save E. waegelei, spec. nov., the mediocaudal row
of spines on the endopodite 1 is reduced.

Fig. 8. Erophiloscia recurvata, spec. nov. Holotype male. Anl: antennula; An2: antenna with detail of apical
organ; Ctf: cephalothorax in frontal view; Cxp: coxal plates with position of noduli laterales; Cx3: coxal plate 3;
Had: habitus in dorsal view; Hal: habitus in lateral view; Tel: pleotelson.

Fig. 9. Erophiloscia recurvata, spec. nov. Holotype male. Mdl: left mandible, with detail of pars intermedia;
Mdr: right mandible; Mxp: maxilliped with detail of endite in rostral view; Mx1: maxillula with detail of apex
of lateral endite in caudal view; Mx2: maxillula.

PEI

Fig. 10. Erophiloscia recurvata, spec. nov. Holotype male. Dac: dactylus 1 in rostral view; PE1-7: pereopods 1, 6,
7 in caudal view, with detail of carpus 1 in rostral view; Scl: ornamental and longest sensory spine of
carpus 1; Sc7: sensory spine of carpus 7; Spl: distal sensory spine of propus 1.

Fig. 11. Erophiloscia recurvata, spec. nov. Holotype male. Gen: genital papilla; PL1-5: pleopods 1-5, rostral view,
with details of endopodite 1 in caudal view.

Erophiloscia acanthifera, spec. nov.
Figs 12-16

Types. Holotype: d, 4mm, Peru, Dept. Huanuco; Distr. Puerto Inca, Rio Yuyapichis, Biological station “Pangua-
na” 9°37'S 74°56'W, altitude 250 m, “A3” leg. W. Hanagarth (SMNS coll. T442). - Paratypes: 258,5? , 1juv., max.
4 mm, same data as holotype (SMNS coll. T443); 485 2.5-4 mm, A2, 1.X11.1975 leg. W. Hanagarth (SMNS coll.
T444); 238, 2722, 3 juv., max 3.5 mm, Wald, X-X1.1975 leg. W. Hanagarth (SMNS coll. T445).

Description

Colour. Dorsally purplish brown with several prominent white spots on tergites, pereonites 5 to
7 with white medial band, continued on pleonite 1, ventrally whitish.

Cephalothorax. Linea and lamina frontalis lacking, linea supra-antennalis and small lateral lobes
present, compound eyes consisting of about 7 ommatidia (Fig. 12, Hal).

Pereon. Tegument smooth and shiny, coxal plates with sulcus marginalis and long flagelliform
nodulus lateralis, nodulus lateralis of coxal plate IV inserting more disatantly from lateral margin
EisTP2ZEp).

Pleon. Retracted from pereon, rather slender, neopleurae very small, pleotelson with almost
straight margins, bearing several tricorn-like setae (Fig. 12, Tel).

Antennula. As in generic diagnosis, medial tuft of aesthetascs consisting of a fewer number than
in other species (Fig. 12, Anl).

Antenna. Rather short, peduncular articles 4 and 5 shorter than in preceding species, flagellum
three-articulate, distal article two times longer than proximal article, apical organ longer than distal
article (Fig. 12, An2).

Mandible. Similar to preceding species (Fig. 13, Mdl/r).

Maxillula. Medial endite with two penicils and apical tip, lateral endite bearing 4+5 teeth, four of
inner set cleft, the other one short, lateral fringe of trichiform setae stepped (Fig. 13, Mx1).

Maxilla. Lateral lobe slightly broader than medial one, lacking setation, medial lobe with about 10
cusps apically (Fig. 13, Mx2).

Maxilliped. Basipodite with sulcus lateralis, palp with two setal tufts apically, proximal article
bearing two setae, endite with small knob-like penicil rostrally and prominent tooth caudally (Fig. 13,
Mxp).

Pereopods. Slender (Fig. 14, PE1-4, 15, PE5-7), carpus and propus of pereopod 1 with antenna-
grooming brush, ornamental sensory spine of carpus 1 double-fringed serrate (Fig. 14 Scl), dactylus
with short inner claw and interungual seta, dactylar seta simple (Fig. 14, Sd3).

Pleopods. Exopodites rather prominent, bearing 5-6 lateral sensory spines in pleopod 3 and 4,
pleopod 5 exopodite triangular, endopodites more or less bilobate (Fig. 16, PL1-5).

Sexual dimorphism. Pereopods ithout sexual dimorphism. Male pleopod 1 exopodite circular,
small, endopodite slender, spermatic furrow distinctly bordered only on lateral side, apex with
rectangular protrusion, terminated by a proximally directed thorn (Fig. 16, PL1). Pleopod 2 exopodite
similar to preceding species, endopodite extraordinary long, flagelliform, surpassing exopodite more
than two times (Fig. 16, PL2). Therefore, pleopod 5 exopodite strongly drawn out for holding pleopod
2 endopodite (Fig. 16, PL5).

Uropod. As in generic diagnosis (Fig. 15, UR).

Genital papilla. Similar to the preceding species (Fig. 16, Gen).

Etymology. The species name is composed of the greek term “acanthos”, which is “thorn” or “hook” and the latin
verb “ferre”, meaning “to bear”, related to the hook on the apex of pleopod 1 endopodite.

Remark. Similar to the preceding species, Erophiloscia acanthifera, spec. nov. lacks a linea frontalis and
a lamina frontalis. It is best recognised by the shape of the male pleopod 1 endopodites with their hook-
bearing apices, looking like a pair of pliers.

Fig. 12. Erophiloscia acanthifera, spec. nov. Holotype male. Anl: antennula; An2: antenna with detail of apical
organ; Ctf: cephalothorax in frontal view; Cxp; coxal plates with position of noduli laterales; Cx3: coxal plate 3;
Had: habitus in dorsal view; Hal: habitus in lateral view; Tel: pleotelson.

Fig. 13. Erophiloscia acanthifera, spec. nov. Holotype male. Mdl: left mandible, with detail of pars intermedia;
Mdr: right mandible; Mxp: maxilliped with detail of endite in rostral view; Mx1: maxillula with detail of apex
of lateral endite in rostral view; Mx2: maxilla.

Dac m

Sd3

Spl
Scl

Fig. 14. Erophiloscia acanthifera, spec. nov. Holotype male. Dac: dactylus 1 in rostral view; PE1-4: pereopods 1-4
in caudal view, with detail of carpus 1 in rostral view; Sc1: ornamental sensory and second longest sensory spine
of carpus 1; Sd3: dactylar seta of dactylus 3; Sp1: distal sensory spine of propus 1.

Fig. 15. Erophiloscia acanthifera, spec. nov. Holotype male. PE5-7: pereopods 5-7 in caudal view; Sc5: sensory
spine of carpus 5; Sm7: sensory spine of merus 7; UR: uropod in rostral view.

HH
100um

Fig. 16. Erophiloscia acanthifera, spec. nov. Holotype male. Gen: genital papilla; PL1-5: pleopods 1-5, rostral
view, with details of endopodite 1 in caudal view.

Andenoniscus
Erophiloscia waegelei

Mm Erophiloscia longistyla

M2 Erophiloscia recurvata

M3 Erophiloscia acanthifera

Fig. 17. Phylogenetic relationships within the genus Erophiloscia Vandel, 1972. M1: male pleopod 5 extraordi-
narily drawn out, no guide slot [male pleopod 5 subtrinagular without distal extension]. M2: caudal row of
spines on male pleopod 1 endopodite reduced [caudal row of spines present]. M3: linea frontalis reduced [linea
frontalis present].

Phylogeny and Biogeography

The genus Erophiloscia Vandel, 1972 is one of several small “philosciid” genera of the Andean region.
Its closest relatives are the genera Andenoniscus Verhoeff, 1941, Xiphoniscus Vandel, 1968 and a hitherto
undescribed genus from Peru, Venezuela and Amazonian Brazil. All these genera are characterized by
the presence of long, flagelliform noduli laterales, with the fourth inserted more distantly from the
lateral margin, the antennula with divergent sets of aesthetascs and the small size with compound eyes
composed of about 6 to 8 ommatidia. Vandel (1972) stated that the long male pleopod 1 endopodite
is a character of generic importance —- an autapomorphy of Erophiloscia — but this is only partially true.
It is comparatively slender. It is the pleopod 2 endopodite which is long and flagelliform. It may reach
caudally beyond the pleotelson. To protect this copulatory device, the medial margin of pleopod 5
exopodite is straight and distally drawn out, holding the endopodite 2 ventrally on this enlargement.
In Erophiloscia recurvata, the mediodistal extension of exopodite 5 is as bent laterally as the pleopod 2
endopodite. The specific structure of the male pleopod 5 exopodite is an autapomorphy of the genus
Erophilosica (Fig. 17, character Ml). In contrast to Vandel’s (1972) statement, half of its species has
pleopod 1 endopodites of unspectacular length. Superficially the structure of pleopod 5 exopodite
resembles the one of several species of Chaetophiloscia Verhoeff, 1908, or Natalscia longistylata Ferrara &
Taiti, 1985, but the fine structure is different: Chaetophiloscia has a guide slot caudally for the endo-
podite 1 resembling the one described by Legrand (1946). This structure is missing in Erophiloscia. These
genera differ in other characters, too: the shape of the antennula, the shape of the ornamental sensory
spine of the carpus 1, the shape of the noduli laterales as can be evidenced from the redefinition of
Chaetophiloscia by Schmalfuß (1990).

In the ground plan of Erophiloscia, the linea frontalis and lamina frontalis is present, as is the row
of small spines on the male pleopod 1 endopodite. They have subsequently been reduced in all species
save E. waegelei. In all the other species, the caudal row of spines on pleopod 1 endopodite is reduced
(Fig. 17, character M2). The type species, E. longistyla is the sister species of an adelphotaxon composed
of E. recurvata and E. acanthifera. The latter have in common a three-tipped distal sensory spine on
carpus 1, the linea frontalis completely reduced, no hyaline lobes on the male pleopod 1 endopodite
(Fig. 17, character set M3).

The two most derived species live in the eastern slopes of the Andes in Peru. The area of
distribution of their common ancestor and of E. longistyla might be fragmented by the uplift of the
Andes in the Tertiary (e.g. Simpson & Haffer 1978). Since E. recurvata and E. acanthifera now occur
sympatrically, their distributional ranges now overlap due to dispersal once speciation had happened.
The distribution of several small philosciids like Andenoniscus and Xiphoniscus within the Andes is,
biogeographically spoken, not “Andean”. The Andean subregion is restricted to the higher parts of the
mountain ranges, the Param6 zone. This area has stong affinities to the temperate southern South
America (Morrone 1992). As can be evidenced from their distributional data (Vandel 1968, 1972,
Verhoeff 1941), they are found in the tropical to subtropical lowlands. These tropical lowlands are part
of the Neotropical realm. This neotropical distribution is not surprising, since their closest relatives, the
members of Prosekia-group are distributed in the Amazon basin and Venezuela. Further collections in
the intervening area will most probably reveal more records or even undescribed species of this genus.

Leistikow (1998b) reported to some material ascribed to Pentoniscus pruinosus Richardson, 1913 from
Costa Rica as a possible undescribed species of Erophiloscia. There is some evidence that this species
could belong to a closely related new genus which is distributed from the Amazon basin north to
Guatemala (pers. obs.). From our point of knowledge, Erophiloscia is purely East-Andean in distribu-
tion, thus occurring in the westernmost parts of the Amazonian biogeographic subregion.

Key to the species

ie lıneaßtrontalisipresentunse. 1 antenne en m a els 2.

nltimeatt:ontalistredueedirn en 3

2. Male pleopod 1 endopodite with caudal row of spines, endopodite 2 with some hooks near the apex
Oboe RE BER LE EEE LE RER re E. waegelei, spec. noV.

— Male pleopod 1 endopodite without caudal row of spines, endopodite 2 without hooks near the
ED E. longistyla Vandel

3. Male pleopod 1 and 2 endopodites and 5 exopodite strongly bent laterally ............neee-
BEER LEBER BEE ERSER ER ÄEESESE Erchten nen teasecen his sssae ann snaetesaiaksuse nen ee E. recurvata, spec. nov.

- Male pleopods straight, endopodites 1 plier-like .................nee E. acanthifera, spec. nov.

Acknowledgements

The author thanks Dr. H. Dalens, Universite de Toulouse, for the loan of the material, the permission to dissect
a specimen and the critical review of the manuscript. He is indebted to Dr. H. Schmalfuss, Staatliches Museum
für Naturkunde for the possibility to examine and describe the species of the Hanagarth collection and Prof. Dr.
J. W. Wägele for his support of this investigation and the possibility to discuss on this work. For manuscript
revision he also is grateful to Dr. A. Ohlers and Dr. S. Taiti.

References

Legrand, J. J. 1946. Les coaptations sexuelles des Oniscoidea. — Bull. biol. France Belg. 80: 241-388

Leistikow, A. 1998a. Redescriptions of terrestrial Isopoda from Chile and Peru (Crustacea: Isopoda: Oniscidea).
— Spixiana 21(3): 215-225

-- 1998b. Considerations about the genus Pentoniscus Richardson, 1913 (Crustacea: Isopoda: Oniscidea) with
description of a new species. — J. Nat. Hist. 32: 1339-1355

Morrone. J. J. 1992. The biogeographical Andean subregion: a proposal exemplified by Arthropod taxa (Arach-
nida, Crustacea, Tracheata). -— Neotropica 42: 103-114

Schmalfuss, H. 1980. A revision of the neotropical genus Ischioscia Verhoeff, with description of four new species.
— Stud. neotrop. Fauna Environm. 15: 125-139

-- 1990. Die Landisopoden Griechenlands. 11. Beitrag: Gattung Chaetophiloscia. —- Rev. suisse Zool. 97: 169-193

Simpson, B. B. & J. Haffer 1978. Speciation patterns in the Amazonian forest biota. - Annu. Rev. Ecol. Syst. 9:
497-518

Vandel, A. 1968. Isopodes terrestres. - in: N. and ]. Leleup (ed.). Mission zoologique belge aux Iles de Galapagos
et Ecuador 84: 35-168

-- 1972. Les isopodes terrestres de la Colombie. — Stud. neotrop. Fauna Environm. 7: 147-172

Verhoeff, K.W. 1941. Land-Isopoden. - in: E. Titschack (ed.). Beiträge zur Fauna Perus. 1 (2. Lieferung): 73-80

Buchbesprechungen

5. Bugledich, E.-M. A.: Diptera: Nematocera. In A. Wells & W. W. K. Houston (eds.) Zoological Catalogue of
Australia. Vol. 30.1. - CSIRO Publishing, Melbourne, Australia, 1999. xiii + 627 pp. ISBN 0-643-06489-3.

Dieser ausgesprochen benutzerfreundliche und informative Katalog behandelt die Nematoceren (Mücken) des
Australischen Kontinents einschließlich der Lord Howe Insel, Norfolk Insel, Cocos Inseln, Christmas Insel,
Ashmore and Cartier Inseln, Macquarie Insel, Heard und McDonald Inseln, und des autralischen Teils der
Antarktis. Für die aus diesem Bereich bekannten Arten bietet er nicht nur alle wünschenswerten taxonomischen
Angaben (Stand: Ende Dezember 1997), sondern auch kurze Angaben zur geographischen Verbreitung und
Ökologie, oft auch weiterführende Literaturzitate.

Für jede Familie gibt es eine kleine Einführung mit Angaben zu Trivialnamen, typischen Merkmalen der
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nung, und ein paar generellen Literaturhinweisen. Bei den spezifischen Angaben sind in der üblichen Weise
innerhalb der Unterfamilien und Triben die Gattungen, und innerhalb dieser die Arten, alphabetisch geordnet.
Synonyme für Gattungen und Arten sind in chronologischer Reihenfolge aufgelistet. Für jeden der verfügbaren
Gattungs- und Artnamen (auch für die Synonyme) ist an Ort und Stelle das vollständige Zitat der Erstveröffentli-
chung angegeben, sowie das Typusmaterial, sein Aufbewahrungsort, und der Locus typicus. Für taxonomische
Entscheidungen, z.B. Synonymisierungen, wird ebenfalls das vollständige Zitat mit Angabe der betreffenden
Seite angeführt. Die geopraphische Verbreitung wird für Arten immer angegeben, für Gattungen nur insofern
sie über die Grenzen des Katalogs hinausgeht. Schließlich folgen für jede Art ein paar Schlagworte zur Ökologie,
teilweise auch umfangreichere Angaben, und manchmal noch eine weiterführende Literaturangabe. Trotz der
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Zusatzinformationen zur Ökologie, bleibt das fest gebundene Buch als solches vom Umfang her noch schön
stabil und handlich.

Drei weitere Dipteren-Bände sind geplant, die Brachycera, Cyclorrhapha: Lonchopteroidea bis Brauloidea,
und Cyclorrhapha: Muscoidea, behandeln sollen. M. Kotrba

6. Detzel, P.: Die Heuschrecken Baden-Württembergs. - Ulmer Verl., Stuttgart, 1998. 580 S., 222 Farbfotos, 132
Verbreitungskarten, 137 Grafiken, 51 Tabellen. ISBN 3-8001-3507-8.

Mit dem vorliegenden Band ist dem Autor unter Mithilfe zahlreicher weiterer namhafter Spezialisten auf dem
Gebiet der Orthopterologie ein sehr umfangreiches Standardwerk gelungen.

Der erste allgemeine Teil des Buches befaßt sich mit Nomenklatur, Zoogeographie, Biologie, Ökologie und
den Lebensräumen von Heuschrecken. Im zweiten, speziellen Teil werden 70 einheimische Heuschreckenarten
(Ensifera und Caelifera) sowie die in Baden-Württemberg einheimische Mantis religiosa ausführlich dargestellt.
Für jede der Arten steht ein eigenes Kapitel mit kurzer morphologischer Beschreibung sowie ihrer Verbreitung
in Eurasien, Deutschland und Baden-Württemberg inkl. Verbreitungskarten. Desweiteren findet man bei den
Einzelartbeschreibungen ausführliche Erläuterungen zur Biologie (Nahrungs- und Fortpflanzungsbiologie) und
zur Ökologie, ihren Gefährdungsstatus (Rote Liste Baden-Württemberg und naturräumliche Rote Liste) mit
Hinweisen zu Schutz- und Pflegemaßnahmen.

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den. Trotz der Spezialisierung auf die baden-württembergische Heuschreckenfauna ist das Werk über die
Grenzen Baden-Württembergs hinaus interessant, da nur wenige, der in Deutschland autochthonen Arten, wie
auf den Alpenraum beschränkte Spezies (Aeropus sibiricus, Bryodema tuberculata, Epacromius tergestinus, Chorthip-
pus pullus, Pholidoptera aptera, Tetrix tuerki) bzw. ausgesprochene Steppenarten (Arcyptera microptera, Gampsocleis
glabra, Platycleis montana, Stenobothrus crassipes) oder die lokal beschränkte Tettigonia caudata nicht näher be-
schrieben sind. Man darf den Autoren zu diesem gelungenen Buch, das seinem Preis wert ist, gratulieren und
kann dem Buch nur noch viele weitere Leser wünschen. M. Breitsameter

SPIXIANA 53-72 München, 01. März 2001 ISSN 0341-8391

Further new and rare species of the genera Fortagonum Darlington
and Collagonum Baehr from New Guinea

(Insecta, Coleoptera, Carabidae, Agoninae)*
Martin Baehr

Baehr, M. (2001): Further new and rare species of the genera Fortagonum Dar-
lington and Collagonum Baehr from New Guinea (Insecta, Coleoptera, Carabidae,
Agoninae). — Spixiana 24/1: 53-72

Fortagonum hornabrookianum, spec. nov., Fortagonum substriatum, spec. nov., Colla-
gonum thoracicum, spec. nov., all from central Papua New Guinea, Fortagonum
insulare, spec. nov. from Japen Island, and Fortagonum laevissimum, spec. nov. and
Collagonum longipenne, spec. nov., both from Star Mountains, westernmost Papua
New Guinea are described. For both genera Fortagonum and Collagonum new keys
are provided that replace the most recent keys to the respective genera (Baehr 1995,
1998). For Collagonum thoracicum, spec. nov. a new subgenus Procollagonum, and for
Collagonum distortum (Darlington) and Collagonum limum (Darlington) a new sub-
genus Paracollagonum is erected. In view of striking differences in external morphol-
ogy, but in particular in shape of the male aedeagus, Procollagonum is the most
plesiotypic member of Collagonum, whereas Paracollagoum combines plesiomorphic
character states in structure of the male genitalia with highly apomorphie character
states in external morphology.

Dr. Martin Baehr, Zoologische Staatssammlung, Münchhausenstr. 21, D-81247
München, Germany.

Introduction

Recently Dr. R. W. Hornabrook (Wellington, New Zealand) kindly sent me a sample of carabid beetles
from New Guinea (mainly Papua New Guinea) for identification. Although they were collected 25-30
years ago, they were no more included in Darlington’s (1968, 1971) monograph about the Carabidae
of New Guinea. The sample includes a number of new records and new species, inter alia two new
species of the genus Fortagonum Darlington and one new species of the genus Collagonum Baehr that
are being described below. Material received recently from Mr. A. Riedel (München), the indefatigable
and unexcelled collector of rare species from Irian Jaya, and from Mr. P. Schüle, (Düsseldorf) included
three additional new taxa of the genus Fortagonum that are added to this paper.

The genus Fortagonum was founded by Darlington (1952). In a supplementary volume, Darlington
(1971) described several additional species of very different shape and structure. Baehr (1992) again
described additional species, and Baehr (1995) redefined the genus and erected a new genus Collagonum
for some species hitherto included in Fortagonum. Most recently Baehr (1998) again described addition-
al species of Fortagonum and gave a revised key to this genus. The most recent key to the species of
Collagonum was given by Baehr (1995).

* In part results of the scientific collections of A. Riedel in New Guinea 1998.

Measurements

Measurements were made under a stereo microscope using an ocular micrometer. Length has been
measured from tip of labrum to apex of elytra, hence, measurements may slightly differ from those of
Darlington. Length of pronotum for width/length ratio has been measured from middle of apex to
base.

Location of types

The types are shared with the Museum of New Zealand, Wellington (MNZ) and Zoologische Staatssa-
mmlung, München (ZSM). Some paratypes are located in the working collection of the author at
Zoologische Staatssammlung (CBM).

Fortagonum insulare, spec. nov.
Fig. 6

Types. Holotype: 9, 5.8.1996 41 Schüle/Stüben West Papua 1000 m Japen Serui nach Ambeidiru km 9 Primär-
wald (ZSM). — Paratype: 17, same data (CBM).

Diagnosis. Distinguished from its nearest relative, F. spinipenne Baehr, by light colour of surface, legs,
and antennae, and by basally wider pronotum that also has less projecting anterior angles.

Description

Measurements. Length: 10.4-10.6 mm; width: 4.10-4.25 mm. Ratios. Width/length of pronotum:
1.48-1.52; width base/apex of pronotum: 1.68-1.74; width pronotum/head: 1.85-1.96; width elytra/
pronotum: 1.64-1.68; length/width of elytra: 1.29-1.37.

Wing-and-seta formula: +w;, 4 4--;-++.

Colour. Dark reddish piceous, lateral margin of pronotum, suture of elytra, labrum, and mandibles
slightly lighter. Mouth parts, antenna, tibiae, and tarsi light reddish, femora slightly darker. Lower
surface piceous.

Head. Rather narrow compared with prothorax. Neck rather wide, somewhat imbedded in pro-
thorax. Eyes fairly large, laterally moderately projecting, orbits distinct, evenly curved. Clypeal suture
distinct. Labrum rectangular, apex feebly concave. Mandibles elongate, straight, but not porrect.
Antenna very slender and elongate, surpassing base of pronotum by about four antennomeres, median
antennomeres almost 5x as long as wide. Both palpi slender and elongate, basal palpomere of
maxillary palpus thickened. Mentum with an elongate, unidentate tooth. No furrow medially of eyes,
though a shallow furrow above antennal base present. Either both supraocular setae present or only
the posterior supraocular seta present, the latter situated at posterior margin of eye. Clypeus and
anterior part of frons with short, shallow, parallel furrow on either side, frons evenly convex, absolutely
smooth. Microreticulation isodiametric, somewhat superficial. Surface glossy.

Prothorax. Comparatively narrow, somewhat conical, widest at posterior third, laterally evenly
though feebly convex, strongly narrowed to apex, moderately narrowed to base. Disk slightly convex,
lateral margins widely explanate though barely separated from disk. Anterior angles rather projecting,
obtuse at apex. Apex regularly and deeply excised. Basal angles rectangular, at apex obtusely rounded.
Base laterally straight, in middle very faintly produced. Disk convex with extremely shallow, v-shaped
sulcus in apical fourth, base near basal margin with a rather deep, oblique, somewaht linear impression
on either side and with a very shallow transverse impression. Median line incomplete, very fine,
ending far from apex and base. Apex distinctly bordered, lateral margin and base not bordered. Both
marginal setae absent. Disk impunctate. Microreticulation very fine, absent on disk, near apex and base
highly superficial, isodiametric. Surface glossy.

at 2" stria. 17-18 marginal setae and 1 preapical seta at 7‘ stria present, humeral group of marginal
series consisting of 6 setae, median and apical pores not much more conspicuous than basal pores,
series slightly interrupted in middle. Intervals impunctate. Microreticulation almost wanting. Surface
highly glossy, rather iridescent. Posterior wings present.

Lower surface. Prosternal process short, posteriorly slightly convex, triangular, ventrolaterally
and posterolaterally bordered. Proepisternum smooth. Mesepisternum coarsely punctate. Metepister-
num moderately elongate, c. 1.5 x as long as wide at anterior border. Epipleura anteriorly moderately
wide, rugose. Abdomen impunctate, though laterally with several fine, elongate wrinkles and shallow
impressions. Microreticulation dense, isodiametric, very superficial. d sternum VII unknown, % ster-
num VII quadrisetose, apex regularly curved.

Legs. Very elongate and slender. 4th tarsomere medially faintly excised. 5th tarsomere asetose
beneath. Vestiture of d anterior tarsus unknown.

d genitalia. Unknown.

? genitalia. Stylomere 2 rather elongate, little curved, with obtuse apex, with 3 fairly small ventral
ensiform setae, a dorsal ensiform seta situated about in middle, and one nematiform seta in a deep
furrow moderately close to apex. Apex of stylomere 1 ventrally with 7-8 setae near base of stylomere
2. Lateral plate with 8-9 setae at or near margin.

Variation. Rather similar though the paratype has slightly wider pronotum and elytra, and it
possesses only the posterior supraorbital seta, which is the first instance of instability of chetaotaxy of
supraorbital setae ever recorded in the genus.

Distribution. Japen Island, western central Irian Jaya. Known only from type locality.

Collecting circumstances. Largely unknown. Both specimens collected in “primary forest” at median
altitude, probably under log or in litter on the ground.

Etymology. The name refers to the occurrence on Japen Island.

Relationships. Certainly this species is very closely related to F. spinipenne Baehr form mainland
western central Irian Jaya that externally looks extremely similar. Unfortunately, the male genitalia are
thus far unknown in both species. Because in the species group to which both species belong aedeagi
usually are highly distinctive, reference to the aedeagi when they are known may settle the question
of full specific or of subspecific status of F. insulare.

Fortagonum substriatum, spec. nov.
Bies0lW7

Types. Holotype: d, Papua NG, Morobe-Pr. Mindik 1400-1500 m 27.4.1998, A. Riedel (ZSM). — Paratype: 1%,
same data (CBM).

Diagnosis. Distinguished by presence of wings, absence of anterior supraocular seta, both pronotal
setae, and anterior discal seta, rather narrow, fairly conical pronotum, narrow and elongate elytra, and
markedly elongate elytral spine opposite 3'% stria. Distinguished from most closely related species
F. spinipenne Baehr and F. insulare, spec. nov. by colouration, longer elytra, shallow, almost impunctate
striation and the many sclerotizations in the internal sac.

Description

Measurements. Length: 11.1-11.7 mm; width: 4.45-4.65 mm. Ratios. Width/length of pronotum:
1.44-1.46; width base/apex of pronotum: 1.60-1.65; width pronotum/head: 1.86-1.90; width elytra/
pronotum: 1.62-1.66; length/ width of elytra: 1.35-1.38.

Wing-and-seta formula: + w;- 4; --;- ++.

Colour. Glossy black, elytra with very faint greenish lustre. Labrum, mouth parts, antenna, and
median and posterior tarsi piceous, three basal antennomeres black, anterior tarsi reddish. Lower
surface black.

Fig. 1. Fortagonum substriatum, spec. nov. d genitalia. Scale: 0.5 mm.

Antenna very slender and elongate, surpassing base of pronotum by more than four antennomeres,
median antennomeres >5x as long as wide. Both palpi slender and elongate, basal palpomere of
maxillary palpus thickened. Mentum with an elongate, unidentate tooth. No furrow medially of eyes,
though a shallow furrow above antennal base present. Only posterior supraocular seta present, at
posterior margin of eye. Clypeus and anterior part of frons with short, shallow, parallel furrow on
either side, frons evenly convex, absolutely smooth. Microreticulation isodiametric, somewhat super-
ficial. Surface glossy.

Prothorax. Comparatively narrow, somewhat conical, widest in posterior third, laterally evenly
though feebly convex, strongly narrowed to apex, moderately narrowed to base. Disk slightly convex,
lateral margins widely explanate though barely separated from disk. Anterior angles rather projecting,
obtuse at apex. Apex regularly and deeply excised. Basal angles rectangular, at apex obtusely rounded.
Base laterally straight, in middle very faintly produced. Disk convex with extremely shallow, v-shaped
sulcus in apical fourth, base near basal margin with a rather deep, circular impression on either side
and with a very shallow transverse impression. Median line incomplete, very fine, ending far from apex
and base. Apex distinctly bordered, lateral margin and base not bordered. Both marginal setae absent.
Disk impunctate. Microreticulation very fine, absent on disk, near apex and base highly superficial,
isodiametric. Surface glossy.

Elytra. Comparatively narrow and elongate, dorsal surface markedly convex, lateral borders in
middle almost straight. Preapical sinuosity extremely feeble. Widest diameter about in middle. Humeri
wide, obtusely angulate but not dentate, apex spinose with elongate spine opposite 3° interval, spines
rather widely separated. Sutural angle with minute denticle. Striae very shallow, deepened towards
apex, extremely finely, almost invisibly punctulate, intervals depressed. Anterior discal seta absent,
both median and posterior setae situated at 2" stria. 17-18 marginal setae and 1 preapical seta at 7" stria
present, humeral group of marginal series consisting of 6 setae, median and apical pores not much
more conspicuous than basal pores, series slightly interrupted in middle. Intervals impunctate. Micro-
reticulation almost wanting. Surface highly glossy, rather iridescent. Wings present.

Lower surface. Prosternal process short, posteriorly slightly convex, triangular, ventrolaterally
and posterolaterally bordered. Proepisternum smooth. Mesepisternum coarsely punctate. Metepister-

num moderately elongate, c. 1.5 x as long as wide at anterior border. Epipleura anteriorly moderately
wide, rugose. Abdomen impunctate, though laterally with several fine, elongate wrinkles and shallow
impressions. Microreticulation dense, isodiametric, very superficial. d sternum VII bisetose, ? sternum
VII quadrisetose, apex regularly curved.

Legs. Very elongate and slender. 4th tarsomere medially faintly excised. 5th tarsomere asetose
beneath. 1-3’ tarsomeres of d anterior tarsus biseriately squamose.

d genitalia. Genital ring narrow, rather parallel, at apex asymmetric. Aedeagus stout, rather
symmetric, lower surface very faintly bisinuate. Apex acute and rather short, lateral margins near apex
faintly concave. Internal sac with four denticulate sclerites: one elongate, about quadridentate rod
apically at bottom of right side, two tridentate plates basally at roof on right and left sides, respectively,
and a small unidentate plate in middle. Both parameres wide, at apex very faintly angulate.

? genitalia. Stylomere 2 rather elongate, little curved, with obtuse apex, with 3 fairly small ventral
ensiform setae, a dorsal ensiform seta situated about in middle, and one nematiform seta in a deep
furrow moderately close to apex. Apex of stylomere 1 ventrally with 6-8 setae near base of stylomere
2. Lateral plate with 6-8 setae at or near margin.

Variation. Apart of some minor differences in shape of pronotum little variation noted.

Distribution. Eastern central Papua New Guinea. Known only from type locality.
Collecting circumstances. Unknown. Presumably collected under log in rain forest at median altitude.
Etymology. The name refers to the light striation of the elytra.

Relationships. This species is presumably most closely related to F. spinipenne Baehr, F. insulare, spec.
nov., and F. subconicolle Darlington, though is distinguished by narrower elytra and very shallow,
almost impunctate elytral striae.

Fortagonum hornabrookianum, spec. nov.
Figs 2, 8

Types. Holotype: d, Garaina Morobe District NE. New Guinea Hornabrook (MNZ,).

Diagnosis. Characterized by presence of wings, absence of anterior supraocular seta, both pronotal
setae, and anterior discal seta, rather narrow, fairly conical pronotum, narrow and elongate elytra, and
markedly elongate elytral spine opposite 3"! stria. Distinguished from most closely related species
F. spinipenne Baehr and F. insulare, spec. nov. by violaceous lustre of elytra, approached elytral spines,
and almost impunctate elytral striae.

Description

Measurements. Length: 11.0 mm; width: 4.4 mm. Ratios. Width/length of pronotum: 1.45; width
base/ apex of pronotum: 1.53; width pronotum/head: 1.86; width elytra/ pronotum: 1.64; length/ width
of elytra: 1.29.

Wing-and-seta formula: +w; - 4 --;- ++.

Colour. Dark piceous black, elytra with slight violaceous lustre. Lateral margins of pronotum
faintly reddish translucent, labrum, mouth parts, antenna, and tarsi dark reddish-piceous, antenna
from 3°! antennomere reddish. Femora and tibiae piceous. Lower surface black.

Head. Moderately narrow compared with prothorax. Neck rather wide, somewhat imbedded in
prothorax. Eyes fairly large, laterally moderately projecting, orbits distinct, evenly curved. Clypeal
suture distinct. Labrum rectangular, apex feebly concave. Mandibles elongate, straight, but not porrect.
Antenna very slender and elongate, surpassing base of pronotum by about four antennomeres, median
antennomeres c. 5x as long as wide. Both palpi slender and elongate, basal palpomere of maxillary
palpus thickened. Mentum with an elongate, unidentate tooth. No furrow medially of eyes, though a
shallow furrow above antennal base present. Only posterior supraocular seta present, at posterior
margin of eye. Clypeus and anterior part of frons with short, shallow, parallel furrow on either side,
frons evenly convex, absolutely smooth. Microreticulation isodiametric, extremely superficial. Surface
highly glossy.

Prothorax. Comparatively narrow, somewhat conical, widest at posterior third, laterally evenly

Fig. 2. F. hornabrookianum, spec. nov. d genitalia. Scale: 0.5 mm.

though feebly convex, strongly narrowed to apex, moderately narrowed to base. Disk slightly convex,
lateral margins widely explanate though barely separated from disk. Anterior angles rather projecting,
obtuse at apex. Apex regularly and deeply excised. Basal angles rectangular, at apex obtusely rounded.
Base laterally straight, in middle very faintly produced. Disk convex with extremely shallow, v-shaped
sulcus in apical fourth, base near basal margin with a rather deep, oblique, slightly longitudinal
impression on either side and with a very shallow transverse impression. Median line incomplete, very
fine, ending far from apex and base. Apex distinctly bordered, lateral margin and base not bordered.
Both marginal setae absent. Disk impunctate. Microreticulation very fine, absent on disk, near apex and
base highly superficial, isodiametric. Surface highly glossy.

Elytra. Rather narrow and elongate, dorsal surface markedly convex, lateral borders in middle
almost straight. Preapical sinuosity extremely feeble. Widest diameter about in middle. Humeri wide,
obtusely angulate but not dentate, apex spinose with elongate spine opposite 3" interval, though rather
approached. Sutural angle with minute denticle. Striae shallow, deepened towards apex, only inner
four striae extremely finely punctulate, puncturation barely recognizable, intervals depressed. Anterior
discal seta absent, both median and posterior setae situated at 2” stria. 18-19 marginal setae and 1
preapical seta at 7" stria present, humeral group of marginal series consisting of 6 setae, median and
apical pores not much more conspicuous than basal pores, series slightly interrupted in middle.
Intervals impunctate. Microreticulation wanting. Surface highly glossy, iridescent. Posterior wings
present.

Lower surface. Prosternal process short, posteriorly slightly convex, triangular, ventrolaterally
and posterolaterally bordered. Proepisternum smooth. Mesepisternum coarsely punctate. Metepister-
num moderately elongate, c. 1.5 x as long as wide at anterior border. Epipleura anteriorly moderately
wide, rugose. Abdomen impunctate, though laterally with several fine, elongate wrinkles and shallow
impressions. Microreticulation dense, isodiametric, very superficial. d sternum VII bisetose, ? sternum
unknown, apex regularly curved.

Legs. Very elongate and slender. 4th tarsomere medially faintly excised. 5th tarsomere asetose
beneath. 1-3" tarsomeres of d anterior tarsus biseriately squamose.

ö genitalia. Genital ring rather parallel, at apex highly asymmetric, apex wide. Aedeagus fairly
elongate, rather symmetric, lower surface very faintly concave. Apex acute, moderately elongate,
straight. Internal sac on right side at bottom near apex with a small, elongate, faintly tridentate
sclerotized plate. Both parameres rather wide, at apex evenly convex.

? genitalia. Unknown.

Variation. Unknown.

Fig. 3. F. laevissimum, spec. nov. d genitalia. Scale: 0.5 mm.

Distribution. Eastern central Papua New Guinea. Known only from type locality.
Collecting circumstances. Unknown. Presumably collected under log in rain forest of median altitude.
Etymology. The name is an acronym in honour of the collector of this and additional species.

Relationships. This species certainly belongs to the bisetosiceps-group, but occupies a somewhat
isolated position due to the markedly approached elytral spines.

Fortagonum laevissimum, spec. nov.
Eies 3,9

Types. Holotype: 8, Busalmin 5000 ft. Star Mts. 16/10/75 Papua New Guinea R. W. Hornabrook (MNZ.).

Diagnosis. Distinguished by presence of wings, absence of anterior supraocular seta, both pronotal
setae, and anterior discal seta, fairly narrow and elongate elytra, and elongate elytral spine opposite
3'4 stria. Distinguished from other species of the bisetosiceps-group by colouration, wide, triangular
pronotum, laevigate elytra, and almost unarmed aedeagus.

Description

Measurements. Length: 11.1 mm; width: 4.6 mm. Ratios. Width/length of pronotum: 1.60; width
base/apex of pronotum: 1.78; width pronotum /head: 2.10; width elytra/ pronotum: 1.59; length/ width
of elytra: 1.24.

Wing-and-seta formula: + w;,-#; --;- ++.

Colour. Reddish-piceous, elytra with distinct greenish lustre. Palpi, antenna, and tarsi light red-
dish, 3° antennomere apically slightly darker, femora and tibae piceous. Lower surface piceous.

Head. Narrow compared with prothorax. Neck rather wide, somewhat imbedded in prothorax. Eyes
fairly large, laterally moderately projecting, orbits distinct, evenly curved. Clypeal suture distinct.
Labrum rectangular, apex feebly concave. Mandibles elongate, straight, but not porrect. Antenna
slender and elongate, surpassing base of pronotum by slightly <4 antennomeres, median antennomeres
c.4.5x as long as wide. Both palpi slender and elongate, basal palpomere of maxillary palpus thickened.
Mentum with an elongate, unidentate tooth. No furrow medially of eyes, though a shallow furrow
above antennal base present. Only posterior supraocular seta present, at posterior margin of eye.
Clypeus and anterior part of frons with short, shallow, parallel furrow on either side, frons evenly

convex, absolutely smooth. Microreticulation extremely superficial, isodiametric. Surface highly glossy.

Prothorax. Wide, markedly triagonal, widest near base, laterally evenly though feebly convex,
strongly narrowed to apex, barely narrowed to base. Disk slightly convex, lateral margins widely
explanate though barely separated from disk. Anterior angles rather projecting, obtuse at apex. Apex
regularly and deeply excised. Basal angles rectangular, at apex obtusely rounded. Base laterally
straight, in middle very faintly produced. Disk convex with extremely shallow, v-shaped sulcus in
apical fourth, base near basal margin with a rather deep, circular impression on either side and with
a very shallow transverse impression. Median line incomplete, very fine, ending far from apex and
base. Apex distinctly bordered, lateral margin and base not bordered. Both marginal setae absent. Disk
impunctate. Microreticulation absent on disk, near apex and base very fine, highly superficial, isodi-
ametric. Surface highly glossy.

Elytra. Comparatively short and wide, dorsal surface markedly convex, lateral borders in middle
almost straight. Preapical sinuosity extremely feeble. Widest diameter about in middle. Humeri wide,
obtusely angulate but not dentate, apex spinose with comparatively short spine opposite 3"4 interval,
spines rather widely separated. Sutural angle with minute denticle. Striae not impressed, extremely
superficial, marked as rows of extremely fine punctures, striae not deepened towards apex, intervals
depressed. Anterior discal seta absent, both median and posterior setae situated at 2nd stria, punctures
somewhat foveate. 16 marginal setae and 1 preapical seta at 7" stria present, humeral group of marginal
series consisting of 6 setae, median and apical pores not much more conspicuous than basal pores,
rather foveate, series slightly interrupted in middle. Intervals impunctate. Microreticulation absent.
Surface highly glossy, rather iridescent. Wings present.

Lower surface. Prosternal process short, posteriorly slightly convex, triangular, ventrolaterally
and posterolaterally bordered. Proepisternum smooth. Mesepisternum slightly punctate. Metepister-
num moderately elongate, <1.5x as long as wide at anterior border. Epipleura anteriorly moderately
wide, rugose. Abdomen impunctate, though laterally with several fine, elongate wrinkles and shallow
impressions. Microreticulation dense, isodiametric, very superficial. d sternum VII bisetose, $ sternum
VI unknown, apex regularly curved.

Legs. Very elongate and slender. 4th tarsomere medially faintly excised. 5th tarsomere asetose
beneath. 1%-3' tarsomeres of d anterior tarsus biseriately squamose.

ö genitalia. Genital ring rather parallel, at apex asymmetric. Aedeagus stout, rather symmetric,
lower surface faintly concave. Apex short and acute, regularly triangular, symmetric. Internal sac
without any sclerotized plates. Left paramere rather wide, at apex evenly convex, right paramere
comparatively elongate.

? genitalia. Unknown.

Variation. Unknown.

Distribution. Westernmost Papua New Guinea. Known only from type locality.
Collecting circumstances. Unknown. Presumably collected under log in rain forest at median altitude.

Etymology. The name refers to the laevigate elytra.

Relationships. This species certainly belongs to the bisetosiceps-group, but occupies a somewhat
isolated position due to the markedly wide pronotum, extremely fine elytral striation, and almost
unarmed aedeagus.

Genus Collagonum Baehr

Baehr, 1995: 30.

Note. The genus Collagonum Baehr was erected for those species of Darlington’s genus Fortagonum that
are characterized by

1. distorted, disk-like pronotum as shown in fig. 11;

2. more or less deep sulcus medially of eye that separates the eye from the frons;

3. characteristically elongate, rod-shaped apex of the aedeagus.

The typical species (of the nominate subgenus) usually also have elongate, not spinose elytra that bear
3 discal punctures. Almost all species are fully winged (see appendix 2).

Subgenus Procollagonum, subgen. nov.

Diagnosis. Without the distorted, disk-like pronotum of genus Collagonum, but with sulcus medially
of eye and with rounded humeri. Elytra short, without discal punctures. Eyes not abruptly prominent.
1%-3'd tarsomeres of d anterior tarsus with few sparse squamae on inner side. Aedeagus with short,
somewhat rod-like apex, fairly similar to that of typical Collagonum, but perceptibly shorter.

Type species. Collagonum thoracicum, spec. nov., by monotypy.

Remarks. The single species C. thoracicum, spec. nov. is included in the genus Collagonum because of
the presence of the sulcus medially of the eye and of its male aedeagus which is fairly similar to those
of Collagonum s. str. in that it bears a somewhat disjoined, rod-like apex which, however, is shorter and
less disjoined. In other characters the species reminds either species of Fortagonum Darlington (not
distorted, disk-like pronotum), or those of the subgenus Paracollagonum (short, ovalish elytra, absence
of discal punctures). Presumably, in certain aspects this is the most basic species of the whole Colla-
gonum-complex.

Collagonum thoracicum, spec. nov.
Figs 4, 10

Types. Holotype: d, Papua NG, Morobe-Pr. Saureri 10 km s. Garaina 1600-1800 m 24.-25.3.1998, A. Riedel (ZSM-
CBM). — Paratypes: 284, Papua NG, Morobe-Pr. Saureri 10 km s. Garaina 1550-1700 m 27.3.1998, A. Riedel
(CBM).

Diagnosis. Distinguished by absence of wings, absence of all fixed setae on head, pronotum, and elytra,
wide, not distorted pronotum with markedly narrowed base, wide and short, unarmed elytra, and
short, rod-like apex of aedeagus.

Description

Measurements. Length: 8.7-8.9 mm; width: 3.7-3.9 mm. Ratios. Width/length of pronotum: 1.46-
1.50; width base/apex of pronotum: 1.50-1.54; width pronotum/ head: 1.90-1.91; width elytra/prono-
tum: 1.36-1.39; length/ width of ae 19-1.21°

Wing-and-seta formula: - w; --; --; - - —

Colour. Black, including four basal antennomeres, femora, and tibiae. Elytra with faint violaceous
lustre. Labrum and mandibles piceous, palpi, antenna, and tarsi reddish. Lower surface black.

Head. Narrow compared with prothorax. Neck rather narrow, elongate behind eyes. Eyes rather
large, moderately protruding, orbits short, obliquely convex. Clypeal suture distinct. Labrum moder-
ately elongate, apex straight. Mandibles moderately elongate, straight. Antenna moderately elongate,
surpassing base of pronotum by c. 2% antennomeres, median antennomeres c. 2.5% as long as wide.
Both palpi elongate, basal maxillary palpomere thickened. Furrow above antennal base and encircling
the eye deep, conspicuous. Both supraocular setae absent. Frons rather evenly convex, impunctate,
laterally with a shallow groove that is crossed by some oblique wrinkles. Microreticulation extremely
fine, very superficial, isodiametric. Surface glossy.

Prothorax. Wide, lateral margin slightly deplanate, especially in posterior half, evenly curved,
slightly more narrowed to apex than to base. Widest diameter a short distance behind middle. Anterior
angles projecting, at apex widely rounded off. Apex deeply excised, excision almost straight. Lateral
margin in posterior half convex, just in front of basal angles very gently concave, basal angles angulate,
c. 100°, at apex obtuse. Base laterally straight, in middle feebly produced. Disk rather convex, lateral
parts slightly deplanate. Anterior transverse depression barely visible, median line fine, almost attain-
ing apex and base, base with a shallow transverse depression. Basal grooves deep, large, irregularly
circular. Apex and lateral margins bordered, base bordered in middle. Both marginal setae absent.
Lateral channel and basal grooves coarsely and irregularly punctate-vermiculate. Disk impunctate,
though with some fine, transverse wirnkles. Microreticulation near apex and base highly superficial,
barely visible, about isodiametric, in middle absent. Surface on disk highly glossy.

Elytra. Short and wide, dorsal surface highly convex, lateral borders almost straight in anterior %,
towards apex evenly rounded. Widest diameter about in middle. Preapical sinuosity rather shallow.
Humeri wide, rounded. Apex separately rounded, without denticle at sutural angle. Striae fairly deep,

Fig. 4. Collagonum (Procollagonum) thoracicum, spec. nov. d genitalia. Scale: 0.5 mm.

impunctate, intervals slightly convex. Discal setae absent. 17-18 marginal setae and 2 apical setae
present, the latter situated near end of 7‘ stria. Intervals impunctate. Microreticulation absent. Surface
markedly iridescent. Posterior wings very short.

Lower surface. Prosternum very short, not surpassing procoxae, rounded off, posteriorly de-
pressed, ventrally bordered. Proepisternum almost impunctate, with dense microreticulation.
Mesepisternum rather densely, though somewhat superficially punctate. Metepisternum short, c. 1.2 x
as long as wide at anterior border. Epipleura anteriorly moderately wide, posteriorly very narrow,
moderately rugose. Abdomen impunctate, though laterally with some fine wrinkles. Microreticulation
distinct, isodiametric. d sternum VII bisetose, ? sternum VII unknown, apex evenly rounded.

Legs. Moderately thin and elongate. 5th tarsomere asetose beneath. 4th tarsomere medially slight-
ly excised. 1-3’ tarsomeres of d anterior tarsus sparsely, asymmetrically squamose.

d genitalia. Genital ring very elongate, with elongate, slightly asymmetric, spoon-shaped apex.
Aedeagus slightly curved, apical part extended to a moderately elongate, strongly sclerotized rod that
is slightly downcurved and slighty widened towards apex. Internal sac without sclerotized plates or
teeth. Both parameres rather elongate, at apex evenly rounded.

? genitalia. Unknown.

Variation. Very little variation noted.

Distribution. Eastern central Papua New Guinea. Known only from type locality.
Collecting circumstances. Unknown. Presumably collected under log in rain forest at median altitude.

Etymology. The name refers to the unusual shape of the pronotum.

Genus Paracollagonum, subgen. nov.

Diagnosis. With external characters of genus Collagonum, e.g. wide, distorted pronotum, deep sulcus
medially of eye, rounded humeri. But aedeagus without the elongate, rod-like apex of typical Colla-
gonum, elytra without discal punctures, eyes abruptly prominent, and pronotum with very wide,
conspicuously upturned margins.

Type species. Fortagonum distortum Darlington, 1971, by present designation.

Remarks. In view of its short, impunctate elytra, abruptly prominent eyes, very wide, conspicuously
upturned margins of pronotum Collagonum limum (Darlington) probably belongs to this subgenus.
However, any final decision is premature, until the male genitalia of this species are recorded which
so far is known only from the slightly damaged female holotype.

Collagonum (Paracollagonum) distortum (Darlington)

Darlington 1971, p. 321, fig. 76 (Fortagonum); Mateu 1977, p. 21, fig. 1 (Fortagonum); Baehr 1992, p. 75 (Fortagonum);
Baehr 1995, p. 31 (Collagonum).

This peculiar species is externally similar to the other species of the genus Collagonum Baehr that was
erected for those species of the genus Fortagonum Darlington that possess a deep sulcus medially of the
eyes, a characteristically wide, distorted pronotum, and a conspicuous, elongate, rod-like apex of
aedeagus. Although in some other character states C. distortum is also different from all other species
of Collagonum except for C. limum Darlington (see above), namely in the distorted head, abruptly
prominent eyes, very wide and upturned lateral margins of pronotum, short ovalish, impunctate
elytra, the main difference is in shape of the aedeagus that does not show the markedly elongate apex
typical for all species of the genus Collagonum that have been examined for this character. As figured
in Mateu (1977), the aedeagus has a fairly short apex, although in those specimens examined by me the
apex is longer and more disjoined than in Mateu’s figure. In spite of this important difference,
C. distortum is yet included in the genus Collagonum, though ranked in a new subgenus Paracollagonum,
because it takes a basic though at the same time highly isolated taxonomic position within the genus
Collagonum.

New records: 1d, Lufa, Mt. Michael, New Guinea, 18.10.72, R. Hornabrook (MNZ); 18, Okapa, Eastern High-
lands, New Guinea, 3.1.1974, R. Hornabrook (MNZ); 18, Daulo Pass, Asarao-Chimbu Divide, New Guinea,
13.3.72, R. Hornabrook (CBM).

Subgenus Collagonum s. str.

Baehr, 1995: 30.
Type species. Fortagonum laticolle Baehr, 1992, by original designation.

Note. The subgenus Collagonum s. str. conforms with the diagnosis as given in the original description
of the genus (Baehr 1995).

Collagonum violaceum Baehr

Baehr 1995, p. 33, figs 18, 21, 22.

New records: 1?, Marawaka, Eastern Highlands, New Guinea, R. Hornabrook (MNZ); 17, Orie, Okapa, Eastern
Highlands, New Guinea, Jan. 1968, R. Hornabrook (MNZ).

Note. This species if known from a rather restricted area in the Eastern Highlands of Papua New
Guinea.

Collagonum longipenne, spec. nov.
Eies’5, 11

Types. Holotype: 3, Bultem 5000 ft. Star Mts. 23/10/75 Papua New Guinea R. W. Hornabrook (MNZ).

Diagnosis. Distinguished from its nearest relative C. riedeli Baehr by longer and narrower elytra and
wider prothorax with less acute and less produced anterior angles.

Fig. 5. Collagonum (s. str.) longipenne, spec. nov. d genitalia. Scale: 0.5 mm.

Description

Measurements. Length: 12.4 mm; width: 4.5 mm. Ratios. Width/length of pronotum: 1.61; width
base/apex of pronotum: 1.39; width pronotum/head: 1.89; width elytra/ pronotum: 1.72; length/ width
of elytra: 1.10.

Wing-and-seta formula: + w;--;- 4; +++.

Colour. The slightly immature holotype has black head and pronotum, and wide, reddish-piceous
margins of pronotum and reddish-piceous elytra. Under the naked eye the elytra bear slight purplish
lustre which vanishes when seen under the microscope. Labrum, mouth parts, antenna, and tarsi
reddish,. 1*-3”! antennomeres, femora, and tibae slightly infuscate. Lower surface piceous.

Head. Narrow compared with prothorax. Neck rather narrow, elongate behind eyes. Eyes rather
large, though but moderately protruding, orbits almost absent. Clypeal suture distinct. Labrum mo-
derately elongate, apex straight. Mandibles moderately elongate, straight. Antenna moderately elon-
gate, surpassing base of pronotum by slightly > two antennomeres, median antennomeres <3 x aslong
as wide. Both palpi elongate, basal maxillary palpomere thickened. Furrow above antennal base and
encircling the eye deep, conspicuous. Both supraocular setae absent. Frons rather evenly convex,
impunctate, laterally with a shallow groove that is crossed by some oblique wrinkles. Microreticulation
extremely fine, very superficial, isodiametric. Surface glossy.

Prothorax. Very wide, laterally very broadly deplanate, evenly curved, slightly more evenly
narrowed to apex than to base. Widest diameter about in middle. Anterior angles remarkably project-
ing, at apex widely rounded off. Apex deeply excised, excision almost straight. Lateral margin convex
to basal angles which bear a very small denticle. Base laterally straight, in middle feebly produced. Disk
fairly convex, lateral parts broadly deplanate, slightly upturned. In anterior third with a shallow,
slightly v-shaped depression, median line distinct, attaining neither apex nor base, base with a shallow
transverse depression. Basal grooves deep, large, about circular. Apex bordered, lateral margins not
bordered, base bordered in middle. Anterior marginal seta absent, posterior marginal seta broken,
though puncture visible, situated right on posterior angle. Lateral channel and basal grooves coarsely
and irregularly punctate-vermiculate, though punctures rather superficial. Disk impunctate, almost
smooth. Microreticulation near apex and base about isodiametric, in middle extremely superficial,
barely visible, consisting of extremely fine transverse lines. Surface on disk highly glossy.

Elytra. Elongate, comparatively narrow, parallel, dorsal surface convex, lateral borders almost
straight in anterior %, towards apex evenly rounded. Widest diameter about in middle. Preapical
sinuosity rather shallow. Humeri wide, rounded. Apex with a short, rounded projection opposite 3rd
interval. Sutural angle with a very small denticle. Striae deep, impunctate, intervals convex. Discal

Figs 6-11. Habitus. 6. Fortagonum insulare, spec. nov. 7. F. substriatum, spec. nov. 8. F. hornabrookianum, spec.
nov. 9. F. Iaevissimum, spec. nov. 10. Collagonum (Procollagonum) thoracicum, spec. nov. 11. Collagonum (s. str.)
longipenne, spec. nov. Lengths: 10.4 mm; 11.0 mm; 11.1 mm; 11.1 mm; 8.7 mm; 12.4 mm.

setae short, inconspicuous, anterior seta near 3'd stria, median and posterior setae near 2" stria. 22-23
marginal setae and 3 apical setae present, two of the latter situated on apical border, one near 7“ stria.
Intervals impunctate. Microreticulation very superficial, barely visible, consisting of extremely fine,
dense, transverse lines. Surface markedly iridescent. Posterior wings fully developed.

Lower surface. Prosternum very short, not surpassing procoxae, rounded off, posteriorly de-
pressed, ventrally bordered. Proepisternum almost impunctate, with dense microreticulation.
Mesepisternum rather densely, though somewhat superficially punctate. Metepisternum elongate,
c.2.5x as long as wide at anterior border. Epipleura anteriorly moderately wide, posteriorly very
narrow, moderately rugose. Abdomen impunctate, though laterally with some fine wrinkles. Microre-
ticulation distinct, isodiametric. d sternum VII bisetose, ? sternum VII unknown, apex evenly rounded.

Legs. Rather thin and elongate. 5th tarsomere asetose beneath. 4th tarsomere medially slightly
excised. 1-3" tarsomeres of d anterior tarsus biseriately squamose.

d genitalia. Genital ring moderately narrow, fairly symmetric, apex narrow, rather short. Aedea-
gus slightly curved, apical part extended to an elongate, strongly sclerotized rod that is downcurved
with a distinct angle. Apex with acute, faintly downcurved tip. Internal sac without sclerotized plates
or teeth. Both parameres rather elongate.

? genitalia. Unknown.

Variation. Unknown.

Distribution. Westernmost Papua New Guinea. Known only from type locality.
Collecting circumstances. Unknown. Presumably collected under log in rain forest at median altitude.
Etymology. The name refers to the remarkedly elongate elytra.

Relationships. This species is presumably most closely related to F. riedeli Baehr, though is distin-
guished by longer and narrower elytra.

Identification
For better identification of the new species the most recent keys to the genera Fortagonum and

Collagonum (Baehr 1995, 1998) are completely updated.

Updated key to the species of the genus Fortagonum Darlington (sensu Baehr 1995)

1. AV\imesspresemten ee EEE D
= Winsslabsent m. eeenesneestee encore ae een eneneese ter nungencanenennne cn ee IS:
2... Both pairsiof supraoeular!setae absentk...........122222c2eseeesnneeee nn aeanen nenne ©
— At least posteniorisupraoeular setaspresenten. nase eenenenane ne 4.

3. Elytra bisetose, elongate, >1.6x as long as wide, <1.33x as wide as pronotum, striae slightly
crenulate, intervals depressed; aedeagus with two dentate sclerites in internal sac. Vogelkop,
extremelwesternIHan Jayal....:.......ce ee see een scerensoharsoscnene tere depressum Baehr

— Elytra unisetose, shorter, c. 1.33% as long as wide, c. 1.5x as wide as pronotum, striae not crenulate,
intervals slightly convex; aedeagus with a single dentate sclerite in internal sac. Western part of

central Isian Jayar.....2. 2: ee ee sinak Baehr
4. ‚Both supraerbital. setae' present. e.....uccesnseseeeeueceereeeeeersantenenan in ecasne esse ee ER >
=s >Anterior supraorbital setarabsent .....2..........00n00eeernencnnreneneeenrane ernennen nenne 6.
5. Pronotum basally much wider than apically. Eastern Irian Jaya .............n.. bisetosiceps Baehr

- Pronotum basally slightly wider than apically (Fig. 6). Japen Island, weastern Irian Jaya ..............
ET ee insulare, spec. nov.

6. Elytra unisetose (only median seta present); prothorax little wider than long. Central eastern Irian

Jayaseeetenneeeenaeseennensenenteeeneneeneenennensensctarseceeneenersensenanaahnerssnnenensensaetnennenetrecheraenetencnetse: denticulatum Baehr

- Elytra bisetose (median and posterior setae present); prothorax considerably wider than long.

DistribuHomaditterem een nretane nenne nahen ne nerrersrerresheasndes L

7. Striae extremely fine, elytra almost laevigate, with greenish lustre. Westernmost Papua New

CUÄTENN, ooasonneneor gen SEHeREeRERRO RT re Eon dRHcHoRRLDnLCRRLRaUR laevissimum, spec. Nov.

- Striae distinct, elytra not laevigate, with bluish or violaceous lustre .......eseenenensenseene 8.

8. Elytra wider and shorter, with short sutural spines (see figs 47, 48 in Darlington 1971) ............. 9

- Elytra narrow and elongate, with elongate sutural spines (Figs 6-8) .....nnneeeesnensnenenn 10.
9. Pronotum wider, sides more straight, anterior angles more protruding. Extreme western Irian

AA nn enenneesssuasacesenecteneneenensontpechnnecutpensentnsensnsshansenersenseseneeossrarnenee subconicolle (Darlington)

- Pronotum narrower, sides more convex, anterior angles less protruding. Central Papua New

STERN ER gern bigemum (Darlington)

10. Sutural spines less widely separated (Fig. 8); elytra with distinct violaceous lustre. Eastern central

BapuauNewa@umean en eeeneueseersneeuenseetsereeneneetarersnnerenensanstetcaneensesennsrneness hornabrookianum, spec. nov.

- Sutural spines widely separated (Figs 6, 7); elytra with bluish or blackish lustre .........u.- DE

11. Striae barely punctulate; elytra with faint bluish lustre; antennae piceous, only median and apical

antennomeres reddish; aedeagus (Fig. 1). Eastern central Papua New Guinea ......nueenenen:

DE er ERROR substriatum, spec. NOV.

- Striae distinctly punctulate; elytra without bluish lustre; antennae largely light reddish; aedeagus

unknown. Western central Irian Jaya'...........eseescscosssensnsoenseeoenescnesenenenenenenensnsnanannnanenecscereresssnnenssensaenene 12%

12. Colour deep black, legs largely blackish, basal antennomeres in part piceous; base of pronotum

narrower, ratio base/apex c. 1.6. Panai Province, mainland of Irian Jaya .............. spinipenne Baehr

- Colour reddish, tibiae and tarsi reddish, antennae including basal antennomeres light reddish; base

of pronotum wider, ratio base/apex c. 1.7. Japen Island ...........eeee insulare, spec. nov.

13. Both’supraocular setaerabsent .............ucueusansansensunensenssnecnsnnonsenesnstneneenstnsnnsnnensensenznrnsenatunnnenssnsenernerseneen 14.

— Posterior supraocular seta present .....useeessesenneenenenenennnnnenenenenensnnnnsnennsesnsssanssenenesenseneenentnesenetenen 16.

14. Elytral striae superficial, intervals depressed, surface slightly iridescent. Western central Irian Jaya

PIERRE ER IRL Een een soeoeslecscessessenesocsnersnanensnaneseresanteskrsnahtererueeen üonenenennerstäneresgsnssrnentene laevigatum Baehr

- Elytral striae deeply impressed, intervals markedly convex, surface not iridescent „nen 15.

15. Anterior angle of pronotum slightly produced laterally, apex obtuse; elytra longer, ratio 1/w >1.32.

@entrallriant]ayal re eeneseseteesteenseussnenese ern an esnerenraserocchnsscnenenensnsnrnananahenefensnererer bufo Darlington

- Anterior angle of pronotum straight, apex acute; elytra shorter, ratio 1/w <1.28. Western central

|Imları JEIyer bs ee RELEASE globulipenne Baehr

16. Elytra usually trisetose, rarely unilaterally unisetose or bisetose; mandibles never straight and very

elongate. Central Papua New Guinea .....uneesnnenenenensenenensnsenenenssnsenenssenenessenenensnsenenennsetnnn 17

- Elytra asetose, or unisetose, or bisetose; either mandibles straight and very elongate, or more or less

fusiform species. Central and eastern Irian Jaya ........nesseneseneneensnnenenensnsenensnsnssnensnsnsssenensnsnenennenn 20.

17. Posterior pronotal seta present .......ueeesenssneneneneesenensenenenensenenenensnsenensnsnsnnssoseesnsnssseensnsnnenensesenenne 18.

— Posterior pronotal seta absent ..........eeeeseessssesesenesenenenessnensnensnsnsnsosnsnsnnnnnnenenensnensnsnensnsnsnsnsnsnsssssssssenssssenenen 19),

18. Margin of pronotum wide; wide, fusiform species. Bulldog Range ............. oodinum Darlington

- Margin of pronotum narrow; rather narrow, barely fusiform species. Mt. Albert Edward .........

Be LS Cnneoskonenuenentessenssentosnnessaecssungsserhangnsanesuner se rehnnueshnesknsenunenunee antecessor Darlington

23.

. Pronotum wider, but less conical; elytra weakly iridescent. Mt. Wilhelm ...... fortellum Darlington
Pronotum narrower, but rather conical; elyira markedly iridescent. Okapa .......... okapa Darlington
MBosteriorpronotal setanpresent,\elytrasumisetoseloribisetoser. rn. EEE 2
Bosterior)pronotal’setalabsent;Telytratasetoser................2222 020 necneee 23:
. Pronotum laterally regularly convex, base as wide as apex, basal angles rounded off, apex very

protruding; elytra bisetose, anterior seta absent. Eastern central Irian Jaya ............ acuticolle Baehr

Pronotum laterally feebly convex, base much wider than apex, basal angles rectangular and obtuse,
apex less protruding; elytra unisetose, only median seta present. Eastern Irian Jaya .............. 223

. Apex of elytra not spinose, though sutural angle faintly denticulate, elytra slightly wider; prono-

tum barely narrowed towards base (Fig. 9). Area east of mountain range to the west of valley of
BOrmealiveRgat seen esse ersaknnn res kan eenlsr eataneeereeheash ehe ur hbereen een unipunctatum Baehr

Apex of elytra elongately spinose opposite 3rd interval, sutural angle not denticulate, elytra slightly
narrower; pronotum distinctly narrowed towards base (Fig. 10). Area west of mountain range to
thexwestsofuyalleysofiborme; River ne nn. eeenneenstane nennen spinosum Baehr

Mandibles not unusually elongate; apex of elytra distinctly spinose opposite 3rd interval; short and
wide, markedlyzfusitorm;speciesa@entral-Irians]ayar2.... nen. curtum Baehr

Mandibles straight and markedly elongate; apex of elytra not spinose; either rather elongate, not
markedly fusiform species, or short and wide species with almost parallel lateral borders of

PEONOBU ee ee REN 24.
. Basal margin of elytra not interrupted at 3rd interval; prothorax <1.8x as wide as head ......... 25,
Basal margin of elytra interrupted at 3rd interval; prothorax >2x as wide as head ................... 26.

. Rather wide, almost parallel species; pronotum >1.25x as wide as long. Central Irian Jaya .........

ee REEL EN En Re EEE ER ne forceps Darlington

Narrow, fusiform species with evenly rounded lateral margins of pronotum; pronotum c. 1.1x as
widerasslong ACentrahlrian Jaya... een ee formiceps Darlington

. Pronotum wider at base, ratio width of base/ width of apex c. 1.8, sides more curved; elytra rather

elonsate. GentralNlniansjaya. een neneneenonenananananunanenenene se seennnsse ne ee cychriceps Darlington

Pronotum narrower at base, ratio width of base/width of apex c. 1.65, sides more parallel; elytra
tathersshort. GentralreasternilnianJaya 2.0... N latum Baehr

Updated key to the species of the genus Collagonum Baehr

Prothorax wide, though not distorted (Fig. 10). Eastern central Papua New Guinea (Procollagonum)
Rn one thoracicum, spec. noV.

Prothorax remarkably. disterted (Eis I)... nenne een en ee ee ee 2
Wings; absent.....enn eu sseessiessensiasnensnssssssneanesnne nennen dose een ee RS 8:
Wingsipresent (Gollaganum!s. Str)... nee en 6.
Eyes laterally abruptly produced; elytra asetose, or (rarely) unilaterally unisetose (Paracollagonum)
EEE ee RE REEL OO TEEHEECLECT0005E0.0000008010600006600066000 4.
Eyes laterally not as abruptly produced; elytra trisetose (Collagonum s. str.) «une 5%

Both supraocular setae present; posterior pronotal seta present; frons conspicuously swollen.
Eentral’Papua New’Guinea......u..2re erregen een enersrmefeerenene distortum (Darlington)

Anterior supraocular seta absent; posterior pronotal seta absent; frons not swollen. Central Papua

New Guinea... amsseuseiteenenresisnsnntenrnsenkenensgsnserarnienhensgessnnrratnnen seht nnnennae ee limum (Darlington)

5. Both supraocular setae present; prothorax narrower, <1.5% as wide as long. Central eastern Irian

VERYEL escoosteneeoegeeesgne Ba I EN RERENE convexum Baehr
- Anterior supraocular seta absent; prothorax wider, c. 1.7x as wide as long. Central Papua New
ENTER: oe ER IEREEREO hornabrooki (Darlington)
DEBothspairsiot. supraoeular!setaerabsent............2...... 020 rRerenesnserertneresnene he sengrereescepereraegenee regte eheene 7
Zw \aleastspesterior:supraoeularisetalpresent Baer ee ee ebenen eeerenenkereheten 10.

7. Eyes laterally abruptly produced; pronotum at apex much narrower than at base. Central Irian Jaya
DOOR NEE RER EU HEHE NEED OHREN ophthalmicum (Baehr)

-— Eyes laterally not as abruptly produced; pronotum at apex only slightly narrower than at base.

NEN ent. nn. uucheslssnsnsuskehtehenucnnngsntisknehnisehgnegiensntägndantsesenssnnntisnsnsntnrtrsnrtäehtinnknenensnserfrsnsehnenssrkchsdee> 8.
SeBothipronotal'setae absent. Eastern Irlan Jayar....unneseeseeeeneeeeraeeenece ee robustum Baehr
- Poser PRIOR EEE I

9. Elytra longer and narrower, ratio length/ width 1.73; prothorax slightly wider, anterior angles less
acute and less protruding (Fig. 11). Westernmost Papua New Guinea ......... longipenne, spec. nov.

- Elytra shorter and wider, ratio length / width 1.60-1.63; prothorax slightly narrower, anterior angles
aeutegand more,protruding” Eastern Irian Jayarzaı..unnenseeeesesearaenenensnsesnenesesteresonerse sense riedeli Baehr

10. Wider species; pronotum wider, laterally more rounded, with shorter, more convex anterior angles.
GentralhapuarNewGuinean.......ete in eennersenekongtnerssrernensuehsereneheshenen re violaceum Baehr

- Narrower species; pronotum narrower, laterally less rounded, with longer, more acute anterior
amelesg@entralkand eastern Irian Jayarı een esse seneneeernensenearaenete ee laticolle (Baehr) 11.

11. Eyes smaller, laterally more abruptly protruding, almost devoid of distinct orbits. Area west of
mountain range to the west of valley of Borme River ...........neeee: laticolle laticolle (Baehr)

-— Eyes larger, laterally less abruptly protruding, with distinct, oblique orbits. Area east of mountain
rangestoßtheswest of valley; of Borme River u.a nennen laticolle macrops Baehr

Remarks

Even with the six species described as new in this present paper the number of species of the genera
Fortagonum and Collagonum actually existing in New Guinea probably is not even approximately
known. Apparently almost all mountain ranges have their own species, or even more than one, and
very unusual species appear as the central ranges of New Guinea are more extensively sampled. Thus,
it becomes more and more difficult to arrange the many differently shaped and structured species, if
we are not willing to erect many new genera to master the high structural diversity.

In trying a proper arangement, the wing-and-setae-formulas that were firstly used by Darlington
(1952) have proved increasingly useful. Hence, an attempt has been made in appendix 2 to arrange all
species according to their wing-and-setae-formula. In addition to the development of wings and to
chetotaxy, for the arrangement some additional character states were included, e.g. presence / absence
of elytral spines, shape of mandibles, structure of aedeagus, shape of elytra and of pronotum.

Evidently these formulas are quite helpful in dividing the genera into (hopefully) natural groups
that in the genus Fortagonum provisionally are called “species-groups”, in the genus Collagonum are
called subgenera. Apparently, in the latter genus the structural differences between the species of the
subgenera Procollagonum and Paracollagonum are much greater than they are between the species-
groups of the genus Fortagonum. But even in Fortagonum, the species-groups are believed to form
monophyletic units, some of which could be raised to subgenera or even to genera when more
evidence, or perhaps, additional species are at hand, that might clear up better the relationships within
the genus.

References

Baehr, M. 1992. On some agonine beetles of the genus Fortagonum Darlington from New Guinea (Coleoptera,
Carabidae, Agoninae). - Mitt. Münch. Ent. Ges. 82: 73-81

-- 1995. New species and new records of the genera Fortagonum Darlington and Collagonum, gen. nov. from
New Guinea (Insecta, Coleoptera, Carabidae, Agoninae). — Spixiana 18: 15-43

-- 1998. New species of the genus Fortagonum Darlington from western New Guinea (Insecta, Coleoptera,
Carabidae, Agoninae). - Spixiana 21: 11-20

Darlington, P. J. Jr. 1952. The carabid beetles of New Guinea. Part 2. The Agonini. — Bull. Mus. comp. Zool. 107:
89-252

-- 1968. The Carabid Beetles of New Guinea. Part III. Harpalinae (Continued): Perigonini to Pseudomorphini.
- Bull. Mus. comp. Zool. 137: 1-253

-- 1971. The carabid beetles of New Guinea. Part IV. General considerations; analysis and history of fauna;
taxonomic supplement. — Bull. Mus. comp. Zool. 142: 129-337

Mateu, J. 1977. A propos du Fortagonum distortum Darlington de la Nouvelle Guinee. (Carabidae Agonini). —
Nouv. Rev. Ent. 7: 21-22

Appendix 1
Alphabetical checklist of the species of the genus Fortagonum Darlington
acuticolle Baehr, 1995

antecessor Darlington, 1971
bigemum (Darlington, 1971)

. Irian Jaya
. Papua New Guinea
. Papua New Guinea

(@ Te) Ko) (a al (ei a)

bisetosiceps Baehr, 1995 . Irian Jaya

bufo Darlington, 1952 . Irian Jaya

curtum Baehr, 1992 . Irian Jaya

cychriceps Darlington, 1952 . lrian Jaya
denticulatum Baehr, 1995 e. Irian Jaya
depressum Baehr, 1995 w. Irian Jaya

forceps Darlington, 1952 e. Irian Jaya

formiceps Darlington, 1971 e. Irian Jaya

fortellum Darlington, 1951 c. Papua New Guinea
globulipenne Baehr, 1998 w. Irian Jaya
hornabrookianum, spec. nov. c. Papua New Guinea
insulare, spec. NOV. Japen I., w. Irian Jaya
laevigatum Baehr, 1998 w. Irian Jaya
laevissimum, spec. noV. w. Papua New Guinea
latum Baehr, 1995 e. Irian Jaya

okapa Darlington, 1971 c. Papua New Guinea
oodinum Darlington, 1971 c. Papua New Guinea
sinak Baehr, 1998 w. Irian Jaya
spinipenne Baehr, 1998 w. Irian Jaya
spinosum Baehr, 1995 e. Irian Jaya
subconicolle (Darlington, 1971) w. Irian Jaya
substriatum, spec. noV. c. Papua New Guinea
unipunctatum Baehr, 1995 e. Irian Jaya

Alphabetical checklist of the species of the genus Collagonum Baehr

convexum Baehr, 1995
distortum (Darlington, 1971)
hornabrooki (Darlington, 1971)
laticolle laticolle (Baehr, 1992)
laticolle macrops Baehr, 1995

. Irian Jaya
. Papua New Guinea
. Papua New Guinea
. Irian Jaya
. Irian Jaya

(GI Te) (o) @)

limum (Darlington, 1952) c. Papua New Guinea
longipenne, spec. nov. w. Papua New Guinea
ophthalmicum (Baehr, 1992) c. Irian Jaya
riedeli Baehr, 1995 . Irian Jaya
robustum Baehr, 1995 . Irian Jaya
thoracicum, spec. Nov. . Papua New Guinea
violaceum Baehr, 1995 . Papua New Guinea

(ay Lo). go fan)

Appendix 2

Compilated wing-and-setae-fomulas of the species of the genera Fortagonum Darlington

and Collagonum Baehr

(+: present; -: absent; — +: variable, or very small)

posterior elytral suborbital pronotal elytral
wings spines setae setae setae

Genus Fortagonum Darlington
acuticolle-group
acuticolle Baehr _ = = En E Re
antecessor Darlington _ _ _ + A er
oodinum Darlington - _ u + re TE ee
okapa Darlington _ = = A = :
fortellum Darlington _ = - 2 en et:
bisetosiceps-group
bisetosiceps Baehr + + + + = = Tip
insulare, spec. nov. + + — + — Een:
spinipenne Baehr + + _ + — u
substriatum, spec. nov. + + - + = een
subconicolle (Darlington) 4 4 = + — et
hornabrookianum, spec. nov. + + — + - = + +
laevissimum, spec. nov. + + = + z > ai a.
bigemum (Darlington) + + = + = ad
denticulatum Baehr + - — + Br ea
depressum Baehr + + = a £ A er
sinak Baehr + + — 2 > lee
unipunctatum-group
unipunctatum Baehr = —+ - + + A ea
spinosum Baehr - + - + + er
bufo-group
curtum Baehr = + = + = a u Rn
formiceps Darlington - — - + = ee
latum Baehr 2 2 en AL = Ar ai 2

bufo Darlington - - =
cychriceps Darlington = = =
forceps Darlington = = =
globulipenne Baehr = = =
laevigatum Baehr - - -

Genus Collagonum Baehr

Subgenus Procollagonum

thoracicum, spec. noV.

Subgenus Paracollagonum

distortum (Darlington)
limum (Darlington)

Subgenus Collagonum s. str.

convexum Baehr

laticolle laticolle (Baehr)
laticolle macrops Baehr
violaceum Baehr
hornabrooki (Darlington)
longipenne, spec. nov.
riedeli Baehr
ophthalmicum (Baehr)
robustum Baehr

++ +++++++

SPIXIANA 73-76 München, 01. März 2001 ISSN 0341-8391

About Andricus polycera (Giraud, 1859) and related forms,
with special remarks on Andricus polycera and A. subterranea

(Insecta, Hymenoptera, Cynipidae)
David Bellido & Juli Pujade-Villar

Bellido, D. & J. Pujade-Villar (2001): About Andricus polycera (Giraud, 1859) and
related forms, with special remarks on Andricus polycera and A. subterranea (Insecta,
Hymenoptera, Cynipidae). — Spixiana 24/1: 73-76

In this paper the status of the polycera group of species is studied. Two new
synonyms are established: A. polycera (= A polycera transversa, syn. nov.) and A. sub-
terranea (= A. trinacriae, syn. nov.) and new characters and problems for the sepa-
ration of the two species are commented. A. marchali is considered as a probable
valid species with uncertain status. Reasons leading to these conclusions are com-
mented. Finally, A. polycera is definitively removed from the Iberian gall wasp fauna.

David Bellido, Juli Pujade-Villar, Universitat de Barcelona. Facultat de Biologia.

Dept. Biologia Animal (Artröpodes), Avda. Diagonal, 645. E-08028-Barcelona.
e-mail address: pujade@porthos.bio.ub.es

Introduction

Andricus polycera and A. subterranea were originally described as two closely related species of the genus
Cynips by Giraud (1859). Mayr (1870) examined Giraud’s material and considered Cynips subterranea
as a mere variety of C. polycera which was followed by posterior authors (Kieffer 1897-1901, Houard
1908, Dalla Torre & Kieffer 1910, Ionescu 1957, Ambrus 1974, Vassileva-Samnalieva 1983). According
to these authors the two forms could be separated by chromatic differences and gall morphology.

Material and methods

Type series of A. polycera and A. subterranea, from the Giraud’s collections deposited in MNHN, Paris
were studied and lectotypes established. All descriptions of these forms in the literature have been
revised and the Vilarrübia collection, deposited in the Zoological Museum of Barcelona, was studied.
SEM pictures have been made without coating and at low voltage to prevent any risk for the specimens.

Results and discussion

Adults of both forms present a high morphological similarity, although A. subterranea individuals are
normally darker, especially in their antenna and tibiae. Colour in cynipids is highly variable, so it is not
rare to find black forms within a normally brown species (Bellido, Melika & Pujade-Villar, in prep.).
Moreover, some cynipids also change their colour depending on the host where they have been reared
from (Pujade-Villar et al, in press) and even from different organs of the same host plant (Pujade-Villar
sg).

Tavares (1931) again raised A. subterranea to species rank and gave additional characters to separate
it from A. polycera: antennal segment lengths different in both species, tarsal claws of A. subterranea with
a longer basal lobe, differences in pubescence of lateral part of the mesosoma and also in median
mesoscutal impression. However, none of these characteristics are sufficient for the separation of the
two forms, and no clear differences are evident following Tavares’ characters. In the same paper
Tavares recorded this species from Spain through material collected by Vilarrubia, in Balenya (Barce-
lona, NE Spain), and this species is also recorded in Vilarubia (1930). However, these records do not
belong to A. polycera, as Nieves-Aldrey (1987) suggested. Andricus polycerus pictures in Tavares’ paper
belong to an A. kollari gall with protuberances, and two galls found at Vilarrubia’s collection in the
Zoological Museum of Barcelona are A. quercustozae galls probably deformed by a parasitoid attack in
their first developmental stages. In north-eastern Iberian Peninsula we have repeatedly observed
similar galls, normally in oaks highly attacked by A. quercustozae. Therefore this species is definitively
excluded from the Iberian gall wasp fauna list.

Galls of A. polycera and A. subterranea are also very similar, although they normally are considered
to be different. A. polycera galls are found in aerial buds of Quercus petraea, Q. robur and Q. humilis,
preferentially on younger, shrub-sized trees (Csöka 1997), while A. subterranea is found on subterrane-
an stems or rhizomae of the same oaks, normally hidden by a thin layer of litter above them. Moreover,
A. subterranea galls are shorter, softer, more irregular and without long protuberances in their apex.
Records of A. subterranea in Ionescu (1957) and Ambrus (1974) are interesting, since galls in the pictures
seem to have appendices in their superior part, like in A. polycera, while in typical galls of A. subterranea
these extensions are not present. Some differences between these two galls could be attributed to their
position. Subterranean galls find moister conditions which probably made them softer, and perhaps the
other differences, like the absence of protuberances or the smaller height, could also be explained by
developmental constraints of their subterranean habitat. On the other hand, height and length of apical
expansions of A. polycera galls are variable, and subterranean galls may only represent an extreme of
this variation.

Other species of cynipids can be found at subterranean and aerial organs, although these misplace-
ments occur more or less frequently, depending on the species. This duality in location has been
observed in Trigonaspis megaptera, Andricus sieboldi (ag. gen.) and Plagiotrochus kiefferianus (ag. gen.)
(Pujade-Villar, pers. obs.). There is also a slight difference of emergence between A. polycera and

A. subterranea: while the first appears from end of October to beginning of November, A. subterranea is
seen in middle of November (Giraud 1859). This variance can be explained by habitat differences and

has been observed in Plagiotrochus kiefferianus (Pujade-Villar, unpublished data).

From all these observations it would seem that these two species are identical, and the galls then
would represent only extremes of variation. However, studies of adults also shows some morpholog-
ical differences and so they do not support this hypothesis, as will be discussed below.

After examination of the type series of both species deposited in the Giraud’s collection from

MNHN, Paris, we have concluded that these two species are very similar morphologically, butwehave

found some differences in propodeum pubescence: In A. polycera the propodeal area (Fig. la) is only

slightly pubescent and normally limited to the superior corners while in A. subterranea the propodeum
(Fig. 1b) normally is densely pubescent and occupies the whole propodeum, reaching always the

nucha. This character is less obvious in some adults but always both forms can be separated. In the type
series of A. polycera there is also a perceptible chromatic variation, including some darker individuals
which are relatively similar to typical A. subterranea adults, but all studied A. subterranea adults are
darker, and although this chromatic aspect is not useful alone, it could be helpful in the separation of
the two forms. Further studies could demonstrate that all variability falls inside the intraspecific rank,

but considering the small differences between other species of cynipids (i.e. in the A. kollari group), we

think it better to maintain the specific status of both forms.

Three other varieties of A. polycera are known: A. polycera transversa Kieffer, A. polycera trinacriae
Stefani and A. polycera marchali Kieffer. Unfortunately, the Stefani collection was lost during 2” World
War (Horn et al. 1990) and the Kieffer collection is very dispersed and the location of many types is
unknown, or they were lost by different reasons, because Kieffer used to return the material to the

collectors. The current status of Kieffer’s taxa should be considered as doubtful, since they can |

represent valid species or forms or not. Tavares (1931) treated A. polycera transversa and A. polycera

trinacriae as a subspecies of A. polycera; he also recorded the high similarity of A. polycera trinacriae galls |

with those of A. subterranea (according to Mayr), and that there were no differences between these

Fig. 1. Propodeal area of Andricus polycera (a) and A. subterranea (b).

forms and the typical one. Therefore, A. polycera trinacriae is probably a synonym of A. subterranea and
A. polycera transversa is probably a synonymic name of A. polycera, given the high variability of galls of
these species. Finally, Tavares considered A. polycera marchali as a different species because of differ-
ences in mesoscutum sculpture, relative length of antenna and hypopygial spine, among others. In this
case, again type material is not available, because the location of Kieffer’s types is unknown, and the
Tavares collection was destroyed during a fire which affected great parts of Lisboa in the last century.
However, according to his paper, the material examined was sent by Marchal, who collected the gall
described by Kieffer, and which came from the same zone. Therefore, we consider that it actually
represents a different species, especially in view of its gall morphology, and that it is rather remotely
related to other forms of A. polycera. In this case Tavares’ characters could be used for the separation
of this species.

Conclusions

- Cynips polycera Giraud, 1859. Lectotype: agamic ?, (deposited in MNHN), here designated (exam-
ined). “Museum Paris 4- C. polycera Aust. G. Coll. Giraud” (white label); “Cynips polycera, typical
series” (white label); “Lectotype” (red label); “Andricus polycera (Giraud), Bellido & Pujade-Villar
det.-1999” (white label). Paralectotypes: 39 agamic ??, 4 galls. Same data of lectotype, emergence
dates: 25" March (1®, extracted from the gall), 20% June (322), 18" August (12 specimens), 28
August (12?P), no additional data (11??, one of them with a white label “4 Cynips polycera G.
Aust. G”); material deposited in MNHN except 5 adults in Barcelona University.

- Cynips polycera var. transversa Kieffer, 1897. Syn. nov. of A. polycera (not examined).

- Cynips subterranea Giraud, 1859. Lectotype: agamic ?, (deposited in MNHN), here designated
(examined). “5 Cynips subterranea G. aust. G” (white label”, “Museum Paris, Coll. Giraud” (white
label), “Cynips subterranea, typical series” (white label), “Lectotype” (red label), “Andricus subter-
ranea (Giraud), Bellido & Pujade-Villar det.-1999” (white label). Paralectotypes: 22 agamic ?9,
2 galls. Same data of lectotype but without the first label; emergence dates: 12" September (8 spec-
imens), 15'" September (2 specimens), 20% September (6 specimens), 4" November (6 specimens),
material deposited in MNHN except 3 adults in Barcelona University.

- Cynips trinacriae Stefani, 1906. Syn. nov. of A. subterranea (not examined).

- Cynips polycera var. marchali Kieffer, 1897. Incertae sedis. Probably a valid species, A. marchali
(Kieffer, 1897)

Key to the valid species of the A. polycera-group
1. Propodeal area normally only slightly pubescent, and hairs restricted to its superior part, never

reaching nucha (Fig. la). Colour variable, but normally brown. Galls in aerial buds of deciduous
oaks, variable in height and in length of apical protuberances. .........nee A. polycera Giraud

- Propodeal area normally strongly pubescent and hairs reaching nucha (Fig. 1b), adults normally
darker, especially in their antenna and tibiae. Subterranean galls shorter, softer, more irregular and
withoutzapiealexpansions....n.r:.senesesseresansanesarsseensntesunseersenssesarerennenaelenasereineees A. subterranea Giraud

Acknowledgements

We remain grateful to Dr. C. Villemant (Curator of Hymenoptera collection of Museum Nat. Hist. Nat., Paris)
and Mr. Oleguer Escola (Curator of Zool. Museum of Barcelona) for loan of material for this study. We also thank
Dr. G. Melika (Systematic Parasitoid Lab., Hungary) for his comments and P. Ros-Farre (Barcelona University,
Spain) for making SEM pictures.

References

Ambrus, B. 1974. Cynipida-Gubacsok-Cecidia Cynipidarum. — Magyarorszäg ällatviläga (Fauna Hungariae),
XII, 2. Akademiai Kiadö, Budapest. 120 pp.

Csoka, G. 1997. Plant galls. - Budapest: Forest Research Institute and Agroinform

Dalla Torre, C. W. & ]J. ]J. Kieffer 1910. Cynipidae. - Das Tierreich 24: 891 pp. + 35 pl. Berlin

Giraud, J. 1859. Signalements de quelques especes nouvelles de cynipides et de leurs galles. - Verh. Zool.-bot.
Ver. Wien 9: 337-374

Horn, W., I. Kahle, G. Friese R. & Gaedike 1990. Collectiones entomologicae. - Akad. Landw. DDR. Vol 1 & 2.
540 pp. Berlin

Houard, C. 1908. Les Zoocecidies des Plantes d’Europe et du Basin de la Mediterranee, Vol. I. - Mus. Hist. Nat.
Labor. Entom. Paris: 1-569

Ionescu, M. A. 1957. Cynipinae. — Fauna Republicii Populaire Romine %(2): 1-246.-Ed. Acad. Republicii Popul.
Rom. (In Romanian)

Mayr, G. 1870. Die mittleleuropäischen Eichengallen in Wort und Bild. - Jahresber. Rossauer Communal-
Oberrealschule Wien 9: 1-34

Nieves-Aldrey, J. L. 1987. Estado actual de conocimiento de la subfamilia Cynipinae (Hym., Parasitica, Cynip-
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Pujade-Villar, J. 1991. Contribuciö al coneixement dels cinipids cecidögens dels arbres i arbusts de Catalunya,
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Barcelona. 1128 pags. (Unpublished)

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SPIXIANA 77-84 München, 01. März 2001 ISSN 0341-8391

A peculiar evolutive lineage of the uncus
in the genus Catada Walker, [1859] 1858

(Lepidoptera, Noctuidae, Hypeninae)
Martin Lödl

Lödl, M. (2001): A peculiar evolutive lineage of the uncus in the genus Catada
Walker, [1859] 1858 (Lepidoptera, Noctuidae, Hypeninae). — Spixiana 24/1: 77-84

A distinct and aberrant lineage of unci within the genus Catada Walker, [1859]
1858 is discussed as one of the rare examples of a continous documentation of the
evolutive development from plesiomorphic to apomorphic features. A brief de-
scription of the genitalic features of the genus is given. The phylogeny of the major
part of the species set of the genus is presented.

Dr. Martin Lödl, Naturhistorisches Museum Wien, 2. Zoologische Abteilung,
Burgring 7, A-1014 Wien, Austria. e-mail: martin.loedl@nhm-wien.ac.at.

Introduction

The genus Catada was described by Walker in [1859] 1858 based on the species Catada glomeralis Walker,
[1859] 1858 in the family “Herminidae”. This species was described in the same publication on a
previous page as Bocana vagalis Walker, [1859] 1858. Both taxa are coming from Sri Lanka.

The exact systematic position of Catada remains unclear, although it is traditionally placed in the
Hypeninae. A redescription and illustration of the type species of the genus Catada and a commented
list of species based on Poole (1989) were given in Lödl (1999b). Derived African species have been
reported in Lödl (2000). The relationship to the genus Nolasena Walker, [1858] 1857 is also under
discussion (Lödl 1999a).

The male genitalia of Catada can be characterised with the following punctation (Fig. 14):

— The uncus region is complex, consisting of an aberrant uncus and a claw like scaphium of the anal
cone.

— The tegumen is slim and slender, forming a swelling from the ventral side.

-— The anellus is a skinny glove covered with spines.

— The valves are skinny and divided into two big lobes at the distal end or simple and wear a bristle
covered lobe in the middle, protruding in the lumen of the genital corpus.

- The sacculus is well developed and forms a giant, flabby eversible tube (coremata!), which is
densely covered with very long hair-like scales. There are three centres of density of hairs: one
ventrally at the base, covered with the longest hairs, one on the dorsal margin in the distal third
and one at the distal tip.

- The vinculum is insignificant, rounded and is not exceeding the length of the remaining genitalia
corpus proximally.

- The aedeagus is a slender, more or less straight tube with a blunt distal end.

Material and Methods

This study was carried out with conventionally dried specimens from the collections of The Natural History
Museum (BMNH) in London, the Naturhistorisches Museum Wien (NHMW) and the Museum national
d’Histoire Naturelle (MNHN) in Paris. Genitalia preparations have been made in the traditional way (mazera-
tion by use of KOH, preparation and isolation of the genitalia tract). The genitalia have been stained with
Chlorazol Black.

The SEM investigations were carried out by using conventionally prepared specimens (alcohol and as a final
stage 99,9 % cooled aceton), coated with gold, on a Jeol 6000/400.

List of species investigated

Base of the investigation is a set of Catada-species (type locality in square brackets; deposit of types in normal
brackets):
Catada antevorta (Viette, 1958) [Madagascar] (MNHN)
Catada canaliferalis (Moore, 1877) [Andaman Islands] (BMNH)
Catada charalis Swinhoe, 1900 [Australia, Queensland] (BMNH)
Catada ndalla Bethune-Baker, 1911 [Angola] (BMNH)
Catada obscura Joannis, 1906 [Mauritius] (BMNH)
Catada phaeopasta Hampson, 1909 [Uganda, Ruwenzori] (BMNH)
Catada renalis (Moore, 1882) [India, Khasia Hills] (BMNH)
Catada transversalis (Moore, 1877) [Andaman Islands] (BMNH)
Catada vagalis (Walker, [1859] 1858) [Sri Lanka] (BMNH)
The type specimens of the species have been examined.

The SEM investigation is based on:
Hypena varialis Walker, [1866] 1865 [Sierra Leone] (NHMW) (SEM study)

Abbreviations
BMNH The Natural History Museum, London

MNHN Museum National d’Histoire Naturelle, Paris
NHMW Naturhistorisches Museum, Wien

sc scaphium

scl length of scaphium
ta tuba analis

te tegumen

u uncus

Results

Normally the decision if the state of a feature of the copulatory system is plesiomorphic or apomorphic
is a very delicate one. A typical hook-shaped uncus as it is found in the genus Hypena Schrank, 1802
is illustrated in fig. 1. The transformation of the uncus from hook shaped to aberrantly helmet shaped
with a lace of sclerotized teeth within the genus Catada is documented in this paper. The hook shaped
state of the uncus in the African species Catada phaeopasta Hampson, 1909 is clearly the plesiomorphic
condition (Figs 2-5).

The transformation of this character follows three steps:

1. A big uncus with two rounded knees, hook shaped with several sclerotized teeth-like setae on the
ventrolateral middle of the uncus. The uncus tip also with some sclerotized teeth. The dorsal
margin is poor in hair-like scales (Figs 2-5).

2. A more or less rounded uncus with a flattened distal part. Two areas of teeth-like setae are found.
One is formed like a tonsure at the distolateral part of the uncus, one is formed as a row of setae
situated ventrolaterally. The hair-like scales of the dorsal margin are confined to the distal part of
the uncus (Figs 6-7, 15).

3. Completely aberrant uncus, like a helmet with a slim and rounded distal part and a blown up basal
part. Both setae areas are linked together to a lace of sclerotized teeth forming a trimming along
the ventrolateral margin of the uncus. The dorsal margin is rich in hair-like scales (Figs 8-13).

There is a clear lineage from the plesiomorphic state “hook-shaped” uncus to an extraordinarily
aberrant “helmet-shaped” uncus. The feature “lace of teeth-like setae” is starting with a few small teeth
in the most primitive form: C. phaeopasta, C. transversalis and C. antevorta (Viette, 1958) represent a clear
intermediate with three separated areas of teeth-like setae on the uncus. One field at the distal area and
two fields at both ventrolateral margins. The mostly evolved species show a completed lace of strong
teeth-like setae. This lace strictly follows the ventrolateral margin of the uncus and forms a distinct
bend at the distal end of the helmet. A phylogeny of the involved Catada-species - mainly based on
characters of the male copulatory system - is given in fig. 16.

Around the uncus occur different accessory or associated structures. These structures are difficult
to define and quite different from species to species. The genus Catada exhibits a well developed
scaphium. According to Tuxen (1956) a scaphium is a sclerotization of the dorsal part of the anal tube.
“Anal tube” is used here as the commonplace term. According to Kristensen (i.l.) the terminal postgen-
ital region which bears the anus as the apical element of the alimentary canal should be referred to as
the “anal cone”. This differentiation may be relevant to point out that the sclerotizations affect the
postgenital wall surrounding the terminal digestive tract and not the digestive tube itself. These
sclerotizations seem to have either a function in protecting the skinny sector of anal cone and anal tube
(s.str.) from fine pointed terminal hooks of the unci and in keeping the uncus in a pocket-knife resting
position (Lödl, in preparation). The scaphium of Catada represents the same interesting evolutive
progression as the uncus does. The primitive state is just a corium-like dorsal surface of the anal cone
with a strong terminal tooth (C. phaeopasta; fig. 5). The intermediate forms (C. transversalis, C. antevorta,
figs 7, 15) exhibit long, strongly sclerotized, claw-like scaphia. The derived species have the claw-like

Figs 2-7. Plesiomorphic states of unci of Catada sp.; scale = 0.1 mm. 2-5. Catada phaeopasta Hampson, 1909.
Holotype, Noctuidae Brit. Mus. slide No. 16508. 2. Uncus (sc scaphium; u uncus), arrow indicates primary teeth-
like setae on the ventral margin of uncus. 3. dto. — enlarged. 4. Tip of uncus with teeth-like sclerotizations.
5. Scaphium. 6-7. Catada transversalis (Moore, 1877). Noctuidae Brit. Mus. slide No. 16503. 6. Uncus and scaph-
ium. 7. Terminal end of uncus with corona of teeth.

11 l L li RAN \ 13

Figs 8-13. Advanced uncus characters of Catada sp., scale = 0.1 mm. 8. Catada vagalis (Walker, [1859] 1858);
Noctuidae Brit. Mus. slide No. 16027. 9. Catada charalis Swinhoe, 1900. Lectotype, Noctuidae Brit. Mus. slide No.
16504. 10. Catada obscura Joannis, 1906. Noctuidae Brit. Mus. slide No. 16512. 11. Catada canaliferalis (Moore,
1877). Noctuidae Brit. Mus. slide No. 16489. 12. Catada ndalla Bethune-Baker, 1911. Noctuidae Brit. Mus. slide

No. 16509. 13. Catada renalis (Moore, 1882). Noctuidae Brit. Mus. slide No. 16507. Abbreviations: sc scaphium;
ta tuba analis; te tegumen; u uncus.

Figs 14-15. Male genitalia of Catada antevorta (Viette, 1958). Holotype, Viette prep. 3283. 14. Genitalia total,
sacculus coremata fully everted, aedeagus in situ. Scale = 0.5 mm. 15. Uncus-scaphium-complex (sc scaphium;
te tegumen, u uncus). Scale = 0.1 mm.

ndalla
renalis
canaliferalis

obscura

charalis
vagalis

antevorta
transversalis

phaeopasta
Fig. 16. Phylogeny of the species of the genus Catada involved in this study.

type (e.g. C. vagalis, figs 8-9) as well as the fork-like type (Figs 11-13). The flat, fork-like type with
rounded pronges inserts with a rounded base directly at the uncus base and can wear a pair of lappets
at the basal third which insert in a skinny bulbus at the articulation of tegumen and uncus. These
lappets are possible insertions of the muscles 9DV1 and 9 VL1. The first is the longitudinal muscle of
the 9% segment arising on the lateral edge of the tegumen and inserting on the sclerotizations around
the anal cone (Eaton 1984). The latter is a ventrolongitudinal muscle of the 9" segment and arises from
the sclerotized, dorsal parts of the diaphragma and also inserts at the sclerotizations around the anal
cone (Eaton 1984). The middle of the “fork” shows a loop-like window in the sclerotization. This area
is weaker and covered by soft tissue. The function is unclear. Form and situation of this loop are of
specific value.

The cooperation between the helmet-shaped uncus of the derived Catada-species and the claw- or
fork-like scaphia during the mating behaviour is highly speculative. A clasping mechanism is also
possible as a pushing mechanism. The author assumes a clear function in controlling the uncus during
the resting behaviour. This is supported by the obvious correlation of the scaphium length and the
uncus length. Even the protection of the skinny anal cone in the derived species seems to be not
necessary, the possession of a scaphium could be a plesiomorphic feature itself. The primitive type
(C. phaeopasta) with long and hook-shaped uncus clearly requires a contrasting feature which is found
in a knob- and teeth-covered sclerotization of the terminal region of the anal cone. The more derived
species with flat and fork-like scaphia with rounded pronges could insert in the extended terminal area
of the helmet-shaped uncus. A prevention of the anal tube from being pierced by a fine-pointed uncus
is clearly not necessary. The corresponding features on the copulatory tract of the females are very
difficult to locate. In contrast to the males, the females have quite average noctuid genitals (Lödl 1999b).
The genus Catada is an Old World genus with a distribution from the tropical Africa to Australia.
Although the distribution pattern and its evolutive development still remain unclear one can assume
the origin of the primitive Catada-taxa in Equatorial Africa. Species with really plesiomorphic states
have not been found in the Oriental and Australian region so far. C. antevorta and C. transversalis as the
intermediate species occur on islands: Madagascar and the Andaman Islands.

Acknowledgements
I am very much indebted to the colleagues of the BMNH who generously helped as ever. In particular I thank
David Carter, Dr. Jeremy Holloway, Martin Honey, Dr. Malcolm Scoble. Thanks are also due to Dr. N. P.

Kristensen, Copenhagen, for his general comments on the “uncus-matter” and Dr. Joel Minet, Paris, for the loan
of material. And I thank Dr. Sabine Gaal-Haszler, Vienna, for reading the manuscript.

References

Eaton, J. L. 1984. Musculature of the Adult Tobacco Hornworm (Lepidoptera: Sphingidae) Abdomen and
Genitalia. - Annls Ent. Soc. Amer. 77(4): 435-441

Lödl, M. 1999a. Notes on Nolasena ferrifervens Walker, [1858] 1857 (Lepidoptera: Noctuidae). - Quadrifina 2: 125-
133

-- 1999b. Redescription of Catada vagalis (Walker, [1859] 1858) and some notes on the genus Catada Walker,
[1859] 1858 (Lepidoptera: Noctuidae: Hypeninae). — Quadrifina 2: 137-144

-- 2000. Catada icelomorpha Bethune-Baker, 1911 syn.n., new junior, subjective synonym of Catada ndalla
Bethune-Baker, 1911 (Lepidoptera: Noctuidae: Hypeninae). -— Quadrifina 3: 23-31

Poole, R. W. 1989. Lepidopterorum Catalogus (new series). Fasc. 118, Noctuidae Pt. 1-3.-E.J. Brill, Fauna & Flora
Publ., Leiden, New York, 1314 pp.

Tuxen, S. L. 1956. Taxonomist’s Glossary of Genitalia in Insects. - Einar Munksgaard, Copenhagen, 1-284.

SPIXIANA 1 85-92 München, 01. März 2001 ISSN 0341-8391

When molecules claim for taxonomic changes:
New proposals on the classification of Old World treefrogs

(Amphibia, Anura, Ranoidea)
Miguel Vences & Frank Glaw

Vences, M. & F. Glaw (2001): When molecules claim for taxonomic changes:
New proposals on the classification of Old World treefrogs (Amphibia, Anura,
Ranoidea). — Spixiana 24/1: 85-92

Recent phylogenetic data, mainly based on mitochondrial DNA analyses, indi-
cate that current classification of neobatrachian anurans is in need of revision. In the
present paper, we review the literature pertaining the molecular and morpholog-
ical phylogeny of Old World treefrogs. The molecular phylogenies indicate that,
among non-hyperoliid Old World treefrogs, one clade is formed by the endemic
genera from Madagascar, and a second one by the Asian and African genera. Both
these lineages are nested within the family Ranidae sensu Blommers-Schlösser
(1993), but their relationships to each other are not unambigously resolved. We
propose to consider the Asian-African lineage as family Rhacophoridae and the
Malagasy lineage as family Mantellidae. Together with the (paraphyletic) family
Ranidae, these two families form the epifamily Ranoidae. Three epifamilies (Arth-
roleptoidae, Microhyloidae, and Ranoidae) form the superfamily Ranoidea. Within
the family Mantellidae, three subfamilies are recognized: Mantellinae (genera
Mantella and Mantidactylus), Boophinae new subfamily (genus Boophis), and Lalio-
stominae new subfamily (genera Laliostoma and Aglyptodactylus). The new classifi-
cation accounts better for the evolutionary relationships of ranoid frogs and fur-
thermore allows for a classification of the involved Malagasy groups in agreement
with their phylogeny. A satisfactory classification of the whole group, however,
will only be possible with increased phylogenetic knowledge, and will probably
include a further partition of the Ranidae.

Miguel Vences, Museum national d’Histoire naturelle, Laboratoire des Reptiles
et Amphibiens, 25 Rue Cuvier, 75005 Paris, France; address for correspondence:
Zoologisches Forschungsinstitut und Museum A. Koenig, Adenauerallee 160,
53113 Bonn, Germany, e-mail m.vences@t-online.de

Frank Glaw, Zoologische Staatssammlung, Münchhausenstr. 21, D-81247 Mün-
chen, Germany, e-mail Frank.Glaw@zsm.mwn.de

1. Introduction

The phylogenetic relationships of Old World treefrogs have been subject of intensive debate during the
past decades. First all classified in a family Polypedatidae (e. g. Ahl 1931, Noble 1931, as opposed to
the unrelated treefrog family Hylidae with a largely Neotropical distribution), Laurent (1951) noted
that one group, which he defined as Hyperoliidae, was osteologically very different from the remaining
taxa (placed into the Rhacophoridae or Rhacophorinae). The important differences between these two
groups were emphasized later in the phylogenetic approaches of Liem (1970), Drewes (1984) and
Channing (1989). However, Laurent (1951, 1986) also noted that, while hyperoliids share several

symplesiomorphies with some African genera formerly considered as basal representatives of the
Ranidae (and today seen as own families, Arthroleptidae and Astylosternidae; see Dubois 1992),
rhacophorines shared synapomorphies with more derived representatives of the extremely diverse and
speciose family Ranidae. Actually, rhacophorids are only distinguished from other ranids by the
presence of an intercalary element between ultimate and penultimate phalanges of fingers and toes,
and by generally (but not consistently) more arboreal habits. Based on the synapomorphies identified
(e.g., presence of a bony sternal style), Laurent (1986), Dubois (1992) and Blommers-Schlösser (1993)
proposed to include the Rhacophoridae as subfamily Rhacophorinae in the family Ranidae. However,
the proposal of Blommers-Schlösser (1993) (i. e. the definition of the family Ranidae as group contain-
ing almost all ranoid taxa with an ossified sternal style) is all but generally accepted by herpetologists.
Most authors continue considering the Rhacophoridae as separate family (e.g., Frost 1985), a view also
shared by internet databases (as the Amphibian Species of the World database, Amphibiaweb, Tree of
life, Genbank; as of 10 November 2000). Furthermore, new names such as “fanged frogs” (Emerson &
Ward 1998, Emerson et al. 2000a) and “boophids” (Richards et al. 2000) have been used to address
ranoid subclades, without a nomenclatural formalization of these groups.

In the last few years, numerous new results on Old World treefrog phylogeny have been published,
several of them referring to the taxa endemic to Madagascar. In the present paper, we will outline the
main conclusions that can be drawn from a comprehensive view of the new results, and propose a
partly modified classification which better reflects the phylogenetic relationships among ranoid frogs
than the previous schemes. We here focus on only a small subset of this speciose group, namely the
non-hyperoliid Old World treefrogs (Rhacophorinae and Mantellinae sensu Blommers-Schlösser 1993).
Further classificatory modifications will successively become necessary with the accumulation of new
data on ranoid groups such as the Hyperoliidae and the Microhylidae.

2. Summary of recently published molecular data on Old World treefrogs

DNA sequences of ranoid frogs have been analyzed in the context of higher-level phylogenies by Hillis
et al. (1993), Hedges & Maxson (1993), Hay et al. (1995), Ruvinsky & Maxson (1996), and Vences et al.
(2000a). More particular aspects were studied by Bossuyt & Milinkovitch (2000), Emerson & Ward
(1998), Emerson et al. (2000a,b), Feller & Hedges (1998), Marmayou et al. (2000), Richards & Moore
(1996, 1998), Richards et al. (2000), and Vences et al. (2000b,c). The dissertation of Vences (1999) contains
a number of crucial results which are partly published (Vences et al. 2000a,b,c) or in progress of
publication (Vences et al. submitted). Wilkinson & Drewes (2000) furthermore undertook a re-analysis
of non-molecular characters of Old World treefrogs. Altogether, the new data allowed for the identi-
fication of a number of well-supported monophyletic goups, while other splits were much less clearly
resolved (compare Fig. 1):

1. Within the derived and monophyletic group of the Neobatrachia (sensu Feller & Hedges 1998), one
clade containing exclusively groups with a firmisternal shoulder girdle is identified as monophylet-
ic by all available molecular data (Hillis et al. 1993, Maxson et al. 1993, Hay et al. 1995, Ruvinsky
& Maxson 1996, Emerson et al. 2000a, Vences et al. 2000a). This clade includes the Ranidae (with
Rhacophorinae and Mantellinae), Microhylidae, Hemisotidae, Hyperoliidae, Arthroleptidae, and
Astylosternidae. It does not include several other Neobatrachian groups which apparently evolved
a firmisternal or pseudofirmisternal shoulder girdle independently: some leptodactylids, “atelopo-
dine” bufonids, sooglossids and dendrobatids (Duellman & Trueb 1986, Hay et al. 1995, Ruvinsky
& Maxson 1996, Graybeal 1997, Vences et al. 2000a).

2. Within the firmisternal clade as defined above, one basal radiation led to the differentiation of the
families Microhylidae, Hemisotidae, Hyperoliidae, Arthroleptidae, Astylosternidae, and Ranidae
(sensu Blommers-Schlösser 1993). Relationships between all these families are not well resolved,
and some (especially Microhylidae and Hyperoliidae) may not be monophyletic (e.g. Emerson et
al. 2000a). However, one group containing all or most forms characterized by a bony style of the
sternum (Ranidae sensu Blommers-Schlösser 1993, including Rhacophorinae and Mantellinae)
appeared as rather well supported monophyletic group in Emerson et al. (2000a) and Vences (1999).

3. A second large and probably explosive radiation led to the differentiation of the more than 980
species in 54 genera (Glaw et al. 1998a) included in the Ranidae sensu Blommers-Schlösser (1993).
Again, relationships between the different clades of this radiation can not yet be unequivocally

resolved (Emerson et al. 2000b, Vences et al. 2000c). However, two treefrog clades are well defined

by molecular synapomorphies:

- A clade endemic to the Madagascan region which is composed of the genera Boophis (Rhaco-
phorinae), Mantella and Mantidactylus (Mantellinae), Laliostoma (formerly Tomopterna; Raninae)
and Aglyptodactylus (Rhacophorinae, Mantellinae or Raninae) (Richards & Moore 1998, Glaw et
al. 1998b, Richards et al. 2000, Bossuyt & Milinkovitch 2000, Vences et al. 2000b, Vences et al.
submitted).

-— A clade with representatives in Africa and Asia which contains the genera classified in the
Rhacophorinae except the Malagasy Boophis (Richards & Moore 1998, Emerson et al. 2000a,
Richards et al. 2000, Vences et al. submitted).

3. Problems in the classification of Malagasy ranoids

The endemic, non-hyperoliid and non-microhylid ranoid frogs endemic to the Madagascan region
have generally been classified in three subfamilies or families (Frost 1985; Blommers-Schlösser & Blanc
1991; Duellman 1993). The phylogenetic results outlined above show that none of the discussed
classifications is fully satisfactory.

— Laliostoma labrosum was included in the genus Tomopterna and the subfamily Raninae or Tomo-
pterninae until recently (Dubois 1992, Glaw et al. 1998b). Molecular studies of Vences et al. (2000b)
revealed that the three geographic groups of Tomopterna (Africa, Madagascar, southern Asia) are
not closely related (see also Bossuyt & Milinkovitch 2000), and should be included in different
genera — Laliostoma being the monotypic Malagasy genus. Richards & Moore (1998) found that
Mantidactylus, Mantella, Boophis, Aglyptodactylus and Laliostoma labrosum are a monophyletic clade,
and suggested that the latter is a rhacophorid. Laliostoma is the only endemic ranoid from Mada-
gascar without an intercalary element.

- Aglyptodactylus has mostly been considered as belonging to the Rhacophorinae (Blommers-Schlös-
ser 1993), but has sometimes also been included in the Mantellinae (Channing 1989). Glaw et al.
(1998b) found that Aglyptodactylus and Laliostoma labrosum are closely related based on non-
molecular characters although both were traditionally classified in different subfamilies. To remove
this classificatory inconsistency they transferred Aglyptodactylus to the Raninae. Extended data sets
of non-molecular characters (Vences et al. in prep.) and molecular data (Vences et al. 2000b and
submitted) confirm close relationships of Aglyptodactylus and Laliostoma.

-— The genera Mantella and Mantidactylus form the subfamily Mantellinae (a third taxon, Laurentoman-
tis, is considered as one out of 12 subgenera of Mantidactylus; Glaw & Vences 1994). This clade is
well defined by ethological synapomorphies (reduction of strong mating amplexus; egg deposition
outside of water). The Mantellinae were either included in the Rhacophoridae (Channing 1989) or
Ranidae (Frost 1985, Blommers-Schlösser 1993, Glaw et al. 1998b), or seen as own family Mantell-
idae (Blommers-Schlösser & Blanc 1991, Dubois 1992).

- The genus Boophis has generally been included in the rhacophorines which are either considered
as subfamily Rhacophorinae of the Ranidae (e.g. Blommers-Schlösser 1993) or as family Rhaco-
phoridae (e.g. Frost 1985).

4. A new classificatory scheme of the superfamily Ranoidea

The data summarized above clearly corroborate Duellman & Trueb’s (1986) statement that ranid
systematics are “in a state of chaos” and demonstrate the need of an update of the classificatory scheme.
The molecular data indicated the presence of multiple para- and polyphyletic taxonomic units within
ranid frogs (e.g. Emerson et al. 2000a, Vences et al. 2000b). The main goal to be achieved in ranid
systematics in the near future is, in our opinion, a classification void of polyphyletic taxa, and without
or with only a small number of paraphyletic taxa.

Several authors have followed a strategy to exclude small, well corroborated monophyletic groups
from large catch-all groups, as a first contribution to a partition of these large groups into monophyletic
units (e.g. Drewes 1985). We here consider the same approach as useful for the case of the family

Ranidae which is large and diverse enough to be partitioned into several families. Largely based on the
scheme of Dubois (1992), we propose:

- to consider the taxa joined in the Ranidae by Blommers-Schlösser (1993) as epifamily (sensu
Dubois 1992) Ranoidae.

- to join the families Arthroleptidae, Astylosternidae and Hyperoliidae in an epifamily Arthro-
leptoidae, and the families Hemisotidae and Microhylidae in an epifamily Microhyloidae. The
monophyly of the Ranoidae appears rather well assessed, while the monophyly of the remain-
ing two epifamilies is questionable at current state (but not contradicted by any relevant data
sets).

- to recognize, within the Ranoidae, a family Rhacophoridae beside the Ranidae. This family
Rhacophoridae contains the Asian and African rhacophorine genera but not the Malagasy
Boophis.

- to recognize, within the Ranoidae and beside the Ranidae and the Rhacophoridae, a family
Mantellidae, which contains the endemic Malagasy genera Mantella, Mantidactylus, Boophis,
Aglyptodactylus and Laliostoma.

- to subdivide the Mantellidae into the following three subfamilies which correspond to three
monophyletic groups (Richards & Moore 1998, Bossuyt & Milinkovitch 2000, Richards et al.
2000, Vences et al. 2000b):

1. Mantellinae Laurent, 1946

Type genus Mantella Boulenger, 1882.

Genera: Mantella Boulenger, 1882 and Mantidactylus Boulenger, 1895.
Distribution: Madagascar and Mayotte Island.

Arboreal, scansoriel, terrestrial or semi-aquatic firmisternal frogs with a bony sternal style and an
intercalary element between ultimate and penultimate phalanges of fingers and toes. Almost all species
with three free tarsal elements, although the third element may be very small in many species and
absent in rare cases. Finger and toe pads with a complete circummarginal groove. First finger shorter
or of similar length as second finger. Males without nuptial pads, and mostly with femoral glands.
Derived reproductive biology (absence of strong mating amplexus, egg deposition outside of water).

2. Boophinae, new subfamily

Type genus Boophis Tschudi, 1838.

Genus: Boophis Tschudi, 1838.

Distribution: Madagascar and Mayotte Island.

Arboreal (some species partly terrestrial) firmisternal frogs with a bony sternal style and an intercalary
element between ultimate and penultimate phalanges of fingers and toes. Two or three free tarsal
elements. Finger and toe pads with a complete circummarginal groove. First finger shorter or of similar
length as second finger. Males with nuptial pads but without femoral glands. Generalized reproductive
behaviour; eggs (no foam nests) are laid into free water (not in leaf axils nor treeholes).

3. Laliostominae, new subfamily

Type genus Laliostoma Glaw, Vences & Böhme, 1998.

Genera: Laliostoma Glaw, Vences & Böhme, 1998 and Aglyptodactylus Boulenger, 1919.
Distribution: Madagascar.

Terrestrial firmisternal frogs with a bony sternal style and with (Aglyptodactylus) or without (Laliostoma)
intercalary element between ultimate and penultimate phalanges of fingers and toes. Two free tarsal
elements. Finger and toe pads without a circummarginal groove. First finger distincly longer than
second finger. Males with blackish nuptial pads (when breeding) but without femoral glands. Gener-
alized reproductive behaviour; eggs are laid into free stagnant water.

5. Discussion

The new classification proposed (Tab. 1, Fig. 1) divides the “ranids” sensu Blommers-Schlösser (1993)
into a number of well-defined monophyletic taxa, and one paraphyletic taxon. The epifamily Ranoidae
corresponds to the monophyletic clade of ranoid frogs morphologically defined by an ossified sternal
style (secondarily reduced in a few taxa). This epifamily consists, according to our classification, of
three families: the monophyletic Rhacophoridae and Mantellidae as defined here, and the paraphyletic
Ranidae. The latter group will certainly be subject of further partitioning in the future; for example, the
African groups defined as Cacosterninae by Blommers-Schlösser (1993) (possibly together with the
genus Tomopterna; Vences et al. in 2000b), and the Petropedetinae, which probably merit recognition
at familial level. The same is true for the enigmatic genera Ptychadena, Hildebrandtia and Lanzarana
(Ptychadeninae sensu Dubois 1992).

Within the Mantellidae, three further monophyletic groups are recognized according to our clas-
sification. These groups are considered as subfamilies. The advantage of this treatment is that the well-
established definition of Mantellinae as a group containing the genera Mantella and Mantidactylus (e.g.
Blommers-Schlösser 1993, Glaw & Vences 1994) remains stable. The classification of the five genera
(Aglyptodactylus, Boophis, Laliostoma, Mantella, Mantidactylus) in three subfamilies (Boophinae, Lalio-
stominae, Mantellinae) of one family replaces their former classification in three subfamilies (Rhaco-
phorinae, Raninae, Mantellinae) of two or three families. The proposed classification is therefore no
exaggerated splitting approach. In addition, it must be kept in mind that two of the involved genera
are very speciose: Mantidactylus currently contains about 70 nominal species in 12 subgenera, and
Boophis contains more than 40 species in seven species groups. At least 15 new species of each genus
are currently in progress of description. Mantidactylus is furthermore paraphyletic (Richards et al. 2000)
and very diverse regarding the morphology, ecology, and reproductive biology of the species included.
We expect that both genera will be partitioned at genus level when phylogenetic information becomes
available to characterize sufficiently the respective lineages.

In a purely cladistic sense, the proposed classification introduces paraphyly into ranid classification
(by accepting a paraphyletic Ranidae beside the Mantellidae and the Rhacophoridae). However, this
situation is already implicitely accepted by recognition of the Rhacophoridae at family level, by the
majority of herpetologists. Furthermore, we consider this paraphyly as a transitory stage, to be

Tab 1. Summary of the ranoid classification of Dubois (1992) and the proposed modifications according to new
phylogenetic data (M monophyletic, P paraphyletic according to present state of knowledge). Subfamilies are
only shown for the Malagasy family Mantellidae.

Classification according to Dubois (1992) Proposed classification
Superfamily Ranoidea Superfamily Ranoidea (M)
Epifamily Arthroleptoidae Epifamily Arthroleptoidae (M?)
Family Arthroleptidae Family Arthroleptidae (M)
Family Astylosternidae Family Astylosternidae (M?)
Family Hyperoliidae Family Hyperoliidae (P?)
Epifamily Dendrobatoidae
Epifamily Hemisotoidae Epifamily Microhyloidae (M?)
Family Hemisotidae Family Hemisotidae (M)
Epifamily Microhyloidae Family Microhylidae (M?)

Family Microhylidae
Family Scaphiophrynidae

Epifamily Ranoidae Epifamily Ranoidae (M)
Family Mantellidae Family Ranidae (P)
Family Phrynobatrachidae Family Rhacophoridae (M)
Family Ranidae Family Mantellidae (M)
[Subfamily Rhacophorinae] Subfamily Mantellinae (M)
[Subfamily Tomopterninae] Subfamily Boophinae (M)
[Subfamily Raninae] Subfamily Laliostominae (M)

Archaeobatrachia Family Dendrobatidae
Family Leptodactylidae
Family Hylidae
Family Bufonidae

etc. (other hyloid families)

Superfam ly Family Hyperoliidae
Hyloidea Eami
amily Arthroleptidae
Neobatrachia 1) Epifam ily Be ;
Arthroleptoidae Family Astylosternidae
Superfamily (3) Epifam ily ie
Ranoidea Microhyloidae Family Microhylidae

Family Hemisotidae

Eh Family Ranidae
Epifamily (4) er
Samaildes (paraphyletic; contains

several subclades which
probably also merit family
rank)

Family Mantellidae (6)

Family Rhacophoridae

Subfamily Boophinae
SO> Subfamily Laliostominae
® Subfamily Mantellinae

Fig. 1. Schematic phylogenetic consensus tree based on recently published molecular data, clade names accord-
ing to proposed scheme. Only selected groups are included in the figure; subfamilies are only shown for the
family Mantellidae. Evidence for the monophyly of numbered clades is, respectively, provided in the papers of
(1) Hay et al. (1995), Feller & Hedges (1998); (2) Hay et al. (1995), Ruvinsky & Maxson (1997), Feller & Hedges
(1998), Vences et al. (2000a); (3) Hay et al. (1995), Feller & Hedges (1998), Vences et al. (2000a), Emerson et al.
(2000a); (4) Emerson et al. (2000a), Marmayou et al. (2000), Vences et al. (2000c), Vences et al. (submitted);
(5) Richards & Moore (1998), Bossuyt & Milinkovitch (2000), Richards et al. (2000), Vences et al. (submitted);
(6) Richards & Moore (1998), Bossuyt & Milinkovitch (2000), Richards et al. (2000), Vences et al. (2000b);
(7) Richards & Moore (1998), Bossuyt & Milinkovitch (2000), Richards et al. (2000), Vences et al. (2000b); (8) Glaw
et al. (1998b), Richards & Moore (1998), Richards et al. (2000); (9) Bossuyt & Milinkovitch (2000), Vences et al.
(2000b).

maintained only until more ranid clades of unquestionable monophyly are identified and considered
as distinct enough to merit familial rank.
6. Acknowledgements

We are grateful to Wolfgang Böhme (Bonn) and Alain Dubois (Paris) for fruitful discussions.

7. References

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Bossuyt, F. & M. C. Milinkovitch 2000. Convergent adaptive radiations in Madagascan and Asian ranid frogs
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Channing, A. 1989. A re-evaluation of the phylogeny of Old-World treefrogs. - S. Afr. J. Zool. 24(2): 116-131

Drewes, R. C. 1984. A phylogenetic analysis of the Hyperoliidae (Anura): Treefrogs of Africa, Madagascar, and
the Seychelles Islands. — Occ. Pap. California Acad. Sc. 139: 1-70

-- 1985. A case of paraphyly in the genus Kassina Girard, 1853 (Anura: Hyperoliidae). - S. afr. J. Sci. 81: 186-
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Dubois, A. 1992. Notes sur la classification des Ranidae (Amphibiens Anoures). - Bull. mens. Soc. linn. Lyon
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Emerson, S. B., R. F. Inger & D. Iskandar 2000a. Molecular systematics and biogeography of the fanged frogs of
Southeast Asia. - Mol. Phyl. Evol. 16(1): 131-142

-- ,C. Richards, R. C. Drewes & K. M. Kjer 2000b. On the relationships among ranoid frogs: A review of the
evidence. — Herpetologica 56(2): 209-230

-—- &R. Ward 1998. Male secondary sexual characters, sexual selection, and molecular divergence in fanged
ranid frogs of Southeast Asia. - Zool. J. Linn. Soc. 122: 537-553

Feller, A. E. & S. B. Hedges 1998. Molecular evidence for the early history of living amphibians. - Mol. Phyl. Evol.
9(3): 509-516

Frost, D. R. (ed.) 1985. Amphibian species of the world: a taxonomic and geographical reference. — Allen Press,
Assoc. Syst. Coll. Lawrence, Kansas, 732 pp.

Glaw, F., J. Köhler, R. Hofrichter & A. Dubois 1998a. Systematik der Amphibien. Liste der rezenten Familien,
Gattungen und Arten. In: R. Hofrichter (ed.), Amphibien. pp. 252-258. - Naturbuch Verlag, Augsburg

-- &M. Vences 1994. A fieldguide to the amphibians and reptiles of Madagascar. 2nd edition. - Vences & Glaw
Verlag, Köln, 480 pp.
‚-- & W. Böhme 1998b. Systematic revision of the genus Aglyptodactylus Boulenger, 1919 (Amphibia:
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Boophis, Mantidactylus, and Mantella). - J. Zool. Syst. Evol. Res. 36(1): 17-37

Graybeal, A. 1997. Phylogenetic relationships of bufonid frogs and tests of alternate macroevolutionary hypo-
theses characterizing their radiation. — Zool. J. Linn. Soc. 119: 197-338

Hay, J. M., I. Ruvinsky, S. B. Hedges & L.R. Maxson 1995. Phylogenetic relationships of amphibian families
inferred from DNA sequences of mitochondrial 125 and 165 ribosomal RNA genes. - Mol. Biol. Evol. 12(5):
928-937

Hillis, D. M., L. K. Ammerman, M. T. Dixon & R. O. de Sa 1993. Ribosomal DNA and the phylogeny of frogs.
— Herp. Monogr. 7: 118-131

Laurent, R. F. 1951. Sur la necessit€ de supprimer la famille des Rhacophoridae mais de creer celle des
Hyperoliidae. — Rev. Zool. Bot. afr. 45: 116-122

-- 1986. Sous-classe des Lissamphibiens (Lissamphibia). Systematique. 594-797 in: P.P. Grasse & M. Delsol
(eds.), Traite de Zoologie, 14, Batraciens, Fasc. 1-B. — Paris (Masson)

Liem, S. S. 1970. The morphology, systematics and evolution of Old World treefrogs (Rhacophoridae and
Hyperoliidae). — Fieldiana Zool. 57: 1-145

Marmayou, J., A. Dubois, A. Ohler, E. Pasquet & A. Tillier 2000. Phylogenetic relationships in the Ranidae
(Amphibia, Anura): Independent origin of direct development in the genera Philautus and Taylorana. - C.R.
Acad. Sci. Paris (Life Sciences) 323: 287-297

Noble, G. K. 1931. The biology of the Amphibia. - New York (Mc Graw Hill), 577 pp.

Richards, C. M. & W. S. Moore 1996. A phylogeny for the African treefrog family Hyperoliidae based on
mitochondrial rDNA. - Mol. Phyl. Evol. 5: 522-532

-- &-- 1998. A molecular phylogenetic study of the Old World Treefrogs, family Rhacophoridae. — Herpet.
J. 8(1): 41-46

-- ,‚R.A. Nussbaum, & C.J. Raxworthy 2000. Phylogenetic relationships within the Madagascan boophids and
mantellids as elucidated by mitochondrial ribosomal genes. — Afr. J. Herpet. 49: 23-32

Ruvinsky, I. & L. R. Maxson 1996. Phylogenetic relationships among bufonid frogs (Anura: Neobatrachia)
inferred from mitochondrial DNA sequences. — Mol. Phyl. Evol. 5: 533-547

Vences, M. 1999. Phylogenetic studies on ranoid frogs (Amphibia: Anura) with a discussion of the origin and
evolution of the vertebrate clades of Madagascar. - Unpublished Dissertation, Universität Bonn

-- ,]J. Kosuch, F. Glaw, W. Böhme & M. Veith (submitted): Molecular phylogeny and biogeography of Old
World treefrogs (Hyperoliidae, Rhacophorinae, Mantellinae). —- Zool. J. Linn. Soc.

-- ,--,S. Lötters, A. Widmer, J. Köhler, K.-H. Jungfer & M. Veith 2000a. Phylogeny and classification of
poison frogs (Amphibia: Dendrobatidae), based on mitochondrial 165 and 12S ribosomal RNA gene
sequences. — Mol. Phyl. Evol. 15: 34-40

-- ,F. Glaw, J. Kosuch, I. Das & M. Veith 2000b. Polyphyly of Tomopterna (Amphibia: Ranidae) based on
sequences of the mitochondrial 165 and 125 rRNA genes, and ecological biogeography of Malagasy relict
amphibian groups. Pp. 229-242 in Lourenco, W. R. & S. M. Goodman (eds.): Diversite et Endemisme de
Madagascar. - Memoires de la Societe de Biogeographie, Paris

-- ,S. Wanke, G. Odierna, J. Kosuch & M. Veith 2000c. Molecular and karyological data on the south Asian
ranid genera Indirana, Nyctibatrachus and Nannophrys (Anura: Ranidae). - Hamadryad 25(2): 9-16, in press

Wilkinson, J. A. & R. C. Drewes 2000. Character assessment, genus level boundaries, and phylogenetic analyses
of the family Rhacophoridae: A review and present day status. - Contemporary Herpet. 2: 1-19

Buchbesprechungen

7. Jahn, I. (Hrsg., unter Mitwirkung von E. Krause, R. Löther, H. Querner, I. Schmidt, K. Senglaub): Geschichte
der Biologie, Theorien, Methoden, Institutionen, Kurzbiographien. 3. neubearbeitete und erweiterte Aufla-
ge, bearbeitet von 21 Autoren, 1998. 1088 S., 227 Abb. 238 Portraits, G. Fischer Verlag, Jena. ISBN 3-437-
35010-2.

Das vorliegende Werk ist eine großartige Zusammenstellung der Geschichte der Biologie, von den Anfängen in
der Vorgeschichte und im Altertum bis in die zweite Hälfte des 20. Jahrhunderts. Die einzelnen Kapitel
behandeln in sich abgeschlossene Themenbereiche und können gut für sich alleine gelesen werden. So kann
jeder Biologe die historischen Wurzeln seines Fachgebietes nachlesen. Der Leser wird dabei die gemeinsamen
Wurzeln seines Spezialgebietes mit anderen Fachgebieten, ihre Verbindungen und gegenseitigen Befruchtungen
sehen und sein eigenes Fach besser verstehen. Besonders interessant und wertvoll sind die 1650 Kurzbiograph-
ien, mit vielen Portraits und einem eigenen Literaturverzeichnis. Jede Kurzbiographie enthält Informationen
zum Bildungs- und Berufsweg sowie biograpische Quellen. Ohne das Werk in seiner Gesamtheit herab setzten
zu wollen, fiel dem Rezensenten auf, dass die neuere (!) Entwicklung der Systematik (phylogenetische System-
atik, Cladistics etc.) nur marginal gewürdigt wurde. Hierin dürfte sich wohl die allgemein zu geringe Wertschät-
zung dieses Teils der Biologie widerspiegeln.
Alles in allem ein monumentales Werk, das uneingeschränkt für Biologen aller Sparten zu empfehlen ist.
K. Schönitzer

8. Weil, A., L. Brown L. & B. Neville: The wentletrap Book. Guide to the Recent Epitoniidae of the world. -
Evolver, Roma, 1999. 246 pp. ISBN 88-8299-002-8.

Published by three distinct enthusiasts, this book treats a well-known and often collected family of caenogastro-
pods (formerly mesogastropods) and fulfills the expectations of the title. It guides through the diversity and
beauty of wentletrap but rests on shells solely.

The benefit of this book lies in the beautiful photographs of an overwhelming majority of epitoniid species.
If specimens of a particular species were not available to the authors, the original plates and figures have been
reproduced. In any case a short description of each species is provided. The arrangement of species follows the
major marine, biogeographic regions from western Atlantic to the eastern Pacific; genera and species are
arranged alphabetically within each chapter. Three appendices are added: (I) a listing of epitoniid names by
author including synonyms; (II) a description (better: diagnoses) of genera and subgenera; and (III) an abbre-
viation list of museums. The book finishes with an extensive bibliography and index.

It is a pity that protoconch morphology, which has become so important for species identification and also
reflects the mode of reproduction, has escaped the interests of the authors as did all anatomical, biological (e.g.
hosts) and ecological knowledge of this fascinating family. Thus, this book is designed (and recommended) for
shellers, although museum curators would also find it very useful to classify and document their collections.

G. Haszprunar

9. Dorresteijn, A. W. C. & W. Westheide (eds.): Reproductive Strategies and Developmental Patterns in
Annelids. Developments in Hydrobiology 142. - Kluwer Academic Publishers, Dordrecht/Boston/London,
1999. xi + 314 pp. ISBN 0-7923-6018-4.

15 years after “Polychaete reproduction” (Fortschritte der Zoologie, G. Fischer Verlag 1984) this volume repre-
sents the proceedings of a symposium near Osnabrück in September 1997. 18 articles by 27 experts cover a
selection of important annelid taxa (including the Pogonophora, but excluding Echiurida and Myzostomida) and
various aspects of reproductive biology and ecology, as well as ontogeny and phylogeny.

The strength of the book lies in the comparative and interactive consideration of many taxa and phenomena
and the overall phylogenetic background. Of particular use are the well-done reviews of relevant literature on
the reproductive biology of various families showing the value of accurate papers some of which are more than
100 years old. However, it is a pity that the whole field of developmental genetics, nowadays called the “evo-
devo-story”, is entirely missing, even Henry and Martindale remain solely at the phenotypic ground. The final
paper by Westheide, McHugh, Purschke and Rouse reviews all current concepts of annelid stem species,
phylogeny and classification. A detailed index at the very end provides help throughout the whole volume.

Despite the lack of evo-devo, which admittedly is clearly stated in the title of the volume, the book is a must
for everyone who is interested in reproductive biology of annelids. G. Haszprunar

Buchbesprechungen

10. Rieppel, ©.: Einführung in die computergestützte Kladistik. - Verlag Dr. Friedrich Pfeil, München, 1999.
ISBN 3-931516-57-1.

Dieses kleine Büchlein ist exakt das, was im Titel angekündigt ist: eine Einführung, nicht mehr, aber auch nicht
weniger. “Einen kurzen”, einführenden Text in die computergestützte Kladistik in deutscher Sprache zu schrei-
ben”, war das Anliegen des Autors, der sich als Paläontologe und Phylogenetiker der Wirbeltiere einen Namen
gemacht hat. Das Vorhaben ist geglückt, wobei insbesondere der wissenschaftsphilosophische Hintergrund der
“Pattern-Cladistics” kritisch beleuchtet wird. Besonders wichtig erscheint mir die Klarstellung, daß das “Parsi-
monie-Prinzip” grundsätzlich nichts mit der Geschichte der Evolution zu tun hat, sondern wissenschaftstheo-
retisch ein Universalprinzip jeder seriösen Untersuchungsmethode darstellt. Sehr wohltuend auch die Aussage:
“Alles empirische Wissen ist und bleibt hypothetisch” (S. 21).

Ich gehe nicht mit allen Aussagen des Autors konform: so ist etwa ein hypothetischer Vorfahre als notwen-
digerweise paraphyletisches Konzept durch das Auffinden einer einzigen Autapomorphie durchaus falsifizier-
bar (S. 27, 29 versus S. 35). Obwohl der Band entsprechende Fachliteratur nicht ersetzen kann und will, ist er
einfach ein “Muss” für alle phylogenetisch interessierten Systematiker - und welcher gute Systematiker wäre
dies nicht —, und sei es nur, um liebgewordene eigene Standpunkte im Lichte dieser schwungvoll und griffig
formulierten Einführung zu überprüfen. Ein sehr geringer Preis macht Rieppels “Einführung” auch für Studie-
rende attraktiv. Ein nach langen Jahren der Stasis deutscher Systematik überfälliges Buch.

G. Haszprunar

11. Wägele, J.-W.: Grundlagen der Phylogenetischen Systematik. ISBN 3-9315162-73-3, Verlag Dr. Friedrich
Pfeil, München, 2000. 315 pp.

Endlich: Die erste umfassende Bestandsaufnahme der neuen phylogenetischen Methoden, die erste ernsthafte
Auseinandersetzung mit der “pattern cladistics” des anglo-amerikanischen Raumes in deutscher Sprache liegt
vor. Die deutsche Systematik und ihre (auch zukünftigen) Vertreter erhalten wieder Anschluss an die laufende
internationale Diskussion im Fach. Das allein wäre schon genug Anlass zur Gratulation an Autor und Verlag.

Es gibt wohl keinen Artikel oder gar ein Buch zu diesem noch immer heftig diskutierten Thema, das allen
gefallen kann. Auch ich fand einige Stellen, wo ich “nein” sagen muß: Phylogenese ist mehr als Artspaltung, der
Begriff “Stammlinienvertreter” gilt m.E. nicht für ausgestorbene Seitenzweige, Analogien sind mehr als zufällige
Ähnlichkeiten. Bezweifeln möchte ich auch, daß das klassische Articulata-Konzept (Annelida & Arthropoda)
versus die vor allem auf molekularen Daten beruhenden Ecdysozoa-Hypothese (Nemathelminthes & Arthro-
poda) die Überlegenheit der (klassischen) Morphologie über die molekulare Sequenzanalyse belegen kann —
eher wird das Gegenteil eintreten. Insbesondere die Paläontologen dürften einige Schwierigkeiten mit den
Wägeleschen Definitionen (z.B. Chronospecies, Stammart, Plesion) bekommen. Schade auch, daß kaum Beispie-
le aus der Botanik angeführt sind, das hätte den Leserkreis sicherlich ad hoc deutlich erweitert.

Trotzdem: Dieses Buch ist immens wichtig, gehört unbestritten ins Regal jedes Systematikers, egal welcher
Teildisziplin. Es hat Lehrbuchcharakter: Man findet alles gut gegliedert, oftmals mit sehr kompakten Zusam-
menfassungen; auch die beschreibenden Appendices zu den einzelnen Methoden sind wirklich gelungen. Ein
sehr guter Index, ein detailliertes Literaturverzeichnis sowie sehr hilfreiche weiterführende Webseiten runden
die hohe Brauchbarkeit dieses Werkes ab. Ein Tip für den Leser: Starten Sie gleich mit dem 2. Kapitel und lesen
Sie die ersten 45 Seiten über die wissenschaftstheoretischen Grundlagen bei Bedarf.

Das Buch ist von der ersten bis zur letzten Seite alles andere als leichte Kost - es will erarbeitet werden;
gerade das macht aber seinen Wert aus. Zusammenfassung für alle Systematiker und jene, die es sein oder
werden wollen: sofort bestellen und so bald wie möglich lesen. G. Haszprunar

Arndt, E.:

Ashe, P.:

oPIXIANA

ZEITSCHRIFT FÜR ZOOLOGIE

herausgegeben von der
ZOOLOGISCHEN STAATSSAMMLUNG MÜNCHEN

Band 23/2000
Verlag Dr. Friedrich Pfeil, München
ISSN 0341-8391

INHALT - CONTENTS

In Memory of Friedrich Reiss (24 December 1937 — 17 August 1999) .......
ANWordktromithesedltorsk ea eeeseheenneensencnnnnsondnsnreren

Microtendipes schuecki Reiss. Hypopygium dorsal. Original drawing by
Frieder Reiss (in SPIXIANA 20/3, 1997) ........uuu.2244srssnnennenennnnesnnneneeennn

Samples of Frieder Reiss’ handwriting from different periods ...............-
Frieder Reiss in the ZSM collection, 1992 .............uuuessnsserssnnnnenenenneeennnns

Larvae of the subfamily Trechinae from the Southern Hemisphere
(Insectam@oleoptera, Carabidae) "rennen ran ann eennneronneenstanuch ans zue:

Reissmesa, nom. nov., a replacement name for Reissia Brundin (Insec-
ta, Diptera, Chironomidae, Diamesinae) ................22042220402200nesnnnenennnenennnn

Ashe, P., J. P. O’Connor & D. A. Murray: Larvae of Eurycnemus crassipes (Panzer) (Diptera:

Baehr, M.:

Birö, K.:

Butler, M. G.:

Caldwell, B. A.:

Cranston, P. S.:

Chironomidae) ectoparasitic on prepupae/pupae of Hydropsyche siltalai
Döhler (Trichoptera: Hydropsychidae), witn a summary of known
chironomid/trichopteran associations ..........uu.2244neesnnnneennnnnnessnnnnnssnnnnneeen

Review of the Pericalus guttatus-complex (Insecta, Coleoptera, Carab-
Idaenlrebinae) ee en asenebedensouesaneenenees
A new species of the leleupidiine genus Colasidia Basilewsky from
New Guinea (Insecta, Coleoptera, Carabidae, Zuphiinae) ..................

Chironomidae (Insecta, Diptera) from Hungary 2. New records of Lipini-
ella moderata Kalugina, 1970 .........uuuessnunneesnnnnensnnnnnennnnnnnnssnnnnnnnnennennnnn

Tanytarsus aquavolans, spec. nov. and Tanytarsus nearcticus, Spec.
nov., two surface-swarming midges from arctic tundra ponds (Insecta,
Biptera,Chironomidae)r 2... eennnenrenenenone neues andeeenendenruns Srokene

First Nearctic record of Neostempellina Reiss, with description of a new
species (Insecta, Diptera, Chironomidae) ...............u..u0urenennenenneennennnn

Austrobrillia Freeman: immature stages, and new species from the Neo-
tropics (Insecta, Diptera, Chironomidae, Orthocladiinae) ......................-

Fahrner, A. & M. Schrödl: Description of Phyllidia schupporum, a new nudibranch species

from the northern Red Sea (Gastropoda, Nudibranchia, Phyllidiidae) ......

Franzen, M. & U. Heckes: Vipera barani Böhme & Joger, 1983 aus dem östlichen Pontus-

Gebirge, Türkei: Differentialmerkmale, Verbreitung, Habitate (Reptilia,
Serpenles m \Vipendae)eren ee NEN Ea Renee neannfreeachesennes

Glaw, F. & M. Vences: A new species of Mantidactylus from northeastern Madagascar

with resurrection of Mantidactylus blanci (Guibe, 1974) (Amphibia,
An lrasaRanidae) ee

Seite
97-100
100

150
162
174

85-91

112

267-274

33-39

41-45

157-158

211-218

163-166

101-111

55-60

61-70

71-83

Gozmäny, L. A.: Two new Holcopogonid species from Africa (Insecta, Lepidoptera,

Holcopogenidae)k........r.. are eneeaianssennneen ade nnor naar rare 279-281
Hinz, R. (t) & K. Horstmann: Die westpaläarktischen Arten von Exephanes Wesmael

(Insecta,Hymenoptera, Ichneumonidae, Ichneumoninae) ....................... 15-32
Jacobsen, R. E. & S. A. Perry: A review of Beardius Reiss & Sublette, with description of a new

species from Everglades National Park, Florida (Insecta, Diptera,

Ghirenemidae) ss 129-144
Kalezie, M. L., G. DZukic, A. Djorovic & I. Aleksic: Body size, age and sexual dimorphism in the

genus Salamandra. A study of the Balkan species (Amphibia, Uro-

dela, Salamandridae)=...:-"e...eeeaeeeee essen ee REN 283-292
Köhler, J.: A new species of Phyllonastes Heyer from the Chapare region of Bolivia,

with notes on Phylionastes carrascoicola (Amphibia, Anura, Lepto-

dactylidae)..eencuueann. ae ee naneanenaennenene ana ae ee an 47-53
Komposch, C.: Trogulus falcipenis, spec. nov., ein neuer Brettkanker aus den Alpen

und dem Dinarischen Gebirge (Arachnida, Opiliones, Trogulidae) ........ 1-14
Kyerematen, R. A. K., T. Andersen & O. A. Szether: A review of Oriental Rheotanytarsus

Thienemann & Bause, with descriptions of some new species (Insec-

ta, Diptera, Chironomidae)) „were..ereneneeeeeeenenanseanenene nennen 225-258
Langton, P. H. & X.-F. Garcia: A review of Cladotanytarsus conversus (Johannsen) with

first records from Europe (Insecta, Diptera, Chironomidae) ..................- 199-206
Lötters, S. & J. Köhler: A new toad of the Bufo typhonius complex from humid montane

forests of Bolivia (Amphibia, Anura, Bufonidae) ..................sussrrsnnerenne 293-303
Makarchenko, E. A.: Cricotopus (Cricotopus) reissi, spec. nov. from Chukchi Peninsula,

northeastern Russia (Insecta, Diptera, Chironomidae) ..........................- 113-116
Messias, M. C.: Oukuriella reissi, a new species of the genus Oukuriella Epler, 1986

(Insecta, Diptera, Chironomidae) ................4usesesnnnnennennnnnnnnsnnnnnnnsennnennennn 159-161
Oliveira, S. J. de: A new, non-marine species of the genus Thalassomya Schiner, 1856

(Insecta, Diptera, Chironomidae, Telmatogetoninae) ...................e....- 117-120
Orendt, C©.: Chironomids of small Alpine water bodies (springs, spring brooks, pools,

small lakes) of the northern Calcareous Alps (Insecta, Diptera, Chiro-

NOMIdae) een erstere Henna nee ae nenne 121-128
Paggi, A. C. & D. A. Suarez: Ablabesmyia reissi, spec. nov., anew species of Tanypodinae from

Rio Negro province, Argentina, with descriptions of the adult female

and preimaginal stages (Insecta, Diptera, Chironomidae) .....................- 259-266
Reiff, N.: Review of the mainly Neotropical genus Caladomyia Säwedal, 1981, with

descriptions of seven new species (Insecta, Diptera, Chironomidae,

Tanytarsın en RE 175-198
Rupp, D. & Ludwig, P.: First record of Steatonyssus noctulus Rybin, 1992 in Central Europe

(Acari, Mesostigmata, Macronyssidae) ..............uuu.rs24sneresnnnnnnensennnnnesennnen 275-278
Sanseverino, A. M. & S. Wiedenbrug: Description of the pupa of Tanytarsus cuieirensis

Fittkau & Reiss (Insecta, Diptera, Chironomidae) ....................22202220000000 207-210
Stur, E.& T. Ekrem: Tanytarsus usambarae, spec. nov. from West Usambara Mts., Tanzania,

East Africa (Insecta, Diptera, Chironomidae) .................2uu.2220002200002000e 000 219-223
Sublette, J. E.& M. S. Mulla: Chironomus strenzkei Fittkau — a new Pan-American distribution,

with a review of recent similar additions to the Nearctic midges

(Insecta, Diptera, Chironomidae) ..............u..-24unrsnnnnneeennnnnnnnsnnnnenennnnnnn 145-149
Trivinho-Strixino, S. & G. Strixino: A new species of Caladomyia Säwedal, 1981, with

description of the female and immature stages (Insecta, Diptera, Chiro-

NOmIdae) LE EERCEN ESS TOEENeL Lee 167-173
Wülker, W. F. & J. Martin: Northernmost Chironomus karyotypes (Insecta, Diptera, Chironom-

daß) ee N 151-156
Büchbesprechungeng , wur=3=-.0::: et ensnuae srasenghesaneere ee near erkenne 40, 46, 54, 84, 92-94, 282, 304
Jahresinhaltsverzeichnislgg Grm Rene neueren neneh 95-96

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SPIXIANA München, 01. März 2001 ISSN 0341-8391

INHALT - CONTENTS

Seite
Tiefenbacher, L.: Umbellula monocephalus Pasternak, 1964, eine seltene Pennatu-
laria aus dem südlichen Westeuropäischen Becken (Anthozoa, Oc-
tocorallia, Pennatularia)........menserreteeene nennen teen ee 1-4
Senz, W.: Eine neue Heteronemertine von der Küste Japans (Nemertini) ...... 5-13
Schwabe, E.: Taxonomic notes on chitons. 1. Trochodochiton de Rochebrune,
1884 — a genus which was fallen into oblivion (Mollusca, Poly-
placophora,.Mopaliidae)®..........e....2: 222204 neeneee nenne ee 15-18
Karanovic, T.: Description of Allocyclops montenegrinus, spec. nov. and a revision
of the genus Allocyclops Kiefer, 1932 (Crustacea, Copepoda, Cyclo-
BO a0) re re ee AED, RRr. ER. A. A 19-27
Leistikow, A.: The genus Erophiloscia Vandel, 1972 - its phylogeny and biogeo-
graphy, with description of three new species (Crustacea, Isopoda,
Oniseidea) mm nn mnn 29-51
Baehr, M.: Further new and rare species of the genera Fortagonum Darlington
and Collagonum Baehr from New Guinea (Insecta, Coleoptera,
Garabidae. \Agoninaß)ilı.aaleeereereereeeee nennen res ssennnee ee 53-72
Bellido, D. & J. Pujade-Villar: About Andricus polycera (Giraud, 1859) and related forms,
with special remarks on Andricus polycera and A. subterranea (In-
sectan Elymenoptera, Gynipidae)i- meer... 000 13-76
Lödl, M.: A peculiar evolutive lineage of the uncus in the genus Catada Walk-
er, [1859] 1858 (Lepidoptera, Noctuidae, Hypeninae) .................... 77-84
Vences, M. & F. Glaw: When molecules claim for taxonomic changes: New proposals on
the classification of Old World treefrogs (Amphibia, Anura, Rano-
KEN cd Aoosedecetiereeererngoeheoneenbench 5 Akinsoochonsssineee 85-92
Büchbesprechunge nn nn... de a Een 14, 28, 52, 93-94
Jahresinhaltsverzeichnis#2000%.... 2.0, mn 2... Veee e EBE 2ede. 95-96

LARRY

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SPIXIANA 97-102 München, 01. Juli 2001 ISSN 0341-8391

A new species of Loxosoma from north-western Finistere, France

(Spiralia, Kamptozoa (= Entoprocta), Solitaria, Loxosomatidae)
Nikola-Michael Prpic

Prpic, N.-M. (2001): A new species of Loxosoma from north-western Finistere,
France (Spiralia, Kamptozoa (= Entoprocta), Solitaria, Loxosomatidae). — Spixiana
24/2: 97-102

A new species of solitary Kamptozoa is described from the intertidal near Saint-
Pol-de-Leon, France. The species is associated with the annelid Petaloproctus terri-
cola and is characterized by long, spine-like processes of the calyx. For the species
the name Loxosoma nielseni, spec. nov. is proposed.

Nikola-Michael Prpic, Institut für Genetik der Universität Köln, Weyertal 121,

D-50931 Köln, Germany.
E-mail: st2224@zi.biologie.uni-muenchen.de

Introduction

The genus Loxosoma Keferstein, 1862 currently contains 25 species (Nielsen 1996) of exclusively marine
kamptozoans characterized by a round sucking disc at the end of the stalk and by a solitary mode of
life. All species live in association with annelids, except for Loxosoma isolata Salvini-Plawen, 1968, which
seems to be living interstitial in middle-coarse sand, and Loxosoma jaegersteni Nielsen, 1966 for which
the adult habitat is unknown. Here, the find of anew and remarkable species of Loxosoma is reported.

Material and Methods

The material was collected by Claus Nielsen at Plage de Pempoul, a beach near Saint-Pol-de-Leon,
France on June 17th, 2000. It comprises one host annelid specimen and about 50 kamptozoans, all
belonging to the new species. Host and epizoa were narcotized with MgC], (7.5 % in distilled water)
and fixed in Bouin’s fixative. Single specimens were then transferred to microscope slides and mounted
in pure Aquatex (Merck). Prior to this treatment live animals were studied in seawater.

Description of the new species

Loxosoma nielseni, spec. nov.
Figs 1-3

Types. Holotype: One specimen of almost maximum size with one bud (Figs 1D, 2A); Plage de Pempoul, Saint-
Pol-de-Leon, France, holotype (on microscopical slide) deposited at Zoological Museum, University of Copen-
hagen, Denmark (registration number: ZMUC-ENT0024). — Paratypes: Nine specimens of different age, from
same locality, also on slides and deposited in ZMUC (registration numbers: ZMUC-ENT0023a-i).

A EB c D E 2007 um

Fig. 1. Ontogenetic stages of Loxosoma nielseni, spec. nov. All panels are to the same scale (scale bar in E: 200 um).
The specimen in E has been deformed during preparation, causing the stalk to be pushed into the calyx. Without
this deformation the specimen would have approximately the same size as the specimen in D. All specimens are
paratypes, except for the one in D, which is the holotype, and all are fixed.

Etymology. The specific epithet refers to Claus Nielsen, in appreciation of his past and current contributions to
our knowledge of the taxonomy, systematics and phylogeny of the Kamptozoa.

Description

The tallest specimens measure about 950 um (300 um stalk and 650 um calyx), while the smallest
specimens are only about 400 um in length. All specimens possess a round sucking disc at the end of
the stalk with which they adhere to the host annelid. This is the basis for their assignment to the genus
Loxosoma. There are always eight short tentacles; even older buds, that already form an atrium, possess
eight tentacle buds (Fig. 2B, arrow). The stomach is almost V-shaped, but has very conspicuous lateral
pouches. On each side of the calyx there are two small structures with a refraction index much different
from that of the remaining body. These possibly unicellular surface epithelial structures look like beads
and are known as “gland cells”, although their function is not well understood (Emschermann 1972).
They have been found in many kamptozoans, so far. Their value for species characterization is deemed
to be low (Emschermann 1972), but in Loxosoma nielseni their number, location and size (see Figs 2A, 3)
is constant. The stalk has a strong musculature consisting of longitudinal and diagonal muscles, that
together form a dense muscle layer which only at the stalk-calyx border dissolves into single muscle
bands.

The most prominent trait of the new species are the lateral spine-like protrusions of the calyx, one
on each side. The smallest individuals do not possess these protrusions and they also lack any signs
of bud formation (Fig. 1A). Only in animals of about 500 um the outgrowths can be seen, but there are
stillno buds (Fig. 1B). Individuals of about 700 um show clearly visible “spines” protruding from small
wing-like extensions of the calyx. Only on one side of the body, right below one of these “wings”, a
small and mouthless bud can be seen in most individuals of this size (Fig. 1C). In the tallest specimens
bud size is increased (Figs 1D,E) and in specimens having very large buds already with an atrium and
tentacle buds, the formation of a second bud on the other side, but otherwise in the identical position
as the first one, can be observed (Fig. 1E). The lateral protrusions are very long in these specimens,
reaching a maximum of 150 um (Fig. 2A, inset).

The different phenotypes are here interpreted as different ontogenetic stages of the new species
(Figs 1A-E). In this interpretation the series of different specimens reveals a very stereotypic way of bud
formation starting when the growing animal reaches a length of 700 um. The site of bud formation is
restricted to a point below and slightly anterior to the base of the lateral processes. Also timing and
sequence of the budding seem to be tightly regulated: budding occurs first only on one side of the calyx.
Budding on the opposite side is initiated only when the first bud is almost ready for detachment. Also

Fig. 2. Morphology of Loxosoma nielseni, spec. nov. A. Composite microphotograph of the holotype. Note the
“gland cells” (arrowheads). Inset in 2A: detail of a lateral process of the specimen shown in IE, demonstrating
the maximum length of this structure. B. Specimen (paratype) with an older bud (arrow), already showing an
atrium and a forming tentacle crown with eight tentacle buds. All panels are to the same scale (scale bar in A:
150 um). All specimens are fixed.

anus
"

"gland cells

lateral
process
u

lateral pockets
of the stomach

stalk muscles

Fig. 3. Schematic representation of the anatomy of Loxosoma nielseni, spec. nov. The figure has been made after
a living specimen. The bases of the oral tentacles are omitted. The digestive tract is shown from the rear aspect
and “semi-transparent” in order to demonstrate the shape of the stomach. Of the ascending branch only the end
and the anus is shown, the descending branch (mouth slit and esophagus) is omitted. The view into the U-turn
can be seen as a heavily ciliated circle in the middle of the living animal. Scale bar: 100 um.

the lateral processes show a constant growth during ontogeny. Although the growth of the body comes
to a rest at approximately 950 um, this point does not mark the end of process growth and bud
formation (compare figs 1D and 1E); it seems that growth of the processes continues until the formation
of the first bud is finished. The function of the processes is not known.

Habitat. Loxosoma nielseni has been found living on a single specimen of Petaloproctus terricola (det.
Claus Nielsen, pers. comm.), collected in the lower intertidal. Substratum: coarse sand and shell debris.

Differential diagnosis. The combination of body size, number of tentacles and the lateral projections
ofthe calyx readily distinguish Loxosoma nielseni from every other species of Loxosoma. Lateral processes
are known from Loxosoma loxalina Assheton, 1912, Loxosoma davenporti Nickerson, 1898 and Loxosoma
saltans Assheton, 1912. Only in the latter species these processes reach a significant length, but are still
visibly shorter than in Loxosoma nielseni. Additionally, Loxosoma saltans has many more tentacles.
Species similar in habitus to Loxosoma nielseni are: Loxosoma claparedei Bobin & Prenant, 1953, Loxosoma
jaegersteni Nielsen, 1966, Loxosoma significans Nielsen, 1964 and Loxosoma agile Nielsen, 1964, all of which
have more tentacles (except for Loxosoma agile) and lack the lateral processes. The host annelid

100

Petaloproctus spec. is reported only for two Loxosoma species (Loxosoma annelidicola Van Beneden &
Hesse, 1864 and Loxosoma spathula Nielsen, 1966), both differing from the new species in lacking lateral
processes and having not the correct number of tentacles. Additionally, Loxosoma annelidicola has a
balloon-shaped calyx and almost no stalk, and therefore not in any way is resembling the new species.

Discussion

The unique trait of the new species are the extremely long processes of the calyx, the function of which
remains to be investigated. Not unique, but exceptional at the least, are the stereotypical mode of
budding and the strictly constant number of tentacles across all developmental stages available for
study.

The scattered range of occurence of Loxosoma species (see map in Nielsen (1996)) suggests that most
existing species of the genus remain to be discovered. The same probably is true for kamptozoans in
general. This is very unfortunate, since kamptozoans play an important role in the discussion about
metazoan phylogeny (Emschermann 1996, Nielsen 1977, 1995, Mackey et al. 1996, Zrzavy et al. 1998).
The knowledge about plasticity of both their bauplan and life cycle, which could shed light on their
affinities to other phyla and on phylogeny in general, critically depends on the discovery, description
and subsequent study of all existing species. Loxosomella brochobola Emschermann, 1993, for example,
possesses extrusive organs that resemble cnidarian nematocysts in appearance and function (Emscher-
mann, 1993). This throws a critical light on the base of the metazoan tree, since it shows, that
nematocyst-like organs can evolve convergently.

Another example for the plasticity of the kamptozoan bauplan may be Symbion pandora Funch &
Kristensen 1995. Although this species is not entoproct, but ectoproct and thus shows some similarities
to Bryozoa (Funch & Kristensen 1995), and although molecular studies seem to advocate affinities to
Rotifera (Winnepenninckx et al. 1998), Symbion pandora in fact may be a highly derived solitary
kamptozoan (see data in Funch & Kristensen (1995) and Funch (1996)). It thus could demonstrate the
impacts on the bauplan and life cycle of solitary Kamptozoa in case of transition from a tube-dwelling,
non-moulting host (polychaete) to a free-living, moulting one (crustacean).

Acknowledgements

The new species was discovered during the TMR Course for Evolutionary and Developmental Biology, held
from May to June 2000 at the Station Biologique, Roscoff. I thank all participants, teachers and organizers for
making the course such a success; hopefully, the discovery of new species will become a trademark of the course.
I especially thank Lars Wittler, who drew attention on the organism in the first place and did all the work
narcotizing and fixing the animals. And an extremely huge thank-you goes to Claus Nielsen, who provided the
knowledge about entoprocts (and many more phyla) and last but not least — he collected the material!

References

Emschermann, P. 1972. Cuticular pores and spines in the Pedicellinidae and Barentsiidae (Entoprocta), their
relationship, ultrastructure, and suggested function, and their phylogenetic evidence. — Sarsia 50: 7-16;
plates 1-11

-- 1993. On Antarctic Entoprocta: Nematocyst-like Organs in a Loxosomatid, Adaptive Developmental Strat-
egies, Host Specificity, and Bipolar Occurrence of Species. — Biol. Bull. 184: 153-185

-- 1996. Kamptozoa (Entoprocta), Kelchwürmer. In: Westheide, W. & R. Rieger (eds.). Spezielle Zoologie, Teil
1, Einzeller und Wirbellose Tiere. - Gustav Fischer Verlag, Stuttgart, Jena, New York

Funch, P. 1996. The Chordoid Larva of Symbion pandora (Cycliophora) Is a Modified Trochophore. — Journ.
Morphol. 230: 231-263

-—- &R.M. Kristensen 1995. Cycliophora is a new phylum with affinities to Entoprocta and Ectoprocta. —
Nature (London) 378: 711-714

Mackey, L. Y., B. Winnepenninckx, R. De Wachter, T. Backeljau, P. Emschermann & J. R. Garey 1996. 185 rRNA
Suggests That Entoprocta Are Protostomes, Unrelated to Ectoprocta. — Journ. Molec. Evol. 42: 552-559

Nielsen, C. 1977. The Relationships of Entoprocta, Ectoprocta and Phoronida. - Amer. Zool. 17: 149-150

101

-- 1995. Animal Evolution. - Oxford University Press, Oxford, New York, Tokyo

-- 1996. Three new species of Loxosoma (Entoprocta) from Phuket, Thailand, with a review of the genus. - Zool.
Scripta 25: 61-75

Winnepenninckx, B. M. H., T. Backeljau & R. M. Kristensen 1998. Relations of the new phylum Cycliophora. —
Nature (London) 393: 636-638

Zrzavy, J., S. Mihulka, P. Kepka, A. Bezd&k & D. Tietz 1998. Phylogeny of the Metazoa Based on Morphological
and 18S Ribosomal DNA Evidence. - Cladistics 14: 249-285

102

SPIXIANA 103-106 München, 01. Juli 2001 ISSN 0341-8391

A new Architectonica from the Philippines

(Mollusca, Gastropoda, Architectonicidae)
Axel Alf & Kurt Kreipl

Alf, A. &K. Kreipl (2001): A new Architectonica from the Philippines (Mollusca,
Gastropoda, Architectonicidae). — Spixiana 24/2: 103-106

A species of the genus Architectonica Röding, 1798 from the southern Philippines
is described as new. Architectonia proestleri, spec. nov. is relatively small for the
genus and lives in deep water. The species is characterized by the presence of a
central midrib on the body whorl and a multi-ribbed base.

Prof. Dr. Axel Alf, University of Applied Sciences Weihenstephan, D-91746
Triesdorf, Germany. E-mail: axel.alf@fh-weihenstephan.de

Kurt Kreipl, Meeresmuseum Öhringen, Höhenweg 6, D-74613 Öhringen, Ger-
many. E-mail: meeresmuseum@t-online.de

Introduction

In summer 2000 local people started dredging at Aliguay Island, Mindanao, Phlippines. During these
dredgings a species of Architectonica was brought up which is similar to Architectonica consobrina Bieler,
1993, but differs in some characteristic features. Therefore it is described here as new.

Architectonica proestleri, spec. nov.
Figs 1-3

Description (in this description the abbreviations for the sculptural elements used by Bieler (1993) are
taken for better comparison, see also figure No. 1).

Shell thin, light of weight. Diameter of Protoconch 1.18-1.23 mm. Teleoconch of medium to small
size, diameter 12-26 mm (5.0-6.25 whorls). Moderately depressed cone shaped with whorls equally
inflated on upper side and base. Umbilicus moderately wide (about 30 % of shell diameter).

Sculpture of upper side. Subsutural rib (SSR) strong and distinctly separated. Mid rib area divided
in three spiral ribs of which the upper mid rib (UMR) and the central mid rib (CMR) usually are broader
than the lower mid rib (LMR). The spaces between the SSR, the UMR, the CMR and the LMR are about
equal and about 50 % of the broadth of the ribs. Upper peripheral rib (UPR) and lower peripheral rib
(LPR) prominent but narrower; of about equal size. Grooves between LMR, UPR and LPR equal and
broader than these between the mid ribs. Upper part of whorl attachment on upper part of lower
peripheral rib (LPR), upper edge of IPR visible in suture. Upper side of shell and periphery crossed by
deeply incised oblique axial grooves which become smooth on body whorl, segments of the mid ribs
more or less corresponding. The sculpture is completely expressed after 0.5 to 1 teleoconch whorlIs.

Base. Infraperipheral rib (IPR) strong but narrow with one more or less fine additional spiral rib
between LPR and IPR. Base with or without 1 to 3 weak spiral ribs near the umbilicus and distinct equal
axial striae which faden towards the periphery. Proxumbilical rib (PUR) narrow but strong; separated
from base by a wide gap. Nodose rib surrounding umbilicus (UC) large and regular. Columellar wall

103

Fig. 1. Generalized cross-section through the bodywhorl of Architectonica proestleri, spec. nov. SSR = subsutural
rib, UMR = upper midrib, CMR = central midrib, LMR = lower midrib, UPR = upper peripheral rib,
LPR = lower peripheral rib, IPR = infraperipheral rib, PUR = proxumbilical rib, UC = umbilical crenae.

forming almost straight inner lip with two plications, deepest groove in UC overhanging umbilicus; no
spiral sculpture on umbilical side of the wall.

Coloration. Entire shell yellowish to light beige. SSR, UMR, UPR, LPR and IPR slightly lighter
(hardly visible in dead collected and light coloured shells) with irregularily brown blotches of different
sizes. Base becoming lighter from the periphery towards the umbilicus, PUR and UC light with
irregular brown to light brown blotches; ribs on the base also with some irregular blotches, sometimes
corresponding to the blotches on PUR and UC.

Operculum and anatomy not available for study.

Types. Holotype: Diameter: 20.7 mm, diameter of protoconch 1.2 mm; height 10.3 mm; 5.5 teleoconch whorls;
probably life found (Field Museum of Natural History, Chicago, IL 60605, U.S.A. no. FMNH 297361).

Paratypes:
No. Diameter Height Diameter of Number of Remarks Collection
[mm] [mm] Protoconch Teleoconch
[mm] whorls
1 26.6 13.5 1.20 6.25 life taken A. Alf, Weidenbach, Germany, lot
no. 227034a
2 18.8 9.9 1.18 5.25 life taken A. Alf, lot no. 227034a
3 17.6 8.8 1.18 5.0 Probably life taken A. Alf, lot no. 227034a
4 17.6 8.8 1.18 5.0 life taken K. Kreipl, Meeresmuseum Öhrin-
gen, Germany
5 16.5 7.6 1.23 4.75 dead taken Senckenbergmuseum Frankfurt,

Germany no. SMF 321180

Type locality. Alltypes were found at Aliquay Island near Dipolog, Western Mindanao, Philippines.
The shells were dredged in a depth of 120-200 fathoms by local people.

Etymology. The species was named after Mr. W. Proestler, Puerto Galera, Mindoro, Philippines who brought
it to our attention.

Discussion

The only similar species to Architectonica proestleri is Architectonica consobrina Bieler, 1993 from which
it differs by its smaller size, the larger diameter of the protoconch (A. consobrina = 1 mm, A. proestleri
= 1.2 mm), the presence of 3 midribs and only weak ribs on the base fading towards the periphery.
Architectonica proerstleri has a few irregular brown blotches on UC, PUR and on the ribs of the base. The
brown blotches on the SSR, UMR, UPR and LPR are quite irregular and of different size. The coloration
of A. proestleri is paler than that of A. consobrina while the blotches are darker.

104

Fig. 2. Holotype of Architectonica proestleri, spec. nov.

Fig. 3. Paratypes of Architectonica proestleri, spec.nov. Paratype 1 (left), Paratype 2 (middle), Paratype 4 (right).

The only other Architectonica with a multi-ribbed base is Architectonica nobilis Röding, 1798 from the
eastern Pacific, as well as subtropical and tropical eastern and western Atlantik, but this differs clearly
from Architectonica consobrina (see Bieler 1993) and also cannot be confused with Architectonica proestleri
because of its much larger and heavier shell, the presence of 2 mid ribs and the absence of blotches on
BE:

105

Adelphotectonica kuroharai (Kuroda & Habe, 1961) and A. nomotoi (Kosuge, 1979) also show a midrib
area which is divided into up to 3 to 4 spiral ribs, but both do not have a PUR.
Acknowledgements
The authors thank Dr. Rüdiger Bieler (Field Museum of Natural History, Chicago) for reading and discussing
the manuscript. All photographs by Mrs. Uschi Damaschke, Möckmühl, Germany.
References

Bieler, R. 1993. Architectonicidae of the Indo-Pacific (Mollusca, Gastropoda). - Gustav Fischer Verlag, Stuttgart,
Jena, New York

106

SPIXIANA 107-110 München, 01. Juli 2001 ISSN 0341-8391

A new species of Turbinidae Rafinesque, 1815
from the northern Red Sea

(Mollusca, Gastropoda)
Kurt Kreipl & Axel Alf

Kreipl. K. & A. Alf (2001): A new species of Turbinidae Rafinesque, 1815 from
the northern Red Sea (Mollusca, Gastropoda). — Spixiana 24/2: 107-110

A new species of the genus Turbo Linnaeus, 1758 from the northern Red Sea is
described. The shell of Turbo marisrubri, spec. nov. is completely different from
those of the three other species of Turbo known to occur in the Red Sea: Turbo
petholatus Linnaeus, 1758, Turbo radiatus Gmelin, 1791 and Turbo (Lunella) coronatus
Gmelin, 1791.

Kurt Kreipl, Meeresmuseum Öhringen, Höhenweg 6, D-74613 Öhringen, Ger-
many. E-mail: meeresmuseum@t-online.de

Prof. Dr. Axel Alf, University for Applied Sciences Weihenstephan, D-91746
Triesdorf, Germany. E-mail: axel.alf@fh-weihenstephan.de

Introduction

The new Turbo species was offered to the senior author by two shell dealers as Turbo pustulatus Brocchi,
1821, respectively as Homalopoma gestroi (Caramagna, 1888). In fact, Turbo pustulatus is not a Turbo, but
a Homalopoma which is a genus of the Turbinid subfamily Colloniinae Cossmann, 1916. Therefore the
correct name for this species is Homalopoma pustulata (Brocchi, 1821). Homalopoma gestroi — originally
described as Collonia gestroi Caramagna, 1888 - is a synonym.

After a close examination of the literature on Red Sea molluscs we found out that our Red Sea Turbo
obviously represents an undescribed species.

Turbo marisrubri, spec. nov.
Fig. 1

Types. Holotype: Northern Sinai, Gulf of Agaba, Egypt, Red Sea; dived in about 30 m (Senckenberg-Museum,
Frankfurt am Main, Germany, no. SMF 321179.) — Paratypes: 6, from same locality.

Description of holotype

Size. Height: 24.1 mm, width: 21.4 mm.

Shell medium-sized, thick and solid; slightly taller than wide (h/w = 1.1).

Apex pinkish-white; teleoconch consisting of 5 whorls, sculptured by very distinct spiral cords.
Early whorls with keeled shoulder bearing minute spines. Body whorl with three strong spiral cords
at midbody with one much weaker cord between them. Subsutural ramp sculptured with four strong
spiral cords of distinct, rounded nodules. Base with seven spiral cords getting weaker towards the
umbilical area. Suture distinct. Columella smooth and evenly curved, heavily calloused; aperture
round; umbilicus completely closed.

107

Fig. 1. Holotype of Turbo marisrubri, spec. nov.

Basic colour deep reddish-brown, with indistinct, rather regularly spaced axial flames of light and
dark brown, particularly on the subsutural ramp. Some of the spiral cords on the base show very small,
alternating light and dark spots. Columella and aperture white, aperture nacreous within; umbilical
area with a small yellowish-orange blotch.

Operculum thick, round, very finely granulate, with a distinct rim along its outer margin, dull
white.

Soft parts not available for study.

Variation. The paratypes 1-6 do not vary much, neither concerning the shape and sculpture nor the
coloration. Paratype 2 is a bit more slender than the holotype specimen and in paratype 3 the nodules

on the subsutural ramp are not as distinct as in the holotype. All paratypes were collected at the type
locality.

108

Fig. 2. Homalopoma pustulata (Brocchi, 1821).

Paratypes:
Paratype remarks height width collection
no. [mm] [mm]

1 with operculum 24.7 20.2 Kurt Kreipl, Meeresmuseum Öhringen, no 7183a
2 with operculum 24.4 18.9 Kurt Kreipl, Meeresmuseum Öhringen, no 7183b
3 with operculum 20.6 16.9 Kurt Kreipl, Meeresmuseum Öhringen, no 7183c
4 16.9 14.2 Kurt Kreipl, Meeresmuseum Öhringen, no 7183d
5 with operculum 23.9 19.5 Axel Alf, Weidenbach, lot no 112115a
6 with operculum 23.1 18.4 Axel Alf, Weidenbach, lot no 112115a

Etymology. Combination of mare (lat.: sea) and ruber (lat.: red). Genitive: maris rubri = of the Red Sea.

Discussion

Turbo marisrubri, spec. nov. cannot be confused with any other Red Sea Turbo. Turbo petholatus Linnae-
us, 1758 reaches a larger size, is completely smooth and its operculum is smooth and coloured. Turbo
radiatus Gmelin, 1791 is much larger, bears distinct scaly protrusions and spines and has a pale
greenish-blue, grey or pinkish-brown operculum. Turbo (Lunella) coronatus Gmelin, 1791 is distinctly
wider than tall (h/w =0.8-1.0; in T. marisrubri h/w =1.1-1.3), low to flat-spired, with 2 spiral rows of
pointed nodes on either side of the periphery.

The only Turbo species which superficially resembles Turbo marisrubri is Turbo cailletii Fischer &
Bernardi, 1856 from the Caribbean but can be easily distinguished by its open umbilicus and the less
numerous spiral cords on the body whorl (about 12 in cailletii; about 16 in marisrubri) which are smooth
in T. cailletii and nodulose in T. marisrubri.

The only other Turbinid species from the Red Sea which shows a certain similarity to T. marisrubri
is Homalopoma pustulata (Brocchi, 1821). This species can be distinguished from T. marisrubri by its
lighter colour (basic colour: white to dirty green) and especially by the row of red blotches on the outer
margin of the aperture. These red blotches often lead into fine red spiral lines visible on the body whorl
near the aperture. Homalopoma pustulata is not a very typical Homalopoma, but on closer examination
shows all features of this genus.

109

|
Acknowledgements
|

We want to thank Mr. Luigi Bozzetti from Milan, Italy, for useful information on Homalopoma pustulata (Brocchi,
1821) and Mr. Domenico Strazzeri, Heidelberg, Germany, for helping us with the translation of Italian texts. All
photographs by Uschi Damaschke, Möckmühl, Germany.

References

Brocchi, G. 1821. Catalogo di una serie di conchiglie raccolte presso la costa Africana del Golfo Arabico dal sig.
G. Forni. - Biblioteca Italiana 24 (October): 73-86

Caramagna, G. 1888. Catalogo delle conchiglie Assabesi. — Boll. Soc. Malacol. Ital. 13: 113-149

Coulombel, A. 1994. Coquillages de Djibouti. - Edisud, Aix-en-Provence. 1-143.

Dekker, H. & Z. Orlin 2000. Check-list of Red Sea Mollusca. - Spirula 47 (supplement): 1-46

Hickman, C. S. & J. H. McLean 1990. Systematic Revision and Suprageneric Classification of Trochacean
Gastropods. — Nat. Hist. Mus. Los Angeles County, Science Series No. 35

Issel, A. 1869. Malacologia del Mar Rosso, ricerche zoologiche e paleontologiche. - Biblioteca Malacologica, Pisa:
1-387

Sharabati, D. 1984.Red Sea shells. - KPI, London, Boston, Melbourne & Henley: 1-128

110

SPIXIANA 111-121 München, 01. Juli 2001 ISSN 0341-8391

Designation of a type species for the genus Prosekia, gen. nov.
from South America

(Crustacea, Isopoda, Oniscidea)
Andreas Leistikow

Leistikow, A. (2001): Designation of a type species for the genus Prosekia, gen.
nov. from South America (Crustacea, Isopoda, Oniscidea). - Spixiana 24/2: 111-121

The neotropical genus Prosekia Vandel, 1968 originally comprised two species
from Venezuela and a third from the Galapagos islands. Since a type species never
was chosen, the genus name is not available according to 8 13a ICZN. With the
redescription of the Venezuelan species Prosekia rutilans (Vandel, 1952) and the
selection as the type of the genus, the name shall be made available for the
systematics of Oniscidea. Additionally, a new comprehensive definition of the
genus is given and its status is discussed in the light of phylogenetic systematics
and its consequences for Oniscidean systematics.

A. Leistikow, Universität Bielefeld, Abteilung für Zoomorphologie und Syste-
matik, Morgenbreede 45, D-33615 Bielefeld / Ruhr-Universität Bochum, Lehrstuhl
für spezielle Zoologie, Universitätsstraßse 150, D-44780 Bochum.

e-mail: Leiste@Biologie.Uni-Bielefeld.de

Introduction

Several distinct species of the family Philosciidae are known from South America. The family Philosci-
idae is a paraphylum because of the characters used for a definition, as the slender runner-habitus (cf.
Schmalfuss 1984) or many characters given in the diagnosis of Vandel (1973): cephalothorax with linea
supra-antennalis, linea frontalis only in primitive species, pleon narrower than pereon, prominent
neopleurae present in primitive species, three-articulate antennal flagellum, endite of maxilliped with
penicil, genital papilla simple. All these characters are present in Alloniscus Dana, 1853 which is thought
to be the most primitive representative of Oniscoidea (Schmalfuss 1989) or even in Ligia Fabricius, 1795
and thus are plesiomorphies of the Philosciidae. Among the Philosciidae, particularly the species
described around the turn of the last century and ascribed to the genus Philoscia Latreille, 1804 are
difficult to determine due to their poor descriptions. It was Vandel (1952, 1968, 1972) who contributed
to our knowledge on the diversity of philosciid Oniscidea from South America on a higher taxonomic
level. Unfortunately, several authors and even Vandel himself obscured the good beginnings by some
inaccuracy. An example is the genus Prosekia Vandel, 1968 which was described to comprise the species
Chaetophiloscia rutilans Vandel, 1952, Chaetophiloscia hamigera Vandel, 1952 and Chaetophiloscia galapagen-
sis Andersson, 1960. Vandel did not designate a type species, so the genus name is unavailable
according to the ICZN. He mentioned in his description only characters which are shared with other
genera. Furthermore, the three species differ considerably in some characters which are important on
a higher taxonomic level. These characters comprise the shape of the cephalothorax, the compound
eyes, the mouth parts, the pereopods and even some details of the pleopods (pers. obs.). Until recently,
the following species were included in Prosekia Vandel, 1968:

111

Prosekia rutilans (Vandel, 1952)

Prosekia hamigera (Vandel, 1952)

Prosekia galapagensis (Andersson, 1960)

Prosekia tarumae Lemos de Castro, 1984

Prosekia silvatica Lemos de Castro & Souza, 1985

Prosekia lejeunei Lemos de Castro & Souza, 1985

Prosekia insularis Lemos de Castro & Souza, 1985

Prosekia albamaculata Lima, 1996
For phylogenetic analysis of the taxon Oniscidea it is necessary to define monophyletic subtaxa. The
reexamination of the members of Prosekia revealed the paraphyly of this genus. Prosekia rutilans was the
first species described in the section on philosciids from Venezuela by Vandel (1952: 124), so this species
is chosen as the type of the genus Prosekia. All the species but Prosekia insularis, which belongs to the
genus Littorophiloscia, recently have been displaced to the genus Androdeloscia Leistikow (Leistikow
1999). P. rutilans from Venezuela and the genus Prosekia gen. nov. are redefined herein.

Genus Prosekia, gen. nov.

Diagnosis. Cephalothorax with linea supra-antennalis and lamina frontalis, faint linea frontalis, com-
pound eyes with about 22 ommatidia in four rows. Antennula and antennal flagellum three-articulate.
Molar penicil of mandibles composed of 5 to 6 branches, maxillula with lateral endite apically bearing
4+6 teeth, 5 of inner set cleft, medial endite with two stout penicils and inconspicuous tip, maxilla
lacking setation, medial lobe half the breadth of lateral lobe, endite of maxilliped without setation and
knob-like penicil, basipodite with sulcus lateralis.

Pereopods long and slender, carpus 1 with transverse antenna-grooming brush and ornamental
sensory spine with serrate double-fringe on apex, sensory spines of considerable length, tricorn-like
setae of basis flagelliform, coxal plates with noduli laterales, on coxal plate IV inserted more medially,
sulcus marginalis present, gland pores not discernible at 400x magnification. Dactylar seta with apex
slightly plumose, no sexual dimorphism. All female and male pleopod 3 to 5 exopodites slightly ovate,
with lateral margin almost straight, no respiratory areas discernible, endopodites of respective pleo-
pods bilobate. Uropod protopodite laterally with groove, endopodite inserting proximally of ex-
opodite.

Type species. Chaetophiloscia rutilans Vandel, 1952, by monotypy, designated herein.

Prosekia rutilans (Vandel, 1952)
Figs 1-6

Material. Lectotype: 3, 7 mm; paralectotypes: ? 9.5 mm (ovigerous), ? 9 mm. Venezuela, El Junquito, leg. G.
Marcuzzi, 2.V11.1950, deposited in Museum National d’Histoire Naturelle, Paris.

Synonymy. Chaetophiloscia rutilans Vandel, 1952.

Description

Colour. Vandel (1952) wrote: “La teinte generale est d’un rouge carmin fonce. Les zones de lineoles
sont bien visibles; on observe une serie de taches plus foncees sur la ligne mediane, et une autre serie
de taches foncees ä la limite des pleurepimeres. Les pleur&pimeres sont pigmentes, a !’exception d’une
tache claire plus ou moins etendue suivant les segments. Le pleon est entierement pigmente, a
l’exception d’une fine ligne mediane. Les pereiopodes sont en grande partie pigmentes; les exopodites
des pl&opodes sont pigmentes.”

Cephalothorax. Linea supra-antennalis and lamina frontalis prominent, linea frontalis inconspic-
uous, slightly bent, medially interrupted, slight lateral lobes, vertex arched, compound eyes composed
of 22 ommatidia in four longitudinal rows (Fig. 1, Ctf).

Pereon. Body rather convex, tegument smooth and shiny, coxal plates with sulcus marginalis and
noduli laterales, inserted more distally from the lateral margin on coxal plate IV, no gland pores.

112

Fig. 1. Prosekia rutilans (Vandel, 1952). 9 paralectotype. Anl: antennula; An2: antenna; Ctf: cephalothorax in
frontal view; Had: habitus in dorsal view; Hal: habitus in lateral view.

1413

Pleon. Narrower than pereon, neopleurae of pleonites 3 to 5 very small, adpressed to pleon.
Pleotelson with slightly concave lateral margins, bearing some tricorn-like setae inserting near cutic-
ular scale.

Antennula. Composed of three articles subequal in length, distal article rather stout, bearing apical
and medial set of aesthetascs, about 8 aesthetascs of medial set directed medially (Fig. 1, Anl).

Antenna. Fairly slender, length ratio of peduncular articles 1:2:2:4:5, flagellum three-articulate,
proximal article the longest, 1.5x longer as articles 2 and 3 each, apical bristle as long as distal article,
all antennal articles bearing tricorn-like setae, aesthetascs on flagellar article 2 and 3 (Fig. 1, An2).

Mandible. Left mandible with pars intermedia densely covered with coniform setae, bearing two
penicils, on right mandible coniform setae somewhat longer, standing more sparsely, only single
penicil, molar penicil composed of 5 to 6 branches, additional plumose seta distally of molar penicil
(Fig. 2, Mdl/r).

Maxillula. Medial endite with two stout penicils and small tip apically, lateral endite terminated
by 4+6 teeth, 5 of inner set cleft, lateral fringe of trichiform setae sinuous, small additional tooth
subapically on rostral surface (Fig. 2, Mx1).

Maxilla. Both lobes lacking setation, medial lobe of half the breadth of lateral, apically bearing
about 10 cusps, medially some setae (Fig. 2, Mx2).

Maxilliped. Basipodite with sulcus lateralis, distal margin only slightly rounded, endite without
setation, caudally with two teeth, knob-like seta and setal tuft of rostral surface lacking, palp with three
setal tufts on medial border, proximal tuft composed of 3 setae, proximal article bearing two long setae
(Fig. 2, Mxp).

Pereopods. Slender appendages with spinose appearance (Figs. 3, PE1-4, 4, PE5-7), particularly
pereopods 5 to 7 with long sensory spines, tricorn-like setae of basis fairly slender, most sensory spines
of pereopod 1 propus and carpus with apical serrate double-fringe, antenna-grooming brush of
carpus 1 medially surrounded by fasciate cuticular scales, dactylus with short inner claw (Fig. 2, Dac),
prominent interungual seta, dactylar seta with inconspicuous plumose apex (Fig. 2, Sd1).

Pleopods. Pleopod exopodites almost rhomboid with lateral margin straight and medial margin
rounded, laterally with four to six sensory spines, exopodite 5 with transverse row of pectinate scales
on caudal surface. Endopodites bilobate, bearing no setation. No respiratory areas areas on exopodites
visible at 400x magnification (Fig. 5, PL1-5, 6, PL1-2).

Sexual dimorphism. In the original description, Vandel (1952) stated, that there is no sexual
dimorphism in the pereopods. Since only female pereopods could be examined by means of a light
microscope, Vandel’s statement could not be verified in detail. At least there are no differences on the
stereoscope level.

Male pleopod 1 exopodite triangular with rounded edges, apex bent laterally, endopodite rather
stout, apex cylindrical, apically rounded laterally serrate, some cuticular striation on rostral surface,
mediocaudal row of spiniform setae proximally terminating at same level as “lateral saw” (Fig. 5, PL1).

Male pleopod 2 exopodite pointed, with sinuous lateral margin bearing five sensory spines,
endopodite straight, surpassing exopodite, apex slightly bulbous (Fig. 5, PL2).

Uropod. Compare with generic diagnosis (Fig. 4, UR).

Genital papilla: Ventral shield pyriform, but more elongate, mouths of ductus ejaculatorii parallel,
surpassing ventral shield considerably (Fig. 5, Gen).

Discussion

The species which were ascribed to the genus Prosekia represented a rather heterogenous assembly, and
at least three different taxa were united in this genus. Vandel (1968) gave a generic diagnosis including
the following characters: “1. Noduli laterales longs, flagelliformes, atteignant le tiers de la longeur du
tergite pereial. 2. Segment terminal de l’antennule portant deux groups distincts d’aesthetascs.
3. Endopodite du premier pleopode mäle court, portant a son extremite des structures complexes et
generalement dentees.” All these characters, which should define the genus Prosekia, are found in
several other South American genera and are therefore no autapomorphies of the genus. Long noduli
laterales are typical for Andenoniscus Verhoeff, 1941, Erophiloscia Vandel, 1972 and Xiphoniscus Vandel,
1968, but they are comparably shorter in P. rutilans (pers. obs.). The short and dentate endopodites of

114

Fig. 2. Prosekia rutilans (Vandel, 1952). ? paralectotype. Mdl: left mandible; Mdr: right mandible; Mx1: max-
illula with detail of apex of lateral endite; Mx2: maxilla; Mxp: maxilliped with detail of endite in rostral view.

1415

Fig. 3. Prosekia rutilans (Vandel, 1952). ® paralectotype. Dac: dactylus in rostral view; PE1-4: pereopods 1-4
caudal view, details in rostral view; Sc1: ornamental sensory spines of carpus 1; Sc2: sensory spine of carpus 2;
Spl: distal sensory spine of propus 1.

116

Sb7 Sau‘

Fig. 4. Prosekia rutilans (Vandel, 1952). ? paralectotype. PE5-7: pereopods 5-7 in caudal view; Sb7: sensory
spine and tricorn-like seta of basis 7; Sd1: dactylar seta 1; Si5: sensory spine of ischium 5; UR: uropod.

1417

UNN/E N
MEN
N 7, "Ay

mm
NA

Fig. 5. Prosekia rutilans (Vandel, 1952). Gen: genital papilla; PL1-2: pleopods 1 and 2 (4 lectotype); PL3-5:
pleopods 3-5 (7 paralectotype).

118

a
200um

Fig. 6. Prosekia rutilans (Vandel, 1952). ® paralectotype. Cx4: coxal plate IV; PL1-2: pleopods 1 and 2; Os4:

oostegite of pereonite 4; Tel: pleotelson.

200um

119

the male pleopod 1 are found in Andenoniscus and Xiphoniscus (pers. obs.), and finally the shape of the
antennule is similar to Andenoniscus (Leistikow 1998), Erophilosica (Leistikow 2000) and Androdeloscia
(Leistikow 1999). The monophylum which is characterized by the shape of the antennula with an apical
pair of aesthetasc and a medial tuft which sticks out is provisorically called the “Prosekia-group”.
Particularly the smaller species are fairly similar to Andenoniscus, they were disposited in the genus
Androdeloscia (Leistikow 1999) due to differences in the shape of the male pleopod 5 exopodite, and the
cephalothorax.

Prosekia insularis Lemos de Castro & Souza, 1986 which was described from eastern Amazonia is
somewhat different. The shape of the maxilliped and the male pereopod 1 are good features for at least
ascribing the species to Littorophiloscia Hatch, 1947 as can be seen in the re-examination of the genus
by Taiti & Ferrara (1986). The shape of male pleopod 1 is quite similar to Littorophiloscia tropicalis Taiti
& Ferrara, 1986 as can be evidenced from the drawings. Unfortunately, the type material of P. insularis
could not be located.

Prosekia rutilans is differing from the above mentioned species of the Prosekia-group by several
characters. The autapomorphies of Prosekia rutilans are:

e faint linea frontalis which is medially even more inconspicuous [linea frontalis present, not inter-
rupted]

°e profrons more level [profrons with two depressions medially of the eyes]

e knob-like penicil of maxillipedal endite reduced [knob-like penicil present]

° sensory spines of the pereopod 1 carpus apically serrate [only one prominent serrate sensory spine
present, other sensory spines with two subapical tips]

° club-like apex of the male pleopod 2 endopodite, “renfle en vesicule ä son extremite” after Vandel

(1952) [endopodite pointed]

Several other characters are plesiomorphies and exclude P. rutilans from a subtaxon of the Prosekia-
group which comprises Andenoniscus, Androdeloscia, Erophiloscia, and Xiphoniscus: The dactylar seta is
apically plumose, a character shared with the Scleropactidae, Ischioscia Verhoeff, 1928 and several
Scyphacidae. The prominent compound eyes with ommatidia arranged in four rows are likewise found
in the above mentioned taxa, whereas all the other taxa of the Prosekia-group have about 10 ommatidia
which do not appear to be arranged in a distinct pattern. In the description of Vandel (1968) the number
of ommatidia refers to the species now in Androdeloscia. The long noduli laterales, the reduction of the
number of branches of the molar penicil are further apomorphic characters Andenoniscus, Androdeloscia,
Erophiloscia, and Xiphoniscus. Thus, P. rutilans is the basalmost representative of the Prosekia-group.

The distant position of nodulus lateralis IV with respect to the lateral margin of the coxal plate is
a character commonly found in many philosciid taxa. This character was used by Vandel (1952) to
define his groupe chaetophiloscien. Since the polarity for this character is not resolved and many genera
are insufficiently known, it is premature to discuss the monophyly of this group. Nonetheless, Prosekia
and its allies may be related to some of the genera of this group.

Interestingly, P. rutilans has a characteristic feature on the male pleopod 1 endopodite: The cuticle
near the apex is forming some hyaline lamellae. There are only few species with this character. Among
these are several members of the Prosekia-group, like Erophiloscia longistyla Vandel, 1972. Therefore, so
this character has to be ascribed at least to the ground plan of the Prosekia-group. No other philosciid
genus from South America bears such a structure and in other taxa it probably evolved independently.
For example, in Southeastern Asia this character can be found in Exalloniscus bicoloratus Taiti & Ferrara,
1988. But this is due to convergence, since the two genera do not have any character in common which
could be evaluated as asynapomorphy of Prosekia and Exalloniscus, nor is there biogeographic evidence
for a close relationship. For more details, the description of E. bicoloratus should be consulted (Taiti &
Ferrara 1988).

With respect to the compound eyes, Vandel (1952) stated that there are about 10 to 12 ommatidia,
in contradiction to this, he figured 14 (Vandel 1952: 123, Fig. 38). The reexamination revealed the
presence of more than 20 ommatidia in the largest specimen, which were arranged in 4 rows.

120

Acknowledgements

The author thanks Dr. H. Dalens, Universite de Toulouse, for the loan of the material, the permission to dissect
a specimen and the critical review of earlier stages of the manuscript. For manuscript revision he also is grateful
to Dr. A. Ohlers and Dr. S. Taiti. He is indebted to Prof. Dr. J. W. Wägele for his support of this investigation
and the possibility to discuss on this work.

References

Andersson, A. 1960. South American terrestrial isopods in the collection of the Swedish State Museum. - Ark.
Zoöl. 12: 537-570

Leistikow, A. 1998. Redescriptions of terrestrial Isopoda from Chile and Peru (Crustacea: Isopoda: Oniscidea).
— Spixiana 21(3): 215-225

-- 1999. Androdeloscia gen. n., anew genus of South American terrestrial isopods with description of 13 new
species (Crustacea: Isopoda: “Philosciidae”). - Rev. suisse Zool. 106: 813-904

-- 2001. The genus Erophiloscia Vandel, 1972 - its phylogeny and biogeography, with description of three new
species (Crustacea: Isopoda: Oniscidea). - Spixana 24(1): 29-51

Lemos de Castro, A. & Souza, L.A. 1986. Tres especes novas de isopodes terrestres do genero Prosekia Vandel
da Amazonia Brasileira. — Revta. Bras. Zool. 46: 429-438

Schmalfuss, H. 1984. Eco-morphological strategies in terrestrial isopods. - Symposia of the Zoological Society,
London 53: 49-63

-- 1989. Phylogenetics in Oniscidea. — Mon. zool. ital. (N.S.) 4: 3-27

-- 1990. Die Landisopoden Griechenlands. 11. Beitrag: Gattung Chaetophiloscia. —- Rev. suisse Zool. 97: 169-193

Taiti, S. & Ferrara, F. 1986. Taxonomic revision of the genus Littorophiloscia with description of six new species.
- J. Nat. Hist. 20: 1347-1380

-- 1988. Revision of the genus Exalloniscus Stebbing, 1911 (Crustacea: Isopoda: Oniscidea). - Zool. J. Linnean
Soc. 94: 339-377

Vandel, A. 1952. Etude des isopodes terrestres r&coltes au Venezuela par le Dr. G. Marcuzzi. - Mem. Mus. civ.
Storia Nat. Verona 3: 59-203

-- 1968. Isopodes terrestres. - in: N. and J. Leleup (eds.) Mission zoologique belge aux Iles de Galapagos et
Ecuador 84: 35-168

-- 1972. Les isopodes terrestres de la Colombie. — Stud. Neotrop.Fauna Environm. 7: 147-172

-- 1973. Les isopodes terrestres (Oniscoidea) de la Melanesie. — Zool. Verhandl. 125: 1-160

121

Buchbesprechungen

12. Denzau, G. & H.: Wildesel. - Thorbecke Verlag Stuttgart (Thorbecke Species 3), 1999. 221 S., zahlr. Abb. ISBN
3-7995-9081-1.

Anders als der bescheidene klingende Titel vermuten läßt, behandelt dieses großformatige und mit Bildern
reichlich ausgestattete Buch nicht nur die eigentlichen Wildesel Afrikas, sondern auch die im deutschen
Sprachgebrauch als “Halbesel” bezeichneten Equidenformen Asiens, also Onager, Dschiggetais, Kulane, Khur
und Kiangs. Alle diese Formen leben in unwirtlichen und schwer zugänglichen Gebieten Afrikas und Asiens.
Viele Aspekte ihres Verhaltens und ihrer Lebensweise sind daher nur unzureichend bekannt. Die Autoren dieses
Buches haben im Verlauf von 15 Jahren die verschiedenen Wild- und Halbeselformen in ihren natürlichen
Lebensräumen beobachten können - Somaliwildesel in der Danakilwüste, Dschiggetais in der Gobi, Kiangs in
Ladakh, die Khur in Nordwestindien und Kulane in Turkmenistan. Als Ergebnis legen sie nun eine umfangrei-
che Monographie der Afrikanischen und Asiatischen Wildesel vor, die viel zur genaueren Kenntnis vom Sozial-
und Territorialverhalten dieser faszinierenden Tiere beiträgt. Daneben setzen sich die Autoren aber auch
ausführlich mit der Stammesgeschichte, der wirtschaftlichen und historischen Bedeutung der Wildesel sowie
ihrer Bestandssituation und ihrem Gefährdungsgrad auseinander. Hervorzuheben ist auch die klare Darstellung
der äußeren Unterscheidungsmerkmale der einzelnen Arten und Unterarten. Ein besonderer Glanzpunkt des
Buches sind die zahlreichen und wirklich meisterhaften Fotografien, die von den Autoren in den Wüsten und
Halbwüsten Afrikas und Asiens aufgenommen wurden. Sie zeigen beeindruckende Momente aus dem Lebens-
zyklus der Tiere, sei es beim Rivalenkampf, bei der Paarung oder auf ihren Wanderungen durch eine scheinbar
lebensfeindliche Natur. Man kann die Bilder, die mehrfach von der BBC prämiert wurden, ohne Übertreibung
als die besten und aussagekräftigsten Aufnahmen wilder Equuiden bezeichnen. Das Buch trägt wesentlich zur
Kenntnis der Wild- und Halbesel bei. Darüberhinaus ist es den Autoren gelungen, etwas von der Sympathie, die
sie für diese meist viel zu wenig beachtete Einhufergruppe empfinden, auf den Leser zu übertragen.

R. Kraft

13. Bergbauer, M. & B. Humberg: Was lebt im Mittelmeer? - Kosmos Naturführer, Stuttgart, 1999, 319 S.,
zahlreiche Farbabbildungen. ISBN 3-440-07733-0.

Der vorliegende Mittelmeer-Naturführer gehört sicherlich zu den besten seines Genres. Das für Taucher,
Schnorchler und Naturinteressierte gleichermaßen empfehlenswerte Buch bietet zunächst eine Einführung in
die Entstehungsgeschichte des Mittelmeeres und in die wichtigsten Großlebensräume. Im Bestimmungsteil
werden, nach Großgruppen geordnet, 368 marine Pflanzen- und Tierarten (darunter 34 Algen, ca. 250 Wirbellose
und 87 Fischarten) mit ansprechenden Farbfotos vorgestellt. Der Schwerpunkt der “umfassenden Zusammen-
stellung häufiger, bekannter und besonderer Arten” liegt verständlicherweise bei optisch auffälligen, größeren
oder farbenprächtigen Arten. Unscheinbare oder versteckt lebende Arten kommen in manchen Gruppen sehr
kurz: So werden z.B. Gehäuseschnecken auf nur 4 Farbtafeln (9 Arten, davon je 2 Kauris und Tritonshörner)
abgehandelt, während immerhin 14 Nudibranchierarten auf 6 Farbtafeln präsentiert werden. Die Auswahl
“besonderer” Arten ist nicht immer nachvollziehbar. So wird neben der heimischen Mittelmeeralge Caulerpa
prolifera auch die jüngst aus dem Roten Meer einwandernde Caulerpa racemosa vorgestellt sowie letztere als
gefährlicher Substratkonkurrent der heimischen Fauna und Flora in der Nähe von Industriehäfen besprochen.
Caulerpa taxifolia hingegen wird zwar als weiterer Einwanderer erwähnt, aber weder mit Bild vorgestellt, noch
werden die durch C. taxifolia in den letzten Jahren verursachten, dramatischen und großräumigen Veränderun-
gen der Flora und Fauna v.a. der französischen Mittelmeerküste angesprochen. Auch sonst bleiben die Leser von
unerfreulichen Abhandlungen über Ursachen und Wirkungen vielfältiger Umweltzerstörungen im Mittelmeer
verschont, das azurblaue Paradies wird mit keinem Wort von Algenblüten getrübt, von Seeigeln kahlgefressen,
von Eiweißfäden oder Teerklumpen verklebt bzw. allsommerlich von Menschenmassen heimgesucht, über-
düngt und leergefischt.

Insgesamt aber vermittelt dieser Naturführer einen guten Überblick über die Fauna und Flora des Mittel-
meeres, der Schnorchlern und Tauchern das Ansprechen vieler Arten mit Hilfe der Fotos und Beschreibungen
ermöglicht. Im Gegensatz zu manch anderem Mittelmeerführer wird jeweils gesondert auf Verwechslungsmög-
lichkeiten und besondere Unterscheidungsmerkmale zu nicht abgebildeten Arten hingewiesen. Für die fachliche
Qualität der Artbestimmungen und sonstiger Angaben garantieren namhafte Wissenschaftler, die als Berater
konsultiert wurden. Für ernsthafte Bestimmungszwecke ist in vielen Gruppen jedoch das Studium spezieller
Fachliteratur nicht zu ersetzen. Kurze Einführungen zur Systematik und Biologie der jeweiligen Großgruppen
wären wünschenswert und übersichtlicher, als die notwendigen Informationen zur Gruppe auf die Beschreibun-
gen der jeweiligen Arten zu verteilen. Erfreulicherweise bietet der Text zu den Fotos sehr viel Wissenswertes
zu Biologie, Lebensweise und Verbreitung der vorgestellten Arten, und damit eine echte Anregung für Unter-
wasserfans zum Weitersuchen und -lesen. M. Schrödl

122

SPIXIANA 123-128 München, 01. Juli 2001 ISSN 0341-8391

Three new species of litter inhabiting spiders
of the genus Scytodes Latreille from northeastern Brazil

(Araneae, Scytodidae)
Cristina A. Rheims & Antonio D. Brescovit

Rheims, C. A. & A. D. Brescovit (2001): Three new species of litter inhabiting
spiders of the genus Scytodes Latreille from northeastern Brazil (Araneae, Scytod-
idae). - Spixiana 24/2: 123-128

Three new species of litter inhabiting Scytodes spiders are described. Scytodes
maresi, spec. nov. from Mata do Pau Ferro, Areia, Paraiba; Scytodes iabaday, spec.
nov. from Estacäo Biolögica de Una, Ilheus, Bahia; and Scytodes hahahae, spec. nov.
from Parque Nacional de Monte Pascoal, Porto Seguro, Bahia, all in Brazil.

Cristina A. Rheims and Antonio D. Brescovit, Laboratörio de Artröpodes, Insti-
tuto Butantan, Av. Vital Brasil, 1500, Butantä, CEP 05503-900, Säo Paulo SP, Brazil.
e-mail: cris.rheims@vol.com.br; adbresc@usp.br

Introduction

Until 1998, the Brazilian Scytodes comprised a group of fifteen species. Most of these were later
synonymized (Brescovit & Rheims 2000) and a total of 10 new species were later described (Brescovit
& Höfer 1999, Brescovit & Rheims 2000, Rheims & Brescovit 2000). As such, the Brazilian scytodid fauna
today still comprises a total of fifteen valid species.

The most commonly known members of this family are large, long legged animals that inhabit
holes and crevices or lie flat against the substrate (Valerio 1981). However, some scytodids are found
inhabiting the ground litter layer. These spiders are very small, measuring between 1.0 and 3.0 mm.
They differ from the larger, long legged species by the presence of characteristic feathery hairs covering
the abdomen and carapace (Valerio 1981, Brescovit & Höfer 1999). Also, the species found in Amazo-
nian non inundated rain forest (“Matas de Terra Firme”) and some restricted areas of the Atlantic Forest
seem to be endemic.

In this paper we describe three new litter inhabiting scytodid species: two from southern Bahia and
one from “Brejo” Forest in Paraiba, in Brazil.

Material and Methods

The material examined belongs to the following collections: IBSP, Instituto Butantan, Säo Paulo (A.D.
Brescovit); MCN, Museu de Ciencias Naturais da Fundagäo Zoobotänica do Rio Grande do Sul, Porto
Alegre (E. H. Buckup); UESC, Universidade Estadual de Santa Cruz, Ilheus, Bahia (M. de Menezes).
Descriptions and terminology follow Brescovit & Höfer (1999). All measurements are in millimeters.
The epigynes were submerged in lactic acid to study internal structures. Micrographs were obtained
with a JEOL (JSM 840A) scanning electron microscope from the “Laboratörio de Microscopia Eletrönica
do Departamento de Fisica Geral do Instituto de Fisica da Universidade de Säo Paulo (USP).”

LHE-IFUSB
_ 2526

25KU

Figs 1-2. Scytodes maresi, spec. nov. 1. Male palp, retrolateral view. 2. Distal area.
Figs 3-4. Scytodes iabaday, spec. nov. 3. Male palp, retrolateral view. 4. Stridulatory pick.

Scytodes maresi, spec. nov.
Figs 1-2, 5-8

Types. Holotype: 3, Mata do Pau Ferro, Areia, Paraiba, Brazil, Nov. 1999, A. D. Brescovit et al. (IBSP 25827). —
Allotype: ?, same data as holotype. — Paratypes: 13 (IBSP 25831); 1? (IBSP 25835); 13, 1? (MCN), all with the
same data as holotype.

Etymology. The specific name honours the former president of the Fundacäo Nacional do Indio (FUNAI),
Carlos Mares, who resigned after the conflicts between indians and the police, which occurred during the 500
years celebrations in Porto Seguro, Bahia, Brazil

Diagnosis. The male of Scytodes maresi, spec. nov. differs from the other litter inhabiting species by a
ventral basal projection in the distal area of the palp (Figs 1-2) with distal triangular portion and basal
rounded area (Figs 6-7). The female differs from the other species by the large size of the seminal
receptacles and the very thin stalks (Fig. 8).

Description

Male (IBSP 25825). Carapace orange with brown margins and brown central pattern as shown on
fig. 5. Pedipalps yellow. Labium and endites yellow. Sternum yellow with brownish margin. Legs
yellow. Abdomen cream colored with brown transversal bands.

Total length 1.66. Carapace arched, 0.94 long, 0.84 wide, covered with short feathery hairs, longer
on ocular area. Eye diameters: PME 0.07, ALE 0.06, PLE 0.07. Lateral eyes on tubercle. Chelicerae with
subapical hyaline keel. Labium 0.07 long, 0.13 wide. Sternum 0.48 long, 0.44 wide. Legs: I - femur 0.82/
patella 0.22 / tibia 0.76 / metatarsus 0.66 / tarsus 0.36 / total 2.82 / II - 0.66 / 0.22 / 0.62 / 0.54 / 0.32 /
2.36 / III - 0.52 / 0.22 / 0.40 / 0.38 / 0.22 / 1.74 / IV - 0.64 / 0.22 / 0.54 / 0.52 / 0.24 / 2.16. Palpal femur

124

8 —— 7

Figs 5-8. Scytodes maresi, spec. nov. 5. Male carapace, dorsal view. 6. Male palp, retrolateral view. 7. Prolateral
view. 8. Female epigynum, dorsal view. Scale lines = 0.05 mm.

presenting stridulatory pick as in S. iabaday, spec. nov. Cymbium with long and slender distal spine.
Bulb 0.18 long, medially constricted (Fig. 7). Distal area presenting dorsal groove and prolateral fold
(Figs 1, 6-7). The prolateral fold with several small semicircular projections only visible under the
electronic microscope (Fig. 2). Abdomen 0.72 long, 0.70 wide, rounded, covered with large feathery
hairs.

Female (IBSP 25825). Coloration as in male.

Total length 1.68. Carapace arched, 0.88 long, 0.82 wide, as in male. Eye diameters: PME 0.06, ALE
0.05, PLE 0.06. Lateral eyes on tubercle. Chelicerae as in male. Labium 0.10 long, 0.14 wide. Sternum
0.46 long, 0.44 wide. Legs: I- femur 0.78 / patella 0.22 / tibia 0.68 / metatarsus 0.54 / tarsus 0.32 / total
2.54 / II-0.56 / 0.22 / 0.48 / 0.50 / 0.28 / 2.04 / III- 0.44 / 0.20 / 0.36 / 0.34 / 0.16 / 1.50 / IV -0.58 / 0.20 /
0.48 / 0.46 / 0.22 / 1.94. Fovea semicircular widely separated. Internal genitalia with short narrow stalks
with sclerotinized adjacent area (Fig. 8). Abdomen 0.80 long, 0.88 wide, as in male.

Variation. Ten 3: total length 1.54-1.70; carapace 0.86-0.96; femur 1 0.72-0.88; bulb 0.16-0.19; five
29: total length 1.62-1.78; carapace 0.88-1.00; femur I 0.68-0.78.

Distribution. Known only from Paraiba, Brazil.

Natural History. The specimens were collected in pitfall traps and Winkler sampling bags. This
species seems to be endemic to the region associated with the so called “Matas de Brejo”, in the state
of Paraiba, Brazil. These areas are defined by Ab’Saber (1999) as humidity islands in the middle of the
“sertöes” (semi-arid areas). These are composed by tropical forests on high grounds, with well irrigated
soils and permanent water courses (Mayo & Fevereiro 1982, Ab’Saber 1999).

Material examined. Brazil, Paraiba: Areia (Mata do Pau Ferro), 13, 3 juv., Sept. 1999, A. D. Brescovit et al. (IBSP
25823); 234, 2 juv., Sept. 1999, A. D. Brescovit et al. (IBSP 25824); 13, 12, 5 juv., Sept. 1999, A. D. Brescovit et al.
(IBSP 25825); 19, 3 juv., Sept. 1999, A. D. Brescovit et al. (IBSP 25826), (06°57' S; 35°44' W), 17, Sept. 1999, A. D.

125

Figs 9-12. Scytodes iabaday, spec. nov. 9. Male carapace, dorsal view. 10. Male palp, retrolateral view.
11. prolateral view. 12. Female epigynum, dorsal view. Scale lines = 0.05 mm.

Brescovit et al. (IBSP 25829), 13, Sept. 1999, A. D. Brescovit et al. (IBSP 25830), 19, Sept. 1999, A. D. Brescovit et
al. (IBSP 25832), 13, Sept. 1999, A. D. Brescovit et al. (IBSP 25833), 17, Sept. 1999, A. D. Brescovit et al. (IBSP
25834), 17, Sept. 1999, A. D. Brescovit et al. (IBSP 25836).

Scytodes iabaday, spec. nov.
Figs 3-4, 9-12

Types. Holotype: S, Reserva Biolögica de Una, Ilheus, Bahia, Brazil, Apr. 23, 1998, A. D. Brescovit et al. (IBSP
18009). — Allotype: ?, same locality as holotype, Apr. 11, 1998, A. D. Brescovit et al. (IBSP 19308). - Paratypes:
14, Parque Nacional do Pau Brasil, Porto Seguro, Bahia, Apr. 23, 1998, A. D. Brescovit et al. (IBSP 17992); 9, same
data as holotype (IBSP 17977).

Etymology. The specific name honours Henrique Iabaday from the Surui Tribe. The indian directly defied the
President of the Brazilian Congress during the 500 Years celebrations.

Diagnosis. The male of Scytodes iabaday, spec. nov. differs from the other litter inhabiting species by
the distal area of the male palp with ventral triangular projection and elevated squared dorsal area (Figs
10-11). The female differs by the spiraled stalk of the pair of seminal receptacles (Fig. 12).

Description
Male (IBSP 18009). Carapace light brown with dark brown pattern in center and along margin (Fig. 9).
Pedipalps yellow with brownish stains at the base of femora. Labium and endites yellow. Sternum
cream colored with brown stains at the base of the coxae. Legs yellow with brownish stains at the base
of femora. Abdomen cream colored with brown transversal bands.

Total length 1.80. Carapace arched, 0.98 long, 0.86 wide, with long feathery hairs on ocular area
(Fig. 9). Eye diameters: PME 0.06, ALE 0.07, PLE 0.07. Lateral eyes on tubercle. Chelicerae with

126

Figs 13-15. Scytodes hahahae, spec. nov. 13. Male carapace, dorsal view. 14. Male palp, retrolateral view.
15. Prolateral view. Scale lines = 0.05 mm.

subapical hyaline keel. Labium 0.06 long, 0.08 wide. Sternum 0.48 long, 0.44 wide. Legs: I - femur 0.64 /
patella 0.22 / tibia 0.78 / metatarsus 0.66 / tarsus 0.38 / total 2.68 / II - 0.64 / 0.18 / 0.60 / 0.52 / 0.36 /
2.30 / III - 0.50 / 0.20 / 0.40 / 0.38 / 0.24 / 1.72 / IV - 0.64 / 0.22 / 0.52 / 0.48 / 0.30 / 2.16. Palpal femur
presenting stridulatory pick long and slender with rounded and projected socket (Fig. 4). Cymbium
with a long and slender distal spine (Figs 10-11). Bulb 0.16 long, distal area with semicircular projec-
tions only visible under electronic microscope (Fig. 3). Abdomen 0.82 long, 0.84 wide, rounded, covered
with large feathery hairs.

Female (IBSP 19308). Coloration as in male.

Total length 2.16. Carapace 0.94 long, 0.90 wide, as in male. Eye diameters: PME 0.07, ALE 0.06, PLE
0.06. Lateral eyes on tubercle. Chelicerae as in male. Labium 0.11 long, 0.12 wide. Sternum 0.41 long,
0.45 wide. Legs: I - femur 0.64 / patella 0.22 / tibia 0.60 / metatarsus 0.56 / tarsus 0.30 / total 2.32 /
II-0.60 / 0.20 / 0.54 / 0.52 / 0.30 / 2.16 / III-0.58 / 0.24 / 0.40 / 0.40 / 0.24 / 1.86 / IV -0.72 / 0.24 / 0.54 /
0.52 / 0.26 / 2.28. Positioning ridge transversal with fovea semicircular widely separated. Internal
genitalia with a pair of oval seminal receptacles and sclerotized area adjacent to base of stalks.
Abdomen 1.22 long, 0.94 wide, as in male.

Variation. Six dd: total length 1.40-1.80; carapace 0.84-1.06; femur I 0.64-0.84; bulb 0.13-0.17; two
?2: total length 1.82-2.16; carapace 0.94-1.00; femur I 0.64-0.90.

Distribution. South of Bahia, Brazil.
Natural History. The specimens were collected in pitfall traps and by manual litter sorting.

Material examined. Brasil. Bahia: Ilheus, 14, F. Dias (UESC); (Campus do CEPLAC), 17, Apr. 11, 1998, A. D.
Brescovit et al. (IBSP 19233); (Reserva Biolögica de Una), 234, F. Dias (UESC), 14, F. Dias (IBSP 25828), 14, F. Dias
(IBSP 25829).

127

Scytodes hahahae, spec. nov.
Figs 13-15

Types. Holotype: 3, Parque Nacional do Monte Pascoal, Porto Seguro, Bahia, Apr. 21-23, 1998, A. D. Brescovit
et al. (IBSP 19460).

Etymology. The specific name honors the hä-hä-häe pataxö6 tribe from southern Bahia, whose chief, Nailton
Patax6, was the leader of the movement against the 500 years celebrations.

Diagnosis. The male of Scytodes hahahae differs from the other litter inhabiting species by the distal area
of the palp with a retrolateral pocket-like projection (Fig. 14) and a pointed projection in the basal area
(Fig. 15).

Description

Male (IBSP 19460). Carapace yellow with brown pattern behind eyes and along the margin (Fig. 13).
Pedipalps yellow. Labium and endites yellow with light brown margin. Sternum yellow with light
brown margin. Legs yellow. Abdomen cream colored with brown transversal bands.

Total length 1.80. Carapace arched, 0.92 long, 0.84 wide, covered with short feathery hairs, longer
on ocular area (Fig. 13). Eye diameters: PME 0.07, ALE 0.07, PLE 0.07. Lateral eyes on tubercle.
Chelicerae with subapical hyaline keel. Labium 0.06 long, 0.09 wide. Sternum 0.46 long, 0.45 wide.
Legs: I- femur 0.84 / patella 0.20 / tibia 0.80 / metatarsus 0.78 / tarsus 0.28 / total 2.90 / II-0.72 / 0.22 /
0.66 / 0.64 / 0.30 / 2.54 / III - 0.58 / 0.22 / 0.44 / 0.44 / 0.20 / 1.88 / IV - 0.64 / 0.22 / 0.58 / 0.56 / 0.26 /
2.26. Palpal femur presenting stridulatory pick as in S. iabaday, spec. nov. Cymbium with a long and
slender distal spine. Bulb 0.18 long, with dorsal groove with prolateral long and slender rim (Fig. 15).
Abdomen 0.88 long, 0.76 wide, rounded, covered with large feathery hairs.

Female. Unknown.

Distribution. Southern Bahia, Brazil.
Natural History. The specimen was collected in manual litter sampling.

Material examined. Only the type.

Acknowledgements

We wish to thank Prof. Pedro Kiyohara and Miss Simone Perche de Toledo (USP) for the scanning electron
micrographs, Hubert Höfer for helpful comments on the manuscript, Maria de F. Dias and Dr. Max Menezes for
the loan of material. This work was supported by CNPq and “Fundagäo de Amparo ä Pesquisa do Estado de Säo
Paulo” (FAPESP No. 96/ 7052-9; 98/11532-1).

References

Ab’Saber, A. N. 1999. Sertöes Sertanejos: uma geografia humana sofrida. - Estudos Avangados 36(13):7-59

Brescovit, A. D. & H. Höfer 1999. Four new species of litter inhabiting Scytodes spiders (Araneae, Scytodidae)
from Amazonia. — Stud. Neotrop. Fauna & Environm. 34: 105-113

-- &C. A. Rheims 2000. On the synanthropic species of the genus Scytodes Latreille (Scytodidae, Araneae) of
Brazil, with synonymies and records of these species in other Neotropical countries. - Bull. Br. Arachnol.
Soc. 11(8): 320-330

Mayo, S. J. & V. P. B. Fevereiro 1982. Mata de Pau Ferro. A pilot study of the Brejo Forest of Paraiba, Brazil. -
Second report to the Winston Churchill Memorial Trust. 29 pp.

Rheims, C. A. & A. D. Brescovit 2000. Six new species of Scytodes Latreille, 1804 (Araneae, Scytodidae) from
Brazil. - Zoosystema 22(4): 719-730

Valerio, C. E. 1981. Spitting spiders (Araneae, Scytodidae, Scytodes) from Central America. - Bull. Am. Mus. Nat.
Hist. 170: 80-89

128

SPIXIANA 129-140 München, 01. Juli 2001 ISSN 0341-8391

Traverhyphes: a new genus of Leptohyphidae
for Leptohyphes indicator and related species

(Insecta, Ephemeroptera)
Carlos Molineri

Molineri, €. (2001): Traverhyphes: anew genus of Leptohyphidae for Leptohyphes
indicator and related species. (Insecta, Ephemeroptera). — Spixiana 24/2: 129-140

Traverhyphes gen. nov. is established and illustrated for two neotropical species:
Traverhyphes indicator (Needham & Murphy, 1924) new comb., and Traverhyphes
pirai, spec. nov. The first is redescribed from imagos of both sexes, nymphs and
eggs collected in Argentina. Traverhyphes pirai is described from male subimagos
from Brazil (Rio de Janeiro). Female imagos, nymphs and eggs of T. indicator are
described for the first time. Traverhyphes, gen. nov. can be distinguished from the
other genera of Leptohyphidae in the male imago by its characteristic genitalia, and
by the opercular gill size and form in the nymphs.

Carlos Molineri, Insue-Conicet, Facultad de Cs. Naturales e Instituto M. Lillo,
Tucumän, Argentina.

Introduction

The family Leptohyphidae is an important component of the neotropical river fauna and is showing
to be a very diverse group of mayflies, with numerous undescribed taxa. Recently, two related new
genera of Leptohyphidae had been described: Allenhyphes Hofmann & Sartori (in Hofmann et al. 1999)
and Yaurina Molineri (2001). In the present paper another genus is proposed: Traverhyphes to include
Leptohyphes indicator Needham & Murphy (1924) and Taverhyphes pirai, spec. nov.

L. indicator was the sixth species described in the genus and even at that time the unusual form of
the penes was remarked using this character to distinguish it from the other species of the genus
(Needham & Murphy 1924: 32). Later Traver (1958) studied the holotype and some subimagos from
Uruguay and redrew the genitalia noting the presence of a pair of “spear-like processes” at the base
of penes that were omitted in the original description. Posteriorly Dominguez (pers. comm.) studied
the holotype discovering the presence of a pair of long posterolateral projections on the styliger plate
that the preceding authors did not mention.

Male imagos of the type species of the genus (L. eximius Eaton) were reared and show genitalia of
the peterseni-type (Molineri, in prep) and it became clear that L. indicator is not congeneric with this
species.

In the present paper a new genus is described for Leptohyphes indicator and a related new species.
New specific description and drawings are given for L. indicator, including for the first time the female,
eggs and nymphs.

Collections from other localities in South America show that Traverhyphes is a widespread group.
Almost all the material was collected in light traps and is represented only by subimagos, one of these
species (Brazil, Rio de Janeiro) is described and illustrated for its interesting male genitalia.

Terms used for thoracie description and discussion are from Kluge (1992).

129

Traverhyphes, gen. nov.
Figs 1-45

Type species. Leptohyphes indicator Needham & Murphy (1924: 33), original designation.
Species included: T. indicator (Needham & Murphy, 1924) comb. nov. and T. pirai, spec. nov.

Etymology. The genus is dedicated to Jay R. Traver whose work is the base for the present paper and other
studies in Leptohyphidae, “-hyphes” for a common termination in generic names of the family.

Description
Imago. Length of male: body: 3.4-3.8 mm; fore wings: 3.3-3.9 mm; hind wings: 0.60-0.78 mm. Length
of female: body: 4.0-5.0 mm; fore wings: 4.3-4.9 mm.

Head (Fig. 2). Eyes of male separated by a distance of 3 x diameter of an eye, eyes of female by 4 x
diameter of an eye; lateral ocelli large, 0.7 x diameter of an eye; median ocellus small; occiput with a
pair of small circular sclerites behind lateral ocelli. Antennae: pedicel 2.5 x length of scape, flagellum
3x length of pedicel. Frontal part of head with a longitudinal crest from median ocellus to venter.

Thorax. Pronotum with sclerites on lateral %, medially membraneous (Fig. 2). Mesonotum: fore
mesonotal transverse invagination (FMI) deep and well marked; anterolateral corners of mesoscutel-
lum darker than the rest, membranous filaments long (Fig. 3). Legs of male: fore and middle femora
of similar length, hind femora 23-32 % longer than fore femora; fore tibiae and tarsi long, 2.2-2.4 x
longer than middle tibiaettarsi and 1.8-2.0 x longer than hind tibiaettarsi. Legs of female: fore and
middle femora of a similar length, hind femora 20-33 % longer than fore femora; fore and middle tibiae
and tarsi of a similar length, hind tibiae and tarsi long, 1.2-1.3 x longer than fore tibiaettarsi. Pair of
tarsal claws of all legs dissimilar, one blunt paddle-like and the other apically hooked, except on fore
legs of male, both blunt. Wings. Fore wings (Figs 1, 4) with fringed posterior margin, vein Icul attached
at base with CuA and CuP by a cross vein, CuP attached at base with A, A and CuP ends very close
on hind margin. Hind wings of male (Figs 5-6) reduced, total length of hind wings 0.17-0.20 length of
fore wings, absent in female. Hind wings of male with fringed posterior margin and two longitudinal
veins, with a long and curved costal projection 0.47-0.60 x length of wing.

Genitalia. Styliger plate with a pair of large posterolateral projections (Figs 8, 10, 11, 15, 38) dorsal
to forceps; an additional pair of smaller projections arise from hind margin, between base of forceps
(Figs 10, 15, 38, 40), from lateral view these projections are acute. Forceps (Figs 8, 10, 11, 15, 38) three-
segmented, segment 1 short and stout, segment 2 long and slender, segment 3 globular and small.
Penes (Figs 9, 13-14, 38, 40) with a deep but unconspicuous apical notch, penes flattened with a pair
of membranous rounded lobes at apex, these lobes with small knobs as in fig. 41; lateral margins of
penes sclerotized; base of penes with a sclerotized ring (Figs 13, 15), and with a pair of relatively short
dorsal spines arising at the base of the apical lobes, apex of spines perforated and directed medially
(Fig. 40). Male terminal filament 3.6 and cerci 2.7 x length of fore wings. Female terminal filament 2 and
cerci 1.6 x length of fore wings.

Nymph (Fig. 37). Length of male: body, 3.5 mm; mesonotum, 1.3 mm; hind femora, 0.85 mm; tails,
3.2 mm. Length of female: body, 3.6-4.1 mm; mesonotum, 1.3-1.4 mm; hind femora, 0.95-1.00 mm; tails
4.2-4.7 mm. Head hypognathous, wider than long. Antennae 3-3.5 x length of head, flagellum with
whorls of fine setae at articulations. Mouthparts: anteromedian emargination of labrum as in fig. 16,
on ventral side with a pair of asymmetrical submedian rows of setae; mandibles as in figs. 21-22, left
prosteca wider at base than right prosteca; molar region of left mandible with a notorious conical tooth
(arrow on fig. 24); maxillae long and slender, suture between galea and lacinia almost absent (Figs 19-
20), palpi small and bisegmented with apical setae; setae and spines as in figs 19-20; hypopharynx with
asymmetrical setation at base of superlinguae, linguae subcuadrate (Fig. 23); labium (Fig. 18) covering
almost completely venter of head, submentum 3 x wider than mentum, palpi three-segmented (Figs
17218)!

Thorax. Pronotum rectangular, slightly concave at posterolateral corners (Fig. 37). Leg proportions:
maximum length/max. width of femora: fore 2.50-2.67, middle 2.73-2.83, hind 2.85-3.08; fore and
middle femora with a similar length, hind femora 23-25 % longer than fore femora; middle tibiae 0.7 %
longer and hind tibiae 43-50 % longer than fore tibiae; fore and hind tarsi of a similar length, middle
tarsi 13-20 % shorter than fore tarsi. Legs long and slender with spines and scattered groups of fine
setae as in figs 32-34; femora slightly bowed; transverse row of spines on dorsum of fore femora as in

130

Figs 1-7. Traverhyphes indicator. Imagos: 1. Female fore wing. 2. Head of male. 3. Posterior half of male me-
sonotum. 4. Male fore wing. 5. Male hind wing. 6. Detail of hindwing. 7. Abdomen of female. Scale =0.1 mm,
except figs. 1, 4 5 = 1 mm.

fig. 32, spines distally flattened as in fig. 31; middle and hind femora with spines on hind margin;
anterior margin of tibiae of all legs with a pair of parallel rows of spines, distal spines of fore tibiae
pectinated (Fig. 35); middle and hind tibiae with an additional row of spines on hind margin (Figs 33-
34); fore margin of tarsi of all legs with a row of spines, cuticle of basal % of all tarsi darker than the
rest. Tarsal claws (Fig. 36) of all legs with a marginal row of 9 denticles at basal %, a pair of distal
asymmetrical rows of denticles (4-6 submarginal denticles on one side and 1 on the other); with a
subapical setae and with a pair of short setae on basal '% of hind margin as in fig. 36.

Abdomen. Segment II laterally expanded forming the articulation of operculate gill; segments III-
VII laterally expanded forming a protective floor for the gills, lateral processes rounded on segments
III-VI but forming a posterolateral projection on VII; posterolateral spines present on segments VIII-IX.
Terga III-VI with few small spines around gill border, terga VII with groups of long spines forming a
diagonal row at each side posteriorly to apex of gills. Gills: gill of abdominal segment II (Figs 25-26)
formed by an opercular dorsal lamella, ovoid and slightly curved distally, and by a pair of smaller
ventral lamellae; these lamellae are dissimilar, the inferior is larger, as long as opercular lamella, and
protect remaining gills from down side; opercular lamella with a pair of dorsal ribs, on ventral margin

131

Figs 8-15. Traverhyphes pirai. Imagos: 8. Genitalia, l.v. (lateral view). 9. Detail of penes, left v.v. (ventral view),
right d.v. (dorsal view). 10. Genitalia, v.v. Traverhyphes indicator: 11. Genitalia, 1.v. 12. Dorsal projection of
penes, d.v. 13. Detail of penes, d.v. at left, v.v. at right. 14. Penes, l.v. 15. Genitalia. v.v. Scale = 0.1 mm.

and near anterolateral border (Fig. 25). Gills III formed by four lamellae (Fig. 27), gills IV-V by three
lamellae (Figs 28-29) and gills VI by two lamellae (Fig. 30). Terminal filament 10-20 % longer than body
and 30 % longer than cerci, both with whorls of long spines at articulations.

132

Figs 16-24. Traverhyphes indicator. Nymph: 16. Labrum. d.v. at left, v.v. at right. 17. Detail of mentum, d.v. at
left, v.v. at right. 18. Labium, v.v. 19. Detail of maxilla, v.v. 20. Maxilla, d.v. 21. Right mandible, d.v. 22. Left
mandible, d.v. 23. Hypopharynx, d.v. 24. Detail of apex of left mandible, v.v. Scale = 0.1 mm.

Eggs. Form: oval, one polar cap present (Fig. 42). Chorionic sculptures: polygonal and semicircular
overlapping plates (Figs 42-43). Attachment structures: a single polar cap and numerous KTC (Knob
Terminated Coiled Threads) distributed around egg surface (Figs 42-44). Micropyle: one per egg,
located near the uncapped pole (Figs 42-43).

138

Figs 25-36. Traverhyphes indicator. Nymph: 25. Gill of abdominal segment II, d.v. 26. Same, v.v. 27. Gill III, v.v.
28. Gill IV, v.v. 29. Gill V, v.v. 30. gill VL, v.v. 31. Fore femoral spine (detail). 32. Fore leg, d.v. 33. Middle leg.
34. Hind leg. 35. Apex of fore tibiae (detail). 36. Fore tarsal claw (detail). Scale = 0.1 mm.

Discussion

Male genitalia of Traverhyphes shows many interesting characters: penes with a pair of conical spines
on the dorsum (Figs 9, 13, 14, 40) near the base and with an accessory dorsal projection between penes
and cerci. This projection is more or less pyramidal with setae at the tip (Figs 12, 39), and seems to
continue with penes base. In the original description and subsequent emendations by Traver (1958)
nothing is said about this pyramidal projection, and the same applies to a pair of long posterolateral
projections of the styliger plate present in the holotype (Dominguez, pers. com.). These features clearly
separate this genus from all other Leptohyphidae known from adults.

The nymphs of Traverhyphes do not match with any of the published nymphal descriptions.
Traverhyphes nymphs share many characters with Leptohyphes edmundsi Allen and a related undescribed
species collected in the same localities (Misiones, Argentina), differing from them mainly in coloration
and opercular gill size and form. Traverhyphes indicator is not congeneric with the other two species
because of the dissimilarity in male genitalia (Molineri, in prep.) and others characters discussed below.

134

Fig. 37. Traverhyphes indicator nymph, dorsal view. Scale = 1 mm.

Kluge (1992) proposed some thoracic characters defining imagos of Leptohyphes. Almost all of them
appear in a similar form in Traverhyphes: mesonotum with distinct transverse mesonotal suture,
posterior scutal protuberances slightly divergent posteriorly and disposition of sutures on lateropost-
notum of mesothorax. These character states are shared by many leptohyphid genera (except Tricory-
thodes) invalidating their use to define the genus Leptohyphes.

The basic structures of the egg of T. indicator are similar to those described for other members of
the family (e.g. Koss 1968, Koss & Edmunds 1974, Kluge 1992), polar cap seems “type I” of Koss &
Edmunds (1974), the chorion is sculptured with overlapping plates and a single micropyle is present
near the uncapped pole. Eggs of Traverhyphes can be distinguished from the other described eggs of the
family by the form and disposition of chorionic plates (Figs 42-43) and form of KTC (length of
pedunculated part and radial disposition of fibers on the terminal knob, figs 42-44).

Imagos of Traverhyphes can be separated from the other members of Leptohyphidae by the follow-
ing combination of characters: 1. posterolateral borders of styliger plate extended posteriorly (Figs 8,
10, 11, 15, 38); 2. hind margin of styliger plate with a pair of acute projections near the base of forceps
(Figs 15, 38, 40); 3. forceps three-segmented (Figs 10, 15, 38); 4. penes almost completely fused,
dorsoventrally flattened and with a pair of dorsal or laterodorsal conical spines (Figs 9, 13, 14, 38, 40);
5. dorsal projection extending from base of penes (Figs 12, 39); 6. basal % of penes forming a ring
distinctly sclerotized (Fig. 13); 7. lateral margins of penes sclerotized (Figs 9, 13); 8. hind wings present
in males, absent in females; 9. membranous processes of mesoscutellum long and slender (Fig. 3).
Nymphs: 1. abdominal gills present on segments II-VI (Figs 26-30); 2. abdominal gill II ovoid and with
a pair of ventral lamellae as in figs 25-26; 3. gills of segments III-VI as in figs 27-30; 4. maxillary palpi
reduced, bisegmented with apical setae (Figs 19-20); 5. submentum enlarged (Fig. 18); 6. fore femora
with a transverse row of relatively long spines (Figs 31-32); 7. middle and hind femora without
transverse row of setae at base (Figs 33-34); 8. tarsal claws as in fig. 36.

Traverhyphes indicator (Needham & Murphy), comb. nov.
Figs 1-7, 11-45

Leptohyphes indicator Needham & Murphy, 1924: 33, pl. 7, figs 77-78 (male); Lestage 1931: 60; Naväs 1931: 322;
Traver 1958: 500, figs 3, 17, 23 (male); Hubbard 1982: 274; Dominguez 1984: 103; Dominguez et al. 1994: 99,
lam. 28, figs 1-3; Molineri 2001.

135

Description

Male imago (in alcohol). Length: body: 3.5-3.8 mm; fore wings: 3.8-3.9 mm; hind wings: 0.65-0.78 mm.
General coloration yellowish light brown. Head whitish yellow shaded with black as in fig. 2. Antennae
yellowish translucent shaded with gray except flagellum translucent yellowish white.

Thorax. Lateral sclerites of pronotum yellowish shaded with black at carinae and lateral margins,
membrane of the median zone whitish translucent shaded with gray; propleurae hyaline, prosternum
yellowish white with brownish margins and shaded slightly with gray. Mesonotum yellowish brown
except anterolateral corners, fore mesonotal transverse invaginaton and lateroparapsidal sutures
brownish; and medioparapsidal sutures, region between posterior scutal protuberances (PSP) and tip
of mesoscutellum yellowish white; shaded with gray on mediolongitudinal line, with a pair of blackish
marks on anterolateral corners of mesoscutellum (Fig. 3). Metanotum yellowish light brown shaded
with gray on hind margin. Pleural sclerites of pterothorax yellowish light brown, membranes whitish
yellow; shaded with black on paracoxal suture. Meso- and metasterna with yellowish brown sclerites,
median membranous zone whitish translucent shaded with gray. Legs. Coxae and trochanters of all
legs yellowish shaded with gray on coxae. Femora of all legs whitish yellow with yellowish margins,
with a small blackish subapical mark on dorsum. Tibiae and tarsi of all legs translucent yellowish
white, shaded completely with gray on fore tibiae and fore tarsi.

Wings (Figs 1, 4-6). Membrane of fore wings hyaline slightly tinged with yellow except C and Sc
areas tinged with brownish yellow, longitudinal veins brownish shaded with gray, cross veins yellow-
ish. Hind wings (Figs 5-6) hyaline with yellowish costal projection.

Abdomen translucent whitish yellow except segments IX-X whitish yellow, shaded with black on
sublateral regions of terga I-IX, heavier on segments I-VI; remaining area of terga shaded slightly with
gray; median line of terga X light brownish (similar to fig. 7). Pleural folds shaded with gray, darker
on segments IV-VI. Abdominal sterna translucent yellowish white. Genitalia (Figs 11-15, 38-41):
styliger plate yellowish white except anterior margin yellowish and lateral margins brownish; poste-
rolateral projections of styliger plate and forceps segment 1 yellowish translucent, remaining segments
of forceps translucent yellowish white. Penes whitish translucent except dorsal pyramidal projection
of penes yellowish. Cerci whitish translucent shaded slightly with gray, darker at articulations;
terminal filament paler.

Female imago (in alcohol). Length: body (abdomen without eggs): 3.0-3.1 mm; fore wings: 4.3-
4.4 mm. General coloration yellowish brown. Head and thorax as in male imago. Fore wings as in fig.
1. Hind wings absent. Legs as male except fore tibiae and fore tarsi, translucent yellowish white,
without shading with gray. Abdomen (Fig. 7) as male imago except sternum IX yellowish white with
yellowish translucent hind margin, slightly excavated apically. Tails whitish translucent.

Female subimago (in alcohol). Length: body (abdomen extended, with eggs): 4.0-5.0 mm; fore wings:
4.3-4.9 mm. Similar to imago.

Mature nymph (in alcohol, fig. 37). Length of male: body: 3.5 mm; mesonotum: 1.3 mm; hind femora:
0.85 mm; tails: 3.2 mm. Length of female: body: 3.6-4.1 mm; mesonotum: 1.3-1.4 mm; hind femora: 0.95-
1.0 mm; tails: 4.2-4.7 mm. General coloration light brownish with black markings.

Head yellowish brown shaded with black among ocelli and around antennae, occiput with grayish
mediolongitudinal band and posterolateral corners. Antennae yellowish translucent. Mouthparts (Figs
16-24) yellowish shaded with gray on median zone of labium.

Thorax yellowish brown shaded widely with gray except on submedian triangular marks of
pronotum; with a pair of notorious blackish ovoid marks between developing wings; developing wings
yellowish translucent with blackish costal margin and black basal sclerite. Pleurae and sterna yellowish
white shaded with gray. Legs whitish yellow with black subapical marks on femora (Figs 32-34); basal
% of tarsi of all legs brownish translucent (Figs 32-34).

Abdomen yellowish brown shaded with black on anterolateral regions of terga II-IX; tergum I
completely shaded with gray; shaded slightly with gray on terga II-VI, less marked on mediolongitu-
dinal band; tergum X shaded with gray on hind margin. Abdominal sterna I-VI yellowish white, VII-
IX yellowish. Gills (Figs 25-30): opercular lamellae light brown with black anteromedian margin, apical
portion with depigmented maculae as in fig. 25, remaining gills translucent yellowish white. Tails
yellowish translucent with whorls of long spines at articulations.

Eggs. Mean length: 182 um; mean width: 79 um. Color: light green, polar cap whitish. Chorion:
polygonal overlapping plates near the uncapped pole, decreasing in number and becoming more or
less semicircular toward the capped pole (Fig. 42). These plates sculptured with small granules on

136

1 i:
A SUN
SC

| \ N
N N

Figs 38-41. Traverhyphes indicator, SEM photographies: 38. Male genitalia, v.v. 39. Dorsal projection of penes,
d.v. 40. Penes and base of forceps, d.v. (pyramidal projection removed). 41. Apex of penes (right apical lobe),
v.v. Scales: 38: 100 u; 39, 40, 41: 10 u.

137

thickest margin (the nearest to the uncapped pole, fig. 43). Attachment structures: a single polar cap
present, formed by numerous knob terminated non-coiled threads (Figs 44-45); few knob terminated
coiled threads attached between chorionic plates and on the smooth chorion rounding polar cap. These
KTC increase in number toward capped pole, and consist in a basal pedunculated part formed by
numerous coiled fibers located radially in the distal part, forming the knob (Fig. 43). Micropyle: one
per egg, located near the uncapped pole (Fig. 42), circular or pentagonal in form and delimited by five
chorionic plates (Fig. 43).

Observations. Posterolateral projections of styliger plate vary in size between subimagos and imagos,
reaching its definitive form after ecdisis to imago. Two male imagos have dark spots on some
longitudinal veins of fore wings, and other two male imagos have posterolateral projections of styliger
plate smaller than the rest.

Life cycle associations. Nymphs and adults are associated by reared nymphs of both sexes.

Material. Holotype © imago, deposited in Cornell University, Ithaca, New York, USA; drawings from wings
and genitalia made by E. Dominguez. - Other material: 534, 19 imagos from ARGENTINA, Misiones, PN
Iguazu, Puerto Canoas, 26-XI-998, at light, Dominguez, Molineri & Nieto Col.; 5034, 25 nymphs from ARGEN-
TINA, Misiones, PP Urugua-i, RP 19, A® Uruzu, 23-24-XI-1998, Dominguez, Molineri & Nieto Col.; 13 imago, 1?
subimago, both reared, from ARGENTINA, Misiones, PP Urugua-i, RP 19, A° Uruzu, 7-XN-1999, Molineri Col.;
929 imagos from ARGENTINA, Misiones, Dpto. San Pedro, Confluencia Rios Alegria y Piray-Guazu, 22-23-XI-
1998, Dominguez et al. Col. Other localities in Misiones: Rio Cuna-Piru, A° Märtires (C’ Azul), Bonpland, San
Vicente, El Soberbio. All the material is deposited in the collections of Instituto- Fundaciön Miguel Lillo,
Tucumän, Argentina.

Discussion. T. indicator can be distinguished from T. pirai by the following combination of characters:
1. abdominal terga shaded with black at lateral margins, median band paler (Fig. 7); 2. penes with a
similar width along their length (Figs 13, 38); 3. origin of peneal spines dorsal (Figs 13-14, 40).

Traverhyphes pirai, spec. nov.
Figs 8-10

Etymology. “Pirai” from the name of the river where the material was collected.

Description

Male subimago (in alcohol). Length: body: 3.4-3.5 mm; fore wings: 3.3-3.6 mm; hind wings: 0.60-
0.73 mm. General coloration yellowish-orange. Head yellowish white shaded completely with gray
except on a pair of small circular marks behind lateral ocelli; venter of head paler, not shaded with gray
except laterally. Antennae: scape and pedicel whitish translucent shaded with gray [flagellum broken
off and lost]. Thorax. Pronotum medially whitish translucent shaded with gray except on a pair of
sublateral longitudinal bands and small scattered dots; lateral Y yellowish-white shaded with gray on
carinae and lateral margins; propleurae and prosternum whitish-translucent shaded with gray. Meso-
notum yellowish-orange, slightly paler between medioparapsidal sutures; anterolateral margins heavy
sclerotized, orangeish-brown; shaded with light gray on carinae and between posterior scutal protu-
berances; anterolateral corners of mesoscutellum with a small grayish mark; membranous filaments
whitish translucent. Mesopleural sclerites yellowish, with whitish membranes, shaded with gray on
carinae; mesosternal sclerites orangeish-yellow, median membrane whitish-translucent shaded with
gray. Metanotum yellowish shaded with gray on carinae and margins; metapleurae and metasternum
whitish-yellow. Legs. Coxae and trochanters of all legs yellowish-white shaded with gray dorsally;
remaining segments of all legs whitish-yellow, femora of all legs with a subapical blackish mark on
dorsum; fore leg shaded completely with gray, remaining legs not shaded. Wings. Membrane of fore
and hind wings whitish translucent, longitudinal veins yellowish white. Abdomen translucent yellow-
ish-white shaded with gray dorsally; shaded more marked on longitudinal submedian bands on terga
II-V and on anterior margin of VI; abdominal sterna shaded with gray only at lateral margins. Genitalia
(Figs 8-10): styliger plate yellowish-white; forceps and penes whitish translucent. Tails whitish-trans-
lucent.

Female and nymph. Unknown.

138

Figs 42-45. Traverhyphes indicator eggs, SEM photographies: 42. General view of egg. 43. Detail with micropyle
and KTC. 44. Basal part of polar cap and KTC near pole. 45. Detail of polar cap. Scales: 42: 100u; 43, 44: 10 u;
45: 1.0 u.

139

Material. Holotype and 5 paratypes © subimagos from: Brazil, Rio de Janeiro, Mun. Rio Claro, Rio Pirai, 8-IV-
1977, CM & OS Flint Jr, Cols. Holotype and three paratypes deposited in National Museum of Natural History,
Smithsonian Institution, Washington D. C., USA; two paratypes deposited in Instituto-Fundaciön Miguel Lillo,
Tucumän, Argentina.

Discussion. Male subimagos of this species can be differentiated from T. indicator by the following
combination of characters: 1. abdominal terga shaded uniformly with gray, darker on a pair of
submedian longitudinal lines on terga I-VI; 2, apical half of penes wider than basal part (Fig. 9);
3. origin of peneal spines dorsolateral (Fig. 9). Subimaginal male genitalia of T. indicator does not change
when molting to imago, except for the relative length of the posterolateral proyections of styliger plate.
For this reason already at the stage of subimago this two species are readily distinguishable.

Acknowledgments

The present paper was completed while the author was supported by a fellowship from the National Council
of Scientific Research of Argentina (CONICET). I want to thanks Eduardo Dominguez for critical reading of the
manuscript.

References

Dominguez. E. 1984. Dos especies nuevas del genero Haplohyphes Allen (Ephemeroptera: Tricorythidae) de la
Argentina. — Rev. Soc. Entomol. Argentina 43(1-4): 103-112

-- ,M.D. Hubbard & M. L. Pescador 1994. Los Ephemeroptera en Argentina. - Fauna de Agua Dulce de la
Republica Argentina 33(1): 142 pp.

Hofmann, C., M. Sartori & A. Thomas 1999. Les Ephemeropteres (Ephemeroptera) de la Guadeloupe (petites
Antilles frangaises). - Mem. Soc. Vaudoise Sci. Nat. 20(1):1-96

Hubbard, M. D. 1982. Catälogo abbreviado de Ephemeroptera da America do Sul. - Pap. Avuls. Zool. 34: 257-
282

Kluge, N. J. 1992. Redescription of Leptohyphes eximius Eaton and diagnoses of the genera Leptohyphes and
Tricorythodes based on the structure of pterothorax (Ephemeroptera: Tricorythidae, Leptohyphinae). —
Opusc. Zool. flluminensia 98: 1-16

Koss, R. W. 1968. Morphology and taxonomic use of Ephemeroptera eggs. - Ann. Entomol. Soc. Am., 61: 696-
721

-- &G.F. Edmunds 1974. Ephemeroptera eggs and their contribution to phylogenetic studies of the order. —
Zool. J. Linn. Soc. 55: 267-349

Lestage, J. A. 1931. Contribution a l’&tude des Ephemeropteres. VIII. Les Ephemeropteres du Chili. - Bull. Ann.
Soc. Entomol. Belg. 71: 41-60

Molineri, C. 2001. A new genus of Leptohyphidae (Insecta: Ephemeroptera). — In: Trends in Research in
Ephemeroptera and Plecoptera (E. Dominguez Ed.), Kluwer Academic/Plenum Publishers, New York.
Pages: 337-345.

Navas, L. 1931. Insectos de la Argentina (7a serie). - Rev. Soc. Entomol. Argentina 3: 317-324

Needham J. G. & H. E. Murphy 1924. Neotropical Mayflies. - Bull. Lloyd Library Botany, Pharmacy Materia
Medica, No. 24, Entomol. Ser. No. 4:1-79

Traver, J. R. 1958. The Subfamily Leptohyphinae (Ephemeroptera: Tricorythidae). - Ann. Entomol. Soc. America
51(5): 491-503

140

SPIXIANA 141-146 München, 01. Juli 2001 ISSN 0341-8391

The larva of Hydropsyche urgorrii Gonzälez & Malicky, 1980

(Insecta, Trichoptera, Hydropsychidae)
Rufino Vieira-Lanero, Marcos A. Gonzälez & Fernando Cobo

Vieira-Lanero, R., M. A. Gonzälez & F. Cobo (2001): The larva of Hydropsyche
urgorrii Gonzälez & Malicky, 1980 (Insecta, Trichoptera, Hydropsychidae). - Spix-
iana 24/2: 141-146

The larva of the hitherto unknown Iberian endemic species H. urgorrii Gonzälez
& Malicky, 1980 is described for the first time and compared with other known
similar Iberian species. The most important features are illustrated and some
zoogeographical and ecological notes are included.

Rufino Vieira-Lanero, Marcos A. Gonzälez & Fernando Cobo, Departamento de
Biologia Animal, Facultad de Biologia, Universidad de Santiago, 15706 Santiago de
Compostela, Spain.

Introduction

According to Gonzälez et al. (1992) the genus Hydropsyche is represented in the Iberian Peninsula by
17 species. Subsequently, Botosaneanu (1999) described H. lagranja Botosaneanu, 1999, from Segovia
(Centre of Spain) and Gonzälez & Malicky (1999) described H. iberomaroccana (previously cited as H. cf.
punica: see Gonzälez et al. 1992) from several localities of southern Spain. Moreover, Malicky (1999)
carried out a review of the European species of the pellucidula-group (with a wide representation of
specimens from Spain) concluding that the Iberian specimens traditionally identified as H. pellucidula
(Curtis, 1834) belongs, in all probability, to H. incognita Pitsch, 1993.

Therefore, 18 Hydropsyche species are known at this moment from the Iberian Peninsula: H. acinoxas
Malicky, 1981; H. ambigua Schmid, 1952; H. brevis Mosely, 1930; H. bulbifera McLachlan, 1878;
H. contubernalis McLachlan, 1865; H. dinarica Marinkövic, 1979; H. exocellata Dufour, 1841; H. iberoma-
roccana Gonzälez & Malicky, 1999; H. incognita Pitsch, 1993; H. infernalis Schmid, 1952; H. instabilis
(Curtis, 1834); H. lagranja Botosaneanu, 1999; H. lobata McLachlan, 1884; H. modesta Navas, 1925;
H. pictetorum Botosaneanu & Schmid, 1973; H. siltalai Döhler, 1963; H. teruela Malicky, 1980; H. tibialis
McLachlan, 1884; and H. urgorrii Gonzalez & Malicky, 1980.

In the last years, several papers have been published about the larval taxonomy of the European
species of this genus (see discussion), and some of these were exclusively devoted to the larvae of some
Iberian species (Millet 1983, Garcia de Jalön 1981, 1983, Zamora-Munoz et al. 1995). So, at this moment
it is possible to identify almost all the larvae of the Iberian Hydropsyche species, and only the larvae of
three Iberian species are unknown: H. acinoxas, H. lagranja and H. urgorrii. These three species are
endemic and belong to the pellucidula-group. The last instar larva of H. urgorrii is described in this paper
for the first time.

141

Material examined and methods

A great number of specimens (346 larvae) of H. urgorrii have been collected from several sample sites
of Galicia (NW Spain). For the specific determination of the aquatic stages, 6 larval exuviae collected
from mature pupae with distinct genitalia were examined thereby ensuring the association between
larval and adult specimens.

We have adopted in this paper the terminology of the larval characters used by Bournaud et al.
(1982), Boon (1978) and De Pietro (1999).

Description of the final instar larva

Mean body length 17 mm (range 16-20 mm, N=22).

Head capsule (Figs 1-3). Almost square in dorsal view, slightly wider at eye level. Mean head
length 2.1 mm (range 1.9-2.4 mm, N =22); mean head width 1.3 mm (range 1.6-2.1 mm, N =22).

The colour pattern (Fig. 1) is quite distinctive: ground colour of the head dorsum dark brown, with
light spots on the frontoclypeal apotome, and a posterior, light oval area on each parietal and around
the eyes. Lateral view with a light, longitudinal band, as in Fig. 3. Head pale brown in ventral view
(Fig. 2), with a dark brown stridulatory band on each gena reaching the oral margin of the head. Muscle
attachment spots ill-defined. Frontoclypeal apotome (Figs 1, 12) roughly pentagonal in shape. Anterior
edge straight, slightly crenulate and similar in width to the posterior third. Lateral margins slightly
narrower, posterior margins concave and the posterior tip pointed. Epistomal sulcus ill-defined but
tentorial pits distinct; often with four muscle attachment spots on the surface, ahead of tentorial pits,
and over 6 lighter spots on the posterior, rounded area. There is a characteristic light, rounded area,
near the posterior tip (the “median portion aboral light spot”), and two light bands just under epistomal
sulcus (the “lateral light spots”); sometimes two ill-contrasted lateral areas can also be distinguished
on the epistomal sulcus (the “lateral portion aboral light spot”). A light transverse ill-defined oral area
(more noticeable when the apotome is isolated) joins the bands below epistomal sulcus in some
specimens (the “oral light spot”). Lateral arms of submentum (Fig. 13) long, narrow, enlarged slightly
at apex. Anterior margin black brown; posterior apex lighter.

Oral pieces (Figs 8, 10, 11). Labrum (Fig. 10) elliptical in dorsal view; the dark colour of the anterior
half extends to the posterior edge as a narrow central band. Labrum surface covered with numerous
short setae, with small bristles between them and with well-developed lateral brushes. Six primary
setae can be recognised. Mandibles (Fig. 8) roughly triangular, dark brown in colour. Left mandible
with an apical and subapical tooth on dorsal blade, ventral blade with an apical and four subapical
teeth, the basal subapical tooth short, rounded triangular, with a brush of long, spiny setae in the
concavity, above basal tooth. Right mandible with a single tooth on dorsal blade and a obliquely row
of small setae dorsally, on the tooth; ventral blade with an apical and three subapical teeth; the basal
subapical tooth very short and rounded triangular. Laterally, on each mandible, there is a deep ridge
with 18-24 hyaline setae and some shorter setae inserted between them. Mentum as in Fig. 11; chestnut
brown in colour, setae grouped on each apical lobe and in an oblique line on each posterolateral corner;
the anterior margin of each apical lobe is rounded and the median channel is straight-sided.

Thorax (Figs 4-7). Pro-, meso- and metanotum (Figs 4-6) lighter than head. Meso- and metanotum
lighter than pronotum. Sclerites with a dense covering of short and strong setae (“scale-like” setae) and
even shorter pointed setae between them. Anterior margin with numerous fine setae. Pronotum with
a rather wide dark dorsal longitudinal band. Each hemipronotum (Fig. 4) subquadrangular in shape.
Meso- and metanotum of uniform colour. The medial regions of the posterior prosternites (Fig. 7) are
strongly pigmented and irregularly squarish; lateral regions lighter in colour, less distinct than the
medial regions and fused with them. Protrochantin (Fig. 9) with numerous setae along ventral portion,
dorsal portion downturned apically.

Abdomen. Abdominal segments I-VII with ventral gills.

142

Figs 1-7. H. urgorrii (last instar larva). 1. Head capsule, dorsal view. 2. Head capsule, ventral view. 3. Head
capsule, lateral view. 4. Hemipronotum, left half. 5. Mesonotum, left half. 6. Metanotum, left half. 7. Proster-
nites.

143

Discussion

We can use several larval keys for the specific identification of the European species of the genus
Hydropsyche (e.g. Lepneva 1964, Hickin 1967, Sedlak 1971, Hildrew & Morgan 1974, Szezesny 1974,
Statzner 1976, Verneaux & Faessel 1976, Boon 1978, Wiberg-Larsen 1980, Edington & Hildrew 1981,
Bournaud et al. 1982, Moretti 1983, Bongard 1990, Pitsch 1993, Waringer & Graf 1997, De Pietro 1999),
and for the North African species (e.g. Dakki & Tachet 1987). However, some of them may be only
partially valid and serious mistakes might result from using these keys in the Iberian Peninsula.

Regarding the Iberian species of Hydropsyche, the larval keys of Garcia de Jalön (1981) and Millet
(1983) include only some of the Iberian species but an updated key has been recently reported by
Zamora-Munoz et al. (1995), including the first larval description of five endemic species. Thus it is
possible to use this key as a starting point for this work. When using the key of Zamora-Munoz et al.
(1995), larvae of H. urgorrii will key out under couplet 15: the characteristics of the frontoclypeal
apotome disagree with those of H. dinarica and H. lobata. The width of anterior edge of the frontoclypeal
apotome of H. dinarica is roughly similar to the width of the posterior third (cf. Garcia de Jalön 1983,
Klima 1989, Waringer & Graf 1997, De Pietro 1999); the posterior margins are not concave, the
epistomal sulcus is not distinct and the lateral light spots under it are rounded, and some specimens
have an ill-defined median portion aboral light spot similar to that of H. urgorrii but fainter. Moreover,
larval size of H. dinarica is greater than that of H. urgorrii.

The frontoclypeal apotome of H. lobata is very much alike to that of H. urgorrii but the former has
a rounded posterior tip and a distinct curved edge - very conspicuous in lateral view - across the
cibarian muscle attachment spots (cf. Garcia de Jalön 1983) that is lacking in H. urgorrii. Moreover, the
lateral portion aboral light spots of H. lobata are ill-defined and the colour of the posterior prosternites
is also different (cf. fig. 2b in Dakki & Tachet 1987): the lateral regions are lighter than the medial ones
in H. urgorrii but concolorous with them in H. lobata.

The frontoclypeal apotome of some specimens of H. urgorrii is very similar in colour to that of
H. brevis (see fig. 3 in Zamora-Munoz et al. 1995), but the posterior tip is not rounded and the lateral
margins are narrower in H. urgorrii; moreover, the coloration of the ventral part ofthe head and thoraeic
sclerites is also different.

Among the eight Hydropsyche species with an overlapped distribution area with H. urgorrii
(H. ambigua, H. dinarica, H. exocellata, H. instabilis, H. lobata, H. incognita, H. siltalai and H. tibialis) only
the frontoclypeal apotome of H. ambigua, H. dinarica and H. lobata can be confused with that of
H. urgorrii. We have already discussed the differences regarding H. dinarica and H. lobata; the larva of
H. ambigua has been described by Zamora-Munoz et al. (1995) and can be clearly distinguished from
that of H. urgorrii by the shape of the frontoclypeal apotome (the lateral margins are more or less
straight up to the epistomal sulcus, the posterior margins are roughly convexe and the posterior third
is similar in width, or even wider, than the anterior margin), the colour pattern of this sclerite (with two
rounded lateral light spots only and with a central, ill-defined, central spot, above them) and the
triangular shape of the submentum (the lateral arms are short and broad).

Habitat and distribution

H. urgorrii is an endemic species of the Iberian Peninsula, where it is confined to the north-western
quarter. In this area, some adults of this species have been recorded from many localities of Galicia and
the north and centre of Portugal (Gonzälez 1988, Gonzalez et al. 1992, Terra 1994) from 15 to 500 m a.s.l.
The typical habitat of this species consists of small and medium waterbeds where the larvae were found
in rapidly flowing stream areas with pebbles and small rocks. In Galicia, H. urgorrii larvae in instars
IV were present from late October to March and adults were recorded from late March to May.

Acknowledgements

We wish to thank Dr. J. Waringer who kindly provided us with specimens of H. incognita from central Europe.
This research was supported by XUGA 20005B98 of Xunta de Galicia.

144

Figs 8-13. H. urgorrii (last instar larva). 8. Mandibles, dorsal view. 9. Right protrochantin, external view.
10. Labrum, dorsal view. 11. Mentum. 12. Frontoclypeal apotome, dorsal view. 13. Submentum.

References

Bongard, T. 1990. Key to the Fennoscadian larvae of Arctopsychidae and Hydropsychidae (Trichoptera). - Fauna
Norv. Ser. B, 37: 91-100

Boon, P. ]. 1978. The use of ventral sclerites in the taxonomy of larval hydropsychids. - In Crichton, M. I. (Ed.).
Proc. 2nd. Int. Symp. Trichoptera. Reading, 1977: 165-173. - Junk. The Hague

Botosaneanu, L. 1999. Notes sur quelques Trichopteres espagnols ou marocains des collections du Museum
national d’Histoire naturelle, Paris (Trichoptera). — Bull. Soc. Ent. France 104(2): 113-116

145

Bournaud, M., H. Tachet & ]. F. Perrin 1982. Les Hydropsychidae (Trichoptera) du Haut-Rhöne entre Geneve
et Lyon. - Annls. Limnol. 18: 61-80

Dakki, M. & H. Tachet 1987. Les larves d’Hydropsyche du Maroc (Trichopteres, Hydropsychidae). - In Bournaud,
M. & Tachet, H. (Eds.). Proc. 5th. Int. Symp. Trichoptera. Lyon, 1986. - Junk. The Hague. Ser. Ent. 39: 25-28

De Pietro, R. 1999. Identification of the larvae of Hydropsyche species from Sicily and peninsular Italy (Tricho-
ptera, Hydropsychidae). - Arch. Hydrobiol. 121/2: 91-117

Edington, J. M. & A. G. Hildrew 1981. A key to the caseless caddis larvae of the British Isles. — Scient. Publs.
Freshw. Biol. Ass. 43: 1-91

Garcia De Jalön, D. 1981. Description of Hydropsyche larvae found in the Iberian Peninsula. — In Moretti, G. P.
(Ed.). Proc. 3rd. Int. Symp. Trichoptera. Perugia, 1980. - Junk. The Hague. Ser. Ent. 20: 87-92

-- 1983. Contribuciön al conocimiento de las larvas del genero Hydropsyche (Trich.) ibericas. - Actas 1° Congr.
Iberico Entomol. Leön: 275-285

Gonzälez, M. A. 1988. Inventario dos Tricöpteros de Galicia (Insecta: Trichoptera). - Cad. Area Cienc. Biol.
(Inventarios), Sem. Est. Gall., II Ocastro-Sada, A Coruna: Ed. do Castro. 45 pp.

-—- & Malicky, H. 1999. Une nouvelle espece de Hydropsyche du groupe pellucidula (Trichoptera, Hydropsychi-
dae). -— Braueria 26: 25-26

-- ,L.S. W. Terra, D. Garcia De Jalön & F. Cobo 1992. Lista faunistica y bibliogräfica de los tricöpteros
(Trichoptera) de la Peninsula Iberica e Islas Baleares. - Asoc. esp. Limnol. (Ed.), Publ. No. 11, 200 pp.

Hickin, N. E. 1967. Caddis larvae. Larvae of the British Trichoptera. - Hutchinson. London, 466 pp.

Hildrew, A. & J. C. Morgan 1974. The taxonomy of the British Hydropsychidae (Trichoptera). — J. Ent. 43: 217-
229

Klima, F. 1989. Hydropsyche dinarica Marinkovik, 1979 (Insecta, Trichoptera) aus dem Rhithral des westlichen
Thüringer Waldes - neu für die Fauna der DDR. - Veröff. Naturhist. Mus. Schleusingen 4: 90-92

Lepneva, S. G. 1964. Fauna of the U.S.S.R. Trichoptera 1, Larvae and pupae of Annulipalpia. - Jerusalem. Israel
program for Scientific Translations (1971). 700 pp.

Malicky, H. 1999. Bemerkungen über die Verwandtschaft von Hydropsyche pullucidula Curtis (Trichoptera,
Hydropsychidade). - Linzer biol. Beitr. 31/2: 803-821

Millet, X. 1983. Les larves del genere Hydropsyche (Insecta: Trichoptera) ä Catalunya. Taxonomia. — Butll. Inst.
Cat. Hist. Nat. 49 (Sec. Zool., 5): 97-103

Moretti, G. P. 1983. Guide per il riconoscimento delle specie animali delle acque interne Italiane. 19. Tricotteri
(Trichoptera). - Cons. Naz. Ricerche AQ 11 (196), Verona, 155 pp.

Pitsch, T. 1993. Zur Larvaltaxonomie, Faunistik und Ökologie mitteleuropäischer Fließwasser-Köcherfliegen
(Insecta: Trichoptera). - Landschaftsentwicklung und Umweltforschung - Schriftenr. Fachber. Landschafts-
entwicklung-Sonderheft S 8. Technische Universität Berlin, Berlin. 316 pp.

Sedläk, E. 1971. Bestimmungstabelle der Larven der häufigen tschechologakisten Arten der Gattung Hydropsyche
Pictet (Trichoptera). - Acta Ent. Bohem. 68: 185-187

Statzner, B. 1976. Zur Unterscheidung der Larven und Puppen der Köcherfliegen-Arten Hydropsyche angustipen-
nis und pellucidula (Trichoptera: Hydropsychidae). - Ent. Germ. 3(3): 265-268

Szczesny, B. 1974. Larvae of the genus Hydropsyche (Insecta: Trichoptera) from Poland. - Pol. Arch. Hydrobiol.
21(3/4): 387-390

Terra, L. S. W. 1994. Atlas provisörio dos Tricöpteros (Insecta. Trichoptera) de Portugal Continental. Instituto
Forestal (Ed.). - Publ. No. 306, 100 pp.

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donnees biologiques et Ecologiques. — Annls. Limnol. 12(1): 7-16

Waringer, J. & W. Graf 1997. Atlas der österreichischen Köcherfliegenlarven: unter Einschluss der angrenzenden
Gebiete. - Wien. Facultas-Univ.-Verl., 286 pp.

Wiberg-Larsen, P. 1980. Bestemmelsesnogle til larver af danske arter af familien Hydropsychidae (Trichoptera)
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Zamora-Munoz, C., J. Alba-Tecedor & D. Garcia De Jalön 1995. The larvae of the genus Hydrosyche (Hydropsych-
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68: 189-210

146

SPIXIANA 2 147-155 München, 01. Juli 2001 ISSN 0341-8391

Tetrigidae from Nepal in the Zoologische Staatssammlung München

(Insecta, Orthoptera, Tetrigidae)
Sigfrid Ingrisch

Ingrisch, S. (2001): Tetrigidae from Nepal in the Zoologische Staatssammlung
München (Insecta, Orthoptera, Tetrigidae). - Spixiana 24/2: 147-155

A list of 20 species of Tetrigidae from Nepal mainly collected during the expe-
ditions of Dr. Dierl, Dr. Forster, and Dr. Schacht in 1964,1967, and 1973 is given
together with some own material. Two species are described as new: Coptotettix
muglingi, spec. nov. and Ergatettix minutus, spec. nov. The material of the Dierl-
expeditions and the types of the new species are deposited in the Zoologische
Staatssammlung München.

Dr. Sigfrid Ingrisch, Eichendorffweg 4, D-34385 Bad Karlshafen, Germany;
e-mail: sigfrid.ingrisch@planet-interkom.de

Introduction

Some years ago, I studied the Orthoptera collected during the expeditions of Dr. Dierl, Dr. Forster, and
Dr. Schacht to Nepal in 1964, 1967, and 1973 (Ingrisch 1990). The Tetrigidae had to be excluded as there
were no reliable tools for identification. The Tetriginae s. str. that represent the majority of the
specimens at hand are not included in the works of Günther (1938a, b, 1939). Some authors even
thought it impossible to identify Tetrigidae of several genera with accuracy (Kevan 1966, see also
Blackith 1988). The taxonomic situation of the Tetrigidae of the Oriental Region is still in a desolate
state. Numerous of the classical taxa described by Bolıvar (1887), Walker (1871), or Hancock (1904, 1907,
1909, 1910, 1912, 1913, 1915) have never been re-examined and from the descriptions alone it is often
impossible to be certain about their identity. A taxonomic revision of the Oriental Tetrigidae based on
multivariate analysis of numerous characters that was projected by Blackith & Blackith (1987) did not
appear. But meanwhile comprehensive works on the Tetrigidae of North India (Shishodia 1991) and
China (Liang & Zheng 1998) are published, and there seems to be some agreement on the identity of
the common species.

The present paper gives a list of species of Tetrigidae collected during the Dierl-Expeditions to
Nepal. Moreover, I use the opportunity to revise some of the Tetrigidae collected during an own
excursion to Nepal (Ingrisch 1987).

The Tetrigidae of the Dierl-Expeditions were collected at light and are all long-winged species.
They are often widespread in North India but several of the species were not yet reported from Nepal.
Information on the collection sites can be found in Dierl (1966). The own collection contained two
undescribed species. All specimens of the Dierl-Expeditions and the types of the new species are
deposited in the Zoologische Staatssammlung München (ZSM); few other specimens are in my own
collection (CI).

147

List of species

Distribution according to Blackith (1992) with supplements by Shishodia (1991), type localities accord-
ing to Blackith (1992) and Otte (1997). Measurements as described in Ingrisch (in press).

Scelimeninae

Eucriotettix annandalei (Hancock, 1915)

Distribution: West Bengal.

Type locality: India: West Bengal, Paresnath, 1100 m.

Locality: 13, Province Bagmati, Kathmandu Valley, Godavari, 1600-1800, 8.V1.1967, Dierl-Forster-
Schacht (ZSM).

Loxilobus assamus Hancock, 1907

Distribution: North East India. Already recorded for Nepal by Chopard & Dreux (1966).

Type locality: India: Assam, Khasi Hills, Cherrapunji.

Locality: 13, 12, Province Sagarmatha, Jubing, 1600 m, 5.V.1964, Dierl-Forster-Schacht. Both speci-
mens are macropronotal forms (ZSM).

Hebarditettix quadratus (Hancock, 1915)

Distribution: India (Darjeeling, Sikkim, Arunachal Pradesh).

Type locality: India: Singla, Darjeeling, 450 m.

Locality: 1?, Syaklung (Sarka Bhanjyang), 700-1000 m, river bed, 23.X.1983, S. Ingrisch (CI), reported
as Systolederus greeni Bolivar, 1892 in Ingrisch (1987).

Discussion. Shishodia (1991) considers that H. quadratus might be a synonym of H. lobatus (Hancock,
1912), as both taxa differ only in the length of the pronotum.

Metrodorinae

Bolivaritettix dubius (Hancock, 1912) [= syn. of Bolivaritettix javanicus (Bolivar, 1909) sensu Blackith 1992]
Distribution: North East India.

Type localities: of Mazarredia dubia Hancock, 1912 = India: Bengal, Lebong; of Mazarredia javanica
Bolivar, 1909 = Java.

Locality: 13, Province Bagmati, Kathmandu Valley, Godavari, 1600-1800, 31.VIl.1967, Dierl-Forster-
Schacht (ZSM); 1°, Kapurgaon - Baglungpani, 1550-2000 m, shrub rich slopes, 21.X.1983, S. Ingrisch
(CI), the latter specimen reported as Bolivaritettix lativertex (Brunner v. W. 1893) in Ingrisch (1987).
Discussion. Bolivaritettix dubius (Hancock, 1912) is regarded to be a doubtful synonym of Bolivaritettix
javanicus (Bolivar, 1909) by Günther (1939), listed as a synonym of the latter species in Blackith (1992)
and Otte (1997), and treated as a separate species in Steinmann (1970) and Shishodia (1991). But so far
nobody compared the types of both taxa.

Tetriginae

Teredorus carmichaeli Hancock, 1915

Distribution: North East India. Already recorded for Nepal by Bei-Bienko (1968).

Type locality: India: Darjeeling Dist., Singla.

Locality: 19, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 9.V.1967, Dierl-Forster-
Schacht (ZSM).

Discussion. Günther (1939) restricts the genus Teredorus Hancock, 1906 (Type species: Teredorus sten-
ofrons Hancock, 1906 from Peru) to American species and includes the Asian species described under
Teredorus in Systolederus Bolivar, 1887, while recent authors follow Hancock (1915) and include also
Asian species in Teredorus (Shishodia 1991, Blackith 1992, Otte 1997, Liang & Zheng 1998). However the
taxonomic relations of both genera are still unresolved.

148

Teredorus frontalis Hancock, 1915

Distribution: North East India.

Type locality: Himalaya: Dharampur, Simla Hills, 1200 m.

Localities: 13, Syaklung (Sarka Bhanjyang), 700-1000 m, river bed, 23.X.1983, S. Ingrisch; 1°, Trisuli
between Fishling - Mugling Bazar, river bed, 26.X.1983, S. Ingrisch (CI), both specimens reported as
Systolederus gravelyi Günther, 1939 in Ingrisch (1987).

Hedotettix attenuatus Hancock, 1904

Distribution: India, Sri Lanka.

Type locality: Sri Lanka: Kesbewa, Colombo.

Localities: 1733, 1692, Province Narayani, Rapti Valley, Jhavani, 200 m, 15.-18.V.1967, Dierl-Forster-
Schacht; 14, 1?, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 10.-11.V.1967, Dierl-
Forster-Schacht; 13, Province Bagmati, Kathmandu — Chauni, 1400 m, 16.1V.1967, Dierl-Forster-
Schacht (all ZSM).

Hedotettix costatus Hancock, 1912

Distribution: India, Bangladesh, Nepal, Sulawesi.

Type locality: India: Bengal, Pusa.

Localities: 694, 92%, Province Narayani, Rapti Valley, Jhavani, 200 m, 15.-18.V.1967, Dierl-Forster-
Schacht; 584, 429, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 10.-11.V.1967,
Dierl-Forster-Schacht (all ZSM).

Hedotettix spec. (near H. attenuatus Hancock, 1904 and H. costatus Hancock, 1912)

Localities: 17, east of Pokhara, 700 m, wasteland within rice fields, 13.-14.X.1983, S. Ingrisch; 24, north
of Ghanpokhara, 2000-2300 m, meadows, 20.X.1983; 13, south of Ghanpokhara, 2000 m, meadows,
20.X.1983, S. Ingrisch; 1?, Ghanpokhara — Kapurgaon, 2000 m, shrub rich cultural land, 21.X.1983,
S. Ingrisch; 334, 52, Baglungpani, 1550 m, meadows, 21.X.1983, S. Ingrisch (all CI).

Discussion. A Hedotettix species lives on subalpine meadows in the Annapurna area of western Nepal
that cannot be assigned with certainty to any of the named species. It was reported as H. costatus in
Ingrisch (1987), but it differs from specimens of H. costatus collected in the Rapti Valley as well as from
H. attenuatus to which it is also similar. It occurs in the macroponotal and brachypronotal morph as well
as in intermediate forms. Additional material should be evaluated before the status of those popula-
tions can be settled with certainty.

Hedotettix gracilis (de Haan, 1843)

Distribution: Sulawesi, Sunda, Java, Sumatra, Bangladesh, India, Myanmar, Sri Lanka, Taiwan, Thai-
land, Vietnam.

Type locality: Java.

Localities: 19, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 10.V.1967, Dierl-
Forster-Schacht; 17, Province Narayani, Rapti Valley, Jhavani, 200 m, 15.V.1967, Dierl-Forster-Schacht
(all ZSM).

Hedotettix grossus Hancock, 1915

Distribution: North East India.

Type locality: Himalaya: Singla, Darjeeling.

Locality: 17, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 10.V.1967, Dierl-Forster-
Schacht (ZSM).

Coptotettix conspersus Hancock, 1915

Distribution: North East India, Sri Lanka.

Type locality: India: Bengal, Siliguri.

Localities: 13, 22%, Province Narayani, Rapti Valley, Jhavani, 200 m, 15.V. 1967, Dierl-Forster-Schacht;
13, 42%, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 6.-11.V.1967, Dierl-Forster-
Schacht; 1?, Province Bagmati, Kathmandu Valley, Godavari, 1600-1800, 31.VII.1967, Dierl-Forster-
Schacht (all ZSM).

149

Coptotettix annandalei Hancock, 1915

Distribution: North East India, North Myanmar, Nepal.

Type locality: India: Darjeeling, Singla.

Localities: 14, east of Pokhara, 700 m, river bed and wasteland within rice fields, 13.-14.X.1983,
S. Ingrisch; 1?, Trisuli, Jugedee, river bed and cultural land, 27.-28.X.1983, S. Ingrisch; 13, Terai:
Chitawan, Gaida-Camp, river bed, savannah, evergreen forest, 28.-29.X.1983, S. Ingrisch (all CI).
Discussion. Recorded already under the same name in Ingrisch (1987), the specimens from Mugling
Bazar however represent a new species (see below). It is possible that this taxon is a mixture of
morphologically similar, sibling species as already Hancock (1915) in the original diagnosis and later
Shishodia (1991) describe some geographic variation. If so, the specimens at hand might represent an
undescribed species, but the material is not enough to solve the problem. The specimens at hand agree
with the descriptions in Hancock (1915) and Shishodia (1991) except for the following points: The
vertex is not even slightly narrowing anteriorly and is 1.4 x wider than one eye in the female and of
equal width with one eye in the males (1.0-1.1 x as wide as one eye) instead of narrower than one eye.
Since no exact values are given in the previous descriptions the relative width of both organs was
probably guessed not measured. The anterior margin of the pronotum is truncate in the male from
Chitawan which agrees with the original diagnosis, while it is a little convex in the male from Pokhara
and in the female. The pronotum bears tubercles between the shoulders which are not very striking in
the male from Chitawan and in the female, but rather striking in the male from Pokhara. The frontal
costa differs somewhat between the three specimens at hand: It is widening ventrad but rather wide
throughout in the male from Chitawan, widening ventrad but rather narrow except at the medial
ocellus in the male from Pokhara, and gradually widening ventrad in the female.

Coptotettix muglingi, spec. nov.
Figs 1-6

Coptotettix annandalei (nec Hancock, 1915) Ingrisch, 1987 partim.

Types. Holotype: @, Nepal: Trisuli river near Mugling Bazar, river bed, 26.X.1983, S. Ingrisch (ZSM). — Para-
types: 2% %, Trisuli river near Mugling Bazar, river bed, 26.X.1983, S. Ingrisch (1°, ZSM; 1?, C]).

Description

Small brachynotal species with small lanceolate tegmina and strongly reduced wings. Integument
granular and slightly tuberculate.

Head with eyes not (female) or little excerted (male). Vertex 0.9 x (female) or 0.8 x (male) as wide
as one eye; anterior margin truncate, not completely reaching anterior margin of eyes; medial carina
distinct, running to end of fossulae; fossulae deep, elongate; lateral carinae raised to dorsal margin of
eyes. Frontal costa in lateral view projecting in front of eyes, rounded together with vertex but slightly
concave above lateral ocelli; in anterior view widened about halfway between dorsal angle and lateral
ocelli, moderately wide dorsal, suddenly more widened between lateral ocelli and antennae, and then
gradually widened ventrad; between antennae 0.8-0.9 x as wide as scapus. Antennae inserted between
ventral margins of eyes such that the middle of the antennal scrobae and the base of the scapus are
placed between the ventral margins of eyes. Pronotum not completely covering abdomen, reaching
about middle of postfemur; surface granular and a little tuberculate; anterior margin truncate or very
faintly convex; posterior margin broad obtuse-angularly rounded; prozonal carinae sharp, almost
parallel (or very little converging posteriorly); medial carina low, a little convex from anterior margin
to behind sulci, substraight (female) or faintly undulate (male) thereafter; internal lateral carinae
absent, lateral margins of disc of pronotal process formed by the external lateral carinae. Paranota with
ventral margin slightly curved laterad; ventral projection distinct with apex roundly truncate; dorsal
projection in male present, its shape short triangularly rounded, in female reduced to a weak convexity;
ventral margin of pronotal process convexly expanded behind tegmen (more expressed in the female
than in the male). Tegmen with free part small, lanceolate, 3.1-4.0 x longer than wide in female, 2.5-
3.6 x longer than wide in male (varying between left and right tegmen of holotype); hind wing
projecting 0.3 mm behind tegmen in male; in female hidden or absent. Femur I and II setose; femur I
stout with dorsal margin convex and indistinctly undulate; femur II compressed and widened (in

150

Figs 1-6. Coptotettix muglingi, spec. nov.: 1-3. Male, holotype. 4-6. Female, paratype. 1,5. Lateral view.
2,6. Dorsal view. 3, 4. Frons.

female 2.7-2.8 x, in male 2.4 x longer than wide), dorsal and ventral margins faintly (in male indistinct-
ly) undulate. Postfemur very thick (2.2 x longer than wide), granular and rugose; dorsal margin
serrulate. Hind tarsus with first segment 1.6-1.7 x (female) or 1.9 x (male) longer than third segment;
posterior metatarsus with pulvilli spinose, variable in length: third pulvillus the longest one, second
pulvillus longer than or of subequal length with first pulvillus. Ovipositor with dorsal valve 2.3-2.6 x
longer than wide.

Measurements (in mm): body length 4 6.89, ? 7.87-7.93; vertex width 3 0.44, ? 0.53; eye width
20.55, ? 0.58-0.59; frontal costa between antenna 4 0.22, ? 0.22-0.24; frontal costa at ventral end 3 0.28,
2 0.31; scapus width 4 0.25, ? 0.27; antennal length 4 3.75, ? 3.75-3.94; pronotum length 4 4.62, ? 5.17-
5.27; pronotum shoulder width 4 2.24, 9 2.6; pronotum lobe width 4 3.15, ? 3.38-3.45; tegmen length
8.0.63, ? 0.78-0.88; femur I length 3 1.63, ? 1.76-1.82; femur Il length 4 1.79, 2 1.95-2.02; femur II width
3.0.75, ? 0.72; postfemur length 4 4.81, ? 5.27; postfemur width 4 2.18, ? 2.37-2.41; hind tarsus I length

151

3.0.89, 2 0.94-0.97; hind tarsus III length 4 0.47, 2 0.56-0.59; dorsal ovipositor valve length 0.88; dorsal
ovipositor valve width 0.34-0.38.

Discussion. The new species differs from most species described under Coptotettix by the strongly
reduced tegmina and wings and the short pronotum reaching only the middle of the postfemur. It is
similar to Paratettix hancockus (Shishodia & Varshney 1987) [= replacement name for Coptotettix parvulus
Hancock, 1912)]. The new species differs from the original description of C. parvulus by the frontal costa
of the head which is distinctly expanded between the lateral ocelli and the antennae (not evenly
widened forward), and the antennae are inserted between the lower margins of the eyes (not between
the lower part of the eyes). From the redescription of P. hancockus in Shishodia (1991), the new species
differs by the vertex which in strict dorsal view does not reach the anterior margin of the eyes (not
reaching front margin), the apex of the pronotum is distinctly obtuse angular (not broadly rounded),
the paranota have two projections (not one projection).

Paratettix hirsutus Brunner v. W., 1893

Distribution: Assam, Myanmar.

Type locality: Burma: Bhamo.

Locality: 13, 22°, Province Narayani, Rapti Valley, Jhavani, 200 m, 15.V. 1967, Dierl-Forster-Schacht
(ZSM).

Discussion. The specimen at hand are smaller than the measurements given in Brunner v. W. (1893)
and Shishodia (1991).

Euparatettix corpulentus Hancock, 1912 [= syn. of Euparatettix variabilis (Bolivar, 1887) ?]
Type locality: of E. corpulentus: India: Bengal, Chapra; of E. variabilis: Philippines.
Localities: 233, 292, Province Bagmati, Kathmandu - Chauni, 1400 m 23.-29.V1.1967, Dierl-Forster-
Schacht; 229, Province Narayani, Rapti Valley, Jhavani, 200 m, 15.V. 1967, Dierl-Forster-Schacht; 1°,
Province Bagmati, Kathmandu Valley, Godavari, 1600-1800, 8.V1.1967, Dierl-Forster-Schacht (all ZSM).
Discussion. This and the following species are currently listed under the synonymy of Euparatettix
variabilis (Bolivar, 1887) (Naskrecki & Otte 1997), a species covering an area from India to the Solomon
Islands. As in my opinion the taxonomic status of the numerous synonyms of E. varlabılis (see
catalogues of Blackith 1992 and Otte 1997) is still unresolved, I prefer for the moment to use Hancock’s
(1912) names for this and the following species, the more as the specimens at hand that should belong
to E. variabilis according to the current synonymy obviously belong to two different species.
Günther (1938b) treats Euparatettix variabilis (Bolivar, 1887) as a synonym of Pseudoparatettix histri-
cus (Stäl, 1861). Shishodia (1991) lists Euparatettix corpulentus Hancock, 1912 under the synonymy of
Euparatettix histricus (Stäl, 1861), Euparatettix tenuis Hancock, 1912 as a separate species. Naskrecki &
Otte (1997) list both E. corpulentus and E. tenuis under the synonymy of E. variabilis (Bolivar, 1887), and
Pseudoparatettix histricus (Stäl, 1861) as a separate species. Blackith (1992) lists E. variabilis twice, as a
separate species and under the synonymy of Pseudoparatettix histricus (Stäl, 1861).

Euparatettix tenuis Hancock, 1912 [= syn. of Euparatettix variabilis (Bolivar, 1887) ?]

Type locality: of E. tenuis: India: Bengal, Pusa; of E. variabilis: Philippines.

Locality: 3? 2, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 9.-12.V.1967, Dierl-
Forster-Schacht (ZSM).

Discussion. See under E. corpulentus.

Ergatettix dorsiferus (Walker, 1871)

Distribution: India, Sri Lanka, Bangladesh, Myanmar, Afghanistan, Iran, China, Taiwan, Central Asia,
Sumatra, Java, Sumba Island.

Type locality: India: Bombay.

Localities: 883, 829, Province Bagmati, Kathmandu - Chauni, 1400 m, 24.V1.1967, Dierl-Forster-
Schacht; 13, Province Bagmati, Kathmandu - Chauni, 1400 m, 29.1V.1967, Dierl-Forster-Schacht; 33g,
222, Province Narayani, Bhainse Dobhan, 730 m, 16.-20.V11.1967, Dierl-Forster-Schacht; 1138, 112%,

Province Narayani, Rapti Valley, Jhavani, 200 m, 15.-19.V.1967, Dierl-Forster-Schacht; 10? 2, Province

Narayani, Rapti Valley, Monahari Khola, Belwa, 350 m, 6.-12.V.1967, Dierl-Forster-Schacht (all ZSM).

152

Discussion. In the series at hand, there are all transient forms from specimens with barely rugose to
such with distinctly rugose pronotum, with or almost without lateral undulating projections in apical
area of pronotum, and with distinct or almost without rugose lateral projections on lateral surface of
postfemora; i.e. from Ergatettix guentheri Steinmann, 1970 (= replacement name for Indatettix nodulosus
(Hancock, 1915)) to Ergatettix dorsiferus (Walker, 1871). Thus the former might be only a synonym of
the latter as already pointed out by Hebard (1929); no further separation between both forms is done.
There are specimens with pale and others with annulated posttibia in the same population (see keys
in Hancock 1915 and Shishodia 1991); thus Ergatettix crassipes (Hancock, 1912), in which the posttibiae
are said to be subinornate, is probably also a synonym of Ergatettix dorsiferus (Walker, 1871) (see also
Shishodia 1991 who lists E. crassipes under the synonymy of E. dorsiferus).

The species was already recorded from Nepal as Ergatettix nodulosus Hancock, 1915 in Bei-Bienko
(1968) and as Ergatettix crassipes (Hancock, 1912) in Ingrisch (1987).

Ergatettix minutus, spec. nov.
Figs 7-10

Types. Holotype: 3, Nepal: East of Pokhara, 700 m, river bed and wasteland within rice fields, 13.-14.X.1983,
S. Ingrisch (ZSM).

Description

Small, brachynotal species with excerted head, greatly reduced tegmina and hind wings; integu-
ment granular and a little tuberculate.

Head with eyes distinctly excerted above pronotum. Vertex 0.7 x as wide as one eye; anterior
margin not completely reaching anterior margin of eyes, slightly convex on both sides of projecting
medial carina; vertex a little depressed with medial and lateral carinae almost raised to dorsal margin
of eyes; fossulae shallow. Frontal costa in lateral view very little projecting between eyes, hardly
concave above lateral ocelli, distinctly projecting between antennae; in anterior view forked about
halfway between dorsal angle and lateral ocelli, gradually widening ventrad; between antennae 0.6 x
the width of scapus. Antennae inserted between ventral margin of eyes thus that the middle of the
antennal scrobae and the scapus base lie in the same level with the ventral margin of the eyes.
Pronotum reaching about middle of postfemur; surface granular and with scattered tubercles; anterior
margin subtruncate; posterior apex broadly rounded; prozonal carinae low, parallel; medial carina
distinct but low, in lateral view faintly convex between sulci and faintly depressed between shoulders,
almost straight. Paranota with two projections, dorsal projection however reduced to a weak convexity;
ventral projection with apex faintly curved laterad, subtruncate (very little convex); ventral margin of
pronotal process concave at tegmen, convexiy expanded behind concavity. Tegmen and hind wings
greatly reduced, almost completely hidden under pronotum. Femur I strong with dorsal margin
convex and indistinctly undulate and faintly serrulate; femur II compressed and widened (2.3 x longer
than wide), with an indistinct preapical constriction, dorsal margin finely serrulate, ventral margin
granular. Postfemur thick (2.5 x longer than wide), granular, dorsal and ventral margins indistinctly
serrulate. Tibia II narrowed ventrad. Hind tarsus with first segment 1.5 x longer than third segment;
posterior metatarsus with third pulvillus longer than first and second pulvilli, all three pulvilli spinose.
Measurements of male (in mm): body length 5.59; vertex width 0.34; eye width 0.48; frontal costa
between antenna 0.16; frontal costa at ventral end 0.23; scapus width 0.25; antenna length 3.66;
pronotum length 3.84; pronotum shoulder width 1.69; pronotum lobe width 2.34; tegmen length 0.41;
hind wings projecting behind tegmen 0.38; femur I length 1.34; femur II length 1.47; femur II width 0.64;
postfemur length 4.03; postfemur width 1.59; hind tarsus I length 0.69; hind tarsus III length 0.47.

Discussion. The new species is similar to Paratettix hancockus (Shishodia & Varshney, 1987). It differs
by the distinctly excerted head, by the vertex being much narrower than one eye, by the anterior margin
of the vertex not reaching the anterior margin of the eyes, and by the paranota of pronotum having two
projections. It looks quite different from two other brachynotal Ergatettix species recently described
from Nepal (Ingrisch in press). From Ergatettix undunotus Ingrisch in press, it differs by the frontal costa
being less concave above the lateral ocelli, by the pronotum being only little rugose and its dorsal
margin almost straight (not distinctly undulate), by the apex of the pronotum being broadly rounded

153

Figs 7-10. Ergatettix minutus, spec. nov., male, holotype. 7-8. Lateral view. 9. Dorsal view. 10. Frons.

(not narrow truncate), by the expansion of the ventral margin of the pronotal process, and by the
extremely reduced tegmina and hind wings. From Ergatettix elevatus Ingrisch in press, it differs by the
head being not extremely expanded dorso-ventrally, by the pronotum with the carinae not so strongly
expressed but the surface distinctly granular (not rather smooth with tubercles) and its apex rounded
(not triangular), by the expansion of the ventral margin of the pronotal process, by the extremely
reduced tegmina and hind wings, and by the femur II being widened (not narrow, elongate).

Bannatettix menghaiensis Zheng, 1993

Distribution: South China.

Type locality: China: Yunnan.

Locality: 17, Province Narayani, Rapti Valley, Monahari Khola, Belwa, 350m, 12.V.1967, Dierl-Forster-
Schacht (ZSM).

154

Acknowledgements

The material on which this study is based was kindly made available by Dr. M. Baehr (München).

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Buchbesprechungen

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15. Dillon, R. T.: The Ecology of Freshwater Molluscs. - Cambridge Univ. Press, Cambridge, 2000. xii + 509 pp.
ISBN 0-521-35210-X.

After more than 20 years this is the first comprehensive review of the voluminous, recent literature on the
subject, i.e. worldwide freshwater molluscs. Ecology is understood in a broad sense: The book starts with two
chapters on autecology of bivalves and gastropods including feeing and digestion, habitat preferences and
reproduction. Life history and intraspecific population dynamics are covered by separate chapters as are
competition, parasitism and predation. Also the problem of neozoans (introduced species) is shortly treated,
whereas endangered species or extinction problems do not receive attention. Most interesting the author
presents a new model of life-history unifying most of the above mentioned aspects.

The reference list is extensive and cites as much as 1.200 papers, most of them published in the last 20 years.
One of the missed “classics” is Frömming'’s (1956) “Biologie der mitteleuropäi-schen Süßwasserschnecken” with
its extremely rich content of original autecological data on many mid-European species, and I also expected the
consideration of the extensive paper by Johnson et al. (in: Streit et al.: Evolutionary Ecology of Freshwater
Animals, Birkhäuser 1997) on the evolution and ecological correlates of uniparental and biparental reproduction
in freshwater snails.

Nevertheless, Dillon’s “Ecology” certainly is aa very good synthesis of a very high number of recent papers,
and thus a must for malacologists, evolutionary genetists, and freshwater biologists, too. G. Haszprunar

16. Beesley, P. L., G. J. B. Ross & C. J. Glasby (eds.): Polychaetes & Allies: The Southern Synthesis. Fauna of
Australia. Vol. 4A: Polychaeta, Myzostomida, Echiura, Sipuncula. — CSIRO Publ., Melbourne/ Australia,
2000. xii + 465 pp. ISBN 0-644-05483-2 (set), 0-643-06571-7.

This second volume on Australian invertebrates in the Fauna of Australia series is much more than an account
of Australian worms. All included taxa are treated extensively concerning biology, phylogeny, taxonomy,
classification, history, ecology, and zoogeography making the volume a real encyclopedic work - certainly a
“must” for everyone who is interested in these groups. Of particular use are the reviews of the smaller groups
Myzostomida and Pogonophora — to my knowledge the first comprehensive treatment since 40 years. In
addition, the volume contains a very detailed description of the morphology, anatomy and biology of 80 % of
polychaete families worldwide.

Phylogeny and consequently classification of all treated taxa is still controversial. The book largely follows
a pragmatic approach, but the various authors correctly mentioned the differing points of view.

The only thing to criticize is the entire lack of scale bars and measurements in all figures of the chapters
Myzostomida and Sipuncula, whereas figures of the remaining chapters are very accurately scaled.

The reader will be very pleased by extensive reference lists of each chapter and an extremely useful glossary
and index at the end of the volume. The beauty of the treated taxa is shown by several marvellous colour tables.
To conclude: Systematics and biology of “Polychaetes & Allies” have done a major step forwards. Congratula-
tions to all people involved. G. Haszprunar

156

SPIXIANA 157-163 München, 01. Juli 2001 ISSN 0341-8391

Description of two new species of Notiobia Perty
from Southern Venezuela

(Insecta, Coleoptera, Carabidae, Harpalini)
Erik Arndt & David W. Wrase

Arndt, E. & D. W. Wrase (2001): Description of two new species of Notiobia Perty
from Southern Venezuela (Insecta, Coleoptera, Carabidae, Harpalini). - Spixiana
24/2: 157-163

Two new species of Notiobia Perty are described from South America. N. varia-
bilis, spec. nov. is one of the largest known species, most similar to N. disparilis Bates
but has equally developed convex elytral intervals in both sexes. It is known from
several localities in the Amazonian lowlands. N. acuminata, spec. nov., known only
from the type locality, is one of the smallest species and is distinguished from the
similar N. flavicincta (Erichson) and N. umbrifera Bates by its peculiar lanceolate
aedeagus. Both species belong to the fruit fall communities in primary rain forests.

Prof. Dr. Erik Arndt, Hochschule Anhalt, Fachbereich LOEL, Strenzfelder Allee
28, D-06406 Bernburg, Germany, e-mail: earndt@loel.hs-anhalt.de

David W. Wrase, Dunckerstr. 78, D-10437 Berlin, Germany,
e-mail: david.wrase@t-online.de

Introduction

Species of the Neotropical subgenus Notiobia Perty of the harpaline genus Notiobia are part of the animal
community associated with fruit fall in tropical forests. Therefore, local and time restricted fruit fall
events in the forests cause complicated ecological adaptations of Notiobia (s. str.) species. All known
species are adapted to specific tree genera (or families). The winged adults arrive in the trees when ripe
fruits are available, at least in part before the fruit fall begins (Paarmann, pers. comm.). After fruit fall,
females lay eggs in the ground of the fruit fall area. Larval Notiobia are spermatophagous and develop
only on fruit fall of specific trees as shown by Arndt et al. (1996) and Paarmann et al. (in press).
Mandibular configuration of larval Notiobia species is strongly associated with the particular seed. The
larvae of N. pseudolimbipennis Arndt and N. flavicincta (Erichson) feed on Ficus (Moraceae). They have
extremely elongated, straight and slender mandibles with 4-5 terebral teeth. Mandibles of a similar
structure occur in the larva of N. nebrioides Perty, which feeds on seeds of Clusiaceae. Larvae which feed
on seeds of Melastomataceae have a short mandible with fewer terebral teeth (Arndt et al. 1996).

At present 24 species of Notiobia (s. str.) are known. The Mexican species were revised by Noonan
(1973). Arndt (1998) revised the species of Brazil including all known taxa from the Amazonian
lowlands. During a research project in Southern Venezuela, two new species were discovered which
are described here.

1157.

Material and methods

Beside the types of the described species, material of the following museum and private collections was
examined:

CMP Carnegie Museum of Natural History (Pittsburgh, U.S.A., R. Davidson).

INPA Instituto Nacional de Pesquisas da Amazönia (Manaus, Brazil, C. R. V. Fonseca)
IRSNB Institut Royal des Sciences naturelles de Belgique (Bruxelles, Belgium, K. Desender)
MPM Milwaukee Public Museum (Milwaukee, U.S.A., G. Noonan)

NMNH National Museum of Natural History (Washington, U.S.A., T. L. Erwin, M. G. Pogue)
ZSM Zoologische Staatssammlung (München, Germany, M. Baehr)

cEA Coll. E. Arndt (Leipzig, Germany)

cKlI Coll. S. Kirmse (Erfurt, Germany)

cWR Coll. D.W. Wrase (Berlin, Germany)

The total body length was measured from the tip of the right mandible to the elytral apex at a
magnification of 10 x using an ocular micrometer in a SM 20 stereobinocular microscope (Carl Zeiss
Jena). Line drawings were prepared using an ocular grid (15 x 15 squares) attached to a SM 20
stereobinocular microscope. Dissections were made with standard techniques. Median lobes were
preserved in Euparal on transparent labels and pinned together with the appropriate specimen.

Description of species

Notiobia (s. str.) variabilis, spec. nov.
Figs 1, 3, 4

Types. Holotype: ©, Venezuela, Amazonas, Orinoco region, Rio Surumoni near La Esmeralda, 03.10N / 65.40W,
12.03.1999, leg. S. Kirmse (ZSM). — Paratypes: 334, (27.02.1999, 01.02.1999, 12.03.1999), 422 (27.10.1997,
13.11.1997, 12.01.1999, 27.01.1999) from the same locality and collector (ZSM, cEA, cKI, cWR).

Other material studied: Brazil, state Amazonas, Lago Janauari near Manaus, 03.205/60.17W (4 specimens,
INPA and cEA); state Rio Grande do Sul, Santo Augusto (1 specimen, CMP); “Chapada” (2 specimens, CMP);
Bolivia, dept. Santa Cruz, Rio Ichilo, Buenavista (6 specimens IRSNB, 1 specimen CMP), Santa Cruz (1 specimen
MPM); dept. Cochabamba, Rio Chapara, 400 m (ZSM), dept. del Sara (1 specimen CMP); Peru, prov. Madre de
Dios, Rio Manu, Pakitza (101 specimens, NMNH); prov. Loreto, Rio Amazonas, Cano Yanamona, 100 m (56
specimens, NMNH), Rio Napo, Rio Sucusari 100 m (26 specimens, NMNH), Rio Samiria (3 specimens, NMNH).

Description

Body length. 11.4-14.0 mm (Holotype 12.6 mm).

Color. Dull, black, or dorsum with slight green metallic lustre.

Head. Labrum straight anteriorly; frons with foveae punctiform; clypeus broadly emarginate; eyes
large and protruding. Mentum with prominent median tooth, mentum and submentum completely
separated by a transverse suture; paraglossa slightly longer than ligula. Microsculpture of isodiametric
meshes, micropunctures present.

Thorax. Pronotum with sides arcuate anteriorly, and straight in the most posterior part; posterior
angles rectangular, base lobed; lateral depression deep, not widened posteriorly; lateral bead complete,
basal and anterior beads only distinct at the sides, absent in the middle. A setigerous puncture in the
middle of lateral groove. Median line present. Microsculpture of very fine transverse meshes and
micropunctures.

Legs. Dorsa of tarsi with single small hairs except the posterior tarsi of females which are glabrous.

Hairs on anterior and median tarsi of males more numerous than those on posterior tarsi, hairs on

anterior tarsi of females more numerous than those of middle tarsi. Anterior and median tarsi of males
strongly expanded laterally.
Elytra. Scutellar striae moderately long, posteriorly turning to stria I, with a basal setigerous

puncture; all intervals equally and comparably convex, subapical sinuation moderate in both sexes;
sutural angles acute. Elytral intervals with micropunctures, all intervals equally microsculptured with |

smooth to finely granulate isodiametric mesh (in the sense of Arndt 1998); interval III with a setigerous

puncture in apical third; interval VII complete, with a small setigerous puncture near apex or not
complete, ending before apex, then setigerous puncture in striae between intervals VII and VI;

158

1 € 2.

Fig. 1. Habitus of Notiobia variabilis, spec. nov. (paratype).
Fig. 2. Habitus of Notiobia acuminata, spec. nov. (paratype).

proximal puncture present. Hind wings fully developed in all examined specimens.

Abdomen. Sternum VI of females evenly rounded apically, with two pairs of setae; sternum VI of
males with one pair of ambulatory setae. Aedeagus with median lobe moderately long with rounded
apex (Figs. 3, 4).

Distribution. Amazonian lowlands in Brazil, South Venezuela, East Peru, Central and East Bolivia.
Etymology. The name was proposed by H. W. Bates for this species.

Discussion. Notiobia variabilis, spec. nov. is a widespread species and present in several museum
collections. Bates already labeled this species with the name N. variabilis (IRSNB) but did not describe
it. N. variabilis is most similar to N. disparilis Bates, but has equally developed convex elytral intervals
in both sexes, whereas N. disparilis has equally convex intervals in males but flat intervals 1, 3,5, and 7
as well as convex intervals 2, 4, and 6 in females. N. variabilis was found to develop on fruit falls of
Goupia glabra (Celastraceae).

159

Figs 3, 4. Aedeagus of Notiobia variabilis, spec. nov. 3. Lateral aspect (holotype). 4. Dorsal aspect (paratype).

Notiobia (s. str.) acuminata, spec. nov.
Figs 2, 5, 6

Types. Holotype: 4, Venezuela, Amazonas, Orinoco region, Rio Surumoni near La Esmeralda, 03.10N / 65.40W,
17.12.1998, leg. S. Kirmse (ZSM). — Paratypes: 2834, 7? 2 from the same locality and collector (13, 12 17.09.1997,
1? 22.09.1997, 11383 15.12.1998, 533 17.12.1998, 18 19.12.1998, 135 1.1.1999, 13 12.1.1999, 13 17.1.1999, 18 21.1.
1999, 238 23.1.1999, 15 25.1.1999, 18, 222 27.1.1999, 12 30.1.1999, 22.2 14.02.1999, 13 24.02.1999, 13 28.12.1999)
(ZSM, cEA, cKlI, cWR).

Description

Body length. 8.9-9.9 mm (HT 9.9 mm).

Color. Dorsum with labrum yellow to red brown, remaining parts of head, pronotum, and elytra
greenish to cupreous with metallic lustre; ventral part of body generally brown; legs and palpi
yellowish; antennae yellowish to piceous.

Head. Labrum straight anteriorly; clypeus broadly emarginate; frons with foveae punctiform,
bearing a distinct clypeo-ocular prolongation to eyes; eyes large and protruding. Mentum with prom-
inent median tooth, mentum and submentum completely separated by a transverse suture; paraglossa
slightly longer than ligula. Microsculpture of fine isodiametric meshes, micropunctures present.

Thorax. Pronotum with sides arcuate anteriorly, and straight in the most posterior part; posterior
angles variable, from slightly concave lateral margin with rectangular angle to straight lateral margin
with obtuse angle; base lobed; lateral depression flat, not widened posteriorly; lateral bead complete,
basal and anterior beads only distinct at the sides, absent in the middle. A setigerous puncture near
widest point of pronotum in anterior half. Median line present. Microsculpture of very fine transverse
meshes and micropunctures.

Legs. Dorsa of anterior tarsi with scattered small hairs, dorsa of middle and posterior tarsi
glabrous. Anterior and median tarsi of males strongly expanded laterally.

160

Figs 5, 6. Aedeagus of Notiobia acuminata, spec. nov. 5. Lateral aspect (holotype). 6. Dorsal aspect (paratype).
Scale bar 1.5 mm (Figs. 3-6).

Elytra. Scutellar striae long, extended to a fourth of elytral length, posteriorly more or less turning
to stria I, with a basal setigerous puncture; inner intervals flat, becoming more convex towards the
lateral margin; subapical sinuation moderate in both sexes; sutural angles acute. Inner elytral intervals
of males green, shining, with fine transverse meshes and micropunctures, whereas lateral two (anterior
part of elytra) to four intervals (posterior part) are more or less pale, covered with granulate micro-
sculpture; pale and granulate region of lateral intervals strongly enlarged in females, covering in some
specimens the whole elytra except the medio-subapical part. Interval III with a setigerous puncture in
apical third; interval VII with a small setigerous puncture near apex and with proximal puncture. Hind
wings fully developed in all examined specimens.

Abdomen. Sternum VI of females evenly rounded apically, with two pairs of setae; sternum VI of
males with one pair of ambulatory setae. Aedeagus with median lobe extremely elongate (Figs. 5, 6).

Distribution. Known only from the type locality (Venezuela, state Amazonas).

Etymology. The name refers to the acuminate and elongate median lobe of aedeagus.

Discussion. N. acuminata, spec. nov. was found on fruit falls of Miconia species (Melastomataceae). The
species resembles N. flavicincta (Erichson) and N. umbrifera Bates in habitus and colour. But it is
distinguished from both species by its peculiar aedeagus, from N. umbrifera also by its extended
granulate and pale region of lateral elytral intervals.

161

162

Key to the adults of Notiobia (s. str.) species from Amazonian lowland

Elytral intervals different in males and females. Males with intervals of elytra convex, dorsum of
males bicolored in most specimens, head and pronotum golden green-cupreous, elytra black with
purple tinge; median lobe of aedeagus with short but wide apex. Females with convex intervals 2,
4 and 6, but flat intervals 1, 3, 5, and 7; dorsum of females uniformly dark colored .......................

Ense aRSunansntanen anenrunensntagehnegntenonsnn the rihhgean sent ehs Hanns enserasneeherene se RR ie rer N. disparilis Bates

Elytral intervals of the same shape in males and females, dorsum not bicolored ...................... 2,

Elytral intervals metallic green, with uniform microsculpture and not granulate; microsculpture of
intervals reticulate or lacking; lateral parts of elytra not yellowish; subapical sinuation of elytra
NEN LEE" | 0R 8) 1 UL ALS AUG fonenfnnAnnRasRnnOsDSGERDERERO Eee oereeoERnERPoroLre7E000000000006090000000064500000900000000000000000000000000 I

Postero-lateral intervals of elytra with a region of distinct granulate microsculpture and/or light
yellowish, remaining median part of elytra shining, with reticulate microsculpture, not granulate,
OR species with elytra black with convex intervals and uniform granulate microsculpture; subap-
jeallsinuation.of elytralpromimentorMOL........2220 ne nseeeeenenenenenne nennen nn ee ER 9:

Subapical sinuation of elytra prominent; dorsum green, bronze or blue-green, lateral intervals in
part pale' yellow-testaceous .....................u.2200s02020200000nsnssensessenaenesnnneetange nennen naar sn. vr ER 4.

Subapical sinuation of elytra not prominent; dorsum of variable color. .............ueeeeeseseseenessenenene 5.

Sternum VI of females produced into a ventrally projected spine ; median lobe of male aedeagus
with short but wide apex; elytra distinctly bicolored, with lateral intervals granulate, pale yellow-
testaceous and inner intervals green-cupreous; in females granulated area in the anterior part
enlarsed,.coverine,thertulllanterionpart orrelytraennn N. virıdula (Dejean)

Sternum VI of females rounded apically; median lobe of male aedeagus with longer and more
narrow apex; elytra not distinctly bicolored in most specimens; granulate area of elytra anteriorly
not expanded to the middle part, dorsal surface more shining, green, cupreous or black with
ereenishWbluishroripurplenlustren RER N. pseudolimbipennis Arndt

Body length 11-15 mm; elytral intervals uniformly colored, black or cupreous ...........eeee 6.

Body length 7-10 mm; lateral intervals of elytra, in some females whole elytra except a preapical
macula, granulate and pale yellow-testaceous, inner intervals green or cupreousS ......neeeeeeee 7%

Body length 11-13 mm; elytra cupreous, rarely black with blue or cupreous lustre, inner intervals
flat and shining, lateral intervals more convex and slightly granulate; median lobe of aedeagus
acuminates Anterioritarsi/ofmales peeuliarlynartower. een N. nebrioides Perty

Body length 13-15 mm; elytra black, dull or with cupreous or bluish lustre, all intervals convex,
more or less distinctly uniformly granulate; aedeagus rounded. Anterior tarsi of males wide......
PEN ee ee NEN ER ER N. variabilis, spec. nov.

Median lobe of aedeagus slender, with rounded apex, apex not elongate. Pale region of lateral
intervals strongly enlarged in females; anterior tarsi of males very wide ............ueeeseseseenesenenennenn

Median lobe of aedeagus elongate, lanceolate or spatula-shaped, not rounded. Pale region of lateral
intervalssextendedrornot..............Keelessehererassaetreeesenen sage nseene user een re ea EEE 8.

Median lobe of aedeagus very slender, apex elongate (Figs. 5, 6); pale region of lateral intervals
enlarged in some females; 8.9-9.9 mm; anterior tarsi of males wider...................uesesessenenenoseeneneneneenenee

Apex of median lobe of aedeagus elongated, spatula-shaped (laterally depressed); pale region of
lateral intervals in both sexes narrow, sometimes indistinct; body length usually 7-8 mm; anterior
tarsirof mMales’nartoWenk een nn aanenesneansnensennanterensupnnnnenhe means nesnsg neuen senesseatere rennen N. umbrifera Bates

Elytra very smooth and shining, microsculpture even in the outer intervals indistinct, intervals not
completely flattened; posterior angles of pronotum not blunt, fairly sharp, subdentate. Median lobe
ofaedeagustalwayssroundedrapicallyrn nn 10.

- All intervals of elytra with equal and distinct microsculpture, elytra dull, intervals very flat,
separated from each other only by a faint row of punctures; posterior angles of pronotum blunt.
Median lobe of aedeagus more or less acuminate. Length 9-10.5 mm. (Note: Larger specimens of
11-13 mm length with shining inner intervals, dull, slightly granulate outer intervals, and narrow
anteriorsmalestarsi,nsee.N. nebrioidestBerty)i........nerenneseneenesnesnneee dee een N. aulica (Dejean)

10. Body length 13-15 mm; median lobe of aedeagus wide and stout; apical part of everted internal sac
withsrreeularly.distributed, 1solated large'spines .........nneennesneneeeeraen anne N. maxima Arndt

- Body length 10-12 mm; median lobe of aedeagus comparatively slender; everted internal sac dorso-
basallyawathsastieldior'5-25)large spimes.n.=...n teen nneneeeeneenee eenaenteneetennn N. glabrata Arndt

Discussion

The number of known species of Notiobia (s. str.) increased to 26 (or 27 respectively, if one follows van
Emden 1953 and includes Anisotarsus concinnus Erichson in Notiobia s. str.). That of known species from
Brazil increased from 11 to 12 with the record of N. variabilis in the Amazonian lowlands. Notiobia
viridula (Dejean), N. disparilis Bates and N. glabrata Arndt were collected with the newly described
species by S. Kirmse at the study site near La Esmeralda in southern Venezuela. All three species are
recorded for the first time in Venezuela.

The newly described species confirm the spermatophagy of larvae and adaptation to fruits of a
single tree family in this subgenus of ground beetles.

Acknowledgements

We thank Susan Kirmse (Institute of Botany, University of Leipzig) for donating the type material of the new
species and Robert L. Davidson (Carnegie Museum, Pittsburgh) for linguistic improvement of the manuscript.
We are very grateful to the curators of the afore mentioned institute collections for sending their specimens and
the support during the stay of the first author in their institutes respectively. We are very indebted to
M. Hornschuh (Berlin) who took the photographs of the Notiobia species.

Zusammenfassung

Zwei Arten der Laufkäfergattung Notiobia Perty werden aus Süd-Amerika beschrieben. Die über weite Teile des
Amazonas-Tieflands verbreitete N. variabilis, spec. nov. ist groß, matt, schwarz oder mit leicht metallischem
Glanz und hat konvexe, granulierte Flügeldeckenzwischenräume. Von der ähnlichen Art N. disparilis Bates
unterscheidet sie sich durch gleichmäßig ausgebildete Flügeldeckenzwischenräume bei den Weibchen.
N. acuminata, spec. nov. gehört zu den kleinsten beschriebenen Arten, mit glänzender grüner bis kupferfarbener
Oberseite mit Ausnahme der helleren, matt granulierten äußeren Flügeldeckenzwischenräume. Diese Art ist
bisher nur vom locus typicus aus Süd-Venezuela bekannt und unterscheidet sich von den beiden ähnlichen
N. flavicincta Erichson und N. umbrifera Bates durch ihren sehr langen, lanzettförmigen Aedeagus-Fortsatz. Beide
neuen Arten wurden auf Fruchtflächen in Primärwäldern gefunden.

References

Arndt, E. 1998. The species of Notiobia Perty (Coleoptera: Carabidae: Harpalini) from Brazil. - Acta Amazonica
28: 285-300

-— ,W.Paarmann & J. Adis 1996. Description of larvae and larval specializations to a specific food in the genus
Notiobia Perty (Coleoptera: Carabidae) from Amazonian lowlands. — Stud. Neotrop. Fauna Environ. 31:
205-216

Emden, F. van 1953. The Harpalini genus Ansiotarsus Dejean (Coleoptera. Carab.). - Ann. Mag. Nat. Hist., Ser.
12, 6: 513-547

Noonan, G. R. 1973. The Anisodactylines (Insecta: Coleoptera: Carabidae: Harpalini): Classification, evolution,
and zoogeography. — Quaest. Entomol. 9: 266-480

Paarmann, W., Adis, J., Stork, N., Gutzmann, B., Stumpe, P., Staritz, B., Bolte, H., Küppers, S., Holzkamp, K.,
Niers, C.& C.R. V. Fonseca (in press). The structure of ground beetle communities (Col., Carabidae) at fig
fruit falls (Moraceae) in a Terra Firme rain forest near Manaus (Brazil). — Ecotropica

163

Buchbesprechungen

17. Pölking, F. & U. Walz: Störche. Leben auf der Kathedrale. — Tecklenborg Verlag, Steinfurt, 1996. 76 S., 83
Farbfotos, geb. ISBN 3-924044-25-2.

In Alfaro - einem kleinen Städtchen am Ebro im Norden Spaniens - sind die Bilder zu diesem Storchenbuch
entstanden. Auf dem Dach der dortigen Kathedrale San Miguel befindet sich die größte Weißstorch-Kolonie der
Erde mit 109 Brutpaaren im Jahr 1996 — das entspricht fast genau dem Brutbestand Bayerns! Die eindrucksvollen
Farbaufnahmen aus dieser Kolonie geben einen Einblick in das Brutgeschäft der Weißstörche und immer wieder
in Übersichtsbildern die für den Mitteleuropäer kaum faßbare Dichte der Horste auf Türmen und Simsen, in
Glockenstühlen und auf Dachschrägen. Für die Qualität der Aufnahmen bürgen die Namen der Autoren - hier
haben zwei der renommiertesten Tierfotografen Deutschlands zusammengearbeitet und ein Ergebnis erzielt,
das einfach begeistert. Beim Blick auf den Text fällt zunächst auf, daß die Abbildungstexte sehr blaßgrau
gedruckt sind und dem Leser einige optische Anstrengung abverlangen. Es wäre angebrachter gewesen, diese
Textteile farbig zu setzen. Der fortlaufende Text ist inhaltlich sehr allgemein gehalten. Die Angaben lassen sich
in fast jedem der in den letzten Jahren erschienenen Storchenbücher nachlesen. Im Bemühen um populäre
Formulierungen wurde manchmal etwas zu dick aufgetragen. Kostprobe: “Dem Freund erlesener Liedbeiträge
aus der Kehle gefiederter Sangeskünstler haben die Störche sehr wenig zu bieten ...”

Der Weißstorch ist übrigens auch nicht der größte europäische Brutvogel, wie auf Seite 25 behauptet wird.
Es ist schade, daß sich der lokale Bezug der Bilder im Text nicht wiederfindet; man erfährt fast nichts über die
Lebensbedingungen in der Umgebung, die eine derart einmalige Konzentration an brütenden Störchen ermög-
lichen. Wegen des Textes lohnt sich die Anschaffung des Buches nicht; sie ist jedoch wärmstens zu empfehlen,
wenn man in der Schönheit der Bilder schwelgen und sich an der Tatsache erfreuen möchte, daß es eine solche
Storchenkolonie überhaupt gibt. R. Pfeifer

18. Flade, J. E.: Die Esel. Haus- und Wildesel. Equus asinus. - Westarp Wissenschaften-Verlags-GmbH Hohen-
warsleben, 2000 (Die Neue Brehm-Bücherei Bd. 638). 122 S., 40 Abb., 20 Tab. ISBN 3-89432-887-8.

Im Mittelpunkt des Buches stehen die Domestikation und Zucht des Hausesels, seine Ausbreitung, sein wirt-
schaftlicher Nutzen und seine kulturgeschichtliche Bedeutung in den verschiedenen Kulturen und Epochen.
Auch die Zucht, Eigenschaften und Verwendung des Maultieres werden relativ ausführlich abgehandelt. Leider
fehlen vergleichbare Angaben zum Maulesel. Weitere, eher kurz gefaßte Kapitel beschäftigen sich mit der
Systematik und Stammesgeschichte der Equidenfamilie allgemein sowie mit dem Körperbau und Verhalten des
Hausesels. Ein abschließendes Kapitel gibt Hinweise zur Haltung von Eseln. Gerade die zoologischen Aspekte
werden etwas kurz und oberflächlich abgehandelt. Auch fallen einige Unstimmigkeiten auf. So wird der Kiang
einmal als eigene Art (mit drei Unterarten) geführt, an anderer Stelle wiederum als Unterart des Onagers
klassifiziert. Auch die Stammartenfrage wird nicht ganz widerspruchsfrei beantwortet. So wird einmal der
Nubische Wildesel als alleiniger Stammvater des heutigen Hausesels genannt, bei der Einteilung der Hausesel-
“Rassen” werden die ägyptisch-arabisch-spanischen Formen jedoch als “Somali-Esel” bezeichnet, wobei aber
unausgesprochen bleibt, ob es sich dabei tatsächlich um Abkömmlinge dieser Unterart des Wildesels handelt.
Verwirrend ist auch, daß Poitou- und Gascogne-Esel zu eben dieser Gruppe, gleichzeitig aber auch zur Gruppe
der europäischen Esel gerechnet werden.

Der kulturgeschichtlich interessierte Leser wird in dem Buch interessante Angaben finden. Der Informati-
onsgehalt der biologischen und systematischen Kapitel ist dagegen eher gering. Insgesamt rechtfertigen Inhalt
und Umfang nicht den relativ hohen Preis des Buches. R. Kraft

164

SPIXIANA 165-169 München, 01. Juli 2001 ISSN 0341-8391

A new species of the genus Lissopogonus Andrewes
from northern Borneo

(Insecta, Coleoptera, Carabidae, Patrobinae)
Martin Baehr

Baehr, M. (2001): A new species of the genus Lissopogonus Andrewes from
northern Borneo (Insecta, Coleoptera, Carabidae, Patrobinae). - Spixiana 24/2: 165-
169

Lissopogonus borneensis, spec. nov. from Sabah and Brunei, northeastern Borneo,
is described. The species is distinguished from the four described species (glabellus
Andrewes, 1923, poecilus Andrewes, 1933, suensoni Kirschenhofer, 1991, and tonkin-
ensis Zamotajlov & Sciaky, 1996) by the combination of uniformly dark colouration,
rather depressed, laterally evenly convex elytra, and differently shaped aedeagus.

Dr. Martin Baehr, Zoologische Staatssammlung, Münchhausenstr. 21, D-81247
München

Introduction

Within carabid material recently collected by W. Schawaller, Staatliches Museum für Naturkunde,
Stuttgart (SMNS) in Sabah, northeastern Borneo, I detected a fairly large series of an apparently
undescribed Lissopogonus species. Later received several specimens through courtesy of Mr. D. Wrase,
Berlin, which were collected in different parts of Sabah and in Brunei.

This genus of rather strangely shaped ground beetles was so far known from four species, namely
L. glabellus Andrewes, 1923 from the southern part of the Himalayas to northern Laos, L. poecilus
Andrewes, 1933 from Java, L. suensoni Kirschenhofer, 1991 from eastern China, and L. tonkinensis
Sciaky, 1996 from North Vietnam. From Borneo, thus far no records were available.

Although originally described as a genus of Pogoninae, Lissopogonus has been recently transferred
by Zamotajlov & Sciaky (1996) to Patrobinae - with fairly good reasons, as I believe. However, as both
authors already stated, its position is quite isolated within Patrobinae, and perhaps the genus requires
an own subgroup.

Perhaps all Lissopogonus are mountain living beetles, though at least the newly described species
mentioned in this paper apparently lives at rather low altitude. Generally, very little is known about
habits and way of life of any species. Not even the altitude range is known, because in the descriptions
rarely any indication to altitude is noted. Most probably, species of Lissopogonus are ground living
inhabitants of the forest floor of montane (rain) forests.

Measurements

Measurements have been made under a stereo microscope by use of an ocular micrometer. length has been
measured from apex of labrum to apex of elytra. Measurements, therefore, may slightly differ from those of other
authors.

165

Fig. 1. Lissopogonus borneensis, spec. nov. Habitus. Length: 5.1 mm.

Material and Methods

Altogether 41 specimens of the new species were available for this study. For the taxonomic treatment standard
methods were used. The genitalia were removed from specimens soaked for a night in a jar under wet
atmosphere, then cleaned for a short while in hot KOH.

The material is shared between Staatliches Museum für Naturkunde, Stuttgart (SMNS), collection D. Wrase,
Berlin (CWR), and the working collection of the author in Zoologische Staatssammlung, München (CBM).

Genus Lissopogonus Andrewes

Lissopogonus Andrewes, 1923: 213; Andrewes 1926: 68; 1933: 275; 1935: 314; Kirschenhofer 1991: 9; Zamotajlov
& Sciaky 1996: 40.

Extensive diagnoses of this genus are to be found in Andrewes (1923, 1935). Zamotajlov & Sciaky (1996)
transferred the genus from Pogonini to Patrobini, though at the same time called in question, whether
simple arrangement within Patrobinae conforms to the many special characteristics of the genus.

Lissopogonus borneensis, spec. nov.
Figs 1-3

Types. Holotype: 4, BORNEO: SABAH, Bingkor N Keningau, 400-500 m, 19.-20.X1.1996, leg. W. Schawaller
(SMNS). — Paratypes: 834, 107%, same data (SMNS, CBM); 13, 42%, BORNEO: SABAH, Bingkor N Keningau,
400-500 m, at light, 20.X1.1996, leg. D. Grimm (SMNS); 734, 52%, BORNEO, BRUNEI, Temburong Kuala
Belalong R. Borcherding leg. / 10.1., 11.IL., 11.-16.3., 13.V., 29.V., VI-VII. 1995 (CBM, CWB); 233, 322, MALAY-
SIA - Sabah prov. Banjaran Crocker Mts. 15 km SW Gunung Alab 4-9.V.1996, alt. 790-850 m M. Strba & R.
Hergovits leg. (CWB).

Diagnosis. Distinguished immediately from L. glabellus Andrewes and L. poecilus Andrewes by ab-
sence of any colour pattern on the elytra; from L. tonkinensis Zamotajlov & Sciaky by regularly convex
elytra with much less distinct striation at apex; and from L. suensoni Kirschenhofer by longer elytra and
less suddenly bent apex of aedeagus.

166

Figs 2,3. Lissopogonus borneensis, spec.nov. d and ? genitalia. 2. Aedeagus from left, parameres. Scale: 0.25 mm.
3. Stylomere 2 and base of stylomere 1. Scale: 0.1 mm.

Description

Measurements. Length: 4.6-5.1 mm; width: 1.9-2.05 mm. Ratios. Width/length of pronotum: 1.13-
1.16; width base/apex of pronotum: 1.09-1.12; width pronotum/head: 1.08-1.12; length/ width of
elytra: 1.42-1.47; width elytra/pronotum: 1.73-1.76.

Colouration. Black, sutural area of elytra more or less distincly dark reddish translucent. Mandi-
bles, palpi, and antennae light brown, legs dark yellowish. Lower surface black or dark piceous.

Head. Slightly narrower than prothorax. Eyes comparatively large, laterally distinctly projecting,
about 1.5 x as long as the oblique orbits. Clypeal suture superficially impressed. Frontal furrows
slightly sinuate, laterally of furrow with a posteriorly widened field that is bounded on both sides by
a carina. Neck constriction rather deep. Labrum transverse, anteriorly very gently excised, 6-setose.
Mandibles moderately elongate, apically suddenly curved. Mentum with distinct, unidentate tooth.
Antenna elongate, almost surpassing middle of elytra, median antennomeres >2 x as long as wide.
Posterior supraorbital seta situated slightly behind posterior border of eye. Surface impunctate, with-
out microreticulation, highly glossy.

Pronotum. Gently cordiform, slightly wider than long, in middle rather depressed, laterally evenly
curved, basal angles rectangular. Widest diameter in anterior third. Base slightly wider than apex.
Apex straight, apical angles feebly projecting, rounded off. Base very gently convex. Marginal channel
narrow throughout, barely widened near basal angles, base and apex not margined. Median line
deeply impressed, basally even deeper and wider. Base laterally with two punctiform impressions on
either side. Basal grooves short, deep. Anterior marginal seta situated slightly in front of anterior third,
posterior marginal seta slightly removed from basal angle. Surface impunctate, without any microre-
ticulation, highly glossy.

Elytra. Moderately elongate, regularly curved, dorsally gently convex, widest at or slightly behind
middle. Shoulders very obtusely dentate. Basal margin strong, sinuate, shortly interrupted near
middle, connected to sutural stria. Scutellar striole and seta wanting. Only sutural stria distinct,
impressed, impunctate. All other striae wanting on disk, or, in some specimens, finest traces of inner
striae visible under high magnification. Short remnants of 2" and 7" striae visible at apex. Marginal
channel narrow throughout. A single setiferous puncture situated at position of 3" interval, in middle.
Marginal pores inconspicuous, about 12 in a row that is slightly interrupted in middle. Surface
impunctate, without any traces of microreticulation, highly glossy. Inner wings present.

Lower surface. Impunctate. Metepisternum c. 1.5 x as long as wide. Sternum VII in Ö bisetose, in
? quadrisetose.

Legs. Without striking features. Two basal tarsomeres of male anterior tarsus slightly expanded
and squamose.

167

& genitalia. Aedeagus moderately elongate, rather strongly though regularly curved, apex wide,
remarkably stout, slightly foliaceous. Internal sac rather simply folded, with several short sclerotized
plates near base. Parameres dissimilar in size and shape, both 4-setose at apex.

2 genitalia. Both stylomeres asetose, very similar to those of L. tonkinensis as figured in Zamotajlov
& Sciaky (1996, fig. 110).

Variation. Rather little variation noted, though distinctness of elytral striae fairly variable.

Distribution. Sabah and Brunei, northeastern Borneo. The few records demonstrate that this species
has a fairly wide distribution.

Collecting circumstances. Barely known, type series collected between 400 m and 850 m.
Etymology. The name refers to the range in northern Borneo.

Remarks. The newly detected species is evidence of a rather wide though still fragmentated distribu-
tion of the genus Lissopogonus in southern and eastern Asia. At present the range of the genus extends
from northern India in the northwest to eastern China in the east, and to Java and Borneo in the south.
Because all species apparently are mountain living, this range must have been achieved by some
mountain hopping, which is highly probable because all five known species apparently are fully
winged. However, no flying activities of any species have been thus far recorded. Generally, habits and
life histories of all species are very inadequately known.

Any considerations about phylogenetic relations and zoogeographic history of this genus seem
premature, as long as the actual status of this enigmatic group is not settled. Even when admitted that
Lissopogonus is better arranged near Patrobinae than in Pogoninae, inclusion into Patrobinae is not
really satisfactory and the erection of a distinct group of same taxonomic level as Patrobinae might
better adjust the true relationships. However, in that case the relations of both, Patrobinae and the
Lissopogonus-group, with Psydrinae sensu latu which probably are closely related to Patrobinae (see
Baehr 1999) have to be settled, before any biogeograpical questions can be started.

Key to the species of the genus Lissopogonus Andrewes

ISSElytraswithrdistinetcolour pattermer.. rn ee EL EEE RRRREEN 2.
= Elytraswathout.anyzcolourspatternen.. ee EN Beh 3%
2. Prothorax little wider than long; elytra with a single puncture. Southern slopes of Himalaya .......

BR n ansehe inananensscannarrnsentndangenanenga ren ae see He reirees Nee see glabellus Andrewes

- Prothorax almost a third wider than long; elytra with two punctures. Java ...... poecilus Andrewes

3. Elytra regularly convex, with indistinct striation at apex ...............22222222022enenenenenenenenenenearensnenznenebensnse 4.

- Elytra reversely oviform, with rather distinct striation at apex. North Vietnam .........eee:
en anssannandsesksehtsosnheecnonsensete ide htine denen ee reäksae lee tonkinensis Zamotajlov & Sciaky

4. Elytra shorter and wider; apex of aedeagus narrower, aedeagus in apical third suddely turned

dewum®EasterniChima@ OR suensoni Kirschenhofer

— Elytra longer and narrower; apex of aedeagus wider, aedeagus more evenly curved, apex not

suddenly turned down (Fig. 2). Northern Borneo .................eeeeseeeeeeseneseneneseenee borneensis, spec. NOV.
Acknowledgements

My thanks are due to Dr. W. Schawaller (Stuttgart) and Mr. D. Wrase (Berlin) for the kind loan of the sample,
to Mr. S. Hine (London) for the kind opportunity to study types of this genus, and to Mr. E. Kirschenhofer
(Vienna) and Dr. R. Sciaky (Milano) for kind information on their recently described species.

168

References

Andrewes, H. A. (1923): Papers on Oriental Carabidae. 10. - Ann. Mag. Nat. Hist. 9 (12): 212-223

-- (1926): On a collection of Carabidae from the Kumaon-Tibetan frontier. - Ent. Monthly Mag. 1926: 65-80

-- (1933): In some new species of Carabidae, chiefly from Java. — Treubia 14: 273-286

-- (1935): The Fauna of British India including Ceylon and Burma. Coleoptera. Carabidae. Vol. II. Harpalinae I.
—- London, Taylor & Francis, pp. 1-323

Baehr, M. (1999): A preliminary survey of the classification of the Psydrinae (Coleoptera: Carabidae). In:
Phylogeny and classification of Caraboidea (ed. Ball, G. E., A. Casale & A. Vigna Taglianti). - Mus. reg. Sci.
nat. Torino 1998: 359-368

Kirschenhofer, E. (1991): Zwei neue Carabiden aus Zentral- und Ostasien (Col., Carabidae, Lebiinae, Pogoninae).
— Zeitschr. Arb. gem. Österr. Ent. 43: 9-12

Zamotajlov, A. & R Sciaky (1996): Contribution to the knowledge of Patrobinae (Coleoptera, Carabidae) from
south-east Asia. - Coleoptera 20: 1-63

169

Buchbesprechungen

19. Platen, H.: Das Rattenbuch. Über die Allgegenwart unserer heimlichen Nachbarn. - Campus Verlag GmbH
Frankfurt/Main, 1997. 262 S., zahlr. Abb. ISBN 3-593-35825-5.

Viele Menschen empfinden beim Anblick oder auch nur bei der Erwähnung von Ratten Abscheu und Ekel. Als
Protagonist zahlreicher Grusel- und Horrorgeschichten werden ihr die abstoßendsten Eigenschaften nachge-
sagt. Ratten gelten aber auch - nicht zu Unrecht - als Überlebenskünstler, die sich trotz ausgefeilter Bekämp-
fungsmaßnahmen nicht aus unseren Städten verbannen lassen. Die Autorin kolportiert in ihrem unterhaltsam
geschriebenen Buch allerlei Mythen und Volkssagen, aber auch zeitgenössische Filme und Romane, in denen die
Ratte als Gruseltier und Inbegriff des Bösen und Unheimlichen dargestellt wird. Dabei werden die tiefenpsycho-
logischen Hintergründe dieser Alptraumphantasien angesprochen, gleichzeitig wird aber auch der Versuch
gemacht, den Überlebens- und Ausbreitungserfolg von Rattus rattus und Rattus norvegicus wissenschaftlich zu
erklären. Manche unglaublich klingende Sensationsmeldung wird relativiert, gewisse physiologische Eigen-
schaften - zum Beispiel die angebliche Resistenz gegen gerinnungshemmende Gifte - ins rechte Licht gerückt.
Ein ganzes Kapitel widmet sich der Geschichte der Pest, an deren Ausbreitung die Hausratte maßgeblich
beteiligt war.

Trotz der vielen furchteinflößenden und unappetitlichen Geschichten über Ratten gewinnt man bei der
Lektüre des Buches den Eindruck, daß die Autorin eigentlich eine Rattenfreundin ist. So kommt natürlich auch
die Rolle der Ratte als Versuchs- und Labortier sowie als Heim- und Kuscheltier nicht zu kurz. Die biologischen
Angaben zu Biologie, Physiologie und Verhalten darf man nicht allzu kritisch unter die Lupe nehmen, das
meiste stammt aus der älteren Literatur und entspricht nicht mehr ganz dem neuesten Wissensstand. Es gelingt
der Autorin jedoch sehr gut, zu zeigen, daß die Ratte seit Jahrhunderten die Phantasie der Menschen beschäftigt
und wie kaum ein anderes Tier zur Reflexionsfläche für allerlei Ängste und zum Feindbild schlechthin wurde.

R. Kraft

20. Hellmann, F., E. Brockmann & (t) P. M. Kristal: I macrolepidotteri della Valle d’Aosta. - Monografie 2,
Museo Regionale di Scienze Naturali Saint-Pierre, Valle d’Aosta, 1999. 284 S., 1 Farbtafel mit 10 Abb.

Die Autoren legen eine komplette Faunenliste aller Großschmetterlinge (im ‘klassischen Sinne’) des Aosta-Tales
vor. Die in italienischer Sprache verfaßte Arbeit ragt mit 1141 nachgewiesenen Arten (!) unter den wenigen
bisher vorliegenden Gesamtbearbeitungen italienischer Regionen in Umfang und Verläßlichkeit deutlich her-
aus. Für die Korrektheit der Determinationen - für Fauneninventare unerläßlich, wenn auch nicht selbstver-
ständlich, wie uns andere Fällen leider lehren - bürgt der Fleiß der Autoren, die es zudem verstanden, eine
Vielzahl von Spezialisten in die Entstehung dieser Faunenliste einzubeziehen. Sieben Arten sind neu für die
italienische Fauna, ca. 60 neu für die Region Val d’Aosta.

Die einleitenden Kapitel behandeln Geographie, Geologie, Klima und Vegetation des Untersuchungsgebie-
tes. Der Leser findet hier auch eine Liste der über 160 Fundorte mit Höhenangaben und genau nachvollziehbarer
Lokalisierung.

Im Hauptteil wird jede Art nach folgenden Kriterien charakterisiert: Phänologie, oft Habitatangaben,
Höhenverbreitung, relative Häufigkeit, detaillierte Verbreitungsdaten innerhalb des Untersuchungsgebietes.
Interessant auch die durchwegs angewandte Einteilung in ‘Corotypen’ nach dem System von Vigna Taglianti.
Auf einer schönen Farbtafel werden zwei endemische Hepialidenarten (Pharmacis anselminae, Pharmacis claudiae)
und eine Lycaenidenart (Polyommatus humedasae) abgebildet.

Das Buch ist unglaublich gut recherchiert und korrekturgelesen, und abgesehen von einer Kleinigkeit - dem
auf der Buchklappe verdruckten Autorennamen Kristall statt Kristal — gibt es hier nichts zu bemängeln. Wenn
es nur mehr solcher Bearbeitungen gäbe! A. Hausmann

170

SPIXIANA 171-175 Meonehen 0112001 ISSN 0341-8391

A new species of the genus Minuthodes Andrewes
from North Queensland, Australia

(Insecta, Coleoptera, Carabidae, Lebiinae)
Martin Baehr

Baehr, M. (2001): A new species of the genus Minuthodes Andrewes from North
Queensland, Australia (Insecta, Coleoptera, Carabidae, Lebiinae). - Spixiana 24/2:
171-175

Minuthodes trimaculata, spec. nov. is described from the lower Cape York Penin-
sula, northern Queensland. The species is closely related to M. froggatti (Macleay)
from North Queensland and far Northern Territory and M. demarzi Baehr from far
Northern Territory. It is distinguished from both species by the different elytral
pattern, presence of a light spot on frons of head, projecting though obtuse basal
angles of pronotum, and scarcely rugose though distinctly microreticulate dorsal
surface of the head. A checklist of the Australian species of the genus Minuthodes
is added.

Dr. Martin Baehr, Zoologische Staatssammlung, Münchhausenstr. 21, D-81247
München, Germany.

Introduction

While working through unsorted and undetermined carabid material in the Coleoptera collection of
the Queensland Museum, Brisbane (QMB), I detected a single specimen of the genus Minuthodes that
at the first glance did not fit any Australian or New Guinean species of this genus known to me. After
comparison with all related species in my working collection it proved to represent a new species that
is, however, closely related to the northern Australian species M. froggatti (Macleay) and M. demarzi
Baehr.

Because the catalogue of Moore et al. (1987) is outdated with respect to the genus Minuthodes, a
checklist of all species recorded from Australia is added to this paper.

Measurements
Measurements have been made under a stereo microscope by use of an ocular micrometer. Length has
been measured from apex of labrum to apex of elytra. Length of pronotum was taken along midline.
Measurements, therefore, may slightly differ from that of other authors, especially Darlington.
Characters
During ample determinating work on Australian and New Guinean Minothodes it turned out that

external characters like shape of pronotum, microstructure (puncturation, striation, microreticulation)
of surface, and even the colour pattern on the elytra are of more value for the distinction of species than

Fig. 1. Minuthodes trimaculata, spec. nov. Habitus. Length: 5.35 mm.

are the male genitalia. These are relatively uniform throughout the genus, while the mentioned external
characters apparently are not only very distinct but also remarkably little variable within each species.
Hence, discrimination of species is possible and even easier by use of external characters than male
genitalia.

Minuthodes trimaculata, spec. nov.
Figs 1,2

Types. Holotype: 4, Musgrave 1.X.1974 G. B. Monteith (OMB, T 93151).

Diagnosis. Relatively large, trimaculate species with elytral pattern similar to M. minima (Macleay),
but body much larger and wider. Distinguished from both most closely related species M. froggatti
(Macleay) and M. demarzi Baehr by presence of a transverse reddish spot on frons, widely interrupted

anterior and posterior elytral spots, less wide pronotum with projecting but distinctly obtuse basal |
angles, and but weakly rugose though markedly microreticulate upper surface of head. Further

distinguished from M. froggatti (Macleay) by more heavily microreticulate and less glossy pronotum
and elytra, and from M. demarzi Baehr by darkened femora, more cordiform pronotum, and far less
rugose and therefore glossier surface of pronotum and elytra.

172

Fig. 2. Minuthodes trimaculata, spec. nov. Male genitalia: aedeagus, parameres, genital ring. Scales: 0.25 mm.

Description

Measurements. Length: 5.35 mm; width: 2.25 mm; ratio width/length of pronotum: 1.78; ratio
width of pronotum/ width of head: 1.15; ratio length / width of elytra: c. 1.45 (elytra somewhat opened).

Colour. Dark piceous-black. Frons posteriorly with two indistinct, transversely arranged, reddish
spots. Elytra trimaculate, with a large, distinct, reddish, irregularly triangular humeral spot that
extends from 3"! stria to 7" stria though is not in contact to any part of the anterior or lateral margins.
In apical half with a common crescent-shaped, reddish spot that laterally extends to 6" stria and is
anteriorly prolonged at position of 4" stria. Both, the humeral spots and the apical spot very widely
separated. Lateral margins of pronotum and elytra narrowly reddish. Palpi and antenna reddish
throughout, three basal antennomeres very faintly lighter than the rest. Femora piceous, tibiae and tarsi
dark reddish, moderately contrasting. Lower surface of pronotum and abdomen laterally piceous, in
middle broadly reddish. Elytral epipleurae anteriorly reddish.

Head. Wide in comparison to prothorax. Frons anteriorly in middle feebly convex, laterally with
very shallow impressions. Impressions with about four to five inconspicuous and rather irregular
longitudinal furrows, in middle with some inconspicuous wrinkles. Base of clypeus also with some
short longitudinal wrinkles. Eyes large, markedly protruding, with small, obliquely convex orbits.
Head but little narrower than prothorax. Clypeo-frontal suture distinct. Anterior margin of clypeus
slightly excised, anterior angles broadly rounded, bisetose. Labrum elongate, considerably longer than
wide, lateral borders oblique, apex straight, behind apex in middle with a small circular groove.
Labrum 6-setose, the proximal seta far removed from apical margin, lateral margins apparently
without additional hairs. Mandibles with some longitudinal furrows on upper surface. Terminal
palpomere of labial palpus as long as penultimate palpomere, apparently impilose. Maxillary palpus
with sparse and very fine pilosity. Mentum with sharp, unidentate tooth. Submentum bisetose, gula
quadrisetose. Antenna short, barely surpassing basal angle of pronotum, median antennomeres but
slightly longer than wide, densely pilose from apex of 4" antennomere, basal antennomeres sparsely
setose. Microreticulation distinct on whole surface including clypeus and labrum, isodiametric. Frons
and clypeus irregularly punctate, surface rather dull, apparently without any pilosity.

Pronotum. Wide, distinctly cordiform. Apex slightly wider than base, feebly concave, anterior
angles broadly rounded. Sides almost evenly rounded though with a very obtuse angle at position of
anterior marginal seta, widest in anterior third. Near basal angle with comparatively elongate sinuo-
sity. Basal angles projecting though obtuse, because the basal margin is considerably curved at angle.
Base in middle gently convex. Base bordered throughout, apex in middle unbordered. Lateral channel
rather narrow throughout, margin slightly upturned. Disk in middle somewhat raised. Median line
distinct, in middle deeply impressed. Basal grooves fairly deep, oblique, prebasal transverse sulcus
distinct. Between median line and lateral margin in middle with a small, circular, moderately deep

173

groove. Anterior marginal seta situated in anterior third, at widest diameter of pronotum, posterior
marginal seta situated at basal angle. Microreticulation im middle slightly superficial, irregularly
transverse, near borders distinct, isodiametric. Puncturation irregular, rather sparse on disk, laterally
and apically coarse and denser. Surface with many shallow, irregularly transverse wrinkles, on disk
moderately glossy, laterally more dull, with moderately dense, short, declined, yellow pilosity.

Elytra. Moderately short and wide, widest behind middle, depressed. Humeri evenly rounded,
sides very feebly convex, apex oblique, deeply sinuate, sutural angles broadly rounded, elytra slightly
dehiscent at suture. Marginal channel slightly widened at anterior third. Striae distinct though shallow,
microreticulation distinct, isodiametric, the whole surface densely punctate and pilose. Pilosity yellow,
rather short, somewhat declined. 3" interval with three discal punctures, the anterior one situated near
base at position of 3'X stria, both posterior punctures situated near 2" stria, 3”! puncture very close to
apex. Punctures and the very short setae arising from the punctures difficult to trace within the dense
puncturation and pilosity. Marginal setae very elongate. Lateral margin extremely finely serrate and
very sparsely pilose in anterior half. Surface rather opaque. Posterior wings fully developed.

Lower surface. Proepisternum impilose, prosternum sparsely pilose. Lower surface of hind body
rather sparsely punctate and pilose. Metepisternum almost 2x as long as wide at apex. Terminal
abdominal sternum of male 4-setose.

Legs. Four basal tarsomeres of male protarsus slightly widened and biseriately squamose, though
4% tarsomere with few squamae only.

Male genitalia. Genital ring elongate, almost symmetric. Basal plate transversely split. Aedeagus
narrow, elongate, fairly curved, lower surface evenly concave, apex straight, elongate. Orificium short,
turned to left side. Internal sac very simply folded, with a small tooth-like sclerite inside orificium.
Parameres very dissimilar, right paramere fairly small, apically widened, left paramere larger.

Female genitalia. Unknown.

Variation. Unknown.

Distribution. Lower Cape York Peninsula, northern Queensland. Known only from type locality.
Collecting circumstances. Not recorded. Holotype captured in October.

Etymology. The name refers to the trimaculate elytral pattern.

Remarks

The genus Minuthodes Andrewes is most speciose and diverse in New Guinea (Darlington 1968, Baehr

1998), whereas for a long time in Australia only a single species was known that occurs in northern
Queensland (Moore et al. 1987). During the last decade, however, additional species were described

from north Queensland as well as from the northern parts of Northern Territory and from northwest- |
ern Australia (Baehr 1990, 1994). At the same time, some species previously included in the genus
Agonocheila Chaudoir by Moore et al. (1987) were arranged in the genus Minuthodes by Baehr (1990).

Hence, at present, in Australia the genus Minuthodes is distributed from eastern South Australia
through Victoria, the Australian Capital Territory, New South Wales, central and eastern Queensland,
far Northern Territory, and to the Kimberley Division in northwestern Australia as far south as Fitzroy
Crossing. For distribution of the involved species see checklist at the end of this paper.

Contrary to New Guinea, in Australia the genus Minuthodes includes a group of very small species
(M. minima Macleay, M. serrata Baehr) that live under bark of eucalypts in open rather than closed

forest. Almost all species of the major group of large, wide species in New Guinea occur-asfarastheir
habits are recorded - in rain forest where they live on the bark of standing trees as well as on logs,
sometimes even in the thick cover of moss on tree trunks. In Australia, M. queenslandica Sloane and

M. walfordi Baehr belong to this group and may possess the same habits. M. froggatti, however, which

likewise belongs to the group of large species, in northern Queensland and in far Northern Territory
has been found by me mainly under bark of eucalypts, e.g. River Gums (Eucalyptus camaldulensis), |

where it lives in a similar way as the small subcortical species mentioned above. No collecting

circumstances are known of M. demarzi Baehr and the new species described herein. From the collecting |

localities, however, I would argue that they likewise live rather under eucalypt bark in open sclerophyll
forest and tropical savannah than in rain forest.

174

In Australia at least, subcorticolous species of open sclerophyll forests and rain forest species of
Minuthodes differ in their elytral pattern. Whereas both rain forest inhabiting species M. queenslandica
Sloane and M. walfordi Baehr possess a pattern consisting of many narrow, light stripes, all sclerophyll
forest inhabiting species possess a bilineate, trimaculate, or quadrimaculate pattern, very similar to
those patterns common in subcorticolous sclerophyli forest living species of other carabid genera. It
seems, hence, that elytral patterns consisting of large spots are better adapted to the conditions in
sclerophyll forests, wheres the multilineate patterns apparently are better adapted to rain forest
conditions. The reasons for this are still unknown, but it may be caused by different predatorial
constraints.

It follows, then, that in Australia several species of Minuthodes apparently have managed to change
their habits of living in rain forest to invade the open sclerophyli forests. As the stock of the genus
Minuthodes certainly was a rather recent immigrant into northern Australia from the north, the genus
represents one of the rare examples of rain forest inhabiting Oriental or Papuan faunal elements that
have successfully invaded the unique Australian habitat of subcortical fissures on eucalypts in open
sclerophyll forests and tropical savannahs. Because this habitat houses a numerous and diverse
subcortical carabid fauna, it would be very interesting to know the way in which certain Minuthodes
species managed to introduce themselves into the new habitat and to escape from competition of the
many species of Psydrinae (Amblytelus, Dystrichothorax), Tetragonoderinae (Scarothrocrepis), Lebiinae
(Agonocheila, Philophloeus, Demetrida, Trigonothops, Phloeocarabus), and Pseudomorphinae (Adelotopus,
Sphallomorpha) that likewise occur under eucalypt bark and actually may be found on the same tree.

Checklist of the species of Minuthodes from Australia
For the benifit of the reader this checklist includes some information about the recorded range of the

species. It was compiled from the literature and from my own collecting and determinating experience
(abbreviations of the states of Australia as usual, e: eastern, n: northern).

demarzi Baehr, 1990 n.NT

froggatti (Macleay, 1888) ne.QLD, n.NT, n.WA
minima (Macleay, 1864) e.SA, VIC, ACT, NSW, QLD
queenslandica (Sloane, 1917) ne.QLD

serrata Baehr, 1990 n.NT, n.WA

trimaculata, spec. nov. ne.QLD

walfordi Baehr, 1994 ne.QLD

Acknowledgements

My sincere thanks are due to Dr. G. B. Monteith, Queensland Museum, Brisbane, for the kind loan of the
material.

References

Baehr, M. 1990. A review of the Australian species of Minuthodes Andrewes, with the description of two new
species (Coleoptera, Carabidae, Lebiinae). — Spixiana 13: 33-41

-- 1994. A new species of Minuthodes Andrewes from Australia (Insecta, Coleoptera, Carabidae, Lebiinae). -
Spixiana 17: 37-41

-- Baehr, M. 1998. Two new species of Minuthodes Andrewes from New Guinea (Insecta, Coleoptera, Cara-
bidae, Lebiinae). — Spixiana 21: 235-240

Darlington, P. J. Jr. 1968. The Carabid beetles of New Guinea. Part III. Harpalinae continued. Perigonini to
Pseudomorphini. — Bull. Mus. Comp. Zool. 137: 1-253

Moore, B. P., T. A. Weir & J. E. Pyke. 1987. Rhysodidae and Carabidae. In: Zoological Catalogue of Australia 4:
17-320. — Australian Government Publishing Service, Canberra

Buchbesprechungen

21. Hürter, H.-A.: Die wissenschaftlichen Schmetterlingsnamen, Herleitung und Deutung. - Verlag Peter Pomp,
Bottrop, Essen, 1998. 482 5. ISBN 3-89355-176-X.

Der Autor legt dem Leser ein ungewönliches, innovatives Nachschlagewerk vor, das — wie kein anderes zuvor
in der Lepidopterologie — die altphilologischen Wurzeln von Schmetterlingsnamen analysiert. Als Zielgruppe
wurden die in den “Schmetterlingen Mitteleuropas” (Forster & Wohlfahrt 1976) genannten Tagfalternamen
gewählt, insgesamt 643 Art- und Gattungsnamen sowie deren wichstigste Synonyme. Da auch bei den Nacht-
faltern oft ähnliche bzw. gleiche Namen auftauchen, kann deren Bedeutung in so manchem Falle auch in diesem,
auf die Tagfalter beschränkten Werk nachgeschlagen werden.

Durchgehend stellt der Autor mehrere Erklärungsversuche nebeneinander, um dann deren Plausibilität in
eingehender Diskussion gegeneinander abzuwägen. Eine Fülle von Fußnoten ergänzt diese Analyse in fundier-
ter Weise.

Die mit Forster & Wohlfahrt (1976) übereinstimmende Nummerierung macht das Werk besonders benut-
zerfreundlich.

Da durch die detaillierten Erklärungen tief in Themenbereiche der Geschichte und der antiken Sagenwelt
(“Liebesgeschichten, Eifersuchtsdramen, Tiergeschichten, Zauberei, Alkoholgeschichten, Bestechung, Mißver-
ständnis mit Todesfolge, Kindesaussetzung, Kannibalismus, Raubmord, politischer Machtkampf”) eingegangen
wird, ist das Buch in vielen Passagen nicht nur sehr informativ, sondern regt durchaus auch zum Schmunzeln an.

Fazit: Ein jedem Lepidopterologen, aber auch jedem humanistisch bzw. altphilologisch Interessierten sehr
zu empfehlendes Werk, das so detailliert auf alle Einzelheiten eingeht, daß bei der Lektüre weder geschichtliches
Vorwissen noch Latein- oder Griechisch-Kenntnisse vonnöten sind.

A. Hausmann

22. Baez, M.: Mariposas de Canarias. — Editorial Rueda S.l., Madrid, 1998. 216 S., 323 Farbfotos. ISBN 84-7207-
1100-3.

Auf 323 durchwegs guten und hervorragend zur Bestimmung geeigneten Farbfotos wird in diesem neuen
Führer mehr als die Hälfte der derzeit von den Kanarischen Inseln bekannten Schmetterlingsarten vorgestellt.
Angenehmerweise behandelt dieses Buch auch die Microlepidoptera ausführlich, so daß man einen ausgewo-
genen Überblick über die Fauna dieser Inselgruppe bekommt.

Obwohl die kanarischen Inseln wegen deren Beliebtheit als Urlaubsziel (und der damit verbundenen
Annehmlichkeiten bei Planung und Durchführung der Reise) schon unzählige Male von Schmetterlingssamm-
lern besucht wurden, so gab es bisher praktisch keine Übersicht über das Fauneninventar, da die publizierten
Daten (v.a. von Klimesch, Rebel und Pinker) in einer großen Zahl von Einzelartikeln in Fachzeitschriften
veröffentlicht wurden.

Der Autor ist ein bekannter Lepidopterologe, der vor Ort an der Universität La Laguna auf Teneriffa
beschäftigt ist. Er stellt dem Buch eine bebilderte, instruktive Übersicht über die verschiedenen Familien voran
und läßt darauf den systematischen Teil folgen. In diesem wird jede Art mit einem Farbfoto und einem
halbseitigen, spanischen Text vorgestellt, der im Wesentlichen aus einer Kurzbeschreibung, einer Charakteristik
der Verbreitung sowie Habitat- und Futterpflanzenangaben besteht.

Den Abschluß des Werkes bilden eine sytematische Liste aller (nicht nur der speziell behandelten!) Arten, |

ein Index der Schmetterlingsarten und ein Index der Raupenfutterpflanzen.

Diese wertvolle Neuerscheinung ist Pflichtlektüre im Gepäck jedes Entomologen, der die Kanarischen |

Inseln bereist, und darüber hinaus ein solides Bestimmungswerk für jeden Naturfreund!
A. Hausmann

176

SPIXIANA 177-190 München, 01. Juli 2001 ISSN 0341-8391

Two new sibling species of Mantidactylus cornutus
from Madagascar

(Amphibia, Anura, Ranidae)
Frank Glaw & Miguel Vences

Glaw, F. &M. Vences (2001): Two new sibling species of Mantidactylus cornutus
from Madagascar (Amphibia, Anura, Ranidae). — Spixiana 24/2: 177-190

A review of the Malagasy frogs which in the past were subsumed under the
name Mantidactylus redimitus, together with new field data, led to the confirmation
of the specific validity of M. cornutus and to the recognition of two new species:
M. tschenki, spec. nov. from Ranomafana is morphologically similar to M. cornutus
but shows distinct differences in advertisement call and has a slightly bilobed
(instead of a roundish) subgular vocal sac. M. tandroka, spec. nov. from higher
elevations of the Marojezy massif is distinguished by a wide head, distinct dorsal
ridges, and colouration from M. cornutus, M. tschenki, and M. redimitus. The discov-
ery of close syntopy of M. redimitus and M. cornutus in central eastern Madagascar
confirms that they represent valid species and that advertisement calls are good
indicators for specific distinctness in Malagasy anurans.

Frank Glaw, Zoologische Staatssammlung, Münchhausenstr. 21, D-81247
München, Germany. E-mail: Frank.Glaw@zsm.mwn.de

Miguel Vences, Museum national d’Histoire naturelle, Laboratoire des Reptiles
et Amphibiens, 25 rue Cuvier, 75005 Paris, France. E-mail: m.vences@t-online.de

Introduction

During the last decades it has become evident that the analysis of advertisement calls is crucial to
understand the species diversity of anurans (e.g. Passmore & Carruthers 1995). This is especially true
for tropical anuran communities like those in Madagascar which are poorly studied and where numerous
new amphibian species are still to be discovered and described. A significant percentage of the recently
discovered new species are morphologically similar to already known species and some of those sibling
species pairs are hardly distinguishable when preserved (Glaw & Vences 2000). However, in all cases
in which anurans with relevant differences in their advertisement calls have been investigated genet-
ically, relevant genetic distances - indicative of reproductive isolation at the species level - have been
found (pers. obs. in more than 20 species pairs from Madagascar, South America, and Asia).

In the present paper we review a complex of species in the Malagasy genus Mantidactylus which
previously (Guibe 1978, Blommers-Schlösser & Blanc 1991) were all subsumed under the name Manti-
dactylus redimitus. Glaw & Vences (1992b) noted the bioacoustic and morphological differences of low-
altitude M. redimitus specimens and those of mid-altitude localities, and described the latter as new
species M. cornutus. We here provide evidence that M. cornutus populations as listed in the distribution
map of Glaw & Vences (1994) are still composed of various species: one new sibling species was
discovered during a bioacoustic survey near the village Ranomafana in south-eastern Madagascar; its
morphology is very similar to M. cornutus but its advertisement calls are rather different. In contrast,
specimens from the Marojezy massif in north-eastern Madagascar differ by distinet morphological
features and are described as second new species, although their advertisement calls are not yet known.

177

Materials and methods

Vocalizations were recorded using portable tape recorders with an external microphone (Vivanco EM 238) and
were analyzed with the MEDAV sound analyzing system Spektro 3.2. The following morphological measure-
ments were taken with a calliper to the nearest 0.1 millimeter: SVL (snout-vent length), HW (head width),
HL (head length), ED (horizontal eye diameter), END (eye-nostril distance), NSD (nostril-snout tip distance),
NND (nostril-nostril-distance), TD (horizontal tympanum diameter), HAL (hand length), FORL (forelimb
length), HIL (hindlimb length), FOL (foot length), FOTL (foot length including tarsus), IMTL, IMTH (length
and height of inner metatarsal tubercle), TIL (length of first toe). Webbing formula is given according to
Blommers-Schlösser (1979). Institutional abbreviations are as follows: BM (Natural History Museum, London);
MNHN (Museum national d’Histoire naturelle, Paris); MRSN (Museo Regionale di Scienze Naturali, Torino);
ZFMK (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn); ZSM (Zoologische Staats-
sammlung, München). Statistical analyses were carried out using SPSS for Windows, version 9. We performed
Mann-Whitney U-tests to test significance of intersexual differences in size and morphometric ratios (relative
tympanum length, ratio TD/SVL; relative size of inner metatarsal tubercle, IMTL /SVL and IMTH/SVL).
Temporal and metric measurements are given as range, with mean + standard deviation in parentheses.

Results
Mantidactylus redimitus (Boulenger, 1889)
Material examined. BM 1947.2.26.55 (holotype; original number BM 89.8.1.24; Madagascar); BM 92.3.7.39-41
(Sahembendrana); BM 1928.5.9.11-12 (Brickaville); BM 1988.593 (Ambatovaky); MNHN 1973.911 (Marojezy, 600-

1300 m altitude); MNHN 1973.937 (Marojezy, 300 m altitude); MNHN 1973.938-939, 1973.941 (Marojezy, 600 m
altitude); ZFEMK 52704-52705 (Nosy Boraha); ZFMK 52716 (Nosy Mangabe, juv.); ZFMK 60073 (An’Ala).

Morphology. Summarizing measurements of the specimens in Table 1 (only adults considered; type

not included due to bad state of preservation) results in a male SVL of 43.4-52.83mm (47.6+3.7mm,
n=6) and a female SVL of 48.1-48.3mm (n=2). Mean male SVL was 99% of mean female SVL. Sexual
size dimorphism was not significant (U-test, P=1). All specimens corresponded to the description in
Glaw & Vences (1994) in having short legs (tibiotarsal articulation reaching at least to the anterior eye

corner, at most between nostril and snout tip), and a rather smooth dorsal skin, without large dermal
spines on the eyes, and with a pair of only faintly expressed tubercles between the eyes. Mean relative
tympanum size did not differ significantly between sexes (U-test, P> 0.6), while intersexual differences
in relative length and height of inner metatarsal tubercle (means of both values larger in males) were
close to significance (U-test, P=0.07). In contrast to the following species, the femoral glands in male

M. redimitus were prominent and always well visible. Their size was 9.7 x2.9mm in MNHN 1973.911,

8.6x2.9mm in MNHN 1973.937, 7.3x3.0mm in ZFMK 52704, 8.4x3.3mm in ZFMK 52705, and |

9.2x3.1mm in ZFMK 60073.

Habitat and habits. Calling activity generally started at dusk, but sometimes single calls were also

heard since the early afternoon (14.30h). Calling males were found in February and March sitting |

horizontally in the vegetation (1-2m above the bottom) along larger brooks (broader than Im). At

An’Ala, where M. redimitus was observed calling syntopically with M. cornutus, the former was only '
found at the edge of a broad brook (>3m) whereas the latter was sitting along a small afflux (not |
broader than 0.5m) only several metres apart. W. Herwig (pers. comm.) photographed a specimen of |

M. redimitus at Vohidrazana on 16 July 2000 at 19.30h. It was sitting on a leaf of a bush about 160 cm

above the bottom. This record indicates that M. redimitus is also active in the comparatively cold and |

dry winter season.

Advertisement calls. Calls were recorded at An’Ala (on 11 February 1995, 17.30h, at 22°C air temper- |
ature) and Marojezy (on 22 February 1995, ca. 21.00h, at 25°C air temperature). They consisted either |
of single notes or note series. Each note (Fig. 1) corresponded to one expiration. At An’Ala, note |

duration was 274-352 ms (309+22 ms, n=9), duration of intervals between notes was 484-717 ms |

(584+75 ms, n=8). Each note was composed of 4-6 pulse groups, each of which contained 2-7 pulses.

The longest note series recorded consisted of 12 notes and had a note repetition rate of 1.1/s. Frequency |

was 900-1400 Hz. At Marojezy, note duration was 315-350 ms (329+10 ms, n=9), duration of intervals
between notes 493-737 ms (597 #106 ms, n=6). Each note consisted of 7-10 pulse groups, each of which |

178

2 Frequenz (kHz)

0 100 200 300 400 500 ms

Fig. 1. Sonagram and oscillogram of one note of Mantidactylus redimitus from Marojezy.

contained 1-10 rather indistinct pulses. Frequency was 900-1500 Hz.

Further recordings from Nosy Boraha were analyzed by Glaw & Vences (1992a,b). The high note
repetition rate (up to 2.5/s) in these recordings differ from the An’Ala and Marojezy data, and may be
due to an exceptional motivation of the corresponding specimen (several specimens were calling rather
close to each other). The low note duration (ca. 100ms) as given by Glaw & Vences (1992b), however,
is an artefact of analysis (probably originated by measuring note duration on the sonagram), the actual
note duration in the Nosy Boraha recordings was similar to that in An’Ala and Marojezy as we
ascertained by re-analysis.

Distribution. Data presented here confirm that M. redimitus is primarily a low-altitude species. At
Marojezy, it has mainly been found at 300-600 m above sea level, only one specimen comes from an
imprecise higher altitude (600-1300 m). The collecting localities at Nosy Boraha and Nosy Mangabe
were close to sea level (<100m altitude), whereas that of An’Ala was at 840m above sea level.
Sahembendrana is also at mid-altitude, while Ambatovaky and Brickaville can be considered as low-
altitude sites. Beside these localities, Glaw & Vences (1994) list one additional locality, Maroantsetra-
Antalaha (that means the path between both cities), which is based on personal observations in 1987
(only photographic voucher available). A further photographic record was made by W. Herwig at
Vohidrazana some 700m above sea level.

Raxworthy & Nussbaum (1996) reported M. redimitus [M. redimitis (sic!)] from altitudes of 650-
1700 m in the Andringitra massif but did not list M. cornutus. Since cornutus-like specimens are known
from Andringitra (see below), it is possible that this record is based on a misidentification. The same
regards their record of M. redimitus from Ranomafana (900-1050 m altitude) which possibly refers to the
new species M. tschenki described below. Two additional records of M. redimitus exists for the “Reserve
Speciale du Pic d’Ivohibe” (1200m altitude) and the corridor to the “Parc National d’Andringitra”,
900m altitude (Raselimanana 1999). Since M. cornutus is not mentioned in this paper and MNHN
material from Pic d’Ivohibe is cornutus-like these records are also likely to refer to M. cornutus or
M. tschenki.

Mantidactylus cornutus Glaw & Vences, 1992

Material examined. ZFMK 52702-52703 (paratypes, Andasibe); ZFMK 53691 (holotype, Andasibe); ZFEMK 59867
(Andasibe); ZSM 573/1999 (paratype, Andasibe, originally ZFMK 53690); ZSM 308/2000 (Vohidrazana, 18°57'
57"S, 48°30' 37"E, 730 m above sea level).

Notes on the type material. In the original description of M. cornutus (Glaw & Vences 1992b: 272) the paratypes
were defined as follows: “Three adult males (ZEMK 53690, 52702, 52702) from Andasibe ...”. The latter number
was a typing error and must be corrected to 52703.

179

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180

b Frequenz (kHz)

0 100 200 300 400 500 ms

Fig. 2. Sonagram and oscillogram of one note of Mantidactylus cornutus from An’Ala.

Morphology. In the studied sample (see Table 1 for measurements), SVL was 37.5-40.1mm
(39.2+1.2mm, n=4) in males and 33.3-48.IJmm (36.1+4.0mm, n=2) in females. Mean male size was
109% of mean female size, no significant sexual size dimorphism was found (U-test, P>0.2). All
specimens largely corresponded to the descriptions of Glaw & Vences (1992b, 1994) in having rather
long legs (tibiotarsal articulation reaching at least nostril, mostly beyond snout tip) and a distinct pair
of blackish tubercles between the eyes. Neither relative tympanum size (mean value smaller in males)
nor relative length and height of inner metatarsal tubercle (means of both values larger in males) were
significantly different between sexes (U-test, P>0.05), but the latter may be due to small sample size.

Habitat and habits. Calling males were found in all months from December to March, indicating a
prolonged mating activity during the rainy season. They were sitting horizontally in the vegetation
(1.5-2.5m above the bottom) along very small (mostly smaller than 0.5m in diameter) and slowly
running brooks, in pristine or degraded forest.

Advertisement calls. Calls were recorded at An’Ala (on 21 March 1995 at 22°C air temperature) and
Andasibe (on 16 January 1995, 19.00h). They were long-lasting regular note series (up to several
minutes). Notes (Fig. 2) were unharmonious. At An’Ala, note duration was 90-113ms (99+7 ms, n=32),
duration of intervals between notes was 616-1280 ms (825 + 140 ms, n=31), note repetition rate was ca.
1/s. One analyzed note consisted of 20 pulses and had a pulse repetition rate of 227/s. Frequency was
1150-2500Hz, dominant frequency 1400-1950Hz. At Andasibe, note duration was 93-111 ms
(102+6ms, n=10), duration of intervals between notes was 536-903ms (720+131ms, n=9), note
repetition rate was ca. 1.3/s. One note was composed of ca. 23 pulses. Frequency was 1000-5100 Hz,
dominant frequency 1300-2100 Hz.

Distribution. The species is reliably known from Andasibe (type locality, ca. 900m altitude), An’Ala
(personal observations and call recordings, ca. 840 m altitude), and most probably from Vohidrazana
(only one female specimen; no calls heard, 730m altitude). The locality Marojezy mountains, above
1300 m altitude (Glaw & Vences 1994), refers to M. tandroka, which is described below. All additional
records of this species (Andreone 1994, Glaw & Vences 1994, Raselimanana 1998, Raxworthy et al. 1998)
are in need of confirmation (see also discussion).

Mantidactylus tschenki, spec. nov.
Figs 3-4
Types. Holotype: ZSM 936 / 2000 (formerly ZFMK 62298), adult male, collected along the road between Ambat-

olahy and Ranomafana, south-eastern Madagascar, on 28 February 1996 by F. Glaw, D. Rakotomalala and
F. Ranaivojaona. — Paratypes: ZFMK 62296 and 62297, adult males, collected close to the village Ranomafana

181

Fig. 3. Mantidactylus tschenki,

spec. nov. (holotype ZSM 936 /2000 in life, ventral view).

Fig. 5. Calling male of Mantidactylus cornutus from Andasibe.

Fig. 6. Calling male of Mantidactylus tschenki, spec. nov. from Ranomafana (Foto: F. Andreone)

(21°14'S, 47°26'E, ca. 550-600 m above sea level), same date and collectors as holotype. MRSN A379, adult male,

collected by F. Andreone at Ranomafana.

Diagnosis. M. tschenki, spec. nov. is characterized as a member of the genus Mantidactylus by presence

of femoral glands and lack of nuptial pads in males, and by its close similarity to Mantidactylus cornutus.
M. tschenki can be distinguished from that species by (a) the very different advertisement calls, (b) a
different shape of the inflated vocal sac in calling males (single subgular in cornutus, slightly bilobed
subgular in tschenki; Figs 5-6), (c) possibly by a slightly lower body size (SVL of adult males 38-40 mm
in cornutus, 35-36 mm in tschenki).

Description of the holotype

Adult male, SVL 36.1 mm. For measurements, see table 1. Body slender; head longer than wide,
slightly wider than body; snout pointed in dorsal and lateral views; nostrils directed laterally, slightly
protuberant, much nearer to tip of snout than to eye; canthus rostralis distinct, concave; loreal region
concave; tympanum distinct, elliptical (slightly higher than wide), 44% of horizontal eye diameter;
supratympanic fold present, straight; tongue ovoid, distinctly bifid posteriorly; vomerine teeth distinct,
in two rounded aggregations, positioned posterolateral to choanae; choanae rounded. Arms slender,
subarticular tubercles single; outer and inner metacarpal tubercles present; fingers without webbing;
relative length of fingers 1<2<4<3, finger 2 distinctly shorter than finger 4; finger disks distinctly

enlarged; nuptial pads absent. Hindlimbs slender; tibiotarsal articulation reaches widely beyond snout

tip; lateral metatarsalia partly connected; inner metatarsal tubercle distinct, outer metatarsal tubercle
not recognizable; webbing formula between toes 1 (1), 2i (2), 2e (1), 3i (2), 3e (1.5), 4i (2.75), 4e (2.5), 5 (1).
Skin on the dorsal surface smooth; back with indistinct and irregular dorsolateral folds; two distinct,
slightly elevated blackish tubercles between the eyes; a number of small granules and dermal spines
above the eyes; no distinct enlarged tubercles in the cloacal region; ventral skin smooth on throat,
slightly granular on belly. Femoral glands very poorly delimited and very indistinct from both external
and internal views; a patch of 10-15 small granules are visible from internal view.

Colour after four years in preservative rather uniformly greyish brown dorsally, with a dark brown
transversal band between the eyes which encloses the pair of blackish tubercles. Limbs with rather
indistinct brown crossbands. Head laterally with two narrow white stripes running from the eye to the
upper lip. Ventrally cream with irregular greyish brown pattern on belly and limbs. Throat (vocal sac)
greyish with a lighter mottling and one discontinuous and irregular median light stripe.

Colour in life. Colour slides are only available of the holotype ZSM 936/2000 (Figs 3-4). Dorsal
colouration and upper flanks brown. Three distinct dark brown crossbands were present on femur and
tibia. The iris was silvery with a reddish brown spot on its posterior edge. The pupil was horizontal.
A distinct white spot between posterior edge of eye and upper lip. The venter was dirty white with
brownish spots on the shoulder girdle, the ventral surface of hindlegs was brownish except for the
yellowish femoral glands. The throat was marbled brown and white.

Variation. The two paratypes correspond morphologically very well to the holotype. The throat lacks
a light median stripe in both specimens. Colouration of ZEMK 62297 is dorsally similar to the holotype
except the darker head sides and a more distinct narrow horizontal dark brown stripe underneath the

canthus rostralis and the supratympanic fold. In ZFMK 62296, a different colour pattern is present

which is also known in M. redimitus and M. cornutus: A distinct beige stripe runs (on each side of the
body) from the snout tip along the canthus rostralis, over the eyes, broadens as dorsolateral band along
the anterior back and finally makes up the whole of the flank colour on the posterior part of the body.

Habitat and habits. Calling males were sitting at night (end of February and beginning of March) on
vegetation ca. 1-2m above the bottom. They were found in primary rain forest and in degraded
vegetation as well. In at least one case no water body was recognized in the vicinity of the calling males.

Advertisement calls. Vocalizations were recorded at the type locality on 2 March 1996 at ca. 22°C air
temperature: Notes (Fig. 7) are unharmonious, distinctly pulsed and emitted in regular series. Tempo-
ral parameters were as follows: Note duration 274-335 ms (293 + 16 ms, n = 20), interval duration 1018-

2076 ms (1393+274ms, n=18). Notes consist of 16-21 pulses (18+1, n=20), the pulse rate is 58-68

(63 +2, n=20) per second. Frequency range is 1300-4000 Hz (dominant frequency 2500-2900 Hz, anoth-
er emphasized frequency band also from 1450-1550 Hz).

184

Frequenz (kHz)

.. ee

= aaunusbunnLruen. -

0 100 200 300 400 500 ms

Fig. 7. Sonagram and oscillogram of one note of Mantidactylus tschenki, spec. nov.

Distribution. Mantidactylus tschenki is reliably only known from the type locality. Several additional
specimens (and thus localities) possibly belong to M. tschenki, but a reliable attribution to either species
is not possible by morphology alone (see discussion).

Etymology. Mantidactylus tschenki is dedicated to Michael Tschenk, in recognition of his generous support to the
biosystematic research at the ZSM.

Relationships. Mantidactylus tschenki appears most closely related to Mantidactylus cornutus. Both
species are very similar by their morphology.

Subgeneric attribution. Mantidactylus tschenki is included in the subgenus Phylacomantis Glaw &
Vences, 1994 based on its similarity to M. cornutus.

Mantidactylus tandroka, spec. nov.
Figs 8-9

Types. Holotype: MNHN 1973.924, adult male, collected by Ch. P. Blanc on 2 July 1972 at the Marojezy massif,
1300 m altitude. — Paratypes: MNHN 1973.922, 1973.926-930, two adult males, three females and one subadult
specimen, same collecting data as holotype. MNHN 1973.912, adult male, collected on 29 November 1972 by Ch.
P. Blanc at the type locality. ZFMK 59894 and ZSM 937/2000 (formerly ZFMK 59895), two adult females,
collected by F. Glaw and O. Ramilison on 28 February 1995 at Marojezy, Campsite 4 (ca. 1300 m altitude).

Diagnosis. M. tandroka is characterized as a member of the genus Mantidactylus by presence of femoral
glands and lack of nuptial pads in males, and by its general similarities to Mantidactylus cornutus,
M. tschenki, and M. redimitus. It is morphologically most similar to M. cornutus and M. tschenki but can
be distinguished from these species by (a) different head proportions, with a shorter snout and a larger
head width relative to SVL (see Fig. 10), (b) presence of distinct pattern of longitudinal folds on the back
which is not found in any specimen of cornutus or tschenki: a pair of folds, absent in the other species,
starts behind the eyes and converges on the anterior back, fading in an area enclosed by the dorsolateral
folds. Some specimens of M. tandroka also remind the sympatric M. leucomaculatus; this species,
however, differs by lack of the tubercle pair between the eyes (black spots may be present but are never
prominent), lack of dorsal ridges, and the presence of laterally blackish vocal sacs in males.

Description of the holotype

Adult male, SVL 38.7 mm. For measurements, see table 1. Body slender; head longer than wide,
wider than body; snout slightly pointed in dorsal view, truncated in lateral view; nostrils directed
posterolaterally, slightly protuberant, much nearer to tip of snout than to eye; canthus rostralis distinct,

185

Fig. 8. Female paratype (ZSM 937 / 2000) of Mantidactylus tandroka, spec. nov. from Marojezy in life (dorsolateral
view).

straight; loreal region concave; tympanum distinct, elliptical (higher than wide), 50 % of horizontal eye
diameter; supratympanic fold very distinct, slightly curved; tongue ovoid, distinctly bifid posteriorly;
vomerine teeth distinct, in two rounded aggregations, positioned posterolateral to choanae; choanae
rounded. Arms slender, subarticular tubercles single; a paired outer and a single inner metacarpal
tubercles present; fingers without webbing; relative length of fingers 1<2<4<3, finger 2 distinctly
shorter than finger 4; finger disks distinctly enlarged; nuptial pads absent. Hindlimbs slender; tibiotar-
sal articulation reaches distinctly beyond snout tip; lateral metatarsalia partly connected; inner meta-
tarsal tubercle distinct, outer metatarsal tubercle very small, almost not recognizable; webbing formula
between toes 1 (1), 2i (2), 2e (1), 3i (2), 3e (1.25), 4i (2.75), 4e (2.5), 5 (1). Skin on the upper surface smooth;
back with many irregular smaller folds, arranged as a discontinuous network; a pair of distinct folds
runs from behind the eyes, converge centripetally onto the anterior back, curves slightly towards the
flanks and fades; laterally from these, a pair of dorsolateral folds runs from ca. 4 mm behind the
supratympanic fold to the inguinal region. Two distinct, blackish tubercles between the eyes; anumber
of granules and small dermal spines above the eyes; no distinct enlarged tubercles in the cloacal region;
ventral skin slightly granular on belly, smooth on throat where the presence of a single vocal sac is
clearly recognizable. Femoral glands very poorly delimited and very indistinct from both external and
internal views; a patch of 16 small granules are visible from internal view.

Colouration dorsally brownish, slightly lighter on the back and head in an area delimited by the
dorsolateral folds. Limbs light brown with brown crossbands: three to four bands on forelimb, four to
five on femur, four to six on tibia, five to six on foot and tarsus. Flanks brown fading into cream towards
the belly. Head laterally brown, with a distinct horizontal dark brown stripe running underneath the
canthus rostralis and the supratympanic fold. Two light vertical stripes run from underneath the eye
to the upper lip. Lower lip brown with five narrow light vertical markings. Ventrally brownish with
irregular light mottling on the throat (vocal sac), cream on the remaining surface, with brown mottling
in the breast region and on the hindlimbs.

186

Sr ER) 3 Fi x ae X x a E; 8 N x
Fig. 9. Female paratype (ZSM 937/2000) of Mantidactylus tandroka, spec. nov. from Marojezy in life (ventral
view).

RER

Colour in life. Colour slides of living specimens are only available of the female paratype ZSM 937 /
2000 (Figs 8-9). The dorsal colouration was light brown to beige, the flanks were light brown fading into
pinkish towards the venter. The limb crossbands were greyish brown. A black stripe below the canthus
rostralis from snout tip to eye and below the supratympanic fold. Tympanum dark brown. The iris was
light yellowish brown in its upper third, dark reddish brown in its two lower thirds. The pupil was
horizontal. The venter was partly transparent with a pinkish shade, more cream coloured in its
posterior part and on the hindlimbs. The throat was dirty white and unspotted.

Variation. Morphologically, the paratypes agree well with the holotype. The general impression of a
broad and relatively short head is very typical in all available specimens. Two specimens (MNHN
1973.927-928) show light stripes from snout tip along canthus rostralis and above eye. These stripes
become broader immediately behind the eyes and along the supratympanic fold, making up the entire
flank colouration from the forelimb insertion on. The distinct and sharp colour border between light
flanks and dark dorsum is situated slightly below the dorsolateral folds. A similar pattern is known in
M. redimitus and M. cornutus and described above for one M. tschenki paratype. In these species,
however, the light colour on the flanks is less extended. The expression of this pattern thus may
constitute another relevant diagnostic character of M. tandroka. The male MNHN 1973.926 shows very
distinct broad cream-white vertical bands running from the anterior and the posterior eye corners to
the upper lip, respectively, reminding the pattern found regularly in M. leucomaculatus. Generally, in
most preserved specimens (also in the females ZFMK 59894, although not in the figured specimen ZSM
937/2000; Figs 8-9) the throat shows an intense brown pattern, as opposed to the greyish colour in
M. redimitus, M. cornutus, and M. tschenki. As far as visible in the preserved males the vocal sac is single
subgular although a slightly bilobed shape as in M. tschenki can not be excluded before observations
of calling males become available. Altogether, male SVL in the type series was 38.7-41.4 mm
(39.3+1.2 mm, n=5), female SVL 39.6-44.7 mm (41.83+2.2 mm, n=5). Mean male size was 94 % of mean
female size, sexual size dimorphism was statistically significant (U-test, P<0.05). Mean relative tym-
panum size did not differ significantly between sexes (U-test, P>0.9), while intersexual differences in

187

ratio HW/SVL

0.36

0.34
<] det. uncertain

@ M. tandroka
A M. tschenki

0.32
& M. cornutus

O M. redimitus
0.8 1.0 1.2 1.4 1.6

ratio END/NND

Fig. 10. Scatterplot of HW/SVL ratio vs. END/NND ratio in Mantidactylus tandroka, M. cornutus and M. tschenki,
and M. redimitus showing separation of M. tandroka from the remaining specimens by head shape parameters.

relative length and height of inner metatarsal tubercle (means of both values larger in males) were
significant (U-test, P< 0.05).

Habitat and habits. ZFMK 59894 and ZSM 937/2000 were collected during the day on the ground in
primary rainforest. Advertisement calls are unknown.

Distribution. Mantidactylus tandroka is only known from the type locality, around 1300 m altitude.

Etymology. Derived from tandroka (Malagasy: horn), referring to the two horn-like tubercles between the eyes
of this and related species (see also the etymology of M. cornutus). The name is considered as invariable noun
standing in apposition to the generic name.

Relationships. Mantidactylus tandroka appears most closely related to Mantidactylus cornutus and
M. tschenki.

Subgeneric attribution. Mantidactylus tandroka is included in the subgenus Phylacomantis Glaw &
Vences, 1994 based on its similarity to M. cornutus and M. tschenki.

Discussion

Vouchers of uncertain attribution. When preserved, M. cornutus and its new sibling species M. tschenki
are virtually indistinguishable by morphological characters. Therefore, the identity of specimens from
four localities (Ranomafana, Andringitra, Col Ivohibe, and Chaines Anosyennes) attributed in Glaw &
Vences (1994) to M. cornutus is uncertain and commented as follows: (1) The Ranomafana specimen
(ZFMK 50593) was defined as paratype of M. cornutus by Glaw & Vences (1992b). Itis a female of rather
large SVL (40.7 mm) in comparison to the four available specimens of M. tschenki (all males). Despite
of this size difference, it may belong to M. tschenki, but further fieldwork is necessary to confirm or
reject the presence of both species in the Ranomafana area. (2) The specimens from the Chaines

Anosyennes, Campsite 4 (MNHN 1972.1471) and Chaines Anosyennes, Ambana (MNHN 1972.1472)

are two males in good state of preservation which are larger than the available specimens of M. tschenki;

furthermore they have a pair of distinct tubercles on the central dorsum lacking in M. tschenki (but

present in the Ivohibe specimens). (3) The three specimens from Col d’Ivohibe, 1400 m altitude (MNHN
1953.74, 1991.2940-2941) are in mediocre state of preservation but agree better with M. cornutus than

188

ratio IMTL/SVL

0.06 on
2 (AN ©)
AR J
GI
females N “ ‘ oO
Br oO \ { U) 6) s®
0.04 HZ \ “DIS
182 4 \ u males <] det. uncertain
_ ° ö \ ® M. tandroka
dee e\ A M. tschenki
A Be © M. cornutus
0.02 — : O M. redimitus
0.01 0.02 0.03 0.04 0.05

ratio IMTH/SVL

Fig. 11. Scatterplot of relative length and height of the inner metatarsal tubercle (ratios IMTL/SVL and IMTH/
SVL) in Mantidactylus redimitus, M. cornutus, M. tschenki, and M. tandroka, showing sexual dimorphism in IMT
size. Original measurements from Tab. 1 (except type of M. redimitus).

M. tschenki regarding body size. (4) The two Andringitra specimens (MNHN 1972.571-572) are rather
small and in bad state of preservation; they are females, and it can not be verified whether they are
actually full-grown adults (only immature oocytes observed after dissection). Summarizing, more
fieldwork is necessary to clarify the identity of these four populations.

Distribution pattern in Phylacomantis. Although one of the new species described in the present
paper (M. tschenki) is only known from south-eastern Madagascar, the new data which have been
gathered during the last years clearly show that the subgenus Phylacomantis has its center of diversity
and endemism in northern Madagascar which together make up less than % of the Malagasy territory.
Including a further new Phylacomantis species from the central east (Andasibe) and Marojezy which is
being described elsewhere, only three out of nine Phylacomantis species (M. cornutus, M. tschenki,
M. corvus) are not known from northern Madagascar and three species appear to be northern endemics
(M. granulatus, M. pseudoasper, M. tandroka). The highest diversity is found in the Marojezy massif
where six species occur: M. redimitus, M. tandroka, M. leucomaculatus, M. granulatus, M. pseudoasper, and
the new species which is being described in a separate paper. This high degree of sympatry is in part
characterized by a restricted altitudinal distribution at Marojezy (e.g. M. tandroka to the high elevations
above 1000 m, M. granulatus mainly at low altitudes). Such an altitudinal segregation may be one factor
favouring the extraordinary anuran diversity in Madagascar and the high degree of range overlap
among closely related species.

Shared characters in Phylacomantis. The new morphological and bioacoustic data presented allow
for the discussion of some general trends in the subgenus Phylacomantis. In two species in which males
and females were available (M. redimitus, M. tandroka), a sexual dimorphism in size of the inner
metatarsal tubercle was noted, males having distinctly larger tubercles than females. Actually, no
overlap between males and females was noted even pooling the data of all four species studied
(Fig. 11). Beside the two mentioned species, the sexual dimorphism of inner metatarsal tubercle within
the genus Mantidactylus is only known in M. granulatus and M. leucomaculatus (Glaw & Vences 1994)
and some species of the subgenus Gephyromantis (pers. obs.), and may be a synapomorphy of this group
of species. On the other hand, sexual dimorphism in tympanum size appears to be absent in the species
studied here. Such a dimorphism is typical for several Mantidactylus, especially species in the brook
edge dwelling subgenera (e.g. Brygoomantis, Chonomantis, Ochthomantis), and is also found in
M. pseudoasper (Glaw & Vences 1994), amember of the subgenus Phylacomantis.

189

There is one other character which seems to be restricted to several species of the subgenus
Phylacomantis and to some members of the subgenus Gephyromantis: The laterally banded morph is only
known to occur in M. redimitus, M. cornutus, M. tschenki and M. tandroka (Phylacomantis), as well as in
M. asper and M. luteus (Gephyromantis) (pers. obs.). All four mentioned Phylacomantis species have also
dermal spines above the eye, a character shared with another undescribed Phylacomantis species, some
species of the subgenus Gephyromantis (M. asper, M. spinifer), and species of the subgenus Spinomantis
(e. g. M. phantasticus).

Advertisement calls of Phylacomantis, as far as known, are composed of a single note type, although
notes can be arranged in groups. The four species studied here share with each other and with
M. granulatus and M. leucomaculatus calls with further structural similarities: They consist of rather
regular series of unharmonious notes with a note duration of at least 50 ms and a pulsed structure,
possibly reflecting relationships between these species which were placed in aM. granulatus group by
Glaw & Vences (1994).

Acknowledgements

We are especially grateful to F. Andreone (Torino) who provided the photo of the calling male of M. tschenki.
D. Rakotomalala, ©. Ramilison and F. Ranaivojaona assisted in the field. W. Herwig provided a photographic
record of M. redimitus from Vohidrazana. Thanks go also to W. Böhme (ZFMK, Bonn) and A. Dubois and
A. Ohler (MNHN, Paris) for the loan of type material. The field work was made possible by a cooperation accord
between the Zoological Institute at the University of Antananarivo (UADBA) and the “Zoologisches For-
schungsinstitut und Museum A. Koenig” (ZFMK,). Financial support was granted both to the field work of F.
Glaw and the work of M. Vences in the MNHN by the “Deutscher Akademischer Austauschdienst” (DAAD).

References

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190

Buchbesprechungen

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24. Laibner, S.: Elateridae of the Czech and Slovak Republics. —- Kabourek, Zlin, 2000. 292 pp. 519 figs, 9 colour
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After a short introduction about morphology of adults and larvae, biology, agricultural significance, collecting,
rearing, and classification, the taxonomic section includes keys to subfamilies, genera, and species. Each taxon
is shortly described, and for species some information about ecology and distribution in the covered area is
presented. The keys are profusely illustrated with many rather simple but clear and informative line drawings.
Most species are figured on one of the 9 colour plates that finish the taxonomic section. A list of species,
references, and an index are added.

A nice piece of work that certainly will be of importance not only to Czech and Slovakian users, though for
all workers who are interested in the central European elaterid fauna. M. Baehr

25. Michener, C. D.: The Bees of the World. - The Johns Hopkins University Press, Baltimore & London, 2000.
XIV + 913 S, zahlreiche Abb. ISBN 0-8018-6133-0.

Dieses grundlegende Werk über die Bienen der Welt hat in seinem ersten Teil ausführliche, einführende Kapitel
von allgemeinerem Interesse zum Beispiel über die Coevolution der Bienen und Pflanzen, ihre soziale Lebens-
weise, die Morphologie, Systematik und Biogeographie der Bienen. Im zweiten, systematischen Teil des Buches
wird auf etwa 700 eng bedruckten Seiten ein umfassender Überblick über die Bienen der Welt gegeben. Es
werden alle Taxa bis hinunter zu den Untergattungen dargestellt und klassifiziert. Dabei werden Bestimmungs-
schlüssel, morphologische Besonderheiten und interessante Hinweise zur Biologie und Verbreitung zusammen-
gestellt. Insgesamt sind etwa 1200 Gattungen und Untergattungen systematisch eingeordnet und behandelt.
Dabei wurden verschiedene taxonomische Änderungen — neue Gattungen, neue Namen und Synonyme -
durchgeführt. Darüber hinaus wird weltweit die wichtigste Literatur zitiert und kommentiert. Unter den
zahllosen Illustrationen bestechen neben vielen Habituszeichnungen besonders die hervorragenden rasterelek-
tronenmikroskopischen Abbildungen, die die morphologische Vielfalt demonstrieren.

Das Buch ist gleichzeitig eine Einführung in die Biologie und Systematik der Bienen und ein umfassendes
Nachschlagewerk. Es ist das Lebenswerk eines der wirklich großen Entomologen des 20. Jahrhunderts und
zweifelsohne ein Meilenstein der Apidologie. K. Schönitzer

191

Buchbesprechungen

26. Turin, H.: De Nederlandse Lookkevers. Verspreiding en Oecologie (Coleoptera: Carabidae). - Nederlandse
Fauna 3.-Nationaal Natuurhistorisch Museum Naturalis, KNNV Uitgeverij & EIS-Nederland, Leiden, 2000.
666 S., 16 Farbtaf., CD-ROM. ISBN 90-5011-136-X.

Dies ist ein monumentales Werk über die Laufkäfer der Niederlande, das eigentlich nicht mehr viele Fragen
offenläßt. Der allgemeine Teil umfaßt allein 124 großformatige Seiten und beleuchtet eine Vielzahl von Aspekten
der Taxonomie, Biologie, Ökologie und Verbreitung der Laufkäfer, aber auch der Fang- und Untersuchungsme-
thoden. Dieser Teil ist umso wertvoller, weil Laufkäfer eine der wichtigsten Gruppen in feldökologischen oder
ökofaunistischen Untersuchungen darstellen. Ein beträchtlicher Teil dieser Untersuchungen wurde überdies in
den Niederlanden durchgeführt - so erhält dieses Buch einen zusätzlichen Wert.

Im speziellen Teil werden alle in den Niederlanden vorkommenden Laufkäfer ausführlich mit ihrer Gesamt-
verbreitung, der Verbreitung in den Niederlanden, ihrer Ökologie und Biologie, sowie ihrer Gefährdung
behandelt, wobei die Gesamtverbreitung in gerasterten Karten, die Verbreitung in den Niederlanden durch
Punktkarten dokumentiert ist. Diagramme zur Habitatbindung sowie für den Fortpflanzungstyp sind gleichfalls
für fast alle Arten beigegeben. Ein umfassendes Literaturverzeichnis, eine ausführliche englische Summary,
sowie ein Register beschließen das Werk. Zusätzliche Informationen können von der beigefügten CD-ROM
entnommen werden. Einige ausgezeichnete Habitusfotos verschiedener Laufkäfer lockern den dicken Band auf.
Außerdem sind von den meisten Gattungen noch ein bis zwei charakteristische Vertreter in vorzüglichen
Schwarzweißzeichnungen dargestellt.

Das Buch kann als Kompendium der Biologie und Ökologie der mitteleuropäischen Laufkäfer angesehen
werden, denn ein beträchtlicher Teil der darin enthaltenen Informationen sprengt den vorgegebenen geographi-
schen Rahmen bei weitem. Das Werk hat nur einen einzigen Nachteil: es ist durchweg in niederländischer
Sprache verfaßt, was die Benutzung für den deutschen Leser doch - jedenfalls zunächst - erschwert. Allerdings
kann man sich verhältnismäßig rasch einlesen.

Dieses Buch ist sicher ein Muß für alle Ökofaunisten, sowie für all diejenigen, die sich in irgendeiner Weise
mit mitteleuropäischen Laufkäfern befassen. Gemessen am Umfang und an der Fülle der enthaltenen Informa-
tionen erscheint der Preis durchaus mäßig. M. Baehr

27. Barnard, P. C. (Hrsg.): Identifying British Insects and Arachnids: an Annotated Bibliography of Key Works.
- Cambridge University Press, 1999. XIII + 353 S., ISBN: 0-521-673241-2.

In diesem Werk werden die verschiedenen Ordnungen von Insekten, die in Großbritannien vorkommen, kurz
vorgestellt und die Literatur zitiert, die zur Bestimmung der jeweiligen Gruppe dient. Sehr hilfreich sind kurze
Anmerkungen, die die Literatur kommentieren. Darüber hinaus enthält das Werk auch je ein Kapitel über
Pseudoskorpione, Opiliones, Acari und Spinnen. Obwohl das Buch auf die britische Fauna ausgerichtet ist, sind
die Angaben oft für ganz Europa von Wert und helfen nicht nur Entomologen, die sich speziell für die britische
Fauna interessieren. Hilfreich und von allgemeinem Wert ist auch die Einführung über die verschiedenen
Informationsquellen in der Entomologie. K. Schönitzer

28. Cole, T. C. H.: Wörterbuch der Tiernamen. Latein - Deutsch — Englisch. Deutsch - Latein - Englisch. —
Spektrum Akademischer Verlag, Heidelberg & Berlin, 2000. 970 S., ISBN 3-8274-0589-0.

Dieses Werk enthält die Namen von 16.000 Tieren in lateinischer, englischer und deutscher Sprache. Es enthält
sowohl wissenschaftlich korrekte Bezeichnungen, als auch sogenannte Trivialnamen und Synonyme. Verständ-
licherweise sind vor allem Säugetiere und Vögel sowie Nutztiere besonders gut vertreten. Aber auch Insekten
und andere Wirbellose kommen nicht zu kurz. Ein unentbehrliches Nachschlagewerk für alle, die Tiere richtig
beim Namen nennen müssen. Das Werk ist auch mit einer CD-Rom erhältlich, die natürlich über noch mehr
Suchmöglichkeiten als die gedruckte Version verfügt. Ein Nachschlagewerk, das in keiner zoologischen Biblio-
thek fehlen darf und das in Zukunft hoffentlich viele Übersetzer und Autoren populärwissenschaftlicher Werke
vor oft so peinlichen Fehlern bewahren wird. K. Schönitzer

192

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SPIXIANA München, 01. Juli 2001 ISSN 0341-8391

INHALT - CONTENTS

Seite

Prpic, N.-M.: A new species of Loxosoma from north-western Finistere, France
(Spiralia, Kamptozoa (= Entoprocta), Solitaria, Loxosomatidae) ..... 97-102

Alf, A. & K.Kreipfl: A new Architectonica from the Philippines (Mollusca, Gastropoda,
Architectonieidae)......2...rt 04:4 ee ee ee anuun: 103-106

Kreipl. K.& A. All: A new species of Turbinidae Rafinesque, 1815 from the northern
Red Sea (Mollusca, Gastropoda) ................u.4nsneennnnnnennnnennennnnnennn 107-110

Leistikow, A.: Designation of atype species for the genus Prosekia, gen. nov. from
South America (Crustacea, Isopoda, Oniscidea) ............................. 111-121

Rheims, C. A. & A. D. Brescovit: Three new species of litter inhabiting spiders of the genus
Scytodes Latreille from northeastern Brazil (Araneae, Scytodidae) 123-128

Molineri, C©.: Traverhyphes: a new genus of Leptohyphidae for Leptohyphes indi-

cator and related species (Insecta, Ephemeroptera) ....................... 129-140
Vieira-Lanero, R., M. A. Gonzälez & F. Cobo: The larva of Hydropsyche urgorrii Gonzälez

& Malicky, 1980 (Insecta, Trichoptera, Hydropsychidae) ................. 141-146
Ingrisch, S.: Tetrigidae from Nepal in the Zoologische Staatssammlung München

(Insecta, Orthoptera, Tetrigidae) .........................2urr2200r2n0nnennonnennnnenn 147-155
Arndt, E. & D. W. Wrase: Description of two new species of Notiobia Perty from Southern

Venezuela (Insecta, Coleoptera, Carabidae, Harpalini) ................... 157-163
Baehr, M.: A new species of the genus Lissopogonus Andrewes from northern

Borneo (Insecta, Coleoptera, Carabidae, Patrobinae) ..................... 165-169
Baehr, M.: A new species of the genus Minuthodes Andrewes from North

Queensland, Australia (Insecta, Coleoptera, Carabidae, Lebiinae). 171-175

Glaw, F. & M. Vences: Two new sibling species of Mantidactylus cornutus from Madagas-
car (Amphibia, Anura, Ranidae) .....................2.u22200000000200002000nnennnnenn 177-190

Büchbesprechungens mem 4... Amuneee een 2... racear ne penneneneneen 122, 156, 164, 170, 176, 191-192

Zeitschrift für Zoologie

SPIXIANA ° Band 24 » Heft3 » 193-288 « München, 01. November 2001 * ISSN 0341-8391

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SPIXIANA 193-202 een NoENorember 2001 ISSN 0341-8391

Proceedings of the FORUM HERBULOT 2001
Neotropical Geometridae: Approaches to a Modern Concept
of the Geometrid System on Genus and Tribe Level
(8.3.-9.3.2001)

Axel Hausmann & Robert Trusch (ed.)

Hausmann, A. & R. Trusch (ed.) (2001): Proceedings of the Forum Herbulot
2001; Neotropical Geometridae: Approaches to a Modern Concept of the Geo-
metrid System on Genus and Tribe Level (8.3.-9.3.2001). — Spixiana 24/3: 193-202

The objectives, a list of participants and a short report on the results of the Forum
Herbulot 2001 is presented emphasizing the great impact that this meeting had for
coordinated, modern research in Geometridology and for creating a worldwide,
IT-based network of scientists working on Geometridae. The abstracts of nine
lectures from the seminary session of the Forum Herbulot are added.

Axel Hausmann, Zoologische Staatssammlung München, Münchhausenstr. 21,
D-81247 München, Germany. E-mail: Axel.Hausmann@zsm.mwn.de

Robert Trusch, Zoologische Staatssammlung München, Münchhausenstr. 21,
D-81247 München, Germany. E-mail: trusch@zsm.mwn.de

Objectives

Research on zoological systematics at the ZSM and its focus on Geometridae:
towards a worldwide network of Geometrid researchers (from the ‘introduction’, shortened).
Axel Hausmann, ZSM

Claude Herbulot (Paris) has been known as the author of an amazing number of publications contrib-
uting valuably to the knowledge of taxonomy and systematics of the Geometridae of the world. His
famous collection is now housed in the ZSM.

The fine collection as well as the productive enthusiasm of Claude Herbulot is a strong incentive for
the pursuit of effective scientific work on Geometridae in the ZSM as regards the systematic order,
taxonomical questions, biogeography and evolution, but also the use of IT in research and collection
management. The FORUM HERBULOT shall focus on problems that can only, or better, or quicker, be
solved by the personal co-operation of the leading specialists of our days. FORUM HERBULOT is
therefore not meant to be just another congress but shall offer an opportunity for Geometridae-
specialists to archieve tangible results that enhance top level progress in the field of Geometridology.

The ZSM houses one of the largest Lepidoptera collections in the world. Many years ago, such
museums ceased to be part of ‘Royal Natural Curiosity Cabinets’, or collections for their own sake that
can be mockingly considered as ‘Zoological Registration Offices’ (Autrum), but have become indis-
pensable partners in research into biodiversity, biogeography, evolution and systematics. In system-
atics (with correlating taxonomy) research is almost exclusively left to the museums; in German
universities at least there are now very few professorships in systematic zoology.

Our responsibility for systematic research is a great challenge. Most museums - as a result of
having far too few personnel — are confronted with serious problems with the arrangement and
curation of the collection (collection management) and with making the collection accessible by modern

IT (cataloguing). On the other hand, computer techniques, methods of molecular biology, and recent,
refined results from geology, climatology and palaeontology offer exciting new possibilities for system-
atic research. Coordination of the limited resources is strongly needed. Under the given conditions,
continuous mutual exchange of opinions and information between specialists on the base of personal
contacts may be the best way to achieve successful progress in systematic research. Neither regional
nor professional frontiers should play a limiting role here.

The work in the Lepidoptera department of the ZSM is particularly and strongly focussed on
Geometridae. For this reason the ZSM was proposed as the organisational framework for a forum for
a worldwide network of Geometridae specialists. The name “Forum Herbulot” has been chosen in
order to express the admiration and the deep respect for the magnificent lifework of a Grand-Master
of our Geometridae guild, but mainly, because the name Herbulot symbolises today in a unique way,
what must be regarded as the most important prerequisite for advanced systematic research, i.e.
profund knowledge of Geometridae in all the continents. The outstanding and beautiful collection
Herbulot with so many types and so many correlated publications forms an extraordinarily fruitful
basis for our work.

I am very glad, that our friend Herbulot was able to accept the invitation. So, his honorary lecture
on South American Eupithecia will be a highlight of the Forum, and we have the opportunity to give
him our warmest congratulations for his 93rd birthday, which he celebrated just 3 weeks ago. I have
to admit, that this factor played a certain role in choosing this date for our Forum.

Participants

Chairman A. Hausmann, ZSM, Munich

Opening address G. Haszprunar, director of the ZSM, Munich

Gunnar Brehm, University of Bayreuth, Germany; Charles V. Covell, University of Louisville, U.S.A.;
Philippe Darge, Clenay, France; Sven-Ingo Erlacher, University of Jena, Germany; Claude Herbu-
lot, Paris, France; Igor Kostjuk, University of Kiew, Ukraine; Martin Krüger, Transvaal Museum
Pretoria, South Africa; Andreas Kunkel, ZSM (Generaldirektion); Michael Miller, ZSM; Vladimir
Mironov, Acad. Sci. Russ., St. Petersburg, Russia; Linda Pitkin, The Natural History Museum,
London, U.K.; Peder Skou, Apollo Books, Stenstrup, Denmark; Manfred Sommerer, München,
Germany; Dieter Stüning, Zoologisches Forschungsinstitut und Museum A. Koenig, Bonn, Germa-
ny; Robert Trusch, ZSM; Janusz Wojtusiak, Jagiellonian University, Kraköw, Poland.

194

Short Report and Results
A. Hausmann, R. Trusch, C. Covell, M. Krüger, L. Pitkin

Aims of the FORUM HERBULOT as outlined by the chairman were agreed upon by the partici-
pants. The need for closer scientific cooperation among geometrid researchers was expressed.
The seminar session highlighted promising possibilities for systematic research. The importance of
Gondwana studies was stressed by M. Krüger. Generic revisions of Ennominae (neotropical) by
L. Pitkin prompted the question of tribal definitions which must be consistent in the other faunistic
regions as well. A tentative tribal classification would seem worth working out and helpful.
Research on Sterrhinae and Eupitheciini could be broadened to an interfaunistic scale (C. Covell;
A. Hausmann; V. Mironov) and thus bridge existing gaps (e.g. Eupithecia in Africa as correlated to
Asia and/or other regions).

Molecular methods (e.g. working groups in Sweden and at the ZSM) need refining to form a more
powerful tool for evolutionary and systematic studies and should supplement the morphological
and ecological context.

A linkage between the GEDIS transects in Ecuador (DFG) and the fascinating altitudinal border-
lined sections of the Erateina-research (J. Wojtusiak) appears very desireable.

A proposal to continue the FORUM HERBULOT at the ZSM in 2003 was welcomed. An internet
facility should be provided at the ZSM as a platform to continue cooperation between researchers
and specialists; A. Hausmann, L. Pitkin, C. Covell, M. Krüger will supervise the goals and ways and
means to achieve them.

Participants expressed their thanks to the organizers and sponsors of the FORUM HERBULOT 2001
and enjoyed the pleasure of attending the ceremony of the awarding of the Spix-Medal to Claude
Herbulot.

195

Abstracts
(Lectures of the Seminar Session)

On Neotropical Eupithecia
Claude Herbulot, 67, rue de la Croix Nivert, F-75015 Paris, France

Eupithecia is a genus with a very large number of species, in fact with the greatest number of species
within the family Geometridae: 1,332 if we only consider the species and ignore the subspecies that
have been listed by Malcolm J. Scoble in his inventory in 1999. However, to be honest I don’t always
agree with his classification of certain taxa as species or subspecies.

The genus Eupithecia is represented throughout the whole world as you can see on the globe on the
transparency where red stripes show all the regions where the genus is known to exist. This picture
is misleading in one respect, in that it fails to show correctly the important facts that the genus is well
represented in Japan, Taiwan and Yunnan but only sparsely in Indochina, Malaysia and in the Sundas
and that there are only two species in Australia and none in New Zealand.

Eupithecia species can be found from sea level to altitudes reaching about 4,000 metres. But, even
though in moderate climates you can find them as well in the low lands as in the mountains, the rule

is that in tropical regions you only find them in higher altitudes (starting at about 1000 metres).

Attempts have been made to divide such a large genus into smaller groups but, it is a very |
homogenous group, and the elements subject to differentiation are merely features of little importance
such as a double areole instead of a single one in the neuration of the wings or differences in the |

structure of the antennae of the males. These variations are, as you know, not strong enough to define
a genus. That is why none of the attempts to split the genus have succeeded.

Up to now, 352 species of Eupithecia are known in the Neotropics, i.e. more than a quarter of all '
known species, their distribution being very scattered: 44 species have been found in South Mexico, 39 |

in Central America, 9 in the Antilles, 9 in Venezuela (including Trinidad, Guyana, Surinam and French
Guiana), 149 in Columbia, Ecuador, Peru and Bolivia (4 countries that I pool because a great many of

the species found there are found in all of these countries), 26 in Brazil, 64 in Chile and 18 in Argentina |

and the Falkland Isles. However, this is far from being the true number of species in the region, as the
investigation in most of the countries is only just starting. This is particularly true for the four central

Andean countries. I estimate that the number of known species in these countries is barely half of all
the species actually present. Strong support for this opinion can be found inmy own collection in which

the species from Ecuador and Peru are well represented and where the number of species I was able

to label with a scientific name is more or less the same as the number of species I was unable to

determine.

The males of some Eupithecia species in the Neotropics show special features which are found |
nowhere else, as you will be able to see on the drawings on the next slides. The hindwings can be |

shortened, cut shorter, or more or less peaked. The forewings, as well as the hindwings, may have
fossules and swellings on their upper surfaces as well as on their undersides. The underside of the
forewings sometimes has long patches with hairs. As a rule the species showing such features seem to
be mainly species from lower and intermediate altitudes, even though I also caught one at 3,200 metres.
Numerous genera have been created for all these species but, as I already told you, they cannot be
considered as valid: the females of these species do not show any special features and the male genitalia
are of the same type as those of all the other species of the genus.

It follows a characterisation of the seven groups of countries to which the author attributed all the |

neotropical Eupithecia species (South Mexico; Central America; Antilles; Venezuela, Trinidad, Guyana,
Surinam, French Guiana; Columbia, Ecuador, Peru, Bolivia; Brazil; Chile; and Argentina, Falkland
Islands).

The genus Eupithecia is well represented in South America, perhaps even better than in the rest of
the world, if one considers the quota of species still to be discovered. Furthermore some neotropical

species show special morphological features which are without parallels in any other region. Can we |

use these findings when searching for the place on earth where the genus originated? I do ask this |

question but I do not dare to answer to it.

196

Suprageneric Classification of the Ennominae: The Neotropical Component
Linda Pitkin, The Natural History Museum, Department of Entomology,
Cromwell Road, London SW7 5BD, U.K. E-mail: Imp@nhm.ac.uk

The subfamily Ennominae is the largest in the Geometridae, comprising approximately 10,000 species
(about half of the total number in the family) in 1100 genera. Many of the genera are

poorly defined, and the classification above that level is still inadequately resolved. Studies in
recent decades, particularly by Holloway ([1994], focussing on the Bornean fauna), Rindge, Scoble, and
other authors) have helped to improve the definition of various tribes and the taxa included in them,
but much more study is needed.

The Neotropical component of the Ennominae is species-rich, with about 3300 species in about 300
genera. These genera have been the subject of a nearly completed review (Pitkin, in prep.), and their
placement within tribes, reviewed in the same work, is considered here, together with some of the
defining characters of those tribes. Study of the Neotropical fauna generally supports the usage of
family-group names within the Ennominae as reviewed by Holloway ([1994]), but a few further
synonyms have been found amongst these names.

Prior to recent decades, many Neotropical genera were dubiously assigned to tribes, or not at all.
Certain tribes have benefited enormously from more recent revision, notably the Nacophorini and
Lithinini (Rindge, 1983 and 1986 respectively), the Palyadini (Scoble, 1995), and the Macariini (Scoble
& Krüger, in prep.). The current review of the Neotropical Ennominae broadens the scope of a number
of tribes by newly assigning many genera to them. This is most marked in the tribes Ourapterygini
(which gains 20 Neotropical genera) and Cassymini (which gains 10 Neotropical genera). The supra-
generic classification of the Ennominae requires further resolution and some of the tribes may not be
monophyletic.

Ennomine tribes represented in the Neotropical Region

Total number of Neotropical genera Newly assigned
Azelinini 2 0
Caberini and Baptini 15 6
Boarmiini 16 %
Melanolophiini [subgroup of Boarmiini] 10 0
Cassymini 14 10
Lithinini 12 3
Macariini 3 0
Nacophorini 43 11
Nephodiini 12 7
Ourapterygini 40 20
Palyadini 6 0
‘Cratoptera group’ 9 9

More than 60 Neotropical genera remain unplaced. Ennomine tribes with Neotropical representation
probably excluded now are Angeronini, Crocallini, and Campaeini.

On the tribal classification of southern African Ennominae (Lepidoptera, Geometridae)
Martin Krueger, Lepidoptera Dept., Transvaal Museum, NFI, Pretoria, South Africa.
E-mail: kruger@nfi.co.za

Since the 1930’s, the geometrid fauna of southern Africa has been considered as relatively well known.
This notwithstanding, revisions published during the past decade have shown that the subregion is
inhabited by an unexpectedly high number of Ennominae, with 545 species in 104 genera thus far
described. The three largest tribes, Macariini, Ennomini, and Boarmiini comprise more than 40 % of
genera and 67 % of species. Examples of less well represented groups include Cassymini, Caberini,
Diptychini, Abraxini, Eutoeini, Lithini, and Gnophini.

The distribution of these taxa on both the species- and genus levels is highly skewed and of
zoogeographic interest. The distribution of Macariini is concentrated in the savannas of the northern

197

and eastern parts of the area. This pattern is in accordance with the utilization of Fabaceae, especially

the species-rich genus Acacia, by most of its representatives. Conversely, boarmiine diversity is |
centered around afromontane forests, although the tribe also enjoys a substantial representation in the |

savanna biome. Perhaps most interesting from a phylogenetic point of view are Ennomini, which

inhabit mainly the macchia-like Fynbos areas of the Western Cape Province and the semi-arid Karoo.

In the northern parts of southern Africa Ennomini are largely montane, which, supported by geological
and phytogeographical data, points towards an ancient origin of this group.

Studies on the Neotropical Sterrhinae (Geometridae)
and Correlation of Species with those in the North American Fauna
Charles V. Covell Jr., University of Louisville, 40292-0001 Louisville (Ky), U.S.A.
E-mail: covell@louisville.edu

Status of North American studies on Sterrhinae:

A. Moths of America North of Mexico series, Fascicle 18.1: progress report

B. Synonymies of species-group names of species in United States and also in Mexico and other
Central and South American countries

C. Un-named or unreported N. American/neotropical species additional to Covell (1983)

D. Currently known Sterrhinae genera and species fauna of North America — Sterrhini (species
numbers in brackets; total species: 100): Eumacrodes Warren (1), Euacidalia Packard (5), Protoproutia
McDunnough (2), Lobocleta Warren (7), Idaea Treitschke (30), Paota Hulst (1), Pigia Guenee (2),
Odontoptila Warren (1), Ptychamalia Prout (1) - Cosymbiini: Pleuroprucha Möschler (2), Cyclophora
Hübner (7), Semaeopus Herrich-Schäffer (5) -— Timandrini: Haematopis Fabricius (1), Calothysanıs
Hübner (1) - Scopulini: Acratodes Guenee (1), Scopula Schrank (24), Leptostales Möschler (8), Lophosis |
Hulst (1). |

|
Status of work on the neotropical Sterrhinae: |
A. Most species are known only from original descriptions, and most were published between 1850 |
and 1950. |
B. Faunal studies have not yet included many records of this subfamily. |
C. Checklist of Sterrhinae of Costa Rica begun 1997 by Covell, but much of it remains incompleted.
D. Geometridae of Ecuador and Galapagos: Projects of Fr. Francisco Pinas R., PUCE, Quito; Lazaro
Roque (Charles Darwin Res. Station), Bernard Landry, and Covell on the “Lepidoptera of the
Galapagos Islands”; and the work of colleagues speaking at this Forum (cf. Gunnar Brehm et al.)
E. Resources at University of Louisville for further systematic studies: 1. Type information & pictures;
2. Literature; 3. Collections
F. Major areas of consideration: 1. Microtaxonomy: Species recognition, descriptions, synonymies,
etc.; 2. Consideration of generic and tribal classification (Holloway’s recent synonymising of |
Anisodes Guenee to Cyclophora Hübner; question of Cyllopodini as Sterrhinae, etc.); 3. Faunal
investigations |
G. Dedication: New species of Cyclophora from Galapagos to be named herbuloti in honor of our
distinguished colleague, Claude Herbulot (C. Covell &L. Raque-Albelo).

Neotropical moths of the genus Erateina (Geometridae, Larentiinae).
Evaluation of morphological and genital characters for the purpose of systematic revision
of the genus
Janusz Wojtusiak, Zool. Mus. Inst. of Zool., Ingardena 6, PL-30060 Kraköw, Poland. |

E-mail: wojt@zuk.iz.uj.edu.pl

Studies on moths of the neotropical genus Erateina Doubleday, 1848, encompassing 89 species and 20
subspecies, were aimed to examine external morphology and genital structures of adults to select
characters of primary importance for establishing possible evolutionary relationships within the group.
These included also SEM method of examination of microstructures on males’ scent organs, which are
developed on hind wings of most of the species. As a result, nine distinctive groups of species can be

198

distinguished within the genus. Some species do not fit well into any of these groups and show
intermediate characters.

Morphological differences between species are unusually high in Erateina. This concerns especially
the variability of hind wing shape, its size and colour pattern. Male genitalia are characterised by wide,
triangular valvae with broadly rounded posterior margin and by a set of tiny rows on internal surface.
Species of all groups bear short, straight or hooked thorn in outer part of the ventral edge of valva,
cristae are well developed, stalked but their size varies from group to group.

Female’s genital structures are characterized by a very large bursa copulatrix which is bulbous in
anterior part and narrow and conical in the posterior. Asymetric sclerotisation inside the bursa is
especially thick at the posterior part and occupies about half of the lenght of the organ.

Scent organs, that seem to play a role in a chemical communication between sexes, also show a
variety of types, from the simplest, bearing only one type of scent scale, as in E. coeruleopicta or
E. meduthina, to the most elaborate ones, as in E. drucei and E. subundulata. The research is being
continued to estimate also the relationship of Erateina with genus Heterusia and Trocherateina with
which it seems to be the most closely related.

Problems in the study of the tribe Eupitheciini (Lepidoptera: Geometridae, Larentiinae)
Vladimir Mironov, Zool. Inst., Universitetskaja nab. 1, RUS-199034 St. Petersburg, Russia.
E-mail: lepid@zin.ru

The tribe Eupitheciini is a most species-rich group of the family Geometridae. According to the
Catalogue of the Geometrid Moths of the World this tribe includes nearly 1,700 species. The majority
of Pug species are small, unattractive grey or brown coloured and can hardly been distinguished from
each other by means of external features. They sometimes show no clear morphological differences.
Therefore, the identification of many species from the tribe Eupitheciini including European species is
extremely difficult. There are some urgent problems connected with the study of the Geometrid Moths
of the tribe Eupitheciini, such as:

1. The composition of the tribe Eupitheciini. — Until now, we have no full information about morpho-
logical signs differing the representatives of this tribe from others. There is no clear knowledge
about how many and which genera form the tribe Eupitheciini. In my opinion, at least 15 genera
belong to this tribe, such as: Axinoptera Hampson, 1893 (13 species); Bosara Walker, 1866 (11 sp.);
Chloroclystis Hübner, [1825] (17+*128” sp.); Glaucoclystis Holloway, 1997 (10 sp.); Gymnoscelis Ma-
bille, 1868 (88 sp.); Eupithecia Curtis, 1825 (1,330 sp.); Eupithystis Holloway, 1997 (1 sp.); Eva Vojnits,
1981 (2 sp.); Micrulia Warren, 1896 (7 sp.); Mnesiloba Warren, 1901 (4 sp.); Nasusina Pearsall, 1908
(4 sp.); Pasiphila Meyrick, 1883 (36 sp.); Pasiphilodes Warren, 1895 (19 sp.); Prorella Barnes &
McDunnough, 1918 (15 sp.) and Ziridava Walker, [1863] (11 sp.).

2. Revision of the genus Eupithecia Curtis, 1825. - The genus includes more than 1,300 species. A great
number of species, subspecies and taxa of infrasubspecific rank have been described on the basis
of single specimens or small series of adults. The descriptions of many species and subspecies do
not correspond to modern requirements. Informations on type-specimens of many Eupithecia
species are still lacking.

3. Revision of the genus Chloroclystis Hübner, [1825]. - The genus Chloroclystis had been expanded to
include a large number of species over many years. According to Holloway (1997), the tropical
representatives of this genus demonstrate a strong morphological diversity. I hope that detailed
study will undoubtedly show that many species previously associated with Chloroclystis should be
excluded from Chloroclystis and included mainly to the genera Gymnoscelis and Pasiphila in revision.

4. Distribution of species. - Modern guides, catalogues and atlases on Eupitheciini are absent for
many countries and broad geographical regions. We have poor knowledge about geographical
variation of many species especially widespread ones. It is necessary to compile the distribution
maps for many species of Eupitheciini of the World.

5. Data on phenology, biology, foodplants of larvae and habitats. - Most of these data are known for
European and north American species of Eupitheciini. Other regions, especially tropical territories
are very poorly studied in these respects.

6. A system of the genus Eupithecia Curtis, 1825. — In the history of Lepidopterology many systems

199

of Eupithecia have been published. The traditional order of species of European Eupithecia in’
particular has become imperfect long ago. Closely related species are often placed far away from |
each other in this system of European Eupithecia. In some European faunistic publications, the
Eupithecia species are arranged totally in disorder. It is necessary to create a new modern system |
of the genus Eupithecia on the basis of more or less clear morphological features, mainly on the basis
of structure of the male and female genitalia.

Estimations about the systematic position of Pseudobiston pinratanai Inoue, 1994
Dieter Stüning, Zool. Forschungsinstitut und Museum Alexander Koenig,
Adenauerallee 150-164, D-53310 Bonn, Germany. E-mail: d.stuening.zfmk@uni-bonn.de

Inoue described this peculiar, large, rather bombycoid moth in the Geometridae, subfamily Geometri-
nae, though he emphasized, on the other hand, the absence of the only known synapomorphy of the '
Geometridae, the characteristic abdominal tympanal organs. He outlined this placement as being |
tentative. However, some of the characters figured in his publication seem to support the proposed‘
position: the presence of a tubular vein M2 in the hindwings, arising close to MI; vein Mi in the |
forewing arising from the commom stalk of R2-R5; large, sclerotized socii in the male genitalia. Other
characters are contradictory to this placement or at least rarely found in the Geometrinae: absence of
tympanal organs; cross-bar between Sc+R1 and cell in the hindwings, R2 arising from the common stalk
of R3+R4 in the forewing (drawn incorrectly in Inoue’s publication); tibia of hindlegs with one pair of
spurs only; shape of socii and furca-arms in male genitalia. |
The aim of this study was to check the given characters and/or to find additional ones supporting or
not the systematic placement proposed by Inoue. The results are still rather preliminary: |
1. Tympanal organs: not reduced or degenerated, but totally absent. Instead there are normal apo-
demes as found in many other families;
2. Tergal phragmata: agreeing with the condition found typical for Ennominae and Geometrinae;
3. Chaetosemata: present;
4. Thoracic tympana: absent (some characters, e.g. venation, shape of antennae, specialized shape of |
scales, small middle- and hind-legs with only one pair of spurs) agree with the conditions found |
in some Notodontidae, but the absence of thoracic tympanal organs contradict a possible relation-

ship).

The (mostly plesiomorphic) characters studied so far neither support nor contradict reliably the
systematic position proposed by Inoue.

Diversity of Geometrid moths along an altitudinal gradient in a Mountain Rainforest
in South Ecuador

Gunnar Brehm, University of Bayreuth, Germany. E-mail: Gunnar_Brehm@yahoo.com

(& Dirk Süßenbach, Christoph Häuser, Konrad Fiedler) |

Herbivorous insects play an important role in terrestrial ecosystems. Therefore, understanding the
extend and determinants of herbivore diversity will be a crucial element of ecosystem analysis. This
is presently aimed by a research group of the German Research Foundation DFG. Its subject is to
investigate functional aspects in a tropical rainforest in South Ecuador, its diversity, dynamic processes |
and potentials of use from an ecosystem viewpoint. We chose two model groups, Geometrid and |
Pyralid moths, in a first step of analysis. Our study area is a mountain rainforest in South Ecuador
(1.800-3.100 m asl). We want to answer the following questions. |

How large is the diversity of Geometrid moths (alpha diversity)?
So far, we found approximately 700 species or morphospecies in the study area. Fisher’s alpha is
a diversity measure which is independent of sample size. The area belongs to the hot spots of
Geometrid diversity in the world because values between 110 and 130 are achieved. A comparable
diversity is thus far only known from SE Asia. |
How do communities change along an altitudinal gradient?

200

So far we investigated an altitudinal gradient between 1,800 m and 2,400 m asl. The distance
between the plots is some 100 m elevation. At lower altitudes, communities are dominated by the
subfamily Ennominae whereas Larentiinae become more important at higher altitudes.

We use the NESS index as a measure of similarity because abundances of the species and the role
of rare species are considered. The ordination method of Multidimensional Scaling (MDS) is used to
visualize results and provide appropriate data for further statistical tests.

Which habitat parameters determine the diversity of Geometrid moths?

MDS allows to test correlations between the diversity of the moths and habitat parameters. We
cooperate with participants of the DFG research group who work in the same study area. They will
provide us with data of vegetation structure, diversity of plant and animal groups and others. These
analyses will be carried out in the next months.

Life Histories of Geometrid moths

So far, little is known about the biology of Neotropical Geometrid moths. Larvae were collected in
the field and reared to adults. Data for more than 20 species of habitus and food plants of the species
can be provided now.

The phylogenetic relationships in Geometrid moths.
An approach using mitochondrial DNA (mtDNA) sequences.
Michael A. Miller, Axel Hausmann & Robert Trusch, Zoologische Staatssammlung,
Münchhausenstr. 21, 81247 Munich, Germany. E-mail: miller@zsm.mwn.de (Miller)

DNA sequence analysis has become a widespread tool for taxonomic identification and reconstruction
of phylogeny. However, molecular genetic approaches to phylogenetic reconstruction face the problem
that the sequence divergence caused by evolutional processes of the genes targeted for sequence
analysis must match the evolutionary pace of the taxon group under consideration. For phylogenetic
interpretation it is necessary to find such loci that can provide appropriate ‘signal sequences’ for
phylogenetic interpretation. In a first approach on Geometrid moths we explored the phylogenetic
signal from DNA-sequences of the mitochondrial ND1 protein-coding gene and the 165 ribosomal
RNA.

Genomic DNA was extracted from 19 specimens of 18 representative taxa of the subfamilies
Ennominae, Larentiinae, and Sterrhinae, focussed on subfamily Sterrhinae with 14 taxa. The subse-

Peribatodes rhomboidaria

Cyclophora puppillaria

. nr Timandra griseata Peribatocbs rhombaicaria
: Timandra comae Bavaria /daea hlicata
Timandıra comae Finland Er
Idaea filicata a
/daea seriata
/daea distinctaria ne
; Idaea aversata Sogpula vigilata
? /daea degeneraria Scopula imitaria
Glossotrophia alba an ee
Scopula marginepunctata
Scopula imitaria Tımancha griseata
Scopula rubiginata Cyclophora pupoillaria
Scopula vigilata & De a
93 r- Xanthorhoe ferrugata Ohloroclystis v-ata
08 Xanthorhoe vidanoi
1 68 Epirrhoe alternata
Chloroclystis v-ata

Fig. 1. Bootstrap 50 % majority-rule consensus tree for 19 Geometrid taxa using 135 informative sites of the
mitochondrial ND1 gene (rooted with Peribatodes rhomboidaria). Settings for tree computation: Heuristic search,
bootstrap with 100 replicates.

Fig. 2. Bootstrap 50 % majority-rule consensus tree of a reduced set of taxa using combined sequences of the
mitochondrial ND1 gene and the 165 rRNA locus (102 parsimony informative characters, rooted with Peribatodes
rhomboidaria). Settings for the analysis see fig. 1.

201

quent PCR amplified a fragment enclosing both genes (primer: ND-2, 5-ACATGATCTGAGT-
TCAAACCGG, Vogler & De Salle 1993; ND-S, 5-TAGAATTAGAAGATCAACCAGG, Weller pers.

comm.). Sequencing was done by an ABI 377 automated sequencer. Sequences were aligned to a

respective sequence of Bombyx mori (GenBank ass. No. NC 002355), gaps and alignment ambiguities in

the stretch of both sequences were excluded for analysis. The data-matrix contained 464 nucleotides

(only ND1) or 779 bp in total (ND1 and 165 combined). Phylogenetic tree reconstruction was done by

parsimony analysis using PAUP 3.1.1 with computation settings indicated in Figs. 1 and 2.

Tree reconstructions show close relations between tribes Timandrini (Timandra griseata, T. comae)
and Cosymbini (Cyclophora puppillaria), supported by relatively high bootstrap values (fig. 1, 2). The
monophyly of the tribe Scopulini, strongly supported by morphological characters, is still unsupported
by the used set of data. The examined Xanthorhoini species (Xanthorhoe spp., Epirrhoe alternata) show
close relations to each other (bootstrap value 98; fig. 1). Certain affinities result as well between
Xanthorhoini (Xanthorhoe vidanoi) and Eupitheciini (Chloroclystis v-ata; bootstrap value 68; figs. 1, 2). The
relationships between the other tribes are not resolved by the present set of data.

We conclude that the preliminary results from our using the two mitochondrial genes ND1 and 168
are not yet satisfactory to tackle all details of subfamily and tribal relationships in the Geometridae by
DNA sequencing. The implementation of more loci should contribute to a better resolution of the tree.
Recently, Abraham et al. (2001) directed this way by running a molecular analysis with three gene
fragments (ND1, D1- and D2- expansion elements of the nuclear 285 RNA) to reconstruct the phylog-
eny of Geometrid familiy with still more encouraging results.

This study, conducted at the Zoologische Staatssammlung München was integral to the “DNA-
TAX project” that aims at establishing insect sequence databases in cooperation with the project
“Inventory of Geometrid Moths of Europe” (A. Hausmann), both part of the “Entomological Data
Information System (EDIS)” initiative, funded by the BMBF.

202

| SPIXIANA 203-206 München, 01. November 2001 ISSN 0341-8391

New data on Anillina of the Oriental Region

(Insecta, Coleoptera, Carabidae, Bembidiini)
Pier Mauro Giachino

Giachino, P. M. (2001): New data on Anillini of the Oriental Region (Insecta,
Coleoptera, Carabidae, Bembidiini). — Spixiana 24/3: 203-206

The author describes the male morphological characters of Argiloborus burckhard-
ti Giachino, 2001, confirming that this species belongs to the “A. thailandicus group”
(sensu Giachino 2001). Argiloborus riedeli, spec. nov. from Salawati Island (Irian
Jaya) is also described. It belongs to the “A. huberi group” (sensu Giachino 2001).
The geographical distribution of the “A. huberi group” is also analysed.

Dr. P.M. Giachino, Museo Regionale di Scienze Naturali, Via Giolitti, 36, 1-10123
Torino, Italy

Introduction

This paper deals with the study of two specimens of Argiloborus Jeannel, 1937 (Coleoptera, Carabidae,
Anillina) from Sumatra and New Guinea that were kindly sent to the author by Martin Baehr, of the
Zoologische Staatsammlung of Munich (Germany). They are the until now unknown male of Argilo-
borus burckhardti Giachino, 2001, and a male of an undescribed species.

The specimens dealt with in this work are preserved in the following collections

MHNG: Museum d’Histoire Naturelle, Geneve

ZSM: Zoologische Staatsammlung, München

Argiloborus thailandicus species group

The “thailandicus species group” is characterized by the following morphological characters: sides of the
pronotum distinctly denticulate in front of the basal angles, aedeagus with a sclerified apical blade,
parameres bearing only one apical seta.

Argiloborus burckhardti Giachino, 2001
Fig. 1

Material examined: Holotype ?, Sumatra: W Sum., # 21, Palopo Nat. Res. N., Bukittinggi, 900 m, 18-20.X1.1989,
Agosti — Löbl - Burckhardt leg. (MHNG); 15, W Sumatra, Batang, Palupu, Bukittinggi, 1400-1500 m, 19.10.1991,
leg. A. Riedel (ZSM).

Diagnosis of the d: The examination of the male specimen of A. burckhardti completely confirms its
attribution to the “Argiloborus thailandicus group” (sensu Giachino 2001) both on the reasons of its
external morphology and for the characters of the aedeagus.

203

Fig. 1. Argiloborus burckhardti Giachino, 2001. Aedeagus, lateral view. Scale: 0.1 mm.

The sizes of the male are as following: total length (from tip of mandibles to end of elytra) 1.15 mm. |
Pronotum slightly less transverse than in the female, with the ratio max. width/ max. length: 1.16; 1.27
in the female. Elytra a little narrower in the male (max. length/max. width ratio: 1.71; 1.57 in the

female).
Protarsi pentamerous and not dilated in the male.

Aedeagus (Fig. 1) small, stocky, with a very developed apical blade, moderately sclerified and '
widely rounded. Internal sac provided, in the median area, with a large ondulated copulatory piece

that continues with obvious muscular bundles slightly sclerified in the preapical area. Parameres

stocky and short, rounded apically and bearing one seta; basal nodules very obvious and well |

chitinised.

Argiloborus huberi species group

The “huberi species group” is characterized by the following morphological characters: sides of prono-
tum distinctly denticulate in front of the basal angles, aedeagus lacking of the sclerified apical blade, |

parameres bearing two apical setae.

Argiloborus riedeli, spec. nov.
Fig. 2

Types. Holotype: 9, Irian Jaya, Sorong-Pr. Salawati Isl., Kalobo, 10-30 m, 19-22.X.1996, leg. A. Riedel (ZSM).

Diagnosis. It is an Argiloborus with features in accordance with the genus (sensu Giachino 2001):
presence of a labial tooth bearing two setae, elytra not emarginate apically, type B umbilicate series
(sensu Jeannel 1963) (the large pores of the umbilicate series are the 2"4, 6", and 9"" ones) with the 7"
pore slightly nearer to the 8" than to the 6" one.

A. riedeli, spec. nov. differs from all other known species of this group by the greater body size as
may be seen hereafter:
. Javanicus Giachino, 2001: 0.79 mm.
. roberti Giachino, 2001: 0.83-0.87 mm.
. huberi Giachino, 2001: 0.91-0.97 mm.
. indonesianus Giachino, 2001: 0.95 mm.
. balinensis Giachino, 2001: 1.08 mm.
. riedeli, spec. nov.: 1.22 mm.

Be gs ge sn

204

Description of the ? holotypus

Total length (from tip of mandibles to apex of elytra)
1.22 mm. Body elongated, depigmented, yellow testa-
ceous, with elytra and abdomen of the same colour; integ-
ument opaque, obvious microsculpture, covered with a
sparse and short pubescence (Fig. 2).

Head robust, hypertrophic, narrower than pronotum,
anophthalmous. Antennae very short, thickened, clearly
moniliform starting from the third antennomere, not
reaching the base of pronotum when stretched back-
wards. Anterior margin of the epistome subrectilinear.
Two supraorbital setae on each side, far from each other
and placed on rows neatly converging backwards, plus a
series of dispersed supranumerary setae not symmetrical
between each other. Mandibles short, simple, without
dorsal crests; labrum provided with 6 anterior marginal
setae; labium transverse, articulated, bearing one obvious
tooth on the anterior margin, epilobes poorly developed;
labial tooth bearing two setae. Maxillary palps with the
penultimate article very big, ovoidal elongated, and the
last one tiny, poorly differentiated.

Pronotum slightly transverse (max. width/max. length
ratio = 1.22), enlarged in front, narrowed at the base, with
maximum width at about the anterior third. Sides poorly
arcuated anteriorly, slightly sinuate in front of the base;
clearly denticulate in front of the basal angles. Anterior
angles widely rounded, not protruding; basal angles right
and marked. Disk scarcely convex, with a short and sparse
pubescence; median groove shallow, hardly marked.
'Marginal groove very wide and flat, enlarged near the
base; anterior marginal setae inserted inside the marginal
groove, at about the anterior sixth; basal setae at the back
angles.

Elytra ovoidal, elongated, not emarginate preapically,
without traces of striae. Disk poorly convex; integument
opaque, with obvious microsculpture and short, sparse
and erect pubescence. Humeri rounded, but well marked;
post-humeral margin denticulate, with a distinct crenula- Fig. 2. Argiloborus riedeli, spec. nov. ? Holo-
tion until the apical third. Marginal groove wide and NP*- Habitus. Scale: 0.1 mm.
obvious until the 9" pore of the umbilicate series.

Chaetotaxis: basal umbilicate pore large, foveate.

Type B umbilicate series (the big pores of the umbilicate series are the 2", 6", and 9" ones (sensu Jeannel
1963)); the first three pores of the humeral group are almost at the same distance from each other, the
4'" pore is much farther and placed almost in the middle between the 3"! and the 5" ones; the 5" pore
is at about the apical third of the elytron and slightly on the elytral disk, the 6" one is nearer to the 5'";
the 7%, 8%, and 9% ones are at about the same distance from each other, the 8" one slightly on the elytral
disk and a little nearer to the 9"; the 7'" one is slightly nearer to the 8" than to the 6". Discal pores
missing.

Legs short and thickset; anterior tarsi pentamerous in the female.

Male unknown.

Derivatio nominis: This new species is dedicated to its collector Alexander Riedel of Stuttgart.

Distribution, ecology: Argiloborus riedeli, spec. nov. is known at present only from the type locality:
Kalobo, on Salawati Island, at the western end of Irian Jaya. This new species was collected in October
at the very low height of only 10-30 m a.s.l.

205

@®_ A. huberi Giachino
® A. javanicus Giachino
® A. roberti Giachino

®_ A. balinensis Giachino
® A.riedeli n. sp.

Fig. 3. Distribution map of the Argiloborus of the “huberi species group”.

Remarks

The Argiloborus species described recently by Giachino (2001), and in particular the one dealt with in

this paper, have a peculiar zoogeographical meaning and further enlarge the presently known distri-

bution area of the whole phyletic lineage of Argiloborus. The genus Argiloborus, in particular, has, as far
as we now know, a very wide distribution that, starting from Madagascar, going through the Mauritius
and Seychelles Isles, Ceylon and southern India, reaches the Malay peninsula and the Islands of
Sumatra, Java, Bali and Salawati, trespassing the Weber line, that marks, between the Islands of Bali
and Lombok, the border between the Oriental Region and the Australian Region for several groups of

living organisms (Jeannel 1942) (Fig. 3). Presently, no Anillina are known from Borneo, Celebes and
New Guinea (if we exclude the new species now known from Salawati Island), whereas in New
Caledonia there is Orthotyphlus Zaballos & Mateu, 1998, of the Zeanillus phyletic series (sensu Jeannel
1963 and Moore 1980).

Acknowledgements

I am very grateful to Dr. Martin Baehr of Zoologische Staatsammlung of Munich for having allowed me to study

the precious material dealt with in this work, and to Prof. Achille Casale of the Dipartimento di Zoologia e
Antropologia Biologica of the University of Sassari for his critical suggestions to the manuscript.

References

Giachino, P.M. 2001. New Anillina (Coleoptera, Carabidae: Bembidiini) of the Oriental Region. In: G. Cuccodoro |

& R. A.B. Leschen (eds): Systematics of Coleoptera: Papers Celebrating the Retirement of Ivan Löbl. - Mem.
Entomol., Intern. Assoc. Publ. (in press)

Jeannel R. 1963. Monographie des “Anillini”, Bembidiides endoges (Coleoptera Trechidae). - Mem. Mus. nat.
Hist. nat., Paris (A) 28: 33-204

-- 1942. La genese des faunes terrestres. Elements de Biogeographie. — Press. Univers. France, Paris: 1-514

|
|

Moore B. P. 1980. A synopsis of the New Zealand Anillini (Coleoptera: Carabidae: Bembidiinae), with des I

tions of new genera and species. — N. Z. Journ. Zool 7: 399-406.
Zaballos J. P. & J. Mateu 1998. Orthotyphlus nom. nov. ne Neotyphlus Zaballos & Mateu, 1997 (Coleoptera,
Carabidae, Anillini). - Nouv. Revue Ent. (N. S.) 15(1): 3

206

| SPIXIANA 207-229 Mann ol November 2001 ISSN 0341-8391

Revision der bisher zu Iselix Förster gestellten
westpaläarktischen Arten von Phygadeuon Gravenhorst

(Insecta, Hymenoptera, Ichneumonidae, Cryptinae)
Klaus Horstmann

Horstmann, K. (2001): Revision of those western Palearctic species of Phyga-
deuon Gravenhorst, which were hitherto placed to Iselix Förster (Insecta, Hymeno-
ptera, Ichneumonidae, Cryptinae). — Spixiana 24/3: 207-229

Those western Palearctic species of Phygadeuon Gravenhorst, which were hither-
to placed to Iselix Förster by some authors, are arranged in seven species groups,
revised and described. Eight species from central and north-western Europe are
described as new. A key is provided for the females of 24 species. Lectotypes are
designated for Ischnocryptus cubiceps Smits van Burgst and Phygadeuon thomsoni
Roman, in order to preserve stability of nomenclature. The following synonymies
are newly indicated: P. geniculatus Kriechbaumer, syn. Platylabus meuseli Lange;
Phygadeuon clotho Kriechbaumer, syn. P. grossae Horstmann; P. atropos Kriechbau-
mer, syn. Ischnocryptus atropos (Kriechbaumer) var. rufifemur Seyrig. Three species
have been reared from their hosts, they are endoparasitoids and koinobionts.

Dr. Klaus Horstmann, Lehrstuhl Zoologie II, Biozentrum, Am Hubland,
D-97074 Würzburg, Germany

Einleitung

Phygadeuon Gravenhorst ist eine der artenreichsten und gleichzeitig am wenigsten bekannten Gattun-
gen der Ichneumonidae. Über die Arten der Westpaläarktis sind in den letzten Jahrzehnten nur
Teilrevisionen kleiner Artengruppen erschienen (Horstmann 1967, 1993). Hier wird eine Revision der
Arten vorgelegt, die traditionell zu Iselix Förster gestellt worden sind. Über den Status dieses Taxons
gehen die Meinungen auseinander: Perkins (1962: 432) und Townes (1970: 103) stellen Iselix als
Synonym zu Phygadeuon, Townes et al. (1965: 147) und Hellen (1967) dagegen führen Iselix als eine von
Phygadeuon getrennte Gattung. Derzeit behandelt die Mehrzahl der Autoren Iselix als Untergattung
oder als Synonym von Phygadeuon (Aubert 1974: 268 f., Yu & Horstmann 1997: 383). Eine Klärung der
anstehenden Fragen wird hier nicht angestrebt; sie kann nur in einer umfassenden Revision von
‚Phygadeuon erfolgen. Auch in Bezug auf den Umfang von Iselix folgt die vorliegende Arbeit der
| ‚Auffassung früherer Autoren. Dagegen werden die bisher zu Iselix gestellten Arten nach der Form des
Legebohrers in sieben Artengruppen aufgeteilt. Auf einige der gefundenen Unterschiede hat bereits
‚Aubert (1974: 269) aufmerksam gemacht. Die Artengruppen austriacus (Gravenhorst), geniculatus
'Kriechbaumer und ponojensis (Hellen) sind vermutlich weder untereinander noch mit den anderen
‚Arten nah verwandt. Die anderen Artengruppen, einzeln (Artengruppe nitidus Gravenhorst) oder zu
| mehreren gemeinsam (Artengruppen brevitarsis Thomson, hercynicus Gravenhorst und lachesis Kriech-
| baumer), können vielleicht später einmal in den Rang von Untergattungen gehoben werden, wenn
‚nähere Kenntnisse zur Lebensweise der Arten vorliegen.

Das Ziel der vorliegenden Arbeit ist es, die Determination der europäischen und insbesondere der
mitteleuropäischen Arten zu erleichtern. Zum einen haben in den letzten Jahrzehnten mehrere Ento-

207

mologen Iselix-Arten aus Cheilosia-Arten (Diptera, Syrphidae) gezüchtet, und die Determination der

Parasiten war ohne eine Revision nicht möglich. Zum anderen befindet sich in der Zoologischen
Staatssammlung (München) ein umfangreiches Material dieser Artengruppe, teilweise aus der Samm-
lung Kriechbaumer (einschließlich einiger Typen), vor allem aber aus der Sammlung Erich Bauer, der
in seinen Hauptsammelgebieten Harz und Oberbayern (siehe Horstmann 1983) viele Iselix-Arten
gefangen hat. Daneben wurde Material aus einigen anderen Museen revidiert (vgl. unten). Die Revi-
sion berücksichtigt alle als Weibchen beschriebenen westpaläarktischen Arten, die von anderen Auto-
ren zu Iselix gestellt worden sind, und enthält dazu Neubeschreibungen von acht Arten. Sie ist aber

keinesfalls umfassend. Sicherlich existieren auch in Mitteleuropa noch weitere unbeschriebene Arten.

Männchen von Phygadeuon sind derzeit in der Regel undeterminierbar. Alle nur im männlichen
Geschlecht bekannten Arten werden hier nicht behandelt, und für Männchen kann auch kein Bestim-

mungsschlüssel entworfen werden. Möglicherweise bleiben deshalb einige Synonymien unerkannt. |

Auch Weibchen können nur sicher determiniert werden, wenn sie vollständig erhalten sind; insbeson-
dere muss ein Fühler und der Legebohrer vorhanden sein. Die Identifikation einiger defekter Typen
war deshalb schwierig, oder sie bleibt unsicher.

Angaben über die Lebensweise liegen nur für drei Arten der Artengruppe hercynicus Gravenhorst |
vor: Sie parasitieren an Arten von Cheilosia Meigen (Diptera, Syrphidae), deren Larven an Carduus- oder

Cirsium-Arten leben.
Das untersuchte Material entstammt folgenden Museen und Sammlungen:
DEIE: Deutsches Entomologisches Institut, Eberswalde

FRI: Sammlung Frilli, Istituto de Defesa delle Piante, Udine
HO: Sammlung Horstmann, Lehrstuhl Zoologie III, Biozentrum, Würzburg
LEW: Laboratorium voor Entomologie, Wageningen

MHNP: Museum National d’Histoire Naturelle, Paris
MPW: Muzeum Przyrodnicze, Wroclaw

MZL Musee Zoologique, Lausanne

NHML: Natural History Museum, London

NMSE: National Museums of Scotland, Edinburgh
NMW: Naturhistorisches Museum, Wien

NRS: Naturhistoriska Riksmuseet, Stockholm

NSF: Naturmuseum Senckenberg, Frankfurt

SMNS: Staatliches Museum für Naturkunde, Stuttgart

SJNÜR Sammlung Stuke, AG Evolutionsbiologie der Universität, Bremen
ZANTE: Zoologiska Institutionen, Lund

ZMH: Zoological Museum, Helsinki

ZSM: Zoologische Staatssammlung, München

Bestimmungsschlüssel für die Weibchen

1. Apicalrand des Clypeus median nicht gezähnt, sondern nur wenig vorgerundet, zwei Zähne
subapical als Oberflächenstruktur erkennbar (Abb. 12); Bohrer dorsal vor dem Nodus mit einer
rundlichen Vorwölbung (Abb. 39); Bohrerklappen 1,5 mal so lang wie das erste Gastertergit

(Artengruppe lachesis Kriechbaumer) ...........eresensenecsecsensenensenseneneenennennennene lachesis Kriechbaumer, 1892

- Apicalrand des Clypeus median deutlich gezähnt, mit einem einzelnen Zahn (Abb. 11), einem
Doppelzahn (Abb. 9) oder zwei deutlich getrennten Zähnen (Abb. 10); Bohrer dorsal vor dem

Nodus gerade (Abb. 32-38 und 42-43) oder mit einem zweiten Zahn (Abb. 40-41); Bohrerlänge

unterschiedlieht. ee DR

2. Bohrer dorsal am Nodus mit einem zahnartigen Vorsprung und davor mit einem weiteren deut-

lichen Zahn (Abb. 40-41)*; Bohrerklappen 0,6-1,3 mal so lang wie das erste Gastertergit (Artengrup- |

pe nitidus Gravenhorst)...........uunseseessensensecnesnsannecnsenssnsetneenssnnennssnnsonsonsennernsensannnenasnnsenssnsecnessennetnesnneensehtechser 3

* Bei P. habermehli liegt der zweite Zahn relativ weit proximal und ist nur bei frei sichtbaren Bohrerstiletten zu

erkennen.

208

——

E:-
| =
1 2 3
SER> ce?

> <> — >

| 4 | | 5 | 6
r
7 8

10 11 12

Abb. 1-8. Dorsalansicht des Kopfes. 1. Phygadeuon bavaricus. 2. P. fraternae. 3. P. praealpinus. 4. P. unidentatus.
‚5. P. laevipleuris. 6. P. nigrifemur. 7. P. atricolor. 8. P. macrocephalus.

Abb. 9-12. Clypeus. 9. P. fraternae. 10. P. praealpinus. 11. P. unidentatus. 12. P. lachesis.

- Bohrer dorsal am Nodus gezähnt oder ungezähnt, davor ohne weiteren Zahn (Abb. 32-38 und
42-45) Bohrerlangesunterschiedliehn:......erceerecneereeeriennesnenneaense ne nneehneeetun dehnen neaeteekeusee cute nt sentenes 8.

‚3. Bohrerklappen 0,6-0,7 mal so lang wie das erste Gastertergit; viertes Fühlerglied etwa zweimal so
lan oe NDTEI ee Renee, habermehli Roman, 1930

- Bohrerklappen 0,9-1,3 mal so lang wie das erste Gastertergit; viertes Fühlerglied bei mehreren
te18279-2,5,malksorlangwierbreit......... en een nnsaieannsieaswens este nee ee 4.

‚4. Schläfen 1,1 mal so breit wie die Augen (von oben gesehen), direkt hinter den Augen etwas
erweitert; Wangenraum 0,9 mal so breit wie die Mandibelbasis; Clypeus dorsal neben der Punk-
tierung etwas quergerunzelt; Hinterfemora 3,6 mal so lang wie hoch. es: aa.
NR forticornis Kriechbaumer, 1892

'— Schläfen höchstens so breit wie die Augen, direkt hinter den Augen parallel oder etwas verengt;
Wangenraum höchstens 0,8 mal so breit wie die Mandibelbasis; Clypeus dorsal neben der Punk-
tierung nicht quergerunzelt; Hinterfemora teilweise gedrungener ..........ueeennnnessensennn 5:

5. Viertes Fühlerglied knapp zweimal so lang wie breit; Hinterfemora 2,9-3,1 mal so lang wie hoch
RESTE ERBE IRBEN SIERT atropos Kriechbaumer, 1892

- Viertes Fühlerglied 2,3-2,5 mal so lang wie breit; Hinterfemora 3,5-3,7 mal so lang wie hoch ... 6.

209

10.

14.

1),

16.

10%

210

Hinterfemora überwiegend oder ganz schwarz; Bohrerklappen 1,3 mal so lang wie das erste,
Gastertersiti....nenceeeehureee neunten nennen ee ee nigrifemur, spec. nov.

Hinterfemora rotbraun; Bohrerklappen 0,9-1,0 mal so lang wie das erste Gastertergit ........... 7
Mesopleuren im Zentrum auf einer größeren Stelle unpunktiert; Area superomedia 0,8-1,0 mal so
langswie breits (Abb»28)2..Walen ann ee laevipleuris, spec. nov.
Mesopleuren im Zentrum überall zerstreut punktiert, Punkte 0,3-0,5 mal so breit wie die Zwischen-
räume; Area superomedia 0,5-0,7 mal so lang wie breit ...........ne.- nitidus Gravenhorst, 1829

Dorsale Kante des Bohrers am Nodus mit einer kleinen Runzel oder einem kleinen Einschnitt, aber
dort nicht deutlich gewinkelt, sondern gerade oder gleichmäßig gerundet (Abb. 32-33 und 35-38)

ES Re EN RENER EC eeEEE ERE EEE EUER EEE SE EEE EEE TEE. 20.00:005006605000005550060000000 98
Dorsale Kante des Bohrers am Nodus deutlich gewinkelt, distal nach ventral abgeknickt (Abb. 34
und»42-48) nk. el. en arena Diergluanhleles SS 20.
Bohrerspitze deutlich aufgebogen (Abb. 33); Bohrerklappen 1,1-1,2 mal so lang wie das erste
Gastertersit(Arteneruppe, brevitarsis Ihomson)e.......... nn EEE 102
Bohrerspitze etwa gerade (Abb. 32 und 35-38); Bohrerklappen häufig länger ........... 1718
Schläfen direkt hinter den Augen parallel; Femora hell rotbraun. ............ brevitarsis Thomson, 1884

Schläfen direkt hinter den Augen erweitert; Femora der Vorder- und Mittelbeine überwiegend, die
dens-Imterbeinesgamzischwarzer.. ne tunetanus Horstmann & Yu, 1999

Fühler 23-24-gliedrig; Bohrerspitze hinter dem Nodus kurz, die dorsale Kante wenig nach ventral
gerundet (Abb. 32); Bohrerklappen 1,7 mal so lang wie das erste Gastertergit (Artengruppe austria-
CUSMLG@TAVENhOLSE))E sn denne austriacus (Gravenhorst, 1829)

Fühler 19-22-gliedrig; Bohrer insgesamt stilettförmig, Spitze hinter dem Nodus lang, die dorsale
Kante etwa gerade (Abb. 35-38); Bohrerklappen 0,6-1,4 mal so lang wie das erste Gastertergit

(Artengrupperhexeynieus Grayenhorst)..en....enenesrreesenaneennnerenereneennr era ERREGER 124

. Schläfen 1,4 mal so breit wie die Augen (Abb. 4); Clypeus apical mit einem spitzen Zahn (Abb. 11);
Areassuperemedianl,2malssollansswie breit ABbY27)e... unidentatus, spec. noV.
Schläfen häufig schmäler; Clypeus apical mit zwei Zähnen (Abb. 9-10); Area superomedia höch-
stenis so lang, wie breit.....:...us.eensnesüanasansansessnesensneneseneneneetnsuepenenennensnere sun eeentne nee ER 1%

. Viertes Fühlerglied 3,3-3,5 mal so lang wie breit (Abb. 15); Hinterfemora 3,8-3,9 mal so lang wie
hoch; Bohrerklappen 0,6-0,7 mal so lang wie das erste Gastertergit.............. praealpinus, spec. nov.
Viertes Fühlerglied höchstens 2,6 mal so lang wie breit; Hinterfemora 2,9-3,7 mal so lang wie hoch; |
Bohrerklappen 0,8-1,4 mal so lang wie das erste Gastertergit............nsenensenensensensnnonsnnennnnsonennensenennener 14.)
Hlinterfemora schwarz oder dunkelbraun gezeichnet. .................2u.2022. 22202220 one 158
Hinterfemorasvollständig; hell’rotbraun „en... sans eaer ee REEEN 17.

Schläfen 1,3-1,4 mal so breit wie die Augen (von oben gesehen); viertes Fühlerglied 2,4-2,6 mal so Ä

lang wie breit; Bohrerklappen 1,3-1,4 mal so lang wie die erste Gastertergit ........eeseseneneneenen |
Ma. re RE Be ne BD AR clotho Kriechbaumer, 1892

Schläfen 1,0-1,3 mal so lang wie die Augen; viertes Fühlerglied 2,0-2,3 mal so lang wie breit; |
Bohrerklappen 0,8-1,2 mal so lang wie das erste Gastertergit ..........nnsnsnsensensensensensensensennennennennenn 16.

Schläfen 1,1-1,3 mal so breit wie die Augen (Abb. 2); Hinterfemora 2,9-3,2 mal so lang wie hoch; |
Bohrerklappen 1,1-1,2 mal so lang wie das erste Gastertergit .......eee fraternae, spec. nov. |
Schläfen 1,0-1,1 mal so breit wie die Augen; Hinterfemora 3,3-3,6 mal so lang wie hoch; Bohrerklap- |

pen 0,8-1,0 mal so lang wie das erste Gastertergit ...........nee hercynicus Gravenhorst, 1829, var. |

Scheitel nach dorsal nicht über das Niveau der Ocellen vorragend; zweites Gastertergit 0,7-0,8 mal |
Sorlang wierbreit ........iucenesensannseessessnsnssenssensenssnasrsenennane nern ernennen Tees ee REN 18.1

Scheitel nach dorsal zumindest etwas über das Niveau der Ocellen vorragend; zweites Gastertergit |
1,0-1,2,mallso lang; wie. breit... lesseteseeenenarne engen rare ERBE 19%

}

| N 1
14
21 /

17 19
23 |

6
22

Abb. 13-20. Fühlerbasis. 13. Phygadeuon bavaricus. 14. P. fraternae. 15. P. praealpinus. 16. P. unidentatus.
17. P. laevipleuris. 18. P. nigrifemur. 19. P. atricolor. 20. P. macrocephalus.
Abb. 21-23. Areola des Vorderflügels. 21. P. bavaricus. 22. P. fraternae. 23. P. atricolor.

his.

DI.

22.

Schläfen direkt hinter den Augen deutlich etwas verengt; Hinterfemora 3,2-3,4 mal so lang wie
hoch; zweites Gastertergit 0,7 mal so lang wie breit... chilosiae Horstmann, 1975

Schläfen direkt hinter den Augen parallel; Hinterfemora 3,1-3,2 mal so lang wie hoch; zweites
@astertersia0,3,malisorlangswienbreit. .................2.0.cn.neteneesnsetasseneneonsaesenesetenn thomsoni Roman, 1925

. Viertes Fühlerglied 2,3-2,4 mal so lang wie breit; Hinterfemora 3,1-3,3 mal so lang wie hoch .......

BR NEN Eh nnanBeneoenchhcasnlaninagstrsgeeertngnigen benkersnessreikenenttrusgehehestahneerennesthäenternehehtetent bavaricus, spec. NOV.

viertes Fühlerglied 2,0 mal so lang wie breit; Hinterfemora 3,6-3,7 mal so lang wie hoch .............
OEL Syetoedkusescsnusesecnessssoncbneetessneitene dene te son eecusheterknne berenenkensneurenerean, hercynicus Gravenhorst, 1829

. Zweites Gastertergit 1,0-1,3 mal so lang wie breit, stellenweise fein gekörnelt und/oder fein

längsgestreift; Bohrerklappen 0,5-0,7 mal so lang wie das erste Gastertergit (Artengruppe genicu-
latusakriechbaumer)kn...r....20 nes eeeseeennenenenveenensessnsnensener ee geenamnneneg ste shanknshentnehrennlann inne beanstandet 21
Zweites Gastertergit etwa 0,7 mal so lang wie breit, glatt oder stellenweise mit sehr feinen Haar-
punkten; Bohrerklappen 0,8-1,6 mal so lang wie das erste Gastertergit (Artengruppe ponojensis

lellem))E ae. ee N NN N RR IRB IR Ro. ouchebsasssnlennesesetersenhneereten 22.
Schläfen direkt hinter den Augen etwa parallel; Punkte auf der Stirn stellenweise breiter als die
Zwischenraume; Körperlänge 6-8 mm ...........u.t2ssusesrenenonstsneeseneseneneneee geniculatus Kriechbaumer, 1892
Schläfen direkt hinter den Augen deutlich verengt; Punkte auf der Stirn höchstens so breit wie die
Zyuischenraume-Konperlange)5. mm... see eneeeereneeeenskenenneee meridionator (Aubert, 1960)
Bohrerklappen 0,8-1,0 mal so lang wie das erste Gastertergit ......enessennenssenenennenen: 23:
Bohrerklappen 1,3-1,6 mal so lang wie das erste Gastertergit ..........eeenennnenssenen: 24.

211

Abb. 24-31. Area superomedia. 24. Phygadeuon bavaricus. 25. P. fraternae. 26. P. praealpinus. 27. P. unidentatus.
28. P. laevipleuris. 29. P. nigrifemur. 30. P. atricolor. 31. P. macrocephalus.

23. Hinterfemora überwiegend schwarzbraun; Area superomedia 0,7 mal so lang wie breit; Bohrer- |
klappen 0,8 mal so lang wie das erste Gastertergit .........eeenen: camargator Aubert, 1982

- Hinterfemora hell rotbraun; Area superomedia 1,0-1,2 mal so lang wie breit; Bohrerklappen so lang
wierdasferstel@astertersutee nen nennen rereretnene oporinus Horstmann & Yu, 1999

24. Schläfen 1,2 mal so breit wie die Augen (von oben gesehen) (Abb. 8); Scheitel nach dorsal etwas
über das Niveau der Ocellen vorragend; viertes Fühlerglied 2,2 mal so lang wie breit; Hinterfemora
2,85mallsoplansswielhocheee ee nn ne tn macrocephalus, spec. nov.

- Schläfen höchstens 1,1 mal so breit wie die Augen; Scheitel nach dorsal nicht über das Niveau der
Ocellen vorragend; viertes Fühlerglied 2,4-2,7 mal so lang wie breit; Hinterfemora 3,4-3,9 mal so
langawielhoche nee ee ene 258

25. Viertes Fühlerglied 2,7 mal so lang wie breit; Geißel ganz schwarz; Hinterfemora 3,9 mal so lang
wie.hech. u. nl in 2 If eeteeri EiPelent atricolor, spec. nov.

- viertes Fühlerglied 2,4-2,5 mal so lang wie breit; Geißelbasis hell rotbraun; Hinterfemora 3,4 mal
so lang: wie Hoch... ee ponojensis (Hellen, 1967)

Artengruppe austriacus (Gravenhorst)

Phygadeuon austriacus (Gravenhorst, 1829) |
Fig. 32

Cryptus austriacus Gravenhorst, 1829: 573 (Frilli 1978: 157).
Iselix elfvingi Hellen, 1967: 91 (Horstmann 1990: 183).

Beschreibung

2: Schläfen 1,1 mal so breit wie die Augen, direkt hinter den Augen ein wenig erweitert, fein und
zerstreut punktiert; Scheitel nach dorsal nicht über das Niveau der Ocellen vorragend; Clypeus basal

212

32 34
33
35 37
36

* —— >>

' Abb. 32-43. Lateralansicht der Bohrerspitze. 32. Phygadeuon austriacus. 33. P. brevitarsis. 34. P. geniculatus.
35. P. bavaricus. 36. P. fraternae. 37. P. praealpinus. 38. P. unidentatus. 39. P. lachesis. 40. P. laevipleuris.
41. P. nigrifemur. 42. P. atricolor. 43. P. macrocephalus.

deutlich und mäßig dicht punktiert, apical mit zwei deutlich getrennten spitzen Zähnen; Wangenraum
' 0,8 mal so breit wie die Mandibelbasis; Fühler 24-gliedrig, das vierte Glied 3,0 mal so lang wie breit,
' Distalhälfte der Geißel nicht deutlich erweitert, Glieder im distalen Drittel der Geißel 1,2-1,3 mal so

lang wie breit; Mesopleuren außerhalb des Speculums fein und sehr zerstreut punktiert; Hinterfemora
3,5-3,8 mal so lang wie hoch; Area superomedia 0,7 mal so lang wie breit; zweites Gastertergit glatt,
0,9 mal so lang wie breit; Bohrer sehr schlank, distal wenig aufgebogen, ohne deutlichen Nodus, ventral
mit feinen Zahnleisten (Abb. 32), Bohrerklappen 1,7 mal so lang wie das erste Gastertergit; Körperlänge
etwa 7 mm; Fühler und Coxen schwarz; Hinterschenkel rotbraun; zweites bis fünftes Gastertergit hell
rotbraun, die folgenden verdunkelt.

Material: 222, Kuopio/FIN (ZMH), Österreich (Coll. Gravenhorst/MPW).

' Variation. 1? von Trifels/ Annweiler / Rheinland-Pfalz /D (NSF) gehört möglicherweise zu dieser Art,
' weicht aber durch folgende Merkmale ab: Schläfen deutlich und mäßig dicht punktiert, Fühler

23-gliedrig; Hinterfemora proximal bis über die Mitte schwarz. Dem $ fehlen allerdings der Bohrer und
die Bohrerklappen.

Artengruppe brevitarsis Thomson

Phygadeuon brevitarsis Thomson, 1884
Fig. 33
Phygadeuon brevitarsis Thomson, 1884: 959 (Frilli 1973: 96 f.).

Roman (1925: 11) hat diese Art fälschlich mit P. hercynicus Gravenhorst synonymisiert.
Beschreibung
2: Schläfen so breit wie die Augen (von dorsal gesehen), direkt hinter den Augen ein kurzes Stück

parallel, fein und zerstreut bis sehr zerstreut punktiert; Scheitel nach dorsal nicht über das Niveau der
Ocellen vorragend; Clypeus basal deutlich punktiert, apical mit zwei getrennten spitzen Zähnen;

213

Wangenraum 0,6-0,7 mal so breit wie die Mandibelbasis; Fühler 21-22-gliedrig, das vierte Glied 2,3-2,4

mal so lang wie breit, Distalhälfte der Geißel etwas keulenförmig erweitert, das breiteste Glied etwa

0,9 mal so lang wie breit; Mesopleuren außerhalb des Speculums deutlich zerstreut bis sehr zerstreut
punktiert, an kleinen Stellen unpunktiert; Hinterfemora 2,8-3,0 mal so lang wie hoch, Area superome- |

dia 0,7-0,9 mal so lang wie breit; zweites Gastertergit glatt, 0,7-0,9 mal so lang wie breit; Bohrer schlank,

distal deutlich aufgebogen, dorsal mit einem sehr schwach angedeuteten Nodus, ventral mit feinen

Zahnleisten (Abb. 33); Bohrerklappen 1,1-1,2 mal so lang wie das erste Gastertergit; Körperlänge 5-6
mm; drittes bis fünftes Fühlerglied rotbraun; Coxen schwarz; Hinterfemora rotbraun; Postpetiolus
caudal-median, das zweite, dritte und die Basis des vierten Gastertergits rotbraun, die folgenden
dunkelbraun, caudal gelb gerandet.

Material: 2122, Emo Park/Leix/IRL, Dulverton/Somerset/GB, Newton Abbot/Devon/GB, Henstead/Suf-
folk/GB (alle NHML); Scäne/S (Coll. Thomson/ZIL), Ringselle / Örebro/S (NRS), Harzburg/ Niedersachsen/D,
Elmau/Bayern/D, Garmisch/Bayern/D (alle ZSM); Weibersbrunn/Spessart/Bayern/D (HO), Podcetrtek /Slo-
wenien (SZM).

Phygadeuon tunetanus Horstmann & Yu, 1999
Ischnocryptus cubiceps Smits van Burgst, 1913: 312 f. - praeocc. in Phygadeuon durch Phygadeuon cubiceps Thomson,
1884 — Lectotypus (?) hiermit festgelegt: “?”, “Smits v. Burgst Omgev. Tunis 3/4 1911” (Coll. Smits van
Burgst/ LEW).

Phygadeuon tunetanus Horstmann & Yu, 1999: 80 — nom. nov. für Ischnocryptus cubiceps Smits van Burgst, 1913.

Beschreibung

9: Schläfen 1,2 mal so breit wie die Augen, direkt hinter den Augen ein kurzes Stück erweitert, fein.
und sehr zerstreut punktiert; Clypeus basal kräftig zerstreut punktiert, apical mit zwei kleinen deutlich

getrennten Zähnen; Wangenraum 0,7 mal so breit wie die Mandibelbasis; Fühler 21-gliedrig, das vierte
Glied 2,3 mal so lang wie breit, Distalhälfte der Geißel etwas keulenförmig erweitert, das breiteste |

Glied knapp so lang wie breit; Mesopleuren außerhalb des Speculums kräftig zerstreut punktiert; |

Hinterfemora 3,1 mal so lang wie hoch; Area superomedia 0,8 mal so lang wie breit; zweites Gaster- |

tergit glatt, 0,8 mal so lang wie breit; Bohrer wie bei P. brevitarsis, Bohrerklappen 1,2 mal so lang wie

das erste Gastertergit; Körperlänge 6 mm; drittes bis fünftes Fühlerglied gelbbraun; Coxen schwarz;
Hinterfemora fast ganz schwarzbraun; zweites bis fünftes Gastertergit hell rotbraun.

Material: 4??, Tunis/Tunesien (LEW).

Artengruppe geniculatus Kriechbaumer

Phygadeuon geniculatus Kriechbaumer, 1892
Fig. 34
Phygadeuon (Ischnocryptus) geniculatus Kriechbaumer, 1892: 343 (Aubert 1974: 269). Von den drei Syntypen dieses
Taxons gehören der Lectotypus (?) von Hochstätt bei Rosenheim und ein Paralectotypus (?) von Hesselo-
he/München zu der hier behandelten Art, ein weiterer Paralectotypus (?) von Hohenschwangau dagegen |
zu P. meridionator (alle ZSM). Die drei von Aubert (l. c.) erwähnten Exemplare vom Tegernsee (ZSM) haben |
keinen Typenstatus. |
Platylabus meuseli Lange, 1911: 540 f. (syn. nov.) (Oehlke & Horstmann 1987: 151). |

Beschreibung |
9: Schläfen etwa so breit wie die Augen, direkt hinter den Augen parallel, sehr wenig erweitert |
oder sehr wenig verengt, kräftig und mäßig dicht bis zerstreut punktiert; Dorsalhälfte der Stirn und
Scheitel mäßig dicht punktiert, Punkte mindestens so breit wie die Zwischenräume; Scheitel nach
dorsal etwas über das Niveau der Ocellen vorragend; Clypeus basal deutlich punktiert, teilweise a |
feinen Querrunzeln, apical mit zwei deutlich getrennten spitzen Zähnen; Wangenraum 0,6-0,7 mal so.
breit wie die Mandibelbasis; Fühler 23-26-gliedrig, das vierte Glied 3,0-3,2 mal so lang wie breit,

214

ı Distalhälfte der Geißel nicht keulenförmig erweitert, Glieder im distalen Drittel der Geißel 1,1-1,2 mal
\ so lang wie breit; Mesopleuren außerhalb des Speculums deutlich und mäßig dicht punktiert, an den
Rändern stellenweise etwas längsgerunzelt; Hinterfemora 3,7-3,9 mal so lang wie hoch; Area supero-
| media 0,6-0,8 mal so lang wie breit; zweites Gastertergit 1,0-1,2 mal so lang wie breit, frontal bis über
‚ die Mitte fein längsgestreift oder mit Körnelreihen (variabel); Bohrer gerade, mit deutlichem Nodus
und deutlichen Zahnleisten (Abb. 34), Bohrerklappen 0,5-0,6 mal so lang wie das erste Gastertergit:
Körperlänge 6-8 mm; Geißelbasis hell rotbraun; Coxen rotbraun, basal unterschiedlich ausgedehnt
verdunkelt; Hinterfemora rotbraun, distal schmal verdunkelt; zweites bis drittes oder viertes Gaster-
tergit rotbraun, die folgenden braun gezeichnet, caudal gelb gerandet, selten auch das fünfte Tergit
ganz rotbraun.

‚ Material: 1722, Großbritannien (aus Coll. Morley) (NHML); Goslar/Niedersachsen/D (ZSM); Düsseldorf /
Nordrhein-Westfalen/D, Worms/Rheinland-Pfalz/D (beide NSF); Garmisch/Bayern/D, München/Bayern/D,
. Rosenheim / Bayern/D, Tegernsee/Bayern/D (alle ZSM); Oberstdorf/Bayern/D, Grindelwald/Bern/CH (bei-
de NHML); Leutasch/Tirol/ A (ZSM); Dundovici/Kroatien (DEIE).

Phygadeuon meridionator (Aubert, 1960), stat. nov.
Iselix geniculatus (Kriechbaumer) meridionator Aubert, 1960: 652.

' Beschreibung
9: Schläfen 0,8 mal so breit wie die Augen, direkt hinter den Augen deutlich verengt, fein und
. zerstreut punktiert; Dorsalhälfte der Stirn und Scheitel fein und zerstreut punktiert, Punkte an vielen
Stellen schmäler als die Zwischenräume (variabel); Scheitel nach dorsal nicht über das Niveau der
Ocellen vorragend; Clypeus basal deutlich zerstreut punktiert, ohne Querrunzeln, apical mit zwei
deutlich getrennten Zähnen; Wangenraum 0,7-0,8 mal so breit wie die Mandibelbasis; Fühler 23-
25-gliedrig, das vierte Glied 2,9-3,1 mal so lang wie breit, Distalhälfte der Geißel nicht erweitert, Glieder
' im distalen Drittel der Geißel etwa 1,1 mal so lang wie breit; Mesopleuren außerhalb des Speculums
' deutlich und mäßig dicht punktiert, an den Rändern auch gerunzelt; Hinterfemora 3,6-3,8 mal so lang
wie hoch; Area superomedia 0,7-1,0 mal so lang wie breit; zweites Gastertergit 1,1-1,3 mal so lang wie
breit, frontal bis über die Mitte fein gekörnelt und/oder längsgestreift; Bohrer wie bei P. geniculatus,
Bohrerklappen 0,6-0,7 mal so lang wie das erste Gastertergit; Körperlänge 4-6 mm; Geißelbasis hell
gelbbraun; Coxen gelbbraun, Hintercoxen basal zuweilen verdunkelt; Hinterfemora rotbraun, distal
schmal verdunkelt; zweites und drittes Gastertergit hell rotbraun, die folgenden braun, caudal gelb
gerandet.

Material: 5??, Gotha/Thüringen/D, Hohenschwangau/Bayern/D, Tegernsee/Bayern/D (alle ZSM); Aygulf/
Var/F (Coll. Aubert/MZL).
Artengruppe hercynicus Gravenhorst

Phygadeuon bavaricus, spec. nov.
Figs 1, 13, 21, 24, 35

Typen. Holotypus: 2, “Ober-Bayern, Ellmau*, ca. 1050 m, 27.VI1.1937. E. Bauer” (ZSM). — Paratypen: 5?? vom
gleichen Fundort; 3?? Garmisch/Bayern/D, 700-1400 m; 1? Ettaler Berg/Oberammergau/Bayern/D, 800 m,
alle im Juli-September gefangen, aus Coll. E. Bauer (ZSM, 1? aus Elmau HO).

Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen 1,2 mal so breit wie die Augen, hinter den Augen

* Bauer hat den Ort Elmau in Oberbayern mit der Schreibweise “Ellmau” angeführt. Dadurch besteht eine
Verwechselungsmöglichkeit mit Ellmau in Nordtirol. An dem letztgenannten Ort hat Bauer aber anscheinend
nicht gesammelt.

215

ein wenig verengt (Abb. 1), mit feinen weit voneinander entfernten Haarpunkten; Scheitel nach dorsal
deutlich über das Niveau der Ocellen vorragend; Augen kahl; Clypeus etwa so lang wie das Gesicht,
von diesem durch eine schwache Furche getrennt, im Profil flach, basal mit kräftigen voneinander
getrennten Haarpunkten, am Apicalrand mit zwei dicht benachbarten spitzen Zähnen; oberer Mandi-
belzahn etwas größer als der untere; Wangenraum 0,6 mal so breit wie die Mandibelbasis; Fühler
20-gliedrig, das vierte Glied 2,3-2,4 mal so lang wie breit (Abb. 13), Distalhälfte der Geißel etwas
keulenförmig, das breiteste Glied etwa 0,8 mal so lang wie breit.

Pronotum lateral zentral an einer kleinen Stelle unpunktiert, an den Rändern zerstreut punktiert,
stellenweise gerunzelt, Epomia kräftig; Mesoscutum median sehr zerstreut punktiert, Seitenlappen
zentral jeweils stellenweise unpunktiert; Mesopleuren außerhalb des großen Speculums überall deut-
lich fein und sehr zerstreut punktiert; Metapleuren ähnlich; Beine kräftig, Hinterfemora 3,1-3,3 mal so
lang wie hoch; Areola etwa regelmäßig fünfeckig (Abb. 21); Nervellus bei 0,7 kräftig gebrochen, etwas
incliv.

Propodeum vollständig gefeldert, in den Feldern kaum strukturiert, Area superomedia 0,7-0,9 mal
so lang wie breit (Abb. 24), Area petiolaris wenig eingesenkt, Seitenecken als etwas breitere Lamellen,
aber nicht spitz vorragend; erstes Gastertergit sehr fein gekörnelt, fast glatt, auf dem Postpetiolus
stellenweise sehr fein längsgerieft (variabel), Dorsalkiele fast bis zu den Stigmen reichend, Dorsolate-
ralleisten bis fast zum Caudalende divergierend; die folgenden Tergite glatt; zweites Gastertergit
1,0-1,2 mal so lang wie breit; Gaster vom dritten Segment an etwas von der Seite zusammengedrückt;
Bohrer schlank, gerade, dorsal ohne deutlichen Nodus, an seiner Stelle eine kleine Runzel, ventral mit |
feinen Zahnleisten (Abb. 35), Bohrerklappen 1,0-1,1 mal so lang wie das erste Gastertersgit. |

Schwarz; Palpen, Mandibeln (Zähne dunkel), drittes bis fünftes Fühlerglied (Ausdehnung etwas
variabel), Tegulae, Trochantellen, Femora, Tibien und Tarsen gelbbraun; Coxen apical unterschiedlich
ausgedehnt gelbbraun gezeichnet; Flügelbasis gelb, Pterostigma mittelbraun, proximal und distal
schmal weißlich, Flügelfläche klar; zweites bis viertes Gastertergit hell rotbraun, die folgenden
schwarzbraun, caudal schmal gelb gerandet. |

Holotypus (?): Kopf 144 breit*; Thorax 250 lang, 122 breit (Mesoscutum); Vorderflügel 565 lang; '
erstes Gastertergit 122 lang; Postpetiolus 52 lang, 55 breit; zweites Tergit 110 lang, 96 breit; Bohrerklap-
pen 135 lang; Körper etwa 780 lang.

d unbekannt.

Variation. Im NRS befindet sich als Paralectotypus von P. thomsoni 12 mit den Etiketten “Gl.” (= Got-
land/S), “Bhn”, das mit dieser Beschreibung gut übereinstimmt, aber nur knapp 6 mm lang und zudem
etwas beschädigt ist. Sein Status ist unklar.

Phygadeuon chilosiae Horstmann, 1975
Phygadeuon chilosiae Horstmann, 1975: 106 ff.

Beschreibung

Die Art ist P. thomsoni sehr ähnlich und vielleicht mit dieser synonym; allerdings ist von beiden
Arten nur sehr wenig Material bekannt. Es bestehen folgende Unterschiede zur Beschreibung von |
P. thomsoni:

$: Schläfen 1,0-1,1 mal so breit wie die Augen, direkt hinter den Augen deutlich etwas verengt; |
Hinterfemora 3,2-3,4 mal so lang wie hoch; zweites Gastertergit 0,7 mal so lang wie breit; Körperlänge |
5 mm. Eine ausführliche Beschreibung findet sich bei Horstmann (I. c.). |

Lebensweise. Als Wirt wurde Cheilosia cynocephala Loew an Cirsium vulgare nachgewiesen. Die Wirte
wurden im Juni als Altlarven im Mark junger Spitzentriebe von C. vulgare gesammelt, die Parasiten
schlüpften im Juli aus den Wirtspuparien. Die Art besitzt also entweder mehr als eine Generation im |
Jahr, oder die Weibchen überwintern als Imagines.

Material: 322, Schlüttsiel/ Husum/ Schleswig-Holstein/D (FRI, HO).

* Alle Maße in Y/ıoo mm.

216

Phygadeuon clotho Kriechbaumer, 1892

Phygadeuon (Ischnocryptus) clotho Kriechbaumer, 1892: 344 f. (Aubert 1974: 269).
Phygadeuon grossae Horstmann 1981: 153 ff. (syn. nov.).

Beschreibung

2: Schläfen 1,3-1,4 mal so breit wie die Augen, direkt hinter den Augen ein kleines Stück erweitert
oder ein längeres Stück parallel, zerstreut bis sehr zerstreut punktiert; Scheitel nach dorsal etwas über
das Niveau der Ocellen vorragend; Clypeus basal fein und dicht punktiert, subbasal mit einigen groben
Punkten, teilweise mit feinen Querrunzeln, apical mit zwei dicht benachbarten spitzen Zähnen; Wan-
genraum 0,6 mal so breit wie die Mandibelbasis; Fühler 22-gliedrig, viertes Glied 2,4-2,6 mal so lang
wie breit, Geißel distal nicht keulenförmig erweitert, das breiteste Glied etwa so lang wie breit;
Mesopleuren außerhalb des Speculums zerstreut bis sehr zerstreut punktiert, ohne größere unpunk-
tierte Bereiche; Hinterfemora 3,3-3,6 mal so lang wie hoch; Area superomedia 0,8-1,0 mal so lang wie
breit; zweites Gastertergit frontal sehr fein gekörnelt, caudal glatt, 0,7-0,8 mal so lang wie breit; Bohrer
wie bei P. bavaricus, Bohrerklappen 1,3-1,4 mal so lang wie das erste Gastertergit; Körperlänge 7-9 mm;
drittes bis fünftes Fühlerglied gelbbraun; Coxen und Hinterfemora schwarz; Gaster hinter dem ersten
Segment hell rotbraun oder das siebente Tergit braun gezeichnet. Eine ausführliche Beschreibung

‚ findet sich bei Horstmann (. c.).

d: Von allen Unterschieden, die zwischen den ?? von P. clotho und P. fraternae gefunden wurden,

| scheint nur die Struktur des Clypeus auch für die dd brauchbar zu sein: Bei P. clotho ist der Clypeus

basal auf größeren Flächen fein punktiert und subbasal grob punktiert und fein quergerunzelt, bei
P. fraternae ist er basal kaum fein punktiert, sondern basal und subbasal grob punktiert und nicht

| quergerunzelt.

[

Lebensweise. Als Wirte wurden nachgewiesen: Cheilosia grossa (Fallen) an Carduus nutans, Cirsium
palustre und C. vulgare, leg. Boldt, leg. Freese, leg. Nurse, leg. Rotheray, leg. Stuke (HO, NHML, NMSE,

' ZSM); Cheilosia albipila Meigen an Cirsium palustre und C. vulgare, leg. Freese, leg. Romstöck, leg. Stuke

. (HO, NMSE). Die Wirte wurden von Juni bis September als Larven gesammelt, die Puparien wurden
‚ im Herbst gebildet, die Parasiten schlüpften von März bis Juli aus den Puparien (bei Treibzucht im
Januar). P. clotho ist also ein Endoparasit und Koinobiont (Rotheray 1988: 22; Freese 1997: 76), im

Gegensatz zu vielen anderen Phygadeuon-Arten.

Material: 2022, 984, Gartlea Farm/Dunbartonshire/ GB (NMSE), Aviemore/Inverness/GB, Staunton/ Glouce-
stershire/GB, Tring/Hertfordshire/GB, Ampton/Suffolk/GB, Blythburgh Wood /Suffolk/GB, Timworth/Suf-
folk/GB (alle NHML); Bremen/D, Stade/Niedersachsen/D (alle HO); Goslar/Niedersachsen/D (ZSM);
Worms/ Rheinland-Pfalz/D (NSF); Bayreuth/Bayern/D (HO, NMSE); München/Bayern/D, Setzberg/Kreuth/
Bayern/D (beide ZSM); Sollenau/Niederösterreich/ A (HO); Rom/I (HO, ZSM).

Phygadeuon fraternae, spec. nov.
Figs 2, 9, 14, 22, 25, 36

Typen. Holotypus: $, “Niedersachsen, Landkreis Stade, Schwingetal WW, Cirsium palustre, ex L, 05.09.1996 leg.
Stuke, 20.5.97, Zucht 74”, “ex (Dipt.) Cheilosia fraterna” (ZSM). - Paratypen: 14 aus der gleichen Zucht (HO); 42%,

18 Gartlea Farm/Dunbartonshire/GB, ex Cheilosia fraterna (Meigen), leg. Rotheray; 12 Portland Hill/Midlothi-

an/GB, ex C. fraterna, leg. Robertson (alle NMSE); 12 Bodendorf/Bayreuth/Bayern/D, ex C. fraterna, leg. Freese
(HO); 12, 18 Braken/Stade/Niedersachsen/D, ex Cheilosia chloris (Meigen), leg. Stuke (HO, STU); 12 Blanken-
burg/Thüringen/D, leg. Schmiedeknecht; 5? Garmisch/Bayern/D, 700-1400 m, Juli-September; 1? Mitten-
wald/Bayern/D, 1000m, 7.7.1924; 1? Elmau/Bayern/D, 1050 m, 3.7.1957; 1? Bad Hall/Oberösterreich/ A,
2.-3.8.1942, alle aus Coll. E. Bauer (ZSM); 1? Podcetrtek/Slowenien, Mai 1938, leg. Jaeger (ZSM).

Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen 1,1-1,3 mal so breit wie die Augen, direkt hinter
den Augen etwas erweitert oder ein kurzes Stück parallel (Abb. 2), sehr zerstreut punktiert oder
stellenweise unpunktiert; Scheitel nach dorsal ein wenig über das Niveau der Ocellen vorragend;
Augen kahl; Clypeus etwa so lang wie das Gesicht, im Profil fast flach, vom Gesicht durch eine schwach

217

entwickelte Furche getrennt, basal mit wenigen kräftigen Haarpunkten, ohne Querrunzeln, am Apical-
rand mit einem Doppelzahn (Abb. 9) oder zwei dicht benachbarten spitzen Zähnen (variabel); oberer
Mandibelzahn etwas größer als der untere; Wangenraum 0,6 mal so breit wie die Mandibelbasis; Fühler
20-gliedrig, das vierte Glied 2,1-2,3 mal so lang wie breit (Abb. 14), Distalhälfte der Geißel etwas
keulenförmig, das breiteste Glied 0,9 mal so lang wie breit.

Pronotum lateral zentral mit einer kleinen unpunktierten Stelle, dorsal relativ dicht punktiert, an
den Rändern gerunzelt, Epomia deutlich; Mesoscutum fein zerstreut punktiert, auf den Seitenlappen
zentral jeweils unpunktiert; Mesopleuren außerhalb des großen Speculums zerstreut bis sehr zerstreut

punktiert; Metapleuren ähnlich; Beine kräftig, Hinterfemora 2,9-3,2 mal so lang wie hoch; Areola mit
dem rücklaufenden Nerven distal der Mitte (Abb. 22); Nervellus bei 0,7-0,8 kräftig gebrochen, deutlich

incliv.
Propodeum vollständig gefeldert, in den Feldern sehr fein oder gar nicht strukturiert, Area supe-

romedia 0,6-1,0 mal so lang wie breit (Abb. 25), Area petiolaris etwas eingesenkt, Seitenecken als
breitere Lamellen, aber nicht spitz vorragend; erstes Gastertergit fein gekörnelt, Postpetiolus teilweise
fein längsgerieft, Dorsalkiele bis zu den Stigmen reichend, Dorsolateralleisten bis fast zum Caudalende
divergierend; die folgenden Tergite glatt; das zweite Gastertergit 0,8-0,9 mal so lang wie breit; Gaster

vom dritten Segment an etwas von der Seite zusammengedrückt; Bohrer gerade, schlank zugespitzt,

dorsal ohne deutlichen Nodus, ventral mit feinen Zahnleisten (Abb. 36), Bohrerklappen 1,1-1,2 mal so

lang wie das erste Gastertergit.

Schwarz; Palpen dunkelbraun; Mandibeln ganz schwarz oder median unterschiedlich breit rot-

braun gezeichnet; drittes bis sechstes oder zweites bis achtes Fühlerglied gelbbraun; Tegulae schwarz,

Flügelbasis gelb, Pterostigma dunkelbraun, proximal breit, distal schmal weißlich, Flügelfläche klar;
Trochantellen teilweise rotbraun gemustert; Vorder- und Mittelfemora proximal schwarz, distal rot-

braun, Hinterfemora schwarz, teilweise distal rotbraun gezeichnet; Tibien und Tarsen der Vorder- und
Mittelbeine rotbraun, Tarsenspitzen dunkel, Hintertibien proximal und distal schwarz, median breit

rotbraun, Hintertarsen schwarzbraun; Gaster hinter dem ersten Segment rotbraun, selten das siebente

oder sechste und siebente Tergit dunkel gezeichnet.
Holotypus (?): Kopf 119 breit; Thorax 209 lang, 99 breit (Mesoscutum); Vorderflügel 480 lang;

erstes Gastertergit 105 lang; Postpetiolus 49 lang, 49 breit; zweites Tergit 91 lang, 110 breit; Bohrerklap-

pen 130 lang; Körper etwa 630 lang.

d: Schläfen durchgehend zerstreut punktiert; Fühler 26-gliedrig, das vierte Glied etwa 2,5 mal so
lang wie breit, Tyloide am 12.-15. Glied, Geißel distal zugespitzt; Mesopleuren im Zentrum unpunk-
tiert; Hinterfemora 4,5 mal so lang wie hoch; Postpetiolus und zweites Gastertergit fein längsgestreift;
Gaster caudal nicht von der Seite zusammengedrückt; Fühler ganz schwarz; sonst etwa wie ?.

Lebensweise. Als Wirte wurden nachgewiesen: Cheilosia fraterna (Meigen) an Cirsium palustre, leg.
Freese, leg. Robertson, leg. Rotheray, leg. Stuke; Cheilosia chloris (Meigen) an Cirsium oleraceum, leg.
Stuke. Die Wirte wurden Ende August bis Anfang Oktober als Larven in den Rosetten der Wirtspflan-
zen gesammelt, die Puparien wurden im Spätherbst gebildet, die Parasiten schlüpften zwischen Mai

und Juli (bei Treibzucht Ende März). Hilpert (1987: 153; als Phygadeuon sp. 1) fing die ?? der Art im
August und September. Die Phänologie und die Parasitierungsstrategie dieser Art stimmen anschei-

nend mit denen von P. clotho überein.

Weiteres Material: 1? wurde in Gartlea Farm/Dunbartonshire/GB an Cirsium palustre aus Cheilosia proxima
(Zetterstedt) gezüchtet (leg. Rotheray, NMSE). Es ist nur 5 mm lang, die Fühler sind 19-gliedrig, und der Gaster

ist hinter der Basis des vierten Tergits dunkelbraun bis schwarz. Sonst stimmt es mit P. fraternae gut überein. |

1? und 5354 wurden auf der Reschenalm/ Vinschgau /Südtirol/I in 2200 m Meereshöhe an Cirsium eriophorum |
aus Cheilosia albipila Meigen gezüchtet (leg. Stuke, HO). Das 2 ist etwas beschädigt, passt aber gut zu P. fraternae |
und gar nicht zu P. clotho, dem anderen Parasiten dieser Wirtsart. 19 aus Halland/S (etwas beschädigt und
ausgebleicht) steckt im NRS unter dem Namen P. hercynicus. Zu P. fraternae gehört auch das von Hilpert (1987: |
153 £.) auf dem Feldberg im Schwarzwald /Baden-Württemberg/D gefangene und unter dem Namen Phyga- |
deuon (Iselix) sp. 1 verzeichnete Material.

218

' Phygadeuon hercynicus Gravenhorst, 1829

Phygadeuon hercynicus Gravenhorst, 1829: 709 f. (Aubert 1968: 181). Dem Holotypus (?) fehlen beide Fühler hinter
dem dritten Glied und die Spitze des Bohrers. Soweit zu erkennen ist, stimmt er mit 1? in Coll. Kriechbau-
mer (ZSM) überein, das Kriechbaumer (1892: 341 f.) aus Coll. Siebold erwähnt (ohne Fundortangabe). Dieses
2, bei dem allerdings auch nur die proximale Hälfte eines Fühlers erhalten ist, wird zur Interpretation der
Art herangezogen.

Die beiden von Kriechbaumer unter dem Namen P. hercynicus erwähnten Exemplare aus München-Isarauen
und vom Tegernsee gehören zu P. laevipleuris (ZSM). P. hercynicus sensu Roman (1925: 11) ist ein Gemisch
von mehreren Arten: P. brevitarsis, P. fraternae, P. nitidus, dazu einige defekte Exemplare (NRS).

Beschreibung
2: Schläfen 1,1 mal so breit wie die Augen, direkt hinter den Augen ein kurzes Stück parallel,
zerstreut bis sehr zerstreut punktiert, stellenweise unpunktiert; Scheitel nach dorsal deutlich etwas
über das Niveau der Ocellen vorragend; Clypeus basal deutlich punktiert, ohne Querrunzeln, apical
/ mit zwei dicht benachbarten spitzen Zähnen; Wangenraum 0,6 mal so breit wie die Mandibelbasis;
‚ viertes Fühlerglied 2,0 mal so lang wie breit (nach dem ? aus Coll. Siebold / ZSM; kein Fühler vollstän-
dig erhalten); Mesopleuren außerhalb des Speculums zerstreut bis sehr zerstreut punktiert; Hinterfe-
"mora 3,6-3,7 mal so lang wie hoch; Area superomedia 0,7 mal so lang wie breit; zweites Gastertergit
| glatt, 1,0-1,1 mal so lang wie breit; Bohrer gerade oder etwas abwärts gebogen, wie bei P. bavaricus,
' Bohrerklappen 0,9 mal so lang wie das erste Gastertergit; Körperlänge 5-6 mm; Geißelbasis gelbbraun;
Coxen dunkelbraun, Vorder- und Mittelcoxen apical rotbraun gezeichnet; Hinterfemora und zweites
‚bis viertes Tergit rotbraun, die folgenden Tergite dunkelbraun gezeichnet, caudal schmal gelb geran-
det. Eine ausführliche Beschreibung des Holotypus findet sich bei Frilli (1974: 132 ff.).

Material: 2?%, Harz/Norddeutschland (Coll. Gravenhorst/MPW), ohne Ort (aus Coll. Siebold) (ZSM).

Variation. Einige ?? weichen von der Nominatform vor allem durch die dunkel gezeichneten und
etwas gedrungeneren Hinterfemora ab. Ihr Status ist unklar. Merkmale: Schläfen 1,0-1,1 mal so breit
wie die Augen, hinter den Augen etwas verengt; Fühler 19-20-gliedrig, das vierte Glied 2,0-2,1 mal so
lang wie breit, Distalhälfte der Geißel etwas keulenförmig, das breiteste Glied 0,9 mal so lang wie breit;
Hinterfemora 3,3-3,6 mal so lang wie hoch, vollständig oder nur median dunkelbraun gezeichnet; Area
superomedia 0,7-0,8 mal so lang wie breit; Bohrerklappen 0,8-1,0 mal so lang wie das erste Gastertergit.
Material: 5??, Plumstad/London/GB, Großbritannien (aus Coll. Morley), Deutschland (aus Coll.
' Ruthe) (alle NHML), Krefeld / Nordrhein-Westfalen/D, Worms/Rheinland-Pfalz/D (beide NSF).

Phygadeuon praealpinus, spec. nov.
Eies3, 10, 15.267357

Typen. Holotypus (?): “Ober-Bayern, Garmisch, 700 m, 25.V11.1954, E. Bauer” (ZSM). - Paratypen: 8?? vom
gleichen Fundort, Juli-August; 7?? Ettaler Berg/Garmisch/Bayern/D, 800 m, Juni-Juli; 2?? Kramer/Gar-
misch/Bayern/D, 850 m, August; 222 Riezlern/ Allgäu/Bayern/D, 1150 m, September, alle aus Coll. E. Bauer
(ZSM, 222 aus Garmisch HO).

Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen so lang wie die Augen, direkt hinter den Augen
ein kurzes Stück parallel (Abb. 3), fein und sehr zerstreut punktiert, stellenweise unpunktiert; Scheitel
nach dorsal wenig über das Niveau der Ocellen vorragend; Augen kahl; Clypeus etwa so lang wie das
Gesicht, von diesem durch eine schwach entwickelte Furche getrennt, im Profil fast flach, basal mit
kräftigen getrennten Punkten, ohne Querrunzeln, apical mit zwei deutlich getrennten spitzen Zähnen
(Abb. 10), oberer Mandibelzahn etwas größer als der untere; Wangenraum 0,6 mal so breit wie die
Mandibelbasis; Fühler 21-gliedrig, relativ schlank, das vierte Glied 3,3-3,5 mal so lang wie breit (Abb.
15), Distalhälfte der Geißel kaum keulenförmig, das breiteste Glied 0,9-1,0 mal so lang wie breit.

Pronotum dorsolateral fein und sehr zerstreut punktiert, stellenweise unpunktiert, in der Furche
schwach gestreift, Epomia deutlich; Mesoscutum fein und sehr zerstreut punktiert, stellenweise un-
punktiert; Mesopleuren außerhalb des großen Speculums fein und sehr zerstreut punktiert, im Zen-

219

trum unpunktiert; Metapleuren sehr zerstreut punktiert; Beine relativ schlank, Hinterfemora 3,8-3,9
mal so lang wie hoch; Areola mit dem rücklaufenden Nerv etwas distal der Mitte (etwa wie Abb. 22); |
Nervellus bei 0,7 kräftig gebrochen, etwas incliv.

Propodeum vollständig gefeldert, in den Feldern wenig strukturiert, Area superomedia 0,8-1,0 mal
so lang wie breit (Abb. 26), Area petiolaris wenig eingesenkt, Seitenecken als etwas breitere Lamellen,
nicht zugespitzt; erstes Gastertergit stellenweise sehr fein gekörnelt und mit einigen feinen Punkten,
auf dem Postpetiolus teilweise sehr fein längsgestreift, Dorsalkiele schwach entwickelt, knapp bis zu
den Stigmen reichend, Dorsolateralleisten in der Caudalhälfte des Postpetiolus parallel; die folgenden
Tergite glatt; zweites Tergit etwa 1,5 mal so lang wie breit; Gaster vom zweiten Segment an schwach,
von dritten an deutlich von der Seite zusammengedrückt; Bohrer gerade, dorsal ohne deutlichen
Nodus, ventral mit feinen Zahnleisten (Abb. 37), Bohrerklappen 0,6-0,7 mal so lang wie das erste
Gastertergit.

Schwarz; Palpen gelblich; Mandibeln median breit rotbraun; zweites bis sechstes Fühlerglied hell
rotbraun, Flagellum median dunkelbraun, distal schwarz; Tegulae gelbbraun, Flügelbasis gelb, Ptero-
stigma mittelbraun, proximal und distal wenig aufgehellt, Flügelfläche klar; Coxen dunkelbraun,
apical gelbrot gefleckt (an den Hintercoxen am wenigsten), Trochanteren gelb und braun gemustert
(variabel), Trochantellen gelblich, Femora, Tibien und Tarsen der Vorder- und Mittelbeine hell rot-
braun, Hinterfemora schwarzbraun, proximal und distal etwas aufgehellt, Hintertibien rotbraun,
proximal und distal schmal verdunkelt, Hintertarsen dunkelbraun; zweites bis viertes Gastertergit
rotbraun, die folgenden dunkelbraun bis schwarz, caudal schmal gelb gerandet.

Holotypus (?): Kopf 135 breit; Thorax 236 lang, 110 breit (Mesoscutum); Vorderflügel 530 lang;
erstes Gastertergit 122 lang; Postpetiolus 57 lang, 42 breit; zweites Tergit 104 lang, 69 breit; Bohrerklap-
pen 88 lang; Körper etwa 660 lang.

d unbekannt.

Phygadeuon thomsoni Roman, 1925

Phygadeuon thomsoni Roman, 1925: 11 f. - Lectotypus (?) hiermit festgelegt: “L-d” (= Lund /Scäne/S), “Hercyni-
cus” (Coll. Thomson/ZIL, unter P. hercynicus). Als Paralectotypen sind in Coll. Thomson 484 vom Fundort
Lund vorhanden, die zur Zeit unbestimmbar sind. Ein Paralectotypus (2) im NRS gehört wahrscheinlich zu '
P. bavaricus. |

Roman (l. c.) hat P. thomsoni als “n. nov.” für P. hercynicus sensu Thomson (1884: 958 f.) nec P. hercynicus '
Gravenhorst eingeführt. Da Thomson keine neue Art beschrieben, sondern Gravenhorst als Autor zitiert hat, |
handelt es sich nicht um ein Nomen novum im Sinne der Nomenklaturregeln, sondern um eine Neube-
schreibung. |

Beschreibung

2: Schläfen 1,1-1,2 mal so breit wie die Augen, direkt hinter den Augen ein kurzes Stück parallel,
zerstreut bis sehr zerstreut punktiert; Scheitel nach dorsal nicht über das Niveau der Ocellen vorra-
gend; Clypeus basal deutlich punktiert, ohne Querrunzeln, apical mit zwei deutlich getrennten spitzen
Zähnen; Wangenraum 0,6 mal so breit wie die Mandibelbasis; Fühler 20-gliedrig, das vierte Glied 2,0
mal so lang wie breit, Distalhälfte der Geißel etwas keulenförmig, das breiteste Glied 0,9 mal so lang
wie breit; Mesopleuren außerhalb des Speculums gleichmäßig zerstreut bis sehr zerstreut punktiert;
Hinterfemora 3,1-3,2 mal so lang wie hoch; Area superomedia 0,8-0,9 mal so lang wie breit; zweites
Gastertergit glatt, 0,8 mal so lang wie breit; Bohrer wie bei P. bavaricus, Bohrerklappen so lang wie das |
erste Gastertergit; Körperlänge 5-6 mm; Geißelbasis gelbbraun; Coxen schwarz; Hinterfemora rot- |
braun; zweites bis viertes Gastertergit rotbraun, teilweise das vierte caudal verdunkelt, die folgenden
dunkelbraun, caudal schmal gelb gerandet. |

Material: 3??, Wicken Fen/Cambridgeshire/GB (NHML); Lund/Scäne/S (ZIL), Krefeld /Nordrhein-Westfa-
len/D (ZSM). |

220

) Phygadeuon unidentatus, spec. nov.
| Figs 4, 11, 16, 27, 38

‚Holotypus (2): “Harz Harzburg, Radautal, 14.1X.1943, E. Bauer” (ZSM).

‚Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen 1,4 mal so breit wie die Augen, direkt hinter den
Augen ein kurzer Stück erweitert (Abb. 4), fein und sehr zerstreut punktiert, stellenweise unpunktiert;
Scheitel nach dorsal deutlich über das Niveau der Ocellen vorragend; Augen kahl; Clypeus etwa so
lang wie das Gesicht, von diesem kaum getrennt, im Profil fast flach, basal mit einigen groben Punkten,
apical mit einem kleinen spitzen Zahn (Abb. 11); oberer Mandibelzahn wenig größer als der untere;
"Wangenraum 0,6 mal so breit wie die Mandibelbasis; Fühler 19-gliedrig, das vierte Glied 2,0 mal so
lang wie breit (Abb. 16), Distalhälfte der Geißel etwas keulenförmig, das breiteste Glied 0,9 mal so lang
wie breit.

Pronotum dorsolateral sehr zerstreut punktiert, in der Furche gestreift, Epomia deutlich; Mesoscu-
tum fein und sehr zerstreut punktiert, auf den Seitenlappen größere Bereiche unpunktiert; Mesopleu-
ren außerhalb des großen Speculums fein und sehr zerstreut punktiert; Metapleuren ähnlich; Beine
kräftig, Hinterfemora 2,9 mal so lang wie hoch; Areola regelmäßig (wie Abb. 21); Nervellus bei 0,7
‚kräftig gebrochen, etwas incliv.

Propodeum lang, fein und vollständig gefeldert, in den Feldern kaum strukturiert, Area supero-
‚media 1,2 mal so lang wie breit (Abb. 27), Area petiolaris wenig eingesenkt, Seitenecken als etwas
breitere Lamellen, nicht zugespitzt; erstes Gastertergit sehr fein gekörnelt, fast glatt, neben den Stig-
men sehr fein längsgestreift, Dorsalkiele nicht bis zu den Stigmen reichend, Dorsolateralleisten fast bis

zum Caudalende divergierend; die folgenden Tergite glatt; zweites Tergit 1,2 mal so lang wie breit;

Gaster vom zweiten Tergit an etwas von der Seite zusammengedrückt; Bohrer schlank, wenig abwärts

gebogen, dorsal ohne deutlichen Nodus, ventral mit feinen Zahnleisten (Abb. 38), Bohrerklappen 1,2
"mal so lang wie das erste Gastertergit.
Schwarz; Palpen gelblich; Mandibeln median breit gelbrot; zweites bis sechstes Fühlerglied hell
‚rotbraun, Flagellum median dunkelbraun, distal schwarz; Tegulae gelbbraun, Flügelbasis gelb, Ptero-
‚stigma mittelbraun, proximal und distal weißlich, Flügelfläche klar; Beine hell rotbraun, Vorder- und
‚ Mittelcoxen basal und Hintercoxen fast ganz dunkelbraun, Hintertrochanteren und Hintertibien pro-
ximal verdunkelt; zweites bis viertes Gastertergit rotbraun, die folgenden dunkelbraun, caudal schmal
gelb gerandet, Postpetiolus caudal-median rotbraun gezeichnet.

Holotypus (?): Kopf 110 breit; Thorax 209 lang, 88 breit (Mesoscutum); Vorderflügel 495 lang;
‚erstes Gastertergit 96 lang; Postpetiolus 42 lang, 42 breit; zweites Tergit 85 lang, 69 breit; Bohrerklappen
‚110 lang; Körper etwa 620 lang.

d unbekannt.

Artengruppe lachesis Kriechbaumer

Phygadeuon lachesis Kriechbaumer, 1892
Figs 12, 39

‚ Phygadeuon (Ischnocryptus) lachesis Kriechbaumer, 1892: 345 f. (Aubert 1974: 269).

ı Beschreibung

| 2: Schläfen 1,1-1,2 mal so breit wie die Augen, direkt hinter den Augen ein kurzes Stück parallel,
. deutlich zerstreut bis sehr zerstreut punktiert, an kleinen Stellen unpunktiert; Scheitel nach dorsal sehr
‚ wenig über das Niveau der Ocellen vorragend; Clypeus basal deutlich punktiert, apical mit einem
‚ lamellenartigen schwach doppelt gewellten Rand, die Zähne nur als subapicale Oberflächenstruktur
‚ erkennbar (Abb. 12); Wangenraum 0,5-0,6 mal so breit wie die Mandibelbasis; Fühler 23-gliedrig, das
| vierte Glied 2,0-2,1 mal so lang wie breit, Geißel in der Distalhälfte etwas keulenförmig erweitert, das
breiteste Glied etwa 0,9 mal so lang wie breit; Mesopleuren außerhalb des Speculums deutlich zerstreut
_ bis sehr zerstreut punktiert; Hinterfemora 2,8-3,1 mal so lang wie hoch; Area superomedia 0,8-1,0 mal
so lang wie breit; zweites Gastertergit glatt, 0,8-0,9 mal so lang wie breit; Bohrer schlank, wenig abwärts

221

gebogen, dorsal vor dem Nodus mit einer rundlichen Vorwölbung, der Nodus selbst nur als kleine
Runzel ausgebildet, ventral mit feinen Zahnleisten (Abb. 39), Bohrerklappen 1,5 mal so lang wie das
erste Gastertergit; Körperlänge 6-7 mm; drittes bis fünftes oder sechstes Fühlerglied rotbraun; Coxen
schwarz; Hinterfemora schwarz, ventral-distal zuweilen rotbraun gezeichnet; Gaster hinter den ersten
Segment hell rotbraun, die letzten Tergite caudal gelb gerandet, der Postpetiolus zuweilen caudal-
median rotbraun gefleckt.

Material: 5%%, Garmisch/Bayern/D, Murnau/Bayern/D, Setzberg/Kreuth/Bayern/D (ZSM, 1? HO).

Artengruppe nitidus Gravenhorst
Phygadeuon atropos Kriechbaumer, 1892

Phygadeuon (Ischnocryptus) atropos Kriechbaumer, 1892: 346 - Lectotypus (?) von Aubert (1968: 193) festgelegt:
“M. Hess. 19.5.86. Krchb.” (= München-Hesselohe). Aubert (1974: 268 f.) führt 1? mit dem Etikett “Teg. |
6.9.56. Krchb.” (= Tegernsee) als “type” an (beide ZSM), ohne seine frühere Festlegung zu berücksichtigen.

Ischnocryptus atropos (Kriechbaumer) var. rufifemur Seyrig, 1927: 79 (syn. nov.) - Holotypus (?): “Vendresse
25.8.26 Benoist” (MHNP).

Aubert (1974: 269) hat P. atropus als jüngeres Synonym zu P. forticornis (Kriechbaumer) gestellt. Zwischen beiden |
Taxa bestehen aber einige Unterschiede (siehe Bestimmungsschlüssel), sie werden deshalb als verschiedene
Arten geführt. Der Holotypus der var. rufifemur Seyrig unterscheidet sich von der Nominatform nur durch
die rotbraunen Hinterfemora, beide werden deshalb hier synonymisiert.

er —

Beschreibung

9: Schläfen so breit wie die Augen, direkt hinter den Augen ein kurzes Stück parallel oder verengt,
sehr zerstreut punktiert, stellenweise unpunktiert; Scheitel nach dorsal nicht über das Niveau der
Ocellen vorragend; Clypeus basal deutlich punktiert, ohne Querrunzeln, apical mit zwei deutlich
getrennten spitzen Zähnen; Wangenraum 0,6-0,7 mal so breit wie die Mandibelbasis; Fühler 19-20-
gliedrig, das vierte Glied 1,9 mal so lang wie breit, Geißel in der Distalhälfte etwas keulenförmig, das
breiteste Glied 0,8-0,9 mal so lang wie breit; Mesopleuren außerhalb des Speculums fein und sehr
zerstreut punktiert, teilweise im Zentrum unpunktiert; Hinterfemora 2,9-3,1 mal so lang wie hoch;
Area superomedia 0,8-1,0 mal so lang wie breit; zweites Gastertergit glatt, 0,7-0,8 mal so lang wie breit;
Bohrer wie bei P. laevipleuris, Bohrerklappen 1,0-1,1 mal so lang wie das erste Gastertergit; Körperlänge '
4-6 mm; Geißelbasis in der Regel nur wenig braun überlaufen, selten das dritte bis fünfte Fühlerglied '
gelbbraun; Coxen schwarz; Hinterfemora schwarz, distal aufgehellt (Nominatform), oder rotbraun
(var. rufifemur Seyrig); zweites bis viertes, fünftes oder sechstes Gastertergit hell rotbraun, die folgen-
den dunkelbraun, caudal schmal gelb gerandet.

Material: 11?2, Newlands/Gloucestershire/GB, Porlock/Somerset/GB, ohne Ort aus Coll. Morley (vermutlich
aus GB) (alle NHML); Mellum/Niedersachsen/D (HO); Babenhausen/Hessen/D (Frankfurt), Garmisch/Bay-
ern/D, München/Bayern/D, Tegernsee/Bayern/D (alle ZSM); Vendresse/ Ardennes/F (MHNP). |

Phygadeuon forticornis Kriechbaumer, 1892

Phygadeuon (Ischnocryptus) forticornis Kriechbaumer, 1892: 344 (Aubert 1974: 269). Da dem Holotypus, dem |
einzigen bisher bekannten Exemplar der Art, die Geißeln fehlen, ist die Interpretation der Art unsicher. |
Beschreibung |
9: Schläfen 1,1 mal so breit wie die Augen, direkt hinter den Augen ein kurzes Stück erweitert, fein |
und sehr zerstreut punktiert; Clypeus basal deutlich punktiert und etwas quergerunzelt, apical mit |
zwei deutlich getrennten spitzen Zähnen; Wangenraum 0,9 mal so breit wie die Mandibelbasis; |
Mesopleuren außerhalb des Speculums fein und sehr zerstreut punktiert; Hinterfemora 3,6 mal so lang
wie hoch; Area superomedia so lang wie breit; zweites Gastertergit glatt, so lang wie breit; Bohrer wie |
bei P. Iaevipleuris, Bohrerklappen 0,9 mal so lang wie das erste Gastersegment; Körperlänge 9 mm; ”
Coxen und Hinterfemora schwarz; Gaster hinter dem ersten Segment rotbraun. |

Material: 12, München/Bayern/D (ZSM).

222.

Phygadeuon habermehli Roman, 1930

Phygadeuon (Ischnocryptus) habermehli Roman, 1930: 6 f. (Townes et al. 1965: 148). Der Lectotypus (?) aus
Kamtschatka ist das einzige untersuchte Exemplar der Nominatform. In Europa kommt eine Morphe vor,
die durch ganz rotbraune Hinterfemora und eine etwas größere Körperlänge abweicht; sie wird ebenfalls
zu P. habermehli gestellt.

Beschreibung

@: Schläfen 1,1 mal so breit wie die Augen, direkt hinter den Augen kaum verengt, fein und sehr
zerstreut punktiert, größere Bereiche unpunktiert; Scheitel nach dorsal nicht über das Niveau der
Ocellen vorragend; Clypeus basal und median grob und zerstreut punktiert, apical mit zwei deutlich
getrennten spitzen Zähnen; Wangenraum 0,6 mal so breit wie die Mandibelbasis; Fühler 20-gliedrig
(bei ?2 aus Mitteleuropa; dem Lectotypus fehlen beide Geißelspitzen), das vierte Glied zweimal so
lang wie breit, Distalhälfte der Geißel etwas keulenförmig, das breiteste Glied 0,8 mal so lang wie breit;
Mesopleuren außerhalb des Speculums zerstreut bis sehr zerstreut punktiert, aber ohne größere
unpunktierte Stellen; Hinterfemora 3,3 mal so lang wie hoch; Area superomedia 0,8 mal so lang wie
breit; zweites Gastertergit glatt, 0,7 mal so lang wie breit; Bohrer wie bei P. laevipleuris, Bohrerklappen
0,6-0,7 mal so lang wie das erste Gastertergit; Körperlänge 5 mm (beim Lectotypus) oder 6 mm (bei ??
aus Mitteleuropa); Geißelbasis nur wenig braun überlaufen; Coxen schwarz; Hinterfemora dunkel-
braun (beim Lectotypus) oder rotbraun (bei ?? aus Mitteleuropa); zweites bis viertes Gastertergit
rotbraun, die folgenden dunkelbraun, caudal gelb gerandet.

Material: 5??, Kamtschatka/Ost-Sibirien (NRS), Goslar/Niedersachsen/D, Garmisch/Bayern/D, ohne Ort
aus Coll. Schmiedeknecht (vermutlich aus Thüringen/D) (ZSM, 1? HO).

Phygadeuon laevipleuris, spec. nov.
Figs 5, 17, 28, 40

Typen. Holotypus: 2, “Weibersbrunn, Spessart, 13.9.67”, “Bayern, Horstm. leg.” (ZSM). — Paratypen: 1? vom
gleichen Fundort und -tag; 2?? Mespelbrunn/Bayern/D, 7.9.1967, leg. Horstmann (alle HO); 3?? Garmisch/
Bayern/D, 700 m, Juli-September; 1? Andechs/Bayern/D, 22.8.1935; 1? Rogaska Slatina/Slowenien, 14.8.1939,
alle aus Coll. E. Bauer (ZSM).

Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen so breit wie die Augen, direkt hinter den Augen
ein kurzes Stück parallel (Abb. 5), mit feinen, stellenweise weit voneinander entfernten Haarpunkten;
Scheitel nach dorsal nicht über das Niveau der Ocellen vorragend; Augen kahl; Clypeus etwa so lang
wie das Gesicht, von diesem durch eine wenig entwickelte Furche getrennt, im Profil flach, basal mit
kräftigen voneinander getrennten Haarpunkten, ohne Querrunzeln, am Apicalrand mit zwei deutlich
getrennten spitzen Zähnen; oberer Mandibelzahn etwas größer als der untere; Wangenraum 0,7 mal
so breit wie die Mandibelbasis; Fühler 20-21-gliedrig, das vierte Glied 2,3-2,4 mal so lang wie breit
(Abb. 17), Distalhälfte der Geißel etwas keulenförmig, das breiteste Glied 0,8 mal so lang wie breit.

Pronotum dorsolateral zerstreut punktiert, an den Rändern gerunzelt, Epomia deutlich; Mesoscu-
tum fein und sehr zerstreut punktiert, Seitenlappen zentral jeweils an kleinen Stellen unpunktiert;
Mesopleuren außerhalb des großen Speculums im Zentrum unpunktiert, an den Rändern zerstreut bis
sehr zerstreut punktiert; Metapleuren sehr zerstreut punktiert; Beine kräftig, Hinterfemora 3,3-3,7 mal
so lang wie hoch; Areola mit dem rücklaufenden Nerven etwas distal der Mitte (etwa wie Abb. 22);
Nervellus bei 0,7 kräftig gebrochen, deutlich etwas incliv.

Propodeum vollständig gefeldert, in den Feldern wenig strukturiert, Area superomedia 0,8-1,0 mal
so lang wie breit (Abb. 28), Area petiolaris wenig eingesenkt, Seitenecken als etwas breitere Lamellen,
aber nicht zugespitzt; erstes Gastertergit sehr fein gekörnelt, stellenweise sehr fein zerflossen längs-
runzlig, Postpetiolus dorsolateral fein längsrissig (variabel), Dorsalkiele bis zu den Stigmen reichend,
Dorsolateralleisten bis fast zum Caudalende divergierend; die folgenden Tergite glatt; zweites Tergit
1,0-1,1 mal so lang wie breit; Gaster vom dritten Segment an etwas von der Seite zusammengedrückt;
Bohrer schlank, gerade oder wenig abwärts gebogen, dorsal am Nodus mit einem zahnartigen Vor-
sprung, davor mit einem weiteren Zahn, ventral mit feinen Zahnleisten (Abb. 40), Bohrerklappen 1,0

223

mal so lang wie das erste Gastertergit.

Schwarz; Palpen braun; Mandibeln und Tegulae schwarzbraun, selten Mandibeln median rotbraun

gezeichnet; drittes bis fünftes Fühlerglied gelbbraun, dann Geißel zunehmend dunkler, median und
distal schwarz; Flügelbasis gelb, Pterostigma schwarzbraun, proximal wenig aufgehellt, Flügelfläche
klar; Coxen und Trochanteren schwarzbraun, Trochantellen, Femora, Tibien und Tarsen rotbraun,
Hintertibien proximal und alle Tarsen distal verdunkelt; zweites und drittes Gastertergit rotbraun,
häufig auch die Seiten des vierten, die folgenden dunkelbraun, caudal schmal gelb gerandet, Postpe-
tiolus median-caudal rotbraun gezeichnet.

Holotypus (?): Kopf 144 breit; Thorax 243 lang, 116 breit (Mesoscutum); Vorderflügel 540 lang;
erstes Gastertergit 121 lang; Postpetiolus 55 lang, 52 breit; zweites Tergit 100 lang, 94 breit; Bohrerklap-
pen 118 lang; Körper etwa 720 lang.

d unbekannt.

Weiteres Material: Zu P. laevipleuris gehören auch die beiden von Kriechbaumer (1992: 342) unter dem Namen

P. hercynicus erwähnten 2? (beide mit gebrochenen Fühlern) aus München-Isarauen/D, 9.9.1884, und vom
Tegernsee/Bayern/D, 11.8.1855 (ZSM).

Phygadeuon nigrifemur, spec. nov.
Figs 6, 18, 29, 41

Typen. Holotypus: ?, “Weibersbrunn, Spessart, 4.9.1967”, “Bayern, Horstm. leg.” (ZSM). - Paratypen: 1? ohne

Ort aus Coll. Morley (vermutlich GB) (NHML); 1? Worms/Rheinland-Pfalz/D, leg. Habermehl (NSF); 12

Garmisch /Bayern/D, 700 m, 30.8.1942, aus Coll. E. Bauer (ZSM); 12 Rojental/ Reschenpass/Südtirol/I, 2200 m,
19.8.1983, leg. Horstmann (HO).

Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen 0,9 mal so breit wie die Augen, hinter den Augen |
verengt (Abb. 6), mit feinen, stellenweise weit voneinander entfernten Haarpunkten; Scheitel nach
dorsal nicht über das Niveau der Ocellen vorragend; Augen kahl; Clypeus etwa so lang wie das

Gesicht, von diesem durch eine wenig entwickelte Furche getrennt, im Profil flach, basal mit kräftigen

voneinander getrennten Haarpunkten, ohne Querrunzeln, am Apicalrand mit zwei deutlich getrenn-

ten spitzen Zähnen; oberer Mandibelzahn etwas größer als der untere; Wangenraum 0,7 mal so breit
wie die Mandibelbasis; Fühler 20-gliedrig, das vierte Glied 2,3-2,4 mal so lang wie breit (Abb. 18),
Distalhälfte der Geißel etwas keulenförmig, das breiteste Glied 0,9 mal so lang wie breit.

Pronotum lateral zentral auf einer kleinen Stelle sehr zerstreut punktiert oder unpunktiert, sonst

zerstreut punktiert, an den Rändern gerunzelt, Epomia deutlich; Mesoscutum überwiegend fein und

sehr zerstreut punktiert, Seitenlappen zentral jeweils stellenweise unpunktiert; Mesopleuren außer-

halb des großen Speculums zerstreut bis sehr zerstreut punktiert, teilweise im Zentrum unpunktiert;

Metapleuren sehr zerstreut punktiert; Beine kräftig, Hinterfemora 3,5-3,7 mal so lang wie hoch; Areola

mit dem rücklaufenden Nerven etwas distal der Mitte (etwa wie Abb. 22); Nervellus bei 0,7 kräftig

gebrochen, etwas incliv.

Propodeum vollständig gefeldert, in den Feldern kaum strukturiert, Area superomedia 0,5-0,7 mal |

so lang wie breit (Abb. 29), Area petiolaris wenig eingesenkt, Seitenecken als etwas breitere Leisten,

aber nicht spitz vorragend; erstes Gastertergit sehr fein gekörnelt, fast glatt, Dorsalkiele fast bis zu den
Stigmen reichend, Dorsolateralleisten bis fast zum Caudalende divergierend; die folgenden Tergite |
glatt; zweites Tergit 0,7-0,9 mal so lang wie breit; Gaster vom dritten Segment an etwas von der Seite |
zusammengedrückt; Bohrer schlank, gerade, dorsal am Nodus mit einem zahnartigen Vorsprung,

davor mit einem weiteren Zahn, ventral mit feinen Zahnleisten (Abb. 41), Bohrerklappen 1,3 mal so
lang wie das erste Gastertergit.

Schwarz; Palpen braun; Mandibeln und Tegulae in der Regel schwarzbraun, selten Mandibeln
median rotbraun gezeichnet; drittes bis fünftes Fühlerglied braun überlaufen (variabel); Flügelbasis
gelb, Pterostigma dunkelbraun, proximal deutlich aufgehellt, Flügelfläche etwas getrübt; Coxen, Tro-

chanteren, Basis der Vorder- und Mittelfemora und die Hinterfemora dunkelbraun bis schwarz,

Vorderfemora zuweilen nicht verdunkelt oder Hinterfemora proximal und distal aufgehellt, Trochan-

tellen rotbraun bis braun, Femora, Tibien und Tarsen sonst hell rotbraun, Hintertibien proximal und

224

|
|

distal schmal verdunkelt, Tarsen distal verdunkelt; zweites bis viertes Gastertergit rotbraun, die
folgenden dunkelbraun, caudal schmal gelb gezeichnet, Postpetiolus median-caudal rotbraun gezeich-
net.

Holotypus (?): Kopf 122 breit; Thorax 200 lang, 108 breit (Mesoscutum); Vorderflügel 440 lang;
erstes Gastertergit 102 lang; Postpetiolus 42 lang, 52 breit; zweites Tergit 79 lang, 97 breit; Bohrerklap-

‚ pen 130 lang; Körper etwa 570 lang.

d unbekannt.

Phygadeuon nitidus Gravenhorst, 1829

Phygadeuon nitidus Gravenhorst, 1829: 708 f. (Aubert 1968: 182). Dem Lectotypus fehlt fast der ganze Bohrer

einschließlich der Bohrerklappen. Er stimmt mit 1? aus Dalecarlia (= Kopparberg/S) im NRS sehr gut
überein; dieses wird zur Interpretation herangezogen. Der Lectotypus trägt die Fundortangabe “Cudowa
VIII” (= Kudowa/PL). Das Etikett stammt nicht von Gravenhorst, sondern wurde später zugefügt, vermut-
lich aufgrund der Beschreibung. Diese nennt aber zwei Fundorte, und es ist unklar, ob dem Schreiber des
Etiketts zusätzliche Informationen vorlagen.

Beschreibung

?: Schläfen etwa so breit wie die Augen, direkt hinter den Augen ein kurzes Stück parallel,
zerstreut bis sehr zerstreut punktiert; Scheitel nach dorsal nicht über das Niveau der Ocellen vorra-
gend; Clypeus basal deutlich punktiert, ohne Querrunzeln, apical mit zwei getrennten spitzen Zähnen;
Wangenraum 0,7 mal so breit wie die Mandibelbasis; Fühler 19-20-gliedrig, das vierte Glied 2,4-2,7 mal
so lang wie breit, Distalhälfte der Geißel etwas keulenförmig, das breiteste Glied 0,9 mal so lang wie
breit; Mesopleuren außerhalb des Speculums gleichmäßig zerstreut bis sehr zerstreut punktiert; Hin-
terfemora 3,5-3,6 mal so lang wie hoch; Area superomedia 0,5-0,6 mal so lang wie breit; zweites
Gastertergit glatt, 0,7-0,8 mal so lang wie breit; Bohrer wie bei P. Iaevipleuris, Bohrerklappen 0,9 mal so
lang wie das erste Gastertergit (nach 1? aus Schweden); Körperlänge 6-7 mm; Geißelbasis gelbbraun
bis dunkelbraun überlaufen; Coxen schwarz; Hinterfemora ganz rotbraun; zweites bis drittes oder
viertes Gastertergit rotbraun, die folgenden dunkelbraun, caudal schmal gelb gerandet. Eine ausführ-
liche Beschreibung des Lectotypus findet sich bei Frilli (1974: 136 ff.).

Material: 772, Dalecarlia (= Kopparberg/S) (NRS); ohne Ort aus Coll. Desvignes (vermutlich GB) (NHML);
Worms/Rheinland-Pfalz/D (NSF); ? Kudowa/PL (siehe oben) (MPW),.

Artengruppe ponojensis (Hellen)

Phygadeuon atricolor, spec. nov.
Figs 7, 19,723, 30, 42

Holotypus: 2, “Ober-Bayern, Garmisch, Kreuzeckweg bei ca. 900 m. 21.VII.1925. E. Bauer” (ZSM).

Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen so breit wie die Augen, direkt hinter den Augen
ein wenig, dann rundlich verengt (Abb. 7), fein und sehr zerstreut punktiert, stellenweise unpunktiert;
Scheitel nach dorsal nicht über das Niveau der Ocellen vorragend; Augen kahl; Wangenraum 0,7 mal
so breit wie die Mandibelbasis; Clypeus knapp so lang wie das Gesicht, basal deutlich punktiert, apical
mit zwei deutlich getrennten spitzen Zähnen; Fühler 20-gliedrig, das vierte Glied 2,7 mal so lang wie
breit (Abb. 19), Distalhälfte der Geißel nicht keulenförmig erweitert, Glieder dort wenig länger als breit.

Pronotum lateral überwiegend relativ dicht punktiert, im Zentrum an einer kleinen Stelle unpunk-
tiert, in der Furche gestreift, Epomia deutlich; Mesoscutum fein und sehr zerstreut punktiert, auf den
Seitenlappen fast ganz unpunktiert; Mesopleuren außerhalb des großen Speculums fein und sehr
zerstreut punktiert, an kleinen Stellen unpunktiert, Metapleuren ähnlich; Beine relativ kräftig, Hinter-
femora 3,9 mal so lang wie hoch; Areola höher als breit (Abb. 23); Nervellus bei 0,7 kräftig gebrochen,
deutlich incliv.

Propodeum vollständig und deutlich gefeldert, in den Feldern auf glattem Grund stellenweise

225

zerflossen gerunzelt, Area superomedia 0,7 mal so lang wie breit (Abb. 30), Area petiolaris etwas
eingesenkt, Seitenecken als breitere Lamellen, aber nicht spitz vorragend; erstes Gastertergit stellen-

weise fein gekörnelt und mit feinen Längsrunzeln, Postpetiolus caudal glatt, Dorsalkiele über die

Stigmen etwas hinausreichend, Dorsolateralleisten bis fast zum Caudalende divergierend; die folgen-
den Tergite glatt: zweites Tergit 0,7 mal so lang wie breit; Gaster vom Caudalende des dritten Segments
an etwas von der Seite zusammengedrückt; Bohrer gerade, dorsal am Nodus deutlich gewinkelt,
ventral mit deutlichen Zahnleisten (Abb. 42), Bohrerklappen 1,6 mal so lang wie das erste Gastertergit.

Schwarz; Palpen gelbbraun; Tegulae schwarzbraun, Flügelbasis gelblich, Pterostigma dunkel-

braun, proximal und distal wenig aufgehellt, Flügelfläche klar; an den Vorder- und Mittelbeinen die '

Trochantellen gelbbraun und dunkelbraun gemustert, die Femora proximal dunkelbraun, median und
distal gelbbraun, die Tibien und Tarsen gelbbraun; an den Hinterbeinen auch die Trochantellen und
Femora schwarzbraun, die Tibien median breit gelbbraun, proximal und distal schmal verdunkelt, die
Tarsen braun; Gaster hinter dem ersten Segment rotbraun, die hinteren Tergite unbestimmt braun
überlaufen, caudal gelb gerandet.

Holotypus (2): Kopf 121 breit; Thorax 204 lang, 110 breit (Mesoscutum); Vorderflügel 495 lang;
erstes Gastertergit 100 lang; Postpetiolus 47 lang, 55 breit; zweites Tergit 77 lang, 104 breit; Bohrerklap-
pen 157 lang; Körper 600 lang.

ö unbekannt.

Phygadeuon camargator Aubert, 1982, stat. nov.
Phygadeuon (Iselix) nitidus Gravenhorst camargator Aubert, 1982: 35.

Beschreibung

?: Schläfen knapp so breit wie die Augen, direkt hinter den Augen kaum, caudal deutlich verengt, |
fein und sehr zerstreut punktiert, kleine Bereiche unpunktiert; Scheitel nach dorsal nicht über das |

Niveau der Ocellen vorragend; Clypeus basal fein und dicht, subbasal kräftig und sehr zerstreut

punktiert, apical mit zwei relativ stumpfen Zähnen; Wangenraum 0,9 mal so breit wie die Mandibel-

basis; Fühler 25-gliedrig, das vierte Glied 2,6 mal so lang wie breit, Distalhälfte der Geißel nicht deutlich

keulenförmig, Glieder im distalen Drittel der Geißel etwa so lang wie breit; Mesopleuren außerhalb des |

Speculums deutlich und mäßig dicht punktiert, ventral relativ dicht punktiert und etwas punktrissig;
Hinterfemora 3,8 mal so lang wie hoch; Area superomedia 0,7 mal so lang wie breit; zweites Gaster-
tergit glatt, breiter als lang; Bohrer wie bei P. atricolor, Bohrerklappen 0,9 mal so lang wie das erste
Gastertergit; Körperlänge etwa 6 mm; Fühler und Coxen schwarz; Hinterfemora proximal und median
schwarzbraun, dann rotbraun, das äußerste Distalende schwarz; zweites bis viertes und dazu die

Frontalhälfte des fünften Gastertergits rotbraun, die folgenden schwarzbraun, caudal gelb gerandet.

Material: 17, Sylvereal/Bouches-du-Rhöne/F (Coll. Aubert/MZL).

Phygadeuon macrocephalus, spec. nov.
Figs 8, 20, 31, 43

Typen. Holotypus: ?, “Ober-Bayern, Garmisch, 700 m, 25.VIl.1940. G. Thelemann” (Coll. E. Bauer/ZSM). -

Paratypus: 1? vom gleichen Ort, Fangdatum 26.8.1950, leg. E. Bauer (ZSM).

Beschreibung

2: Kopf und Thorax mit glattem Grund; Schläfen 1,2 mal so breit wie die Augen, direkt hinter den
Augen ein wenig erweitert (Abb. 8), sehr zerstreut punktiert, stellenweise unpunktiert; Scheitel nach
dorsal deutlich über das Niveau der Ocellen vorragend; Augen kahl; Clypeus wenig länger als das
Gesicht (dieses auffällig kurz), basal kräftig punktiert, apical mit zwei voneinander getrennten spitzen
Zähnen; Wangenraum 0,5 mal so breit wie die Mandibelbasis; Fühler 20-gliedrig, das vierte Glied 2,2
mal so lang wie breit (Abb. 20), Distalhälfte der Geißel wenig keulenförmig erweitert, Glieder dort etwa
so lang wie breit.

226

Pronotum lateral kräftig und dicht punktiert, in der Furche gerunzelt, Epomia deutlich; Mesoscu-
tum sehr zerstreut punktiert, Seitenlappen zentral jeweils an kleinen Stellen unpunktiert; Mesopleuren
außerhalb des großen Speculums deutlich, aber zerstreut bis sehr zerstreut punktiert; Metapleuren
deutlich und relativ dicht punktiert; Beine kräftig, Hinterfemora 2,8 mal so lang wie hoch; Areola
regelmäßig oder mit dem rücklaufenden Nerven etwas distal der Mitte (variabel); Nervellus bei 0,7

kräftig gebrochen, deutlich incliv.

Propodeum vollständig, aber relativ fein gefeldert, in den Feldern auf glattem Grund zerflossen

‚ punktiert, nur die vorderen Seitenfelder fast glatt, Area supromedia 0,8-0,9 mal so lang wie breit (Abb.

31), Area petiolaris wenig eingesenkt, Seitenecken als breite Lamellen, aber nicht spitz vorragend;
erstes Gastertergit fein gekörnelt, kaum gestreift, Dorsalkiele fein, knapp bis zu den Stigmen reichend,
Dorsolateralleisten bis fast zum Caudalende divergierend; die folgenden Tergite glatt; zweites Tergit
0,6-0,7 mal so lang wie breit; Gaster vom Caudalende des dritten Tergits an etwas von der Seite
zusammengedrückt; Bohrer gerade, dorsal am Nodus deutlich gewinkelt, ventral mit deutlichen
Zahnleisten (Abb. 43), Bohrerklappen 1,3-1,4 mal so lang wie das erste Gastertergit.

Schwarz; Palpen gelbbraun; Mandibeln median gelbbraun überlaufen; drittes bis fünftes Fühler-
glied rotbraun (teilweise etwas dunkel überlaufen); Tegulae rotbraun bis dunkelbraun, Flügelbasis
hellgelb, Pterostigma dunkelbraun, proximal und distal schmal weißlich, Flügelfläche klar; Coxen
schwarzbraun, apical aufgehellt; Trochanteren dunkelbraun und gelbbraun gemustert; Beine sonst hell
rotbraun, Hintertibien proximal und distal und Hintertarsen dunkelbraun gezeichnet; zweites, drittes
und die Basis des vierten Gastertergits rotbraun, die folgenden dunkelbraun, caudal gelb gerandet.

Holotypus (?): Kopf 122 breit; Thorax 206 lang, 102 breit (Mesoscutum); Vorderflügel 470 lang;
erstes Gastertergit 91 lang; Postpetiolus 44 lang, 50 breit; zweites Tergit 75 lang, 111 breit; Bohrerklap-
pen 124 lang; Körper etwa 630 lang.

& unbekannt.

Phygadeuon oporinus Horstmann & Yu, 1999

Phygadeuon autumnalis Schmiedeknecht, 1905: 673 und 699 (Horstmann 1990: 49 £.) -— praeocc. durch Phygadeuon
autumnalis Provancher, 1882.
Phygadeuon oporinus Horstmann & Yu, 1999: 80 — nom. nov. für Phygadeuon autumnalis Schmiedeknecht, 1905.

Beschreibung

2: Schläfen wenig breiter als die Augen, direkt hinter den Augen nicht, dann rundlich verengt, fein
und sehr zerstreut punktiert, stellenweise unpunktiert; Scheitel nach dorsal nicht über das Niveau der
Ocellen vorragend; Clypeus basal deutlich zerstreut punktiert, apical mit zwei feinen, deutlich ge-
trennten Zähnen (entgegen der Angabe von Schmiedeknecht, 1. c.); Wangenraum 0,8 mal so breit wie
die Mandibelbasis; Fühler 20-21-gliedrig, das vierte Glied 2,4-2,7 mal so lang wie breit, Distalhälfte der
Geißel nicht keulenförmig erweitert, Glieder dort so lang wie breit oder wenig breiter als lang;
Mesopleuren im Zentrum glatt, an den Rändern fein zerstreut punktiert; Hinterfemora 3,7-3,8 mal so
lang wie hoch; Area superomedia 1,0-1,2 mal so lang wie breit; zweites Gastertergit glatt, 0,7 mal so
lang wie breit; Bohrer wie bei P. atricolor, Bohrerklappen so lang wie das erste Gastertergit; Körperlänge
etwa 5 mm; Geißelbasis mehr oder weniger stark gelbbraun überlaufen; Coxen rot, Hintercoxen basal
verdunkelt; Hinterfemora rotbraun; zweites und drittes Gastertergit gelbbraun, die folgenden bräun-
lich (variabel).

Material: 322, Thüringen/D (NMW, SMNS).

[58]
158)
N

Phygadeuon ponojensis (Hellen, 1967)
Phyzelus ponojensis Hellen, 1967: 94 (Horstmann 1975: 104).

Beschreibung

?: Schläfen 0,9 mal so breit wie die Augen, direkt hinter den Augen deutlich verengt, fein und
zerstreut punktiert; Scheitel nach dorsal nicht über das Niveau der Ocellen vorragend; Clypeus basal
kräftig zerstreut punktiert, apical mit zwei kleinen Zähnen; Wangenraum 0,9 mal so breit wie die
Mandibelbasis; Fühler 20-gliedrig, das vierte Glied 2,4 mal so lang wie breit, Distalhälfte der Geißel
kaum keulenförmig erweitert, Glieder dort 1,1 mal so lang wie breit; Mesopleuren außerhalb des
Speculums fein und sehr zerstreut punktiert; Hinterfemora 3,4 mal so lang wie breit; Area superomedia
0,8 mal so lang wie breit; zweites Gastertergit glatt, 0,7 mal so lang wie breit; Bohrer wie bei P. atricolor,

Bohrerklappen 1,5 mal so lang wie das erste Gastertergit; Körperlänge etwa 4 mm; Geißelbasis rot-

braun; Coxen und Hinterfemora schwarz; zweites bis viertes Gastertergit rotbraun, die folgenden
dunkelbraun, caudal gelb gerandet.

Material: 1?, Ponoj (= Ponoi/Kola/Russia) (ZMH).

Danksagung

Für ihre Hilfe beim Entleihen von Typen und anderem Sammlungsmaterial danke ich: A. Albrecht (Zoological
Museum, Helsinki), J.-F. Aubert und M. Sartori (Musee Zoologique, Lausanne), J. Casewitz Weulersse und
C. Villemant (Museum National d’Histoire Naturelle, Paris), R. Danielsson (Zoologiska Institutionen, Lund),
E. Diller (Zoologische Staatssammlung, München), L. Ficken und M. G. Fitton (Natural History Museum,
London), M. Fischer und S. Schödl (Naturhistorisches Museum, Wien), M. Kak und M. Wanat (Muzeum
Przyrodnicze, Wroclaw), J.-P. Kopelke (Naturmuseum Senckenberg, Frankfurt), T. Kronestedt (Naturhistoriska
Riksmuseet, Stockholm), J. Oehlke (Deutsches Entomologisches Institut, Eberswalde), T. Osten (Staatliches
Museum für Naturkunde, Stuttgart), G. E. Rotheray und M. R. Shaw (National Museums of Scotland, Edin-

burgh), J.-H. Stuke (AG Evolutionsbiologie der Universität, Bremen) und K. W. R. Zwart (Laboratorium voor

Entomologie, Wageningen).

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229

Buchbesprechungen

29. Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol.
8: Mollusca, Part 1 (1998/2000), Vol. 9: Mollusca, Part 2 (1996). - Santa Barbara Museum of Natural History,
Santa Barbara, CA. ISBN 0-936494-21-2 (14 Volume Set). vii + 250 pp. and vii + 228 pp.

The Santa Maria Basin and Western Santa Barbara Channel is a remarkable biogeographic region of the Eastern
Pacific with a long tradition in faunistics particularly concerning malacology. The two volumes provide a very
valuable compilation of the molluscan fauna of this important area elaborated by specialists of the respective
taxa. The chapter on the two aplacophoran classes also includes the formal description of no less than five (six)
species new to science; unfortunately these are solely based on hard parts and external morphology, thus the
systematic position of the two solenogastres species remains somewhat tentative. Also two new bivalve and
seven new gastropod species are formally described. Clear identification keys are provided and accurate

descriptions and (mostly black & white) figures of nearly all species included are presented. Several “Genus sp.”
in all molluscan classes demonstrate that more work is necessary for a complete inventory of the Santa Barbara |

species, a hopefully stimulating tool for professional scientists or skilled amateurs.

Strength of the two volumes lies in the detailed data on synonyms, the material examined by the authors,

and the location of types of the respective species, reflecting the immense value of accurate museum collections.
I found the extensive reference list of each chapter very complete and accurate as is the index of each part. The
extensive glossary will certainly help beginners and students. Figures and printing are of medium quality, but
this is acceptable, since it is reflected by a low price, which makes the two volumes available for a broad

readership. No doubt that the “Atlas” will prove its value in the future of biodiversity research in the Santa Maria
Basin and Western Santa Barbara Channel and may serve as a core of further Eastern Pacific taxonomic

initiatives. G. Haszprunar

30. Harper, E. M.,J. D. Taylor & J. A. Crame (Eds.): Evolutionary Biology of the Bivalvia. - Geological Society
Special Publication 177, 2000. - ISBN 1-86239-076-2 (hbk), ISSN 0305-8719. Alden Press, Oxford, vii + 494 PP-

This volume includes the proceedings of The Biology and Evolution of the Bivalvia meeting held in Cambridge in

September 1999. Accordingly, my first congratulation to the editors and authors because they have succeeded
to produce this multiauthor book within a year. Three main topics are concerned: (1) Molecular phylogeny of
the Bivalvia and its subgroups. Two teams once more demonstrate that available sequence data lead to
reasonable results within the major taxa but largely fail to clear up the interrelationships of the major groups or
monophyly or the sister-group of the Bivalvia as a whole. (2) Several articles concern various aspects of the

extensive bivalve fossil record, and phylogenetic as well as functional considerations are presented. (3) Also
morphological data based on electron microscopy or functional anatomy still contribute significantly to our

understanding of bivalve phylogeny and evolution aside from eco-functional aspects - if well elaborated and
analysed. I particularly appreciate the obvious interaction of all three fields mentioned in the various discussion
parts; this makes the present volume a very well-done and important source of information on the current state
of the art of phylogeny and evolutionary history of the Bivalvia. Also price and content have a reasonable
relation; thus, if you are interested in bivalves — go and buy this book. G. Haszprunar

31. Magerl, C. & D. Rabe (Hrsg.): Die Isar. Wildfluss in der Kulturlandschaft. - Verlag Kiebitz Buch, Vilsbiburg,
1999, Großformat, geb., 192 S., viele Farb- und sw. Fotos. ISBN 3-9804048-5-4.

Der Fluss vor unserer Haustür: gezähmt oder wild, Erholungslandschaft oder Kloake? Eine Vielzahl von
Autoren beleuchtet die Isar von allen Seiten: Flößer, Energiewirtschaft, Landschaftsplanung und Renaturierung,
Fische, Botanik, Auwälder, die geschützten Abschnitte, die wichtigsten Städte, das Erholungspotential und nicht
zuletzt die Vogelwelt, alles in 43 Kleinkapiteln dargestellt und von unserem AIB-Redakteur mit herausgebracht.

Das Buch besticht weniger durch den Text, der aber genügend und korrekte Sachinformation liefert, als durch

die schönen Farbfotos, die Heimatliebe erwecken, wobei oft historische Vergleiche, garniert mit seltenen
Schwarzweiß-Dokumenten, herbeigezogen werden.

Über die Vögel der Isarstauseen lassen sich M. Schötz und für Ismaning U. & P. Köhler aus, die auch die
Problematik des zu sehr gereinigten Isarwassers für das Fortbestehen des Speichersees als Ramsargebiet |

erläutern.

Der Band wurde nach dem Pfingsthochwasser 1999 geschrieben, und es scheint, als hätten die Behörden
dazugelernt, denn Baumaßnahmen zur Rück(?)führung in einen naturnahen Zustand sind allenthalben unter-
wegs. Bleibt zu hoffen, dass davon unsere Raritäten wie auch die Artenvielfalt profitieren können. Als Ge-

schenkband und Diskussionsgrundlage gut geeignet. T. Mischler

230

SPIXIANA 231-233 NE TEN er emner2001 ISSN 0341-8391

Araucoscia Verhoeff, 1939 is a juniour synonym
of Pseudophiloscia Budde-Lund, 1904

(Crustacea, Isopoda, Oniscidea)
Andreas Leistikow

Leistikow, A. (2001): Araucoscia Verhoeff, 1939 is a juniour synonym of Pseudo-
philoscia Budde-Lund, 1904 (Crustacea, Isopoda, Oniscidea). - Spxiana 24/3: 231-233

The two philosciid genera Araucoscia Verhoeff, 1939 and Pseudophiloscia Budde-
Lund, 1904, both recently reexamined by their type material have to be considered
synonyms. New material from Chile, where both genera were reported from, is
attributed to Pseudophiloscia inflexa Budde-Lund, 1904. The characters found in this
species match the characters found in Araucoscia. Thus, Araucoscia is considered a
juniour synonym of Pseudophiloscia.

Andreas Leistikow,Universität Bielefeld, Abteilung Zoomorphologie und Syste-
matik, Morgenbreede 45, D-33615 Bielefeld, Germany
E-mail: leiste@biologie.uni-bielefeld.de

Introduction

There are only few species of terrestrial isopods reported from Chile, among them some genera and
species representing the evolutionary level of the philosciid facies. These genera are Araucoscia Verho-
eff, 1939 and Pseudophiloscia Budde-Lund, 1904, both were subject to a reexamination of the type
material (Leistikow 1998 a, b). For both taxa, autapomorphies were found but it appeared that the
boundaries of Araucoscia were more restrictive than those of Pseudophiloscia and the suspicion arose that
Arausoscia might be included in Pseudophiloscia (Leistikow 1998b). New material from Isola Wellington
off the coast of southern Chile revealed the fact that the slight differences are even more inconspicuous
to consider both species members of different genera. Moreover, some amendments to the diagnosis
of Pseudophiloscia Budde-Lund, 1904 shall be given, particularly with respect to the mouth parts and
noduli laterales, the diagnosis has to be expanded.
For the Museo Zoologico della Universitä Firenze, the acrconym MZUF is used.

Systematic section
Pseudophiloscia Budde-Lund, 1904

Paraphiloscia Stebbing, 1900 in part
Araucoscia Verhoeff, 1939

Diagnosis. Cephalothorax with linea supra-antennalis, no linea frontalis and only slight lateral lobes,
compound eye with 15 ommatidia, pleon retracted from pereon. Antennula three-articulate, antennal
flagellum three-articulate, apical bristle shorter than distal article, with moderately long free sensilla
inserting basally. Mandible bearing molar penicil consisting of 10 branches, each arising separately,
lacinia mobilis of left side bulky, lateral endite of maxillula with up to 10 simple teeth, medial endite

231

without apical point, maxilla with lateral lobe three times broader than medial one, maxilliped with
endite lacking knob-like penicil, basipodite with sulcus lateralis.

Pereopods slender, inner claw of dactylus short, dactylar seta simple, carpal antenna-grooming
brush distinct, ornamental sensory spine with handlike apex, coxal plates narrow, lacking sulcus

marginalis and gland pores, three rows of noduli laterales, on coxal plate VII, five noduli laterales.
Pleopods without respiratory areas, lateral margin bearing sensory spines, male endopodite of pleopod |

1 acute. Uropod with lateral groove, endopodite inserting proximally of exopodite.

Species included: Pseudophiloscia inflexa Budde-Lund, 1904, P. angusta (Dana, 1852), P. chilenica |

(Verhoeff, 1939) comb. nov.

Pseudophiloscia inflexa Budde-Lund, 1904
Figs 1-5

Material examined. 19, 12, Chile, Isola Wellington, Puerto Eden, leg. Daccordi, 12.2.1988, MZUF no. 3183; |

Microscopic slides of 15 same data as above.

Additions to description

The species was described by Budde-Lund (1904), figuring some of the mouth parts and the |
habitus, a re-examination of the syntypes was performed by Leistikow (1998b). Since the material was

incomplete to some extent, information on the characters not observed will be given:
Colour. Dorsally chestnut with light spots, on coxal plates two light areas separated by darker

chestnut, paramedian line darker than remaining tegument, cephalothorax chestnut with small light

spots, pereopods chestnut, pleopods light yellowish brown.

Pereon. Tegument shiny, covered with slender tricorn-like setae, three noduli laterales on coxal
plates I-VI, coxal plate VII bearing five noduli laterales, four along the distal margin, the most medial

one in paramedian position on pereonite (Figs 1, 4)
Antenna. Apical bristle as long as distal flagellar article, free sensilla near basis almost as long as
apical bristle (Fig. 3)

Maxillula. Medial endite rather bulbous, subapical tip very inconspicuous, penicils stout, lateral |

endite with a small tooth paralaterally on rostral surface (Fig. 5).
Pereopods. Male pereopod 7 with small setose area on ischium (Fig. 2).

Discussion. As stated previously (Leistikow 1998a,b), the three species Araucoscia chilenica Verhoeff,
1939, Pseudophiloscia angusta (Dana, 1852) and P. inflexa Budde-Lund, 1904 form a monophyletic group
characterized by the following autapomorphies:

° Two rows of noduli laterales per side [only one row of noduli laterales per side]
° Coxal plates very narrow [coxal plates of normal breadth]
° Coxal plates without sulcus marginalis [coxal plates with sulcus marginalis]

° Lateral endite of maxillula with about ten simple teeth not fitting the 4+6-pattern [maxillula with |

4+6 teeth, five of inner set cleft]

The inclusion of the species chilenica into the genus Pseudophiloscia Budde-Lund, 1904 aims at the
establishment of amonophylum separated from other so-called genera by a gap in character states, i.e.,
the traditionally accepted characters for the characterisation of genera (Taiti & Ferrara 1980) should
unite species in monophyletic taxa, called genera, which are separated from other genera by distinct
character states not allowing a continuum from one character state to another. Consequently, all the
species of “philosciid” Oniscoidea bearing the above mentioned characters are placed in Pseudophiloscia.
The slight differences in the morphology of the maxillula are only of interest on the species level.

Acknowledgements

The author is indebted to Prof. Dr. T. Bartolomaeus for the possibility to use the facilites of the department of
zoomorphology. Thanks are to Dr. S. Taiti and Dr. F. Ferrara for the providance with the new material of
Pseudophilsocia inflexa.

232

Figs 1-5. Pseudophiloscia inflexa Budde-Lund, 1904, male mounted on microsopic slides. 1. Coxal plate VII, scale
bar=400 um. 2. Male pereopod 7, rostral view, scale bar =400 um. 3. Articles 2 and 3 of antennal flagellum with
detail of apical organ, scale bar =200 um and 100 um, respectively. 4. Scheme of position of noduli laterales.

ı 5. Maxillula in rostral view, scale bar = 100 um.

References

Budde-Lund, G. 1904. A revision of “Crustacea Isopoda terrestria”, with additions and illustrations. 2. Sperillon-
inae. 3. Armadillo. - Copenhagen: 33-144

Leistikow, A. 1998a. Redecriptions of terrestrial Isopoda (Crustacea: Oniscidea) from Chile and Peru. — Spixiana
21(3): 215-225

-- 1998b. The genus Pseudophiloscia Budde-Lund, 1912 in South America. — Mitt. Mus. Naturkde. Berlin, zool.
Reihe 74(2): 233-241

Taiti, S. & F. Ferrara 1980. The family Philosciidae in Africa, south of the Sahara. - Mon. zool. ital. (n. s.) 13: 53-98

Verhoeff, K. W. 1939. Von Dr. G.H. Schwabe in Chile gesammelte Isopoda terrestria, Diplopoda und Chilopoda.
— Zeitschr. f. wiss. Zool. 8: 301-324

233

Buchbesprechungen

32. Bibikow, D. I.: Die Murmeltiere der Welt. - Westarp-Wiss., Magdeburg; Spektrum Akad. Verlag, Heidel-
berg. 2., völlig neu bearb. und erw. Aufl., 1996. (Die Neue Brehm-Bücherei; Bd. 388). 228 S., 64 Abb., 20 Tab,
1 Farbtaf. ISBN 3-89432-426-0.

Schulze, G.: Die Schweinswale. Familie Phocoenidae. — Westarp-Wiss., Magdeburg. 2. überarb. Aufl., 1996.
(Die Neue Brehm-Bücherei; Bd. 583). 191 S., 114 Abb., 4 Tab. ISBN 3-89432-379-5.

Wagenknecht, E.: Der Rothirsch. Cervus elaphus. - Westarp-Wiss., Magdeburg; Spekrum Akad. Verl,
Heidelberg. 3. überarb. Aufl., 1996. (Die Neue Brehm-Bücherei; Bd. 129). 156 S., 50 Abb., 6 Tab., 1 Farbtaf.

ISBN 3-89432-500-3.

Die “Neue-Brehm-Bücherei”, ehemals vom Ziemsen-Verlag in Wittenberg-Lutherstadt herausgegeben, schien
nach Wende und Wiedervereinigung zum Untergang verurteilt. Dem Verlag Westarp Wissenschaften ist es zu

verdanken, dass die bewährte Reihe nun doch bestehen bleibt. Einige ältere Bände wurden seither neu aufgelegt,

teilweise in neuer Bearbeitung, in jedem Fall aber in etwas größerem Format und in besserer Druckqualität.

Der Band über die Schweinswale entspricht inhaltlich bis auf geringe Ergänzungen der 1. Auflage aus dem

Jahr 1986, doch wurden die Zahl der Abbildungen vermehrt. Das Buch über die Murmeltiere wurde dagegen
völlig neu konzipiert und wesentlich erweitert. Dabei wurden insbesondere neuere Originalarbeiten des Autors
über die Soziobiologie und Ethökologie zentralasiatischer Murmeltierarten berücksichtigt. Die Abbildungen der
ersten Auflage, die von relativ schlechter Qulaität waren, wurden durch wesentlich bessere Aufnahmen ersetzt.

Inwieweit sich der Band über den Rothirsch von der verhergehenden Auflage unterscheidet, kann der
Rezensent nicht beurteilen, da ihm diese nicht zur Verfügung stand. Doch scheint zumindest das Literaturver- '

zeichnis nicht wesentlich aktualisiert worden zu sein und enthält überwiegend ältere Arbeiten.

Die Neuauflagen folgen dem bewährten Konzept der Reihe, dem Leser eine kompakten, aber wissenschaft- '

lich seriösen Überblick über eine Tierart oder -gruppe zu geben, wobei die Ergebnisse der verschiedenene
biologischen Teildisziplinen - Morphologie, Anatomie, Physiologie, Ethologie und Tiergeographie - berücksich-
tigt werden.

Die wesentliche Verbesserung von Druckqualität, Format und Ausstattung ist in jedem Fall zu begrüßen,
zu kritisieren ist allerdings der relativ hohe Preis. R. Kraft

33. Eisenberg, J. F.& K. H. Redford: Mammals of the Neotropics. Vol. 3: The Central Neotropics: Ecuador, Peru,
Bolivia, Brazil. - The University of Chicago Press Chicago and London 1999. 609 S., zahlr. Abb. u. Tab. ISBN
0-226-19541-4.

Mit dem vorliegenden dritten Band wird eine vielbeachtete Handbuchreihe über die Säugetiere der Neotropis
abgeschlossen. Nachdem in den beiden vorangegangenen Bänden (erschienen 1989 bzw. 1992) die nördlichen
bzw. südlichen Staaten dieser Faunenregion behandelt wurden, umfaßt der Geltungsbereich diesmal die in der
Mitte des südamerikanischen Kontinents gelegenen Staaten Ecuador, Peru, Bolivien und Brasilien. Auch die
Säugetiere der Galapagos-, Falkland- und anderer Inseln sind eingeschlossen. Der Aufbau entspricht weitge-
hend dem der ersten beiden Bände: Im Hauptteil werden Merkmale, Verbreitung, Lebensraum und Lebenswei- |
se der einzelnen Arten beschrieben. Verbreitungskarten, Skizzen mit Schädel- und Zahnmerkmalen sowie
Farbtafeln mit Habituszeichnungen ergänzen den Text. Zu jeder Ordnung gibt es ein umfangreiches Literatur-
verzeichnis. Besondere Erwähnung verdienen die einleitenden Kapitel über fossile Säugerfaunen Brasiliens und
Perus. Sie zeigen sehr anschaulich, wie sich die neotropische Säugetierfauna im Verlauf von Tertiär und Quartär |
aus autochthonen und allochthonen Elementen zusammengesetzt hat und tragen dadurch wesentlich zum |
Verständnis ihrer Komplexität und ihres Artenreichtums bei. Ein abschließender Teil behandelt aktuelle ökolo-
gische und populationsbiologische Themen. Insbesondere Biodiversität und Gefährdungsgrad neotropischer |
Lebensräume werden hier angesprochen.

Die dreibändige Reihe ist das Ergebnis jahrzehntelanger Literatur- und Feldstudien der beiden Autoren. |
Entsprechend informativ und authentisch sind die Angaben. Es ist überaus zu begrüßen, daß nun - erstmals seit
dem nicht mehr ganz aktuellem Standardwerk von CABRERA & YEPES aus dem Jahr 1960 - ein vollständiges,
übersichtlich gegliedertes und zudem sehr ansprechend gestaltetes Kompendium der süd- und mittelamerika- |
nischen Säugetierfauna vorliegt - und dies zu einem erstaunlich günstigen Preis. R. Kraft |

234

SPIXIANA 3 235-240 Venen oO NovemBer2001 ISSN 0341-8391

Contribution to the taxonomy of European Poronota 1.
Oribatella and Anachipteria

(Acari, Oribatida)
Gerd Weigmann

Weigmann, G. (2001): Contribution to the taxonomy of European Poronota
(Acari, Oribatida) I. Oribatella and Anachipteria. — Spixiana 24/3: 235-240

In the slide collection of Oribatida from Carl Willmann and from L. Kneissl,
stored in the Zoologische Staatssammlung in Munich, some species have been
found which need to be revised. One of this slides refer to “Oribatella meridionalis”
(det. Willmann); the revision of it resulted in synonymization with Oribatella super-
bula (Berlese, 1904). A related new species from Germany, Oribatella similesuperbula,
spec. nov. is described. Further on a slide with the lable “Tectoribates latitectus” (det.
Willmann) turned out to be a new species, described as Anachipteria dubia, spec.
nov., which is compared with A. howardi Berlese, 1908, the senior synonym of
A. latitectus Berlese, 1908.

Prof. Dr. Gerd Weigmann, Institute of Biology, Lab. Soil Zoology and Ecology,
Free University Berlin, Grunewaldstr. 34, D-12165 Berlin

Introduction

Within revisional studies on central european Oribatida some new species have been found; in some
further species the older descriptions need some additions to be comparable with modern ones. This
contribution deals mainly with material from the Willmann collection in the Zoologische Staatssa-
mmlung in Munich. A slide with “Oribatella meridionalis Berlese, 1908” has been reexaminated in the
light of a revisional publication of Berlese’s Oribatella species (Bernini 1975), the nomen novum
O. willmanni Subias & Gil-Martin, 1995, for this slide material, and a closely related german species with
similar characters, Oribatella similesuperbula, spec. nov.

A second part of this contribution deals with “Tectoribates latitectus (Berlese, 1908)” in the sense of
Willmann (1931), belonging to Anachipteria in modern taxonomy. All european collections of this
american species and the related A. howardi need to be reexaminated. Willmann’s material does not
belong to Berlese’s species and is described as Anachipteria dubia, spec. nov., below.

Oribatella superbula (Berlese, 1904)

According to the revision by Bernini (1975) the italian species Oribatella meridionalis Berlese, 1908, is a
junior synonym of O. superbula (Berlese, 1904). Willmann (1931) illustrated an “O. meridionalis” in his
keys in the group of species with 3 claws. But the reexamination of his slide brougth the result that the
two mounted specimens from italian origin have 2 claws, indeed, as described for superbula. In fig. la
the dorsal aspect of one of the mounted specimens is presented, the claws of legs are sketched, only.
As far as visible the ventral characters agree with O. superbula after Bernini (1975). It seems that
Willmann did not collect “O. meridionalis” in Germany, himself, but he gave the diagnosis after Sellnick

235

EN a

} FE EN A eh, We

Fig. 1a. Oribatella superbula (Berlese), dorsal aspect with indication of the claws (slide of Willmann collection, |
Munich). b. Oribatella similesuperbula, spec. nov., dorsal aspect.

(1928), who notes no origin of his material. Cited by Bernini (1975), in Southern France a 2-clawed |
“O. meridionalis” has been found by Lions (1972). All these specimens refer to O. superbula.
In Spain one specimen of Oribatella has been found and cited by Subias & Gil-Martin (1995) that |
resembles O. superbula in the sense of Bernini (1975), but it differs by legs with 3 claws each. Subias & '
Gil-Martin assumed that the specimen must be identical with Willmann’s “3-clawed O. meridionalis”,
and therefore it was renamed as “O. willmanni Subias & Gil-Martin, 1995” whithout further details. But, '
as reported above, Willmann’s specimens of “O. meridionalis” have 2 claws and origin from Italy, and |
they belong to O. superbula. The spanish Oribatella is a species inquirenda; the nomen novum for |
Willmann’s O. meridionalis is based on the erroneous assumption that Willmann’s specimens would
have 3 claws at the legs, also. In the following, the description of a new species from Germany with
3 claws at the legs is presented which is related to O. superbula. |

Oribatella similesuperbula, spec. nov.
Figs 1b, 2a-c

of lamella with long internal and external spicular teeth. Sensillus long and only scarcely fusiform |
thickened. Rostral border with 2 small lateral teeth, in the middle undulating. Epimeral setation "
formula normal (3-1-3-3); 4c very strong and moderately prolonged, 3c moderately strong and pro- |
longed; other setae thin. Legs with 3 claws. |

|
|
Diagnosis. Small body size, length about 320-380 um. Prodorsum with interlamellar tubercle; cuspis
|

Description |
General characters. Total body length (without lamellae and tutoria) of females 340-380 um, males '
320-340 um. Colour reddish-brown, cuticle smooth.
Prodorsum. Frontal border of rostrum with no real incision, between small lateral teeth the border |

236

Fig. 2a. Oribatella similesuperbula, spec. nov., epimeral region. b. rostrum with genal teeth and tutorial tips in
dorso-frontal aspect. c. rostrum of an other specimen. d. Oribatella superbula (Berlese), rostrum with genal teeth
and tutorial tips in dorso-frontal aspect (after Bernini 1975).

has two or three incurvations (Figs 2b, c); dorsal part of rostrum with carina. Normal shape of lamellar
cuspis: inner cuspis teeth mostly shorter than outer teeth, the latter at the lateral side with 0-3 little
teeth; with small interlamellar (“translamellar”) tubercle between the cuspides; lamellar and interla-
mellar setae long and strong. Sensillus thick setiform (Fig. 1b). Tutorium long and broad, with distal
teeth, as usually (Fig. 2b). Genal teeth not very broad (smaller than in O. superbula). Rostral setae long,
with setulae. Pedotectum I very large.

Notogaster. Cuticle smooth with fine puncture, with striation at the anterior part of the ptero-

_ morphs. 10 pairs of granulated notogastral setae of moderate length, anterior setae up to 35 um, 2 pairs

at the posterior border very small; 4 pairs of small areae porosae; pteromorphs of normal form, as

_ typical in the genus (Fig. 1b).

Ventral region. Epimeral region (Fig. 2a) with the usual setation 3-1-3-3. Some setae short and thin
(la, 1b, 2a, 3a); others longer and thin (1c, 3b, 4a, 4b); 3c longer and moderately thick, 4c thick and
prolonged, about 40 um, reaching to the apodeme between epimers II and III. Aggenital setae of normal
small size, as 6 pairs of genital setae, 3 pairs of adanal setae and 2 pairs of anal setae.

Legs. All legs with 3 claws (heterotridactyl). Genu I with ventral tooth, genu IV without tooth.

Origin of the specimens. (1) Some specimens have been found by the author in moss and decaying
wood of a trunk. Forest near Milseburg, Rhön Mountains, Germany; sample 813, date 4.6.1994. The
type and a syntype is deposited in Zoologische Staatssammlung, Munich. Further specimens are in the
collection of the author. (2) Three slides in the Kneissl collection, stored in Zoologische Staatssammlung
in Munich, belong to this species, most probably. The specimens in the slides “Oribatella berlesei” K1194,

K1195 and K1196 are damaged, but the visible details refer to the new species. Origin is Oberalting,

Bavaria, leg. Kneissl 7.8.1910 from decaying wood (“Mulm”).

Discussion: O. similesuperbula, spec. nov. is closest related to O. superbula. Common characters are:
anterior border of rostrum small, without pad; strong epimeral setae 3c and 4c; middle body size. Main
differences are: legs with three claws (superbula with 2 claws); seta 4c of epimers (Fig. 2a) less prolonged
than in superbula, in which the seta 4c reaches the apodeme between epimers I and II; genal teeth
smaller; frontal border of rostrum undulated (Figs 2b, c; with incision in superbula: fig. 2d); body length

, somewhat larger with 320-380 um (in superbula 290-350 um).

At today’s knowledge the 3-clawed spanish species Oribatella willmanni Subias & Gil-Martin, 1995
must not be identical with O. similesuperbula, spec. nov. and it requires careful reexamination (cf.
discussion in the section on O. superbula). But, in the case of specific identity the name O. willmanni
would be senior synonym of O. similesuperbula, spec. nov.

237

Fig. 3a. Anachipteria howardi (Berlese), dorsal aspect of a syntype, Berlese collection (after Norton & Kethley
1989). b. Anachipteria, dubia spec. nov., dorsal aspect. c. Sacculus Sa with notogastral seta.

Anachipteria howardi (Berlese, 1908) and A. latitecta (Berlese, 1908)

The taxonomical literature on Anachipteria presents a confusing picture. There are different opinions on

the definition of the families Achipteriidae and Oribatellidae. The Anachipteria species have been put

into the family Oribatellidae by some authors (cf. Perez-Inigo 1993; see discussion in Bernini 1973),
partly as Tectoribates species, because Anachipteria, in contrast to other achipteriid genera, lacks the
knifelike anterior projection at the pteromorph. On the other hand, the morphology of the lamellar
complex and the tutoria are more or less the same than those in typical achipteriid genera, as Achipteria,

Parachipteria and Pseudachipteria, which have the typical knifelike anterior projection at the pteromorph. |
The systematic position seems to be solved definitely by Seniczak (1977): the larvae and nymphs of |
Anachipteria show the typically folded notogaster and other characters of Achipteria juveniles, contrast- |

ing the juveniles of Oribatella. Anachipteria is a member of Achipteriidae.

In two very short descriptions, without figures, in the same paper Berlese (1908) established
Sphaerozetes howardi and Sphaerozetes latitectus, both originating from Columbia, North America. The
differences should be in the sensillus shapes. Later on Berlese changed the lable of a “latitectus” slide
by adding “howardi” with pencil. It seems that Berlese regarded both species as synonymous, as

indicated in his unpublished catalogue (see Norton & Kethley 1989). The reexamination of Berlese’s "

slides led Norton & Kethley (1989) to the same assumption of synonymy of both species with the

priority of howardi, described firstly (cf. Marshall et al. 1987; cf. Mahunka & Mahunka-Papp 1995). It |

belongs to Anachipteria in modern literature.
European records of both species, A. howardi and “A. latitecta”, need to be reexamined. Grandjean

(1932) discusses the species in connection with his description of Anachipteria deficiens Grandjean, 1932. |
Willmann (1931) describes and figures “Tectoribates latitectus” very cursorily, a specimen collected in !
north-western Germany, near Friesoythe. Seniczak (1977) figures the adult of “A. latitecta” together !
with juveniles, but it seems to be A. deficiens, more probably. The redescription of “A. howardi” from

Hungaria by Mahunka (1996) is comparably good standard, but the sensillus shape is somewhat

238

smaller; it indicates the possibility that the species occurs in North America and in Europe, also. A
reexamination of the Berlese slides by Norton & Kethley (1989; as by the author) resulted in the
drawing fig. 3a of “Sphaerozetes latitectus” (Berlese’s slide 73/16; conspecific with syntype slide 73/15
with the pencil mark “tipico = Howardi”). An originally designated type specimen of S. howardi has not
been found. Supposed, that the sensillus shape has some variability, the redescriptions of Norton &
Kethley (1989), fig. 3a, and Mahunka (1996) refer to the same species, A. howardi. A. latitectus (Berlese,
1908) is regarded as junior synonym.

Diagnosis of Anachipteria howardi (Berlese, 1908): Pteromorphs dorsally with a moderate sinus at
the bothridia; Areae porosae of notogaster large, the largest is Aa; anterior notogastal setae c, moder-
ately large (about 30 um), posterior setae small; cuspis of lamellae somewhat truncated with a nearly
transverse frontal border, at the outer side of the cuspis a small but distinct tooth; sensillus fusiform,
_ mostly asymmetrical, with distinct tip, granulated; body length about 385-440 um.

Anachipteria dubia, spec. nov.
Fig. 3b,c

. This species exists up to now as two specimens, only. The type specimen is mounted in a microscopical
' slide in the Willmann collection in the Zoologische Staatssammlung in Munich and is labled as
“Tectoribates latitectus”. It is the specimen which has been illustrated by Willmann (1931: p. 181).
Willmann indicates it as the single german species of the genus Tectoribates; he refers that the lamellae
look like that of “Notaspis” (now Achipteria and related). Because of sacculi on the notogaster the species
is different from Anachipteria howardi, which is the senior synonym of Sphaerozetes latitectus Berlese,
1908, as discussed in the last section.

The type locality after Willmann (1931) and the slide lable is: Moss of a bog; Rolfsort, Wolfstange
bei Friesoythe (Oldenburg); type specimen in M134 - 13.8.27; the locality is in north-western Lower
' Saxonia, Germany (a second slide is M135). The slides are deposited in Zoologische Staatssammlung
in Munich.

The mounted status of the species does not allow differentiated description, especially of the
_ ventral morphology and the lateral aspect.

Diagnosis: Pteromorphs dorsally with a moderate sinus at the bothridia, without a knifelike anterior
projection at the pteromorph; notogaster with 4 pairs of sacculi instead of areae porosae; anterior
notogastal setae c, moderately large (about 30 um), posterior setae small; cuspis of lamellae at the
frontal border obliquely cut, with the corner at the outer side of the cuspis, as typical for most species
of the genus; sensillus very small fusiform with distinct tip; body length about 470 um. As far as visible
the ventral morphology shows no specific characters. Legs with 3 claws.

Discussion: A. dubia, spec. nov. is the first species within Anachipteria with sacculi instead of areae
porosae on the notogaster. The special form of the pteromorphs and all other characters, as far as visible
in the slide, fit into the genus. But, the sacculi seem to me an insufficient argument to split off a new
genus. The very similar achipteriid genera Achipteria and Parachipteria differ only by notogastral sacculi
in the first and notogastral areae porosae in the latter genus, a poor argument for separating genera.
For instance, I know a Peloptulus species with sacculi on the notogaster (the common species have areae
porosae). As known by oribatologists, both homologous structures (see Norton et al. 1997) occur in
many poronotic families, from case to case. It has no or minor worth for phylogenetic argumentation,
but it seems to me being a typological character, only. The species name A. dubia spec. nov. refers to
the doubtful systematic position within Anachipteria, regarding the genus definition, up to now.

References

Berlese, A. 1908. Elenco di generi e specie nuove di Acari. — Redia 5: 1-15
Bernini, F. 1973. Notulae Oribatologiae 8: Sur le genre Tectoribates Berlese 1910 (Acarida, Oribatei). - Acarologia
15: 540-554

239

-- 1975. Notulae Oribatologicae XIII. La famiglia Oribatellidae (Acarida, Oribatei) nell’ Arcipelago Toscano.
— Lav. Soc. Ital. Biogeogr. (N. S.) 5: 429-507

Grandjean, F. 1932. Observations sur les Oribates (3e serie). —- Bull. Mus. nat. Hist. natur. (2) 4: 292-306

Lions, J. C. 1972. Ecologie des Oribates (Acariens) de la Sainte Baume (Var). - These, Universite de Provence:
1-549

Mahunka, S. 1996. Oribatids of the Bükk National Park (Acari: Oribatida). - In: The fauna of the Bükk National
Park: 491-532

-- & L. Mahunka-Papp 1995. The oribatid species described by Berlese (Acari). - Hung. Nat. Hist. Mus.,
Budapest: 323 pp.

Marshall, V.G.,R.M. Reeves & R. A. Norton 1987. Catalogue of the Oribatida (Acari) of continental United States

and Canada. -— Mem. Ent. Soc. Canada 139: 1-418

Norton, R. A. & J. B. Kethley 1989. Berlese’s North American Oribatid mites: historical notes, recombinations,
synomymies and type designations. — Redia 67: 421-499

-- ,G. Alberti, G. Weigmann & S. Woas 1997. Porose integumental organs of oribatid mites (Acari, Oribatida).
1. Overview of types and distribution. — Zoologica 48,146: 1-31

Perez-Inigo, C. 1993. Acari: Oribatei, Poronota. — In: Ramos, M. A. (ed.), Fauna Iberica. 3. Mus. Nac. Cien. Natur.,
Madrid: 1-320

Sellnick, M. 1928. Formenkreis: Hornmilben, Oribatei. — In: Brohmer, P., P. Ehrmann & G. Ulmer (eds), Die

Tierwelt Mitteleuropas 3, 4. Lief. (Teil 9). Quelle & Meyer, Leipzig: 1-41

Seniczak, S. 1977. The systematic position of moss mites of the genus Anachipteria Grandjean, 1935 (Acarina, '

Oribatei) in the light of ontogenetic studies. — Acarologia 18: 740-747

Subias, L. S. & J. Gil-Martin 1995. Nuevas citas oribatologicas (Acari, Oribatida) para la fauna espanola. - Boln |

Asoc. esp. Ent. 19: 25-51

Willmann, €. 1931. Moosmilben oder Oribatiden (Cryptostigmata). — In: Dahl, F. (ed.), Die Tierwelt Deutsch-

lands. 22. Fischer, Jena: 79-200

240

| spvaana | 2 | 3 | 241-244 ee OIENovember 2001 ISSN 0341-8391

Oenospila kopperi, spec. nov.,
eine neue grüne Geometride aus Sumatra

(Insecta, Lepidoptera, Geometridae, Geometrinae)

Axel Hausmann & Manfred Sommerer

Hausmann, A. & M. Sommerer (2001): Oenospila kopperi, spec. nov., eine neue
grüne Geometride aus Sumatra (Insecta, Lepidoptera, Geometridae, Geometrinae).
— Spixiana 24/3: 241-244

Diagnostic differences in the male genitalia distinguish the new Sumatran spe-
cies O. kopperi from the externally similar congeners in the Himalayas (O. strix) and
in Sundaland (O. altistrix, O. gemmans, O. microstrix). The new species was found in
montane forest habitats. The larva and the host-plants remain unknown. Holotype
Sin ZSM.

Axel Hausmann, Zoologische Staatssammlung München, Münchhausenstr. 21,
D-81247 München, Germany. E-mail: Axel.Hausmann@zsm.mwn.de

Manfred Sommerer, Volpinistr. 72, D-80638 München, Germany

Material

Oenospila kopperi, spec. nov.
Figs

Typen. Holotypus d, SUMATRA sept., Dairi mts. 30 km E Sidikalang, 1600 m, 20.11.1999, leg. U. Buchsbaum.
— Paratypen: (9) in ZSM: SUMATRA sept., 15, Deli, Dolok Merangir, 150 m, 8.V1.1967, leg. Dr. E. Diehl; 13, id.,
180 m, IX.1970-1.1971; 12, Simalungun, HW 2a, 28 km SW P.Siantar, 98°59'E, 2°46'N, 1050 m, 27.11.1992, leg. Dr.
E. Diehl; 13, 522, Dairi mts. 30 km E Sidikalang, 1600 m, 20.11.1999, leg. U. Buchsbaum; (28) in coll. Sommerer
(alle leg. Dr. E. Diehl soweit keine andere Angabe): SUMATRA sept., 12, Dairi mts. 30 km E Sidikalang 1600 m,
31.11.1984; 12, id., 1800 m, 27.1V.1986; 12, Tele W Tobasee 1600 m, 9.IX.1975; 336, id., 10.11.1984 (leg. Kobes);
13, id., 12.VIL.1992 (Gen.Praep.MS 1996-72); 23, Sitahoan E Prapat 1450 m, 26.V1.1990; “Holzweg 3”, 1150 m;
12, id., 18.1.1982 (Gen.Praep.MS 1999-172); 15, id., 25.1.-6.11.1982; 15, id., 30.X.1982 (Gen.Praep.MS 1996-70); 1%,
id., 14.X1.1982; 18, id., 28.-30.X1.1982 (Gen.Praep.MS 1996-71); 12, id. 12.IV.1983; 253,18, id., 15.-28.V.1983; 1%,
id., 15.VIL.1983; 12, id., 2.-10.X11.1983; 32%, id., 25.-31.1.1984; 18, id., 4.X1.1990 (Gen.Praep.MS 1999-11); 222, id.,
21.11.1993; 1, id., 3.V.1997; 12, “Sipirok III” 16 km NE Sipirok 1300 m, 25.X11.1995; 15, (Tapanuli sel.), “Sipirok
3”, 10 km NE Sipirok 1300 m, 29.1.1995.

Das Typenmaterial wurde dankenswerterweise von Mitgliedern der HETEROCERA SUMATRANA SOCIETY e. V. (Göt-
tingen), die zur ZSM eine enge Kooperation unterhält, für die wissenschaftliche Bearbeitung zur Verfügung
gestellt. Die faunistische Erforschung Sumatras im Kontext der Faunen des Sundalands entspricht den sat-
zungsmäßigen Zielen dieses Vereins.

Beschreibung

Habitus und äußere Morphologie (Abb. 1). Flügelspannweite 26-29 mm. Flügelfärbung hellgrün,
Vorderflügelcosta weiß. Postmedianlinie braun, deutlich, stark gezackt. Mittelpunkt schwärzlich. An
der Vorderflügelcosta auf der Höhe des Mittelpunktes ein weiterer schwärzlicher Punkt. Hinterflügel-

241

Abb. 1. Oenospila kopperi, spec. nov., Paratypus, d, Sumatra sept., “Sipirok 3”, 10km NE Sipirok 1300 m,
29.1.1995 (R. Kühbandner pinxit).

Analrand mit schwarzem Dreiecksfleck in der Mitte. Aderenden an der Fransenbasis durch deutliche
schwarzbraune Punkte markiert. Stirn grün. Palpen & 1,2facher, ? 2,5facher Augendurchmesser,
letztes Palpenglied beim ? glatt beschuppt, länger als der Augendurchmesser. Fühler beim d auf % der
Gesamtlänge doppelt gekämmt, längste Fühlerkammzähne 6-8fache Geißelbreite. d Hintertibia etwas
verbreitert, Terminalsporen fehlend, Proximalsporen von extrem ungleicher Länge. Hinterbein mit
etwas verkürztem Tarsus, Tarsus : Tibia = 2,5 : 3,5 mm.

ö Genitalapparat (Abb. 2). Valvencosta stark sklerotisiert und zentral mit einem spitzen Fortsatz

bewehrt. Sacculus-Anhänge schwach sklerotisiert, distal mit einem gebogenen, fingerförmigen Fort-

satz. Aedoeagus terminolateral stark sklerotisiert und mit winzigen Dornen besetzt.

? Genitalapparat (Abb. 3). Gut mit der generischen Diagnose in Holloway (1996: 249; fig. 266)
übereinstimmend. Im Vergleich mit der Typusart, O. flavifusata (Walker, 1861), mit auffälliger sklero-
tisierter Verbreiterung zwischen Ductus Bursae und Corpus Bursae. Signum-Ring groß.

Diagnose. Die neue Art gehört zu den südostasiatischen Geometrini im Genus Oenospila, das indo-
australisch verbreitet ist. Habituell zeichnet sie sich durch einen dunklen Dreiecksfleck im Hinterflügel
aus, wo die Postmediane auf den Analrand trifft. Da sie dieses Merkmal mit weiteren Arten wie den
gleich großen O. strix (Butler, 1889), O. gemmans Prout, 1935 und O. altistrix Holloway, 1996 sowie der
kleineren O. microstrix Holloway, 1996 teilt, ist eine sichere Unterscheidung nur an Hand der Genital-
morphologie des d möglich: Valvencosta bei O. strix und O. altistrix ohne spitzen Fortsatz bzw. ein
solcher nur als kurzer Zahn entwickelt. Sacculus-Anhänge bei O. strix breit und kurz, bei altistrix

zungenförmig ausgeprägt und stark sklerotisiert, bei O. gemmans zungenförmig, schwach sklerotisiert, '

mehr ventral anliegend und schräg stehend (Holloway 1996: 251).

242

Abb. 3. Oenospila kopperi, spec. nov., Paratypus, ® Genitalapparat (photo R. Trusch).

243

Drei vorliegende Falter (19, 22?) aus West-Malaysia aus der Sammlung Herbulot (ZSM) äußerlich
strukturell und genitaliter der neuen Art nahestehend mit etwas längeren Palpen, jedoch ist weiteres
d-Material nötig, um die taxonomische Stellung dieser Populationen zu klären.

Biologie. Die ersten Stände und die Futterpflanzen der Raupe sind nicht bekannt. Hinweise zur
Biologie von O. flavifusata (Walker, 1861), der Typusart der Gattung Oenospila, die auch in Sumatra
vorkommt, finden sich bei Holloway (1996: 250).

Verbreitung. O. kopperi ist bisher nur von Sumatra bekannt und wurde dort in Urwaldbiotopen
zwischen 1100 m und 1800 m gefunden. Sie könnte eine Schwesterart der aus vergleichbaren Bergwäl-
dern Borneos beschriebenen und bisher nur von dort bekannten O. altistrix darstellen.

Diskussion. Biogeographisch sind die - leider dramatisch schwindenden - Urwaldbiotope der Sun-
dainseln von höchstem Interesse. Sie liegen einerseits im Schnittpunkt großräumiger Entwicklungsli-
nien der biologischen Diversität, die vom australasischen, vom südchinesischen und vom himalaya-
nisch-indischen Raum ausgehen. Andererseits zeigen wesentliche Komponenten unserer paläarkti-
schen Fauna noch Verwandtschaft zu südostasiatischen Elementen, was mit den erdgeschichtlichen
Ereignissen seit dem Tertiär zusammenhängen dürfte. Für das Verständnis der Prozesse der Artbil-
dung und der biologischen Diversität in der nördlichen Hemisphäre spielt daher die Erforschung der
südostasiatischen Faunen eine wichtige Rolle.

Die Gemeinsamkeiten und die Verschiedenheiten in den Faunen der einzelnen großen Sundainseln
und des angrenzenden Festlands ergeben ein komplexes Bild, das - im Wettlauf mit der rasanten
Naturzerstörung — dringend der weiteren Ergänzung in wissenschaftlicher Zusammenarbeit mit den
Institutionen vor Ort (wie dem Museum Zoologicum Bogoriense, Java) bedarf. Die neue Geometriden-
Art Oenospila kopperi mit ihren nächsten “Verwandten” im Himalaya (O. strix), auf Borneo (O. altistrix
und O. microstrix), in Sumatra (O. microstrix ) und in Java und Bali (O. gemmans) ist ein schönes Beispiel
für die subtilen Ausprägungen der faunistischen Diversität in Südostasien. Auf die Anmerkungen von
Holloway (1996) zu den einzelnen Arten wird verwiesen.

Derivatio nominis. Die neue Art wird nach Hilmar Kopper (Vorsitzender des Aufsichtsrats der Deutschen
Bank AG) benannt.

Literatur

Holloway, J. D. 1996. The Moths of Borneo part 9 Geometridae (Oenochrominae, Desmobathrinae, Geometri-
nae). -— Malay. Nat. Journ. 1996: 249-251

244

| | SPIxIaNA 245-247 Mitindhen, II. Ntorramibren Ali ISSN 0341-8391

A new Oocorys from Western Australia

(Mollusca, Gastropoda, Cassidae)
Kurt Kreipl & Axel Alf

Kreipl, K. & A. Alf (2001): A new Oocorys from Western Australia (Mollusca,
Gastropoda, Cassidae). — Spixiana 24/3: 245-247

A new species of the family Cassidae Swainson, 1832 is described from deep
water off Western Australia. The new species belongs to the genus Oocorys Fischer,
1883 in the subfamily Oocorythinae Fischer, 1883. Type species of the genus is
Oocorys sulcata Fischer, 1883. The Oocorythinae live in deep to very deep water
(120 m to more than 4500 m).

The new species is compared with the similar looking Oocorys lussii Bozzetti,
1990 from the Western Indian Ocean now is regarded as a form of Oocorys verrilli
(Dall, 1889) (Alan Beu, pers. comm.).

Kurt Kreipl, Meeresmuseum Öhringen, Höhenweg 6, D-74613 Öhringen, Ger-
many

Prof. Dr. Axel Alf, Dr. Müller Str. 9, D-91746 Weidenbach, Germany

Introduction

In December 2000 the senior author received three specimens of an unidentified Oocorys from the
Australian shell dealer and collector Hugh Morrison. After a close examination of the shells and an
intensive discussion with Dr Alan Beu, Lower Hutt, New Zealand we decided to describe this species
as new to science.

Oocorys morrisoni, spec. nov.
biesal22

Types. Holotype: 160 naut. miles off Broome, Western Australia, on sand and mud in 450-500 m; collected by
scampi trawlers. Size: height: 39.7 mm; width: 28.5 mm; adult, with operculum. Western Australian Museum,
Perth, Western Australia, coll. no. WAM S 1365.

Paratypes: 1, in coll. Hugh Morrison, Kingsley, Western Australia. Size: height: 32.2 mm; width: 22.0 mm;
adult, with operculum. - 2, in coll. Kurt Kreipl, Meeresmuseum Öhringen, Germany coll. no. 8400. Size: height:
32.5 mm; width: 20.9 mm; slightly subadult, with operculum.

Description of holotype

Shell ovate, small for the genus; thin-shelled but solid. Protoconch of 3 whorls, teleoconch of 2%
whorls. Teleoconch whorls sculptured with strong, regularly-spaced spiral ridges (22 on the body
whorl) crossed by numerous fine growth lines forming strong beads at points of intersection. Suture
not incised. Very distinctive sutural ramp on the shoulder area. Outer lip thickened and reflected,
bearing 15 rather low but distinct teeth along its inner edge. Parietal shield very weakly developed with
spiral sculpture showing through. Columellar margin with 4 distinct teeth in the upper part of the
aperture.

245

Fig. 1. Oocorys morrisoni, spec. nov. Holotype. a. Dorsal view. b. Ventral view.

b. Ventral view.

Colour bright cream to soft orange with a moderate gloss. Outer lip and lower part of the columella
white.
Operculum corneous, paucispiral; yellowish-brown.

Discussion: Oocorys morrisoni, spec. nov. at first glance very much resembles Oocorys verrilli form lussii
(Bozzetti, 1990) from the western Indian Ocean (South Africa to Madagascar and Reunion), but differs
in having more spiral cords on the body whorl (22 in morrisoni; 18-20 in verrilli form lussii) and more
teeth on the inner edge of the outer lip (15 in morrisoni; 12 in verrilli form lussii). Oocorys verrilli form

‚ lussii does not have the distinct sutural ramp of morrisoni. The basic colour of verrilli form lussii is pale
, beige whereas morrisoni shows a bright cream to soft orange colour. The very distinct sutural ramp and

particularly the coloration of Oocorys morrisoni, spec. nov. are the most distinguishing features of this
new species as most other Oocorys are plain white, or at most pale cream or pale beige.

Etymology. We name this new species after the Western Australian shell dealer and collector Hugh Morrison
who brought it to our attention.

Acknowledgements
For very useful information we want to thank Dr. Alan Beu, Institute of Geological and Nuclear Sciences, Lower
Hutt, New Zealand. All photographs by Uschi Damaschke, Möckmühl, Germany.

References

Bozzetti, L. 1990. A new Oocorys from South Africa. - La Conchiglia 23(256): 46
Kreipl, K. 1997. Recent Cassidae. — Verlag Christa Hemmen, Wiesbaden: 1-151

Quinn, J. F. jr. 1980. A new genus, species and subspecies of Oocorythidae (Gastropoda: Tonnacea) from the
Western Atlantic. —- Nautilus 94(4): 149-157

247

Buchbesprechungen

34. Reid, F. A.: A field guide to the mammals of Central America and Southeast Mexico. — Oxford University
Press New York, 1997. 334 S., 48 Farbtaf., zahlreiche Verbreitungskarten. ISBN 0-19-506401-1.

Das Buch beschreibt Merkmale, Lebensweise, Verbreitung und Bestandssituation der Säugetiere Mittelameri-

kas. Der Geltungsbereich erstreckt sich vom Isthmus von Tehuantepec durch Mittelamerika bis Panama und

deckt damit ein Gebiet ab, das viele naturkundlich interessierte Touristen anlockt. Durch das Zusammentreffen

nordamerikanischer und neotropischer Faunenelemente findet sich in dieser Region eine besonders arten- und

formenreiche Säugetierfauna. Es ist eine bewundernswerte Leistung der Autorin, alle Arten dieses Gebietes —
F Q ö . . 9 I
immerhin mehrere 100 - in einem Buch zu vereinen, wobei alle Gruppen mit der gleichen Sorgfalt und

wissenschaftlichen Seriosität abgehandelt werden. Sogar Wale und Seekühe sind eingeschlossen. Doch nicht nur

die Zoologin, sondern auch die Illustratorin Fiona A. Reid verdient Anerkennung: Mit ganz wenigen Ausnah-
men werden alle Arten in farbigen Zeichnungen abgebildet. Diese Abbildungen sind nicht nur ansprechend, '
sondern auch sehr informativ und wirklichkeitsnah und zeigen, daß sie nicht nach Museumsbälgen, sondern im
Freiland “nach dem Leben” gezeichnet wurden. Auch die Merkmalsbeschreibungen im Text sind sehr detailliert
und differenziert und zeugen von dem zeichnerisch geschultem Auge der Autorin. Alle taxonomischen und

biologischen Angaben sind gut belegt und entsprechen dem aktuellen Kenntnisstand. Das Buch ist eine bemer-
kenswerte Neuerscheinung und ergänzt bisherige Werke, die Nord- oder Südamerika getrennt behandelt.

R. Kraft

—Z<2ZZALALCZnZnnmnA CI —

35. Sartori, M. & P. Landolt: Fauna Helvetica 3: Atlas de Distribution des Ephemeres de Suisse (Insecta, '

Ephemeroptera). - Schweizerische Entomologische Gesellschaft Neuchatel, 1999, 214 S. ISBN 2-88414-014-X. |

In der bekannten Reihe ist nach dem Bestimmungsbuch der schweizerischen Eintagsfliegen von 1992 nun auch

ein Verbreitungsatlas erschienen, der im allgemeinen Teil zweisprachig (Französisch, Deutsch), und im spezi- |
ellen Teil, der die einzelnen Arten vorstellt, in französischer Sprache ausgeführt ist. Dieser Teil dokumentiert die

Höhenverbreitung, die Flugzeit, die Typologie, d.h. die von den Larven besiedelten Lebensräume, den Lebens-
zyklus, die Ökologie, die Verbreitung und den Status, d.h. die zeitliche und räumliche Entwicklung der

jeweiligen Art in der Schweiz, und enthält eine Verbreitungskarte der 85 nachgewiesenen Arten. Der vorange-

stellte allgemeine Teil beschreibt kurz die Biologie, die Bedeutung dieser Insektengruppe, die Geographie und
Hydrologie der Schweiz mit Karten und Tabellen ebenso wie die die Besiedlung beeinflussenden physischen
Faktoren, die Biogeographie, die nomenklatorischen Änderungen seit 1992, die Anzahl der zugrundeliegenden

Datensätze und die Entwicklung der Artenzahl. Es folgt eine allgemeine Information zur Handhabung der

Artenvorstellung und der verwendeten Kürzel. Den Schluß bildet eine Diskussion, die Angaben zur Faunistik

und Ökologie unter Einbeziehung des Einzugsgebietes, der Höhenverbreitung, der Phänologie, der Entwick- |
lungszyklen, wiederum der Typologie und dem Status der Arten enthält. Daran schließt sich noch ein Kapitel '
über die Besonderheiten der Eintagsfliegenfauna und deren Gefährdung an. Sieht man von einigen Ungenau- |

igkeiten der deutschen Übersetzung ab, so ist dieser Band der Fauna der Schweiz ein wesentliches Werkzeug,
die Verbreitung und Ökologie dieser Insektengruppe zu erfassen. E.-G. Burmeister

36. Jacquemin, G. & Boudbt, J.-P.: Les Libellules (Odonates) du Maroc. — Societe Francaise d’Odonatologie, Bois

D’Arcy, France,1999, 150 S. ISBN 2-9507291-3-4.

Nach den Erfassungen der Libellenfauna Morokkos von Lieftinck (1966) und Dumont (1972) ist eine Reihe neuer
Arten, aber besonders auch Fundlokalitäten hinzugekommen, die in dem vorliegenden broschierten Heft

dokumentiert sind. Dieses enthält neben Kurzfassungen zur Biologie und Ökologie der Arten eine Bestim- |

mungstabelle der Imagines, eine Darstellungsfolge der Flügeladerung und Zeichnung der bisher nachgewiese-
nen 58 Arten sowie eine Liste der Fundorte mit deren graphischer Umsetzung. Die Artenliste ist ausgedehnt auf
die Region des Maghreb (Marokko, Algerien, Tunesien). Die folgende artspezifische Dokumentation enthält
Angaben der Fundlokalitäten, der Zoogeographie im allgemeinen und speziell bezogen auf Marokko, wobei die
populationsspezifischen Charaktere im Vergleich hervorgehoben werden. Dem folgen Angaben zu Flugzeiten
und eine Verbreitungskarte in Marokko. Einer Zuordnung der Arten zu den biogeographischen Regionen folgt
wiederum ein Vergleich mit dem Arteninventar der Maghreb-Staaten sowie eine erste Rote Liste der Libellen
Morokkos mit Erläuterungen. Die umfangreiche Bibliographie erweist sich als unentbehrliches Hilfsmittel, um
Einzelinformationen zu den Funden zu erhalten. Den Abschluß dieser neuen gelungenen Zusammenfassung
zur Libellenfauna dieses nordafrikanischen Landes bilden Fotos von Fundlokalitäten und Freilandaufnahmen
der 58 Libellenarten. E.-G. Burmeister

248

| SPIXIANA 249-256 München, 01. November 2001 ISSN 0341-8391

A new species of Paroedura Günther from northern Madagascar

(Reptilia, Squamata, Gekkonidae)
Frank Glaw, Miguel Vences & Kathrin Schmidt

Glaw, F., M. Vences & K. Schmidt (2001): A new species of Paroedura Günther
from northern Madagascar (Reptilia, Squamata, Gekkonidae). - Spixiana 24/3: 249-
256

Paroedura lohatsara, spec. nov. is described from Montagne des Francais, a decid-
uous dry forest on a karstic underground in the far north of Madagascar. The new
gecko species is relatively large (up to 80.6 mm snout-vent length and 156.1 mm
total length) and has blackish markings on the head which can form a vermiculated
pattern. It is further characterized by having the nostril excluded from contact with
the rostral scale, distinctly enlarged and spinous tubercles on the dorsal surface,
and specific colouration of juveniles and adults. The relationships of the new
species are obscure: Based on the nostril position, P. lohatsara belongs to the phe-
netic Paroedura picta species group which was hitherto considered as largely re-
stricted to southern Madagascar. Other morphological and chromatic characters
indicate a closer relationship of P. lohatsara with the syntopic P. stumpffi which is a
member of the P. sanctijohannis species group. Two further species of Paroedura,
P. stumpffi and P. karstophila, are recorded from Montagne des Frangais for the first
time.

Frank Glaw, Zoologische Staatssammlung, Münchhausenstr. 21, D-81247 Mün-
chen, Germany. E-mail: Frank.Glaw@zsm.mwn.de

Miguel Vences, Museum national d’Histoire naturelle, Laboratoire des Reptiles
et Amphibiens, 25 rue Cuvier, F-75005 Paris, France. E-mail: m.vences@t-online.de

Kathrin Schmidt, Zoologisches Institut der Universität, Luisenstr. 14, D-80333
München, Germany. E-mail: kschmidt@zi.biologie.uni-muenchen.de

Introduction

The genus Paroedura Günther, 1879 comprises nocturnal geckos which are endemic to Madagascar and
the Comoro islands (Guibe 1956, Dixon & Kroll 1974), although fossil remains are also known from the
Aldabra Atoll (Arnold 1976). The genus was recently reviewed by Nussbaum & Raxworthy (2000).
According to these authors, Paroedura currently contains 14 species and can be divided into two
phenetic species groups. The sanctijohannis group is defined by having the nostril in contact with the
rostral scale whereas the picta group is defined by having the nostril excluded from contact with the
rostral scale. The picta group was hitherto considered as restricted to the dry southern and western
Madagascar, whereas species of the sanctijohannis group generally occur in less dry regions.

In this paper we describe a new species of Paroedura from northern Madagascar which belongs to
the picta group according to the nostril position, but also shares similarities with a species of the
P. sanctijohannis group.

249

Material and methods

Specimens were anesthetized by injection with chlorobutanol, fixed with 96 % ethanol and stored in 70 %
ethanol. To make comparisons easier, the terminology and abbreviations of characters largely follow Nussbaum
& Raxworthy (2000). Abbreviations used: UADBA = Universite d’Antananarivo, Departement de Biologie Ani-
male; ZEMK=Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn; ZSM = Zoologische
Staatssammlung, München. SVL=snout-vent length; TL=tail length; HL=head length; HW=head width, at
widest point; SL=snout length, anterior edge of eye to tip of snout; ED=horizontal eye diameter; EO=ear
opening diameter; AGL=axilla-groin length; Forelimb = forelimb length, from axilla to tip of longest finger;
Hindlimb=hindlimb length, from groin to tip of longest toe; Supralab=number of supralabial scales;
Infralab=number of infralabial scales; Sdim=number of subdigital lamellae on digits I-V of manus;
Sdlp=number of subdigital lamellae on digits I-V of pes. Counts are listed left-right. All measurements were
done with a caliper to the nearest 0.1 mm by the same person (FG). Material of 11 of the 14 Paroedura species
(all except P. maingoka, P. vahiny and P. homalorhina) from the ZSM and ZFMK collections was available for
comparison.

Paroedura lohatsara, spec. nov.
Figs 1-3

Types. Holotype: ZSM 985/2001, adult male, collected 14-21 March 2000 at Montagne des Francais (between
12°19'17"S, 49°20'13"E, 174 m altitude and 12°19'34"S, 49°20'09"E, 334 m altitude), northern Madagascar, by
F. Glaw, K. Schmidt & M. Vences. Captured as adult and kept for more than one year in captivity before
preserved. — Paratypes: ZSM 529/2000 (adult male), ZSM 807/2001 (adult female, captured as adult and kept
for about one year in captivity before preserved), ZSM 530/ 2000 (juvenile), and three further uncatalogued adult
specimens (one male and two females which are still kept alive in the vivarium), all collected 14-21 March 2000
at the same locality by the same collectors as the holotype. ZSM 986/ 2001, just hatched juvenile, and ZSM 987 /
2001, 7-14 days old juvenile, both captive bred, being offspring of the holotype and of one of the two uncata-
logued female paratypes.

Diagnosis. Paroedura lohatsara, spec. nov. is a relatively large-sized species (maximum SVL 80.6 mm,
maximum total length 156.1 mm) with prominent dorsal tubercles which are arranged into distinct
longitudinal rows. It differs from the species of the sanctijohannis group (P. gracilis, P. homalorhina,
P. karstophila, P. oviceps, P. masobe, P. sanctijohannis, P. stumpffi, P. tanjaka and P. vazimba) in having the
nostril excluded from contact with the rostral scale by interposition of a large prenasal scale. P. lohatsara
shares the nostril position with the species of the picta group (P. maingoka, P. bastardi, P. picta, P. vahiny
and P. androyensis), but differs from P. androyensis and P. vahiny in much larger size (80.6 compared to
47 mm maximum SVL); from P. maingoka and P. picta by distincly larger dorsal tubercles which are
arranged into obvious longitudinal rows. It differs from P. bastardi (including the recently described
subspecies P. b. ibityensis, see Rösler & Krüger 1998) by mainly tetrahedral dorsal tubercles (mainly
trihedral in P. bastardi), a relatively longer and thinner tail, and the shape of the postmental scales
(distinctly longer than wide in P. lohatsara versus regular hexagonal in P. bastardi). Furthermore,
P. lohatsara differs from all other Paroedura species by its distinct adult colouration and from P. bastardi,
P. maingoka, P. picta, and P. stumpffi by juvenile colouration (the juvenile colourations of the other
species are still undescribed).

Description of the holotype

Measurements and counts in tab. 1. Well preserved, with complete original tail. Hemipenis extruded,
head wider than neck, about as wide as body. Snout angled downward to tip, slight depression
between prominent canthal ridges. Ear opening is a vertical slit. Tail longer than snout-vent length,
nearly round in cross section, with sharply pointed tip; ventral pygal section with pair of postcloacal
sacs. Digits moderately expanded at tips. Rostral scale rectangular, wider than tall, as wide as mental.
Nostril in contact with large prenasal anteriorly, and four further scales, but not with first supralabial.
First supralabial largest, labials smooth. Snout and interorbital scales juxtaposed, some raised, scales
in front of orbits tuberculate, as are larger lateral occipital scales. Dorsolateral neck and body scales
very heterogeneous with about 12 distinct longitudinal rows at midbody of enlarged, spiny, mainly
tetrahedral tubercles; enlarged tubercles separated mostly by small flat scales and smaller tubercles.
Dorsal scales of forelimbs flat or tuberculate and weakly imbricate. Dorsal scales of hindlimbs large and

250

Paroedura lohatsara,

spec. nov. Female paratype (uncatalogued).

strongly tuberculate, much smaller above kneejoint. Ventral scales of forelimbs and hindlimbs slightly
smaller than surrounding ventral scales of the body. Dorsal pygal scales like dorsal body scales;
lateroventral pygals tuberculate but less spinous. First 19 postpygal tail segments each with transverse
row of spiny tubercles dorsolaterally; first four rows with 10-12 tubercles, gradually decreasing to the
last rows with about four tubercles; posterior tail with flat scales. Mental triangular, bordered poste-

riorly by a pair of elongate, irregular hexagonal postmentals. Postmentals contact mental, first infral- |

abial, one enlarged lateral gular, one smaller posterolateral gular, and one slighty larger central gular.
First three infralabials significantly larger than others. Gulars small, granular. Ventrals of chest and
abdomen flat, posterior abdominals largest. Proximal subdigitals in rows of 2-3, distally to narrow leaf-
like rows of scales followed by enlarged row supporting terminal pads. Pair of squarish, terminal pads,
each pad about 1 mm across. Claws curving downwards between terminal pads of digits.

Colour after one day in alcohol virtually identical to that in life (Fig 1). Head dorsally beige with
more or less symmetrical blackish markings. A black band from the second supralabial to anterior eye

and from posterior eye to a point above the ear opening. A beige mid-dorsal stripe (of 2 mm diameter

at midbody) runs from the neck to the pygal portion of tail. Neck and dorsum beige with blackish spots
which are arranged into four irregular longitudinal rows; two rows border the mid-dorsal stripe, the

other two rows run more dorsolaterally to the pygal portion of the tail. Additional black spots are
present on the flanks. Many of the enlarged tubercles on the flanks are whitish. Dorsal surface of

forelimbs and hindlimbs beige with dark brown markings. Postpygal tail dorsally with about nine
whitish and nine black alternating transverse bands which are partly not well delimited in the proximal
portion of the tail. Throat, chest, venter, pygal tail portion and ventral parts of forelimbs and hindlimbs
whitish; ventral side of tail light brownish. Tongue dark grey at its distal tip. A yellow ring around the

eye. Iris silvery-golden with small veins in life, not recognizable when preserved since pupil was much

enlarged after preservation.

Tab. 1. Morphometric and meristic variation among the holotype and two adult paratypes of Paroedura lohatsara. '

Measurements in mm.

Collection number ZSM 985/ 2001 ZSM 529 / 2000 ZSM 807/ 2001
Status holotype paratype paratype
Sex male male female
Maturity mature mature mature
SVL 70.6 69.0 72.8
TE 75.8 45.8* 65.0
HL 25.1 24.0 25.6
HW 17.5 16.3 17.0
Sb IT, 33 9.6
ED 57 5.4 5.5
EO 2.7x0.5 2.2x0.5 3.1x0.8
AGL 30.4 30.1 33.0
Forelimb 237, 25.0 25.9
Hindlimb 36.3 34.4 32.0
Supralab 12-11 10-10 13-12
Infralab 11-12 9-9 11-11
Sdlm I 8-8 10 9-9
Sdim II 10-9 9 10-11
Sdim III 11-10 12 11-11
Sdim IV 12-12 11 11-11
Sdlm V 10-10 11 10-10
Sdip I 8-9 8-9 99
Sdip II 10-11 9-9 10-11
Sdlp II 12-13 12-13 11-12
Sdip IV 14-14 13-13 13-13
Sdip V 13-14 14-12 12-11

* tail regenerated

252

Variation. Morphometric and meristic variation of two paratypes (ZSM 529/2000 and ZSM 807 / 2001)

‚ are summarized in table 1. ZSM 529/ 2000 is an adult male with extruded hemipenes; the right forelimb

was removed for future DNA studies. The regenerated tail has no spinous tubercles and is irregularly

‚ marbled with brown and white. The general colouration is similar to the holotype whereas that of the
' female ZSM 807 / 2001 is more contrasting: the largely symmetrical blackish markings on the head form
' a vermiculated pattern and the black spots on the dorsum are less clearly arranged into longitudinal
‚ rows. A distinct white band bordered by a thin blackish band on each side runs from the anterior
‚ dorsum to the anterior insertion of arm. The original and complete tail has 11 alternating dark and 10

light bands and is distinctly thinner than in the holotype. The whitish scales on the flanks are very

‚ prominent. The three living paratypes (all with original tail) largely agree with the preserved types.
‚ Their size (measured 12 July 2001 in life) was 73.9 mm SVL + 82.2 mm TL (male), 80.6 mm SVL +
' 72.5 mm TL (female, Fig. 2), and 79.7 mm SVL + 57.1 mm TL (female, last tail tip missing). TL is shorter
‚ than SVL in the three females, but longer than SVL in the two males with original tail. The SVL of the
three adult males (69.0-73.9 mm SVL) is slightly shorter than in the three females (72.8-80.6 mm), but

total length appears similar in both sexes. All type specimens and all further captive-bred specimens
agree in having distinct dark markings on the head. The juvenile paratypes (ZSM 530/2000, 32.4 mm
SVL + 31.3 mm TL; ZSM 986 / 2001, 32.0 mm SVL + 26.8 mm TL; ZSM 987 / 2001, 31.2 mm SVL + 29.8 mm

_ TL) have a distinct juvenile colouration which was also typical for the other juveniles which we reared
in captivity (Fig. 3): Four distinct whitish transverse bands on the dark brown back and flanks.

Laterally, the most anterior band is distinctly narrower than the two following bands and ends pointed
slightly anterior of the insertion of the forelimbs. The two bands between forelimbs and hindlimbs are

' not pointed laterally and have the same width on the entire flanks as on the back. The posteriormost
' band, positioned between the hindlimbs, is restricted to the back. There is no light mid-dorsal line. The

upper surface of the tail is banded with beige and brown in the preserved specimens, but bright orange
with dark crossbands in life. Dark symmetrical markings on the head are already well recognizable.
The head is relatively broad and short, especially in comparison with similar sized juveniles of

P.stumpffi and P. bastardi. About three months after hatching the juvenile colouration gradually
converts into the adult colouration. Subadults still have more or less distinct transversal bands on the

back whereas in older adults these bands are poorly or not recognizable. A light mid-dorsal stripe is
present in subadults and adults.

Distribution and conservation. Paroedura lohatsara is only known from the Montagne des Francais in
the far north of Madagascar. Numerous animal and plant species appear to be endemic to this karstic
massif, among them a still undescribed snake species of the genus Heteroliodon (pers. obs.). It is
therefore likely that Paroedura lohatsara is a further endemic species of the Montagne des Frangais
massif. In this case the new species may be considered as vulnerable due to its small range although
its habitat is apparently not immediately threatened by destruction. Regarding the numerous endemics
in Montagne des Frangais this area should be protected as nature reserve.

Habitat and habits. In nature, the new species was only observed at night in dry forest on a karstic

_ underground, several days after heavy rains. The geckos were mainly climbing on rocks and branches

_ up to two metres above the ground. In captivity, juveniles and adults were able to feed on relatively

large insects. Nussbaum & Raxworthy (2000) described a prominent vertical or anteriorly curved tail
display in disturbed Paroedura maingoka which they interpreted as defensive behaviour. A similar
behaviour was sometimes observed by us in disturbed captive-bred subadult P. lohatsara and
P. stumpffi, e.g. when the specimens were faced with torchlight at night. However, it was also observed

in undisturbed P. lohatsara in the vivarium, indicating that it may also serve for intraspecific commu-

nication. A detailed analysis of this and other behavioural traits in Paroedura picta was provided by
Brillet (1986, 1993).

Sympatric species. Two other species of Paroedura were found in sympatry with P. lohatsara. Both are
hereby recorded for the first time at Montagne des Frangais: P. karstophila (ZSM 531/2000 and 532/
2000) was found in close syntopy with P. lohatsara in the limestone massif whereas Paroedura stumpffi
(ZSM 635/ 2000) was only encountered on the slope between the massif and the sea, outside the karstic
underground.

253

Reproduction. One male and two females of P. lohatsara, captured as adults at Montagne des Frangais

in March 2000, were kept and bred in a vivarium (1mx0.5mx0.8m) together with a couple of
P. stumpffi from the same locality. As usual in other Paroedura species (Schröder 1987, Rösler 1998), a

clutch generally consisted of two single eggs which were neither glued to each other nor to the

underground. The white eggs had a hard calcareous shell and were buried singly few millimetres into

sandy ground. In comparison to P. stumpffi, eggs of P. lohatsara were generally deposited in drier places
and were always distinctly larger. 15-20 days after egg deposition eggs measured 13.2x 10.3 mm and

13.4 x 10.4 mm (n=2) in P. lohatsara and 10.8 x 8.2 mm to 11.6 x 8.6 mm (n=5) in P. stumpffi (a similar size |
difference was observed but not measured in numerous other cases). Eggs were incubated differently, '
either at a constant temperature of 27°C or at 30°C during the day and 23°C at night. Juveniles |

hatched 70-82 days after the discovery of the eggs (at 27 °C) and after 85-87 days under the variable
incubation temperatures, indicating that the total time between egg deposition and hatching was ca.

70-90 days. Hatchlings of P. lohatsara (total length 59-60 mm, n=2) were distinctly larger than those of |
P. stumpffi (total length 50-52 mm, n=3). All juveniles were identified either as P. stumpffi or P. lohatsara,

indicating that hybridization did not occur in the vivarium.

Only little is known about reproduction of P. lohatsara in nature. The total length of a juvenile
paratype (ZSM 530/2000), collected 21 March 2000, is only 4 mm longer than that of just hatched
juveniles, indicating that it was only few weeks old when captured. Assuming an approximate egg

incubation time of 70-90 days (as found in captivity) the corresponding egg was laid at the beginning |

of the rainy season as is typical in many Madagascan reptiles (Glaw & Vences 1996).

The first eggs laid by the F1 generation were found on 30 June 2001, ten months after hatching of
the first juveniles (22 August 2000), indicating that sexual maturity in captivity was reached less than |

ten months after hatching. To document the adult size at first reproduction, the largest male and female
of the FI-group were measured after the first eggs were found: The largest male measured 111 mm total
length and the largest female 106 mm total length.

According to our experience, egg laying females of P. lohatsara, P. tanjaka and P. bastardi apparently
have an enormous need of calcium and are vulnerable to rachitis. Despite of regular addition of several

pulverized Calcium products to the food, the faeces sometimes contained a remarkable amount of sand
which was apparently actively ingested to take in calcium. After much pulverized limestone was added |

to the vivarium the faeces contained less sand. The high need of calcium may explain why several

Paroedura species like P. karstophila, P. tanjaka, and P. lohatsara seem to be restricted to limestone habitats.

This restriction may be true for other oviparous Malagasy reptiles with hard-shelled eggs as well.

Etymology. loha (Malagasy) means head; tsara (Malagasy) means beautiful or good. The specific name lohatsara

refers to the beautiful head colouration of the new species and is considered as invariable noun in apposition

to the generic name.

Available names. Several available names in the genus Paroedura are considered as junior synonyms |

of valid species names, all of them belonging to species having the nostril separated from the rostral.
These names need to be considered as possible earlier names for P. lohatsara. Diplodactylus porogaster
Boulenger, 1896 (type locality: “south-western Madagascar”) is considered as synonym of Paroedura

androyensis (Angel 1942, Guibe 1956), and Diplodactylus robustus Boulenger, 1896 (type locality: “south- |

western Madagascar”) as synonym of P. picta (Angel 1942, Guibe 1956). The type specimens of both taxa
are deposited in the Natural History Museum at London and are not available at present. However,

based on the chromatic and morphological characters given in the original descriptions (Boulenger |

1896) it seems likely that these synonymies are correct. In addition, the type locality of both taxa is in

south-western Madagascar whereas P. lohatsara is only known from a karstic massif at the northern tip |
of the island. The type locality (“Tullear”) of Phyllodactylus madagascariensis Mocquard, 1894 is in south- |

western Madagascar as well. This taxon is considered as synonym of P. picta (Angel 1942) which is
common around Tulear according to our observations. It is based on a single poorly preserved
specimen with a SVL of 40 mm (Mocquard 1895) and the characters given in the original description

seem to fit to those of juvenile P. picta. Paroedura guibene Dixon & Kroll, 1974 was shown to be a junior |
synonym of P. bastardi by Nussbaum & Raxworthy (2000). Summarizing, none of the synonyms is |

available as earlier name for P. lohatsara.

Relationships. As already noted above, the genus Paroedura can be divided into two phenetic species |
groups based on the position of the nostril (Nussbaum & Raxworthy 2000). The species of the

254

Fig. 3. Juveniles of Paroedura bastardi (left above), P. lohatsara (middle) and P. stumpffi (right below).

sanctijohannis group are widely distributed in different climatic regions in western (P. tanjaka, P. va-
zimba), eastern (P. gracilis, P. masobe) and northern Madagascar (P. homalorhina, P. oviceps, P. stumpffi,
P. karstophila), and even on the Comoro Islands (P. sanctijohannis). The species of the picta group are
thought to be largely restricted to arid southwestern Madagascar (P. maingoka, P. bastardi, P. picta,
P. vahiny and P. androyensis). P. lohatsara does not support this distributional pattern: It clearly belongs
to the picta group based on the position of nostrils, but is only known from extreme northern
Madagascar. Although the range of P. lohatsara is widely separated from the other species of the picta
group, this fact does not necessarily argue against the monophyly of this phenetic group since several
animal and plant taxa of the Montagne des Frangais have affinities to those of dry western and southern
Madagascar. Further research is therefore necessary to clarify whether the two phenetic species groups
which are defined by a single morphological character (nostril position) represent natural clusters or
not.

Another remarkable morphological character in Paroedura is the prominence of the dorsal tubercles.
These are very distinct and arranged in obvious longitudinal rows in P. bastardi, P. stumpffi, and
P. lohatsara but less prominent (and arranged in obvious longitudinal rows in only some taxa) in the
other species (gracilis, oviceps, karstophila, masobe, sanctijohannis, androyensis, picta, vazimba, vahiny,
homalorhina, tanjaka, maingoka). The similarity of P. stumpffi and P. bastardi regarding this character was
possibly the reason why Dixon & Kroll (1974) considered their P. guibene (which was synonymized with
P. bastardi by Nussbaum & Raxworthy 2000) as “mainland form” of P. stumpffi. Most of the prominent
dorsal tubercles of P. bastardi can be described as trihedral whereas those of P. stumpffi and P. lohatsara
are mainly tetrahedral. Further similarities between the latter two species in (1) the shape of the tail
(relatively long and always thin in P. lohatsara and P. stumpffi versus relatively short and often rather
thick in P. bastardi), (2) the shape of the postmental scales (distinctly longer than wide in P. lohatsara
and P. stumpffi versus regular hexagonal in P. bastardi), (3) several characters of adult and juvenile
colouration, and (4) geographic distribution (P. stumpffi and P. lohatsara in the north, P. bastardi in the
west and south) may indicate that P. lohatsara is perhaps more closely related to P. stumpffi than to
P. bastardi.

Acknowledgements

We are grateful to Marion Kotrba (ZSM) who improved the English and to Wolfgang Böhme (ZFMK) who
allowed the loan of specimens for comparison. The research in Madagascar was made possible by a cooperation
accord between the UADBA and the ZSM. Malagasy authorities are acknowledged for collection and export
permits. The research was supported by the “Deutsche Forschungsgemeinschaft” DFG (GL 314/1 to F. G.) and
the “Deutscher Akademischer Austauschdienst” (to M. V.).

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256

t
h

Two new species of the genus Oreophryne

from Irian Jaya, Indonesia

(Amphibia, Anura, Microhylidae)

Rainer Günther, Stephen J. Richards & Djoko Iskandar

Günther, R., S. J. Richards & D. Iskandar (2001): Two new species of the genus
Oreophryne from Irian Jaya, Indonesia (Amphibia, Anura, Microhylidae). - Spixiana
24/3: 257-274

Two new species of Oreophryne from western Irian Jaya (Papua Barat = West
Papua) are described. They are small (SVL of males < 24mm) and distinguished
from all congeners by having prominent, angular snouts and dark brown or black
head sides and throats. Both taxa are scansorial, calling from leaves between about
0.5 m and 2.5 m above the ground in rainforest habitats. The advertisement calls of
both species consist of long trains of distinctly pulsed notes, but calls differ from
one another in several structural features including the number of pulses per note.

Rainer Günther, Institut für Systematische Zoologie, Museum für Naturkunde
der Humboldt-Universität zu Berlin, Invalidenstr. 43, D-10115 Berlin, Germany

Stephen ]. Richards, Vertebrate Department, South Australian Museum, North
Terrace, Adelaide, SA 5000, Australia

Djoko Iskandar, Laboratory of Biosystematics and Genetics, Jurusan Biology,
Faculty of Mathematics and Natural Sciences, Institute of Technology, Bandung 10,
Jalan Ganesha, Bandung 40132, Indonesia

Introduction

The most recent overviews of New Guinea amphibians recognise 14 species of the microhylid genus
Oreophryne from mainland New Guinea and surrounding islands (Frost 1985, 1998-1999, Zweifel &
Tyler 1982). Since those summaries only one additional species, the extremely small O. minuta, has been
described from the region (Richards & Iskandar 2000). However recent surveys in Indonesian New
Guinea (Irian Jaya) by S. Richards and D. Iskandar, and independently by R. Günther, have revealed
a number of new Oreophryne species. Two of these, one from the Wondiwoi Mountains at the base of
the Wandammen Peninsula, and the other from the Wapoga River basin, appear to be very closely

_ related to one another but distinct from all previously described Oreophryne. Here we describe the new
_ species and present brief observations on their advertisement calls and natural history.

Material and methods

Frogs were generally collected at night when they were easily detected by their distinctive advertisement calls.
Several individuals of both new species were photographed in life and all specimens were anaesthetised with
chlorobutanol and fixed in 2 % formalin (RG) or 70 % ethanol (SJR, DI). Tissue samples were extracted from some
specimens and stored in 70 % ethanol before the animals were fixed in formalin, to allow later DNA studies. All
specimens were preserved in 75% ethanol upon return to the laboratory.

257

Advertisement calls were recorded with a Sony Walkman TCD-D100 Digital Audio Tape (DAT) and
Sennheiser microphone MKE 300 (RG) or a Sony WMD-6C Professional Walkman and ECM-Z 200 microphone

(SJR). Calls were analyzed with Avisoft-SAS Lab software. Two paratypes of the new species from Wandammen
Peninsula were cleared and stained using a method modified from Dingerkus & Uhler (1977). Snout-urostyle

length (SUL), tibia length (TL), length of 4th toe (LAT) and length of 3rd toe (L3T) — from tip of the toes to

proximal end of inner metatarsal tubercle, tarsus length (TaL), and distance between the supratympanic folds
immediately behind eyes (FD), were measured with a vernier calliper; length of the first toe (LIT) from tipto
distal end of inner metatarsal tubercle, length of the inner metatarsal tubercle (LMT), horizontal diameter ofthe

disc of the 4th toe (T4D) and that of the 3rd finger (F3D), head length from tip of snout to posterior margin of
tympanum (HL), head width taken in the ear region (HW), distance from anterior corner of orbital opening to

center of naris (END), internarial distance between the centres of the nares (IND), distance from anterior to
posterior corner of orbital opening (ED), and horizontal diameter of tympanum (TyD) were measured with an

ocular micrometer in a binocular dissecting microscope.

Type specimens are deposited in the Museum für Naturkunde, Berlin (ZMB), the South Australian Museum, |

Adelaide (SAMA) and the Museum Zoologicum Bogoriense, Bogor (MZB).
Photographs in Figs 14 and 15 were taken by S. Richards, all others by R. Günther.

Oreophryne atrigularis, spec. nov.
Figs 1-13

Types. Holotype: ZMB 62226, adult male, collected by R. Günther on 30.VII.1998 at an altitude of 610 ma.s.l,

Wondiwoi Mountains at the base of the Wandammen Peninsula, about 8 km west of the coastal village of
Yeretuar, 2°56'S, 134°36'E, Nabire district, Irian Jaya, Indonesia. — Paratypes: 22 adult males with inventory
numbers ZMB 62162, 62164-67, 62182, 62215, 62217-20, 62222-25, 62641-42 and 62644-45, MZB. Amph. 7361-62
and SAMA R55924; one adult and one subadult female (ZMB 62640 and 62163); and two juvenile specimens ZMB

62216 and 62646. Two adult males (ZMB 63291 and 63292), were cleared and double stained as skeletal !
preparations and are stored in glycerine. All paratypes were collected between 350-750 m a.s.l. on the eastern |
slopes of the Wondiwoi Mountains west of the coastal village of Yeretuar, ZMB 62214-26 between 28.VIL- |
1.VII.1998, ZMB 62162-66, 62182 between 21.-29.VIII.1999 and ZMB 62167, 62640-46 between 7.-12.V.2000. |

Collectors were R. Günther, S. Marani, G. Mareku and I. Tetzlaff.

Diagnosis. O. atrigularis is assigned to the genus Oreophryne on the basis of the structure of its shoulder
girdle: clavicles and procoracoids are present but reduced. The former are small ossified bones which

are broadly separated and reach laterally only to the middle of the coracoids. The cartilaginous '
procoracoids nearly meet the anterior tip of the sternum, and extend laterally no further than the

middle of the coracoids. A combination of the following characters separates the new species from all
hitherto known congeners: (1) snout strongly protruding and canthus rostralis sharply edged, (2)
throat and sides of head black, (3) no webbing between toes, (4) fifth toe shorter than third toe.
Advertisement call a long series of notes, each between 171 and 262 milliseconds in length.

Description of the holotype

An adult male with the following measurements (in mm) and ratios: SUL 22.0, TL 11.7, LAT 9.9, LST
7.2, ED 6.7, Tal 7.2, LIT 1.7, LMT 1.0, T4D 1.0, EBD1.1, HL 6.8, FW 83, END2.0,IND2TZED2ZZI,D
0.8; TL/SUL 0.53, HL/SUL 0.309, HW /SUL 0.377, END/SUL 0.091, ED/SUL 0.109, HL/HW 0.819 and
F3D/SUL 0.050. Skin smooth, snout tapering and projecting, canthus rostralis sharply defined; no
maxillary or vomerine teeth; two palatal ridges, anterior ridge very low and with indistinct lobes,
posterior ridge with 14 small but distinct whitish denticles; tongue elongated, without indentation and

free posteriorly; one rather long slit-like vocal sac opening on each side of the tongue, with a single |

internal subgular vocal sac. Loreal region straight and angled outwards; tympanum covered by skin
and scarcely visible. Head broader than long; legs rather long and slender, all fingers and toes
expanded distally into discs with circummarginal grooves, discs on fingers somewhat broader than
those on toes, terminal disc of 3rd finger about twice as broad as penultimate phalanx, first finger and
first toe conspicuously reduced. Fingers in order of decreasing length 3>4>2>1; toes 4>3>5>2>1.
Subarticular tubercles and metatarsal tubercle poorly developed. Upper side of head, dorsum and |
limbs beige with some spots and marbling. A conspicuous whitish mid-dorsal line is bordered by

concentrations of blackish pigments along its whole length. A similar line extends from heel to middle
of posterior side of tibia. A w-shaped mark in the scapular region surrounds two whitish spots. A black '

258

Fig. 1. Lateral view of a paratype (ZMB 62167) of Oreophryne atrigularis, spec. nov.

\ ni EN TUN

ne 2

Fig. 2. Ventral view of a paratype of Oreophryne atrigularis, spec. nov.

Waazı

Fig. 3. Ventral view of the right hand (left) and the right foot (right) of a paratype (ZMB 62644) of Oreophryne
atrigularis, spec. noV.

band from upper arm to tip of snout contrasts sharply with the beige dorsal colouration but grades into
the blackish throat ventrally. Anterior of belly speckled with dark brown, posterior of abdomen cream
coloured without spots. Concealed parts of hind limbs unpigmented, yellowish in preservative but
reddish in life.

Variation in the type series. Snout-urostyle length of 22 adult males ranged from 20.2 to 22.5 mm
(mean 21.5 mm, SD 0.65). One female was adult at 26.3 mm, and another female appears to be subadult
at 22 mm, with oocytes in the early stages of development. Two specimens of 15.8 and 18 mm SUL
respectively show no signs of sexual maturity. Selected body proportions of the type series (except
juveniles and cleared and stained specimens) are listed in table 1. Most specimens beige dorsally, but
some with greyish-brown ground colour in life, retained in ethanol. Shape of pupil is oval in all
specimens, but colour in life varied from yellowish or silvery to brown. Margin of eyelid was white in
all specimens in life. Tympana in all specimens more

or less covered by skin. Dorsum of some specimens Tab. 1. Body proportions of the type series (n=23
with minute dark spots (visible under magnifica- males and 2 females) of Oreophryne atrigularis;
tion), but others with larger spots and areas of mot- sp=Standard deviation. ö

tling on dorsum and extremities. Thirteen out of 27
specimens have a thin whitish mid-dorsal line ex-

tending from snout to end of urostyle, and in most Ratio Ne an Pe

specimens a similar line extends from heel onto pos- TL/SUL 0.54 0.019 0.48-0.57
terior side of tibia (Fig. 1). About 50 % of animals HL/SUL 0.30 0.010 0.28-0.32
exhibit a w-shaped mark in scapular region that en- HW/SUL 0.37 0.016 0.33-0.40
compasses two whitish spots. Commonly a few HL/HW 0.82 0.037 0.74-0.89
ED/SUL 0.110 0.005 0.098-0.120

small warts on dorsum and/or dorsal side of hind

limbs. Concealed surfaces (while sitting) of hind EINIDY/SIOIL 0 US ST
: - : : 5 F3D/SUL 0.053 0.004 0.043-0.060
limbs ininearlysall'speeimens without‘ dark pigement mn

260

1
[

Fig. 4. Cleared and double stained paratype (ZMB 63291, older specimen) of Oreophryne atrigularis, spec. nov.
Dorsal view of the whole skeleton except distal parts of extremities.

cream coloured in fixative and reddish or red in life (Fig. 2). A black band from insertion of upper arm
through lower part of tympanum to tip of snout, including eye and sides of snout, is conspicuous in
all specimens. This black stripe contrasts sharply with beige colour of dorsum, but merges with
blackish colour of the throat. Throat pale grey (not blackish) in only two specimens (ZMB 62640, female;
ZMB 62641, male). Black colour of throat may extend to chest and anterior of belly, but generally grades
posteriorly into dark brown or dark grey marbling. Some individuals with very small white spots on
dark parts of venter, and some with an inconspicuous whitish cross on chest. Size and shape of toes
and fingers as described for the holotype (Fig. 3). Ventral side of extremities more or less spotted. Skin
smooth dorsally and ventrally in most specimens, but some with belly granular. Black coloration on
throat and laterally on head is consistent among males, females and juveniles demonstrating that this
coloration is not a secondary sexual character. However conspicuous black colour of head, evident in
living specimens during the day and in preserved specimens as well, is more or less faded, and in some
cases invisible, in specimens actively calling at night.

261

Fig. 5. Premaxillaries of Oreophryne atrigularis, spec. nov. (ZMB 63291).
Fig. 6. Prevomers of Oreophryne atrigularis, spec. nov. (ZMB 63291).

Fig. 7. Hyoid plate of Oreophryne atrigularis, spec. nov. Anterior process of processus posterolateralis is bifurcate

in this older specimen (ZMB 63291) but has only a single process in the younger cleared specimen.
Fig. 8. Shoulder girdle of Oreophryne atrigularis, spec. nov. (ZMB 63291).
White: bones; stippled: cartilago.

Osteology (based on two cleared and double stained paratypes, ZMB 63291 and 63292): Otoccipitals
(sensu Trueb 1973) broad and fused with frontoparietals, sutures well marked, frontoparietals paired,
long and broad, with a very small interspace and covering most of the sphenethmoid; paired nasals
somewhat broader than long, about one quarter as long as frontoparietals plus otoccipitals (Fig. 4);
processus paraorbitalis of the nasal well developed and venterolaterally oriented. Each premaxillary
with a strongly bent alary process (Fig. 5). Maxillary without teeth and only marginally overlapping
premaxillary; ventral ramus of squamosal long, its zygomatic ramus very small and its otic ramus well
developed; prevomers (sensu Trueb 1973) do not meet medially and nearly completely include the
choanae (Fig. 6); pars lateralis of parasphenoid broad and long, pars medialis (cultriform process) long,

nearly reaching posterior border of palatine bones; cartilaginous hyoid plate has large antero-lateral |

processes as well as bifurcate postero-lateral processes (Fig. 7), anterior process of postero-lateral

process with bifurcate tip in the older specimen (ZMB 63291, see Fig. 7) but with a single tip in the

younger one (ZMB 63292); hyalia without processes, postero-medial processes completely ossified,

slightly bent and each showing a small postero-medial crista. Coracoids slightly angled, procoracoids

cartilaginous, club-shaped and reaching laterally only to middle of coracoid, a thin rod-shaped, ossified

clavicula is attached to the anterior margin of each procoracoid (Fig. 8). Sternum voluminous, omos-

ternum lacking, xiphisternum with two lateral lobes. In the obviously younger specimen (ZMB 63292)
the sternum is mostly cartilaginous (only corpus sterni and manubrium sterni with a few bone
substances). In the older specimen (ZMB 63291) corpus sterni and manubrium are completely ossified

and only lateral lobes are cartilaginous. Both specimens show two other differences regarding ossifi- |
cation of bones: (1) in the younger specimen there is a cartilaginous “bridge” between coracoid and |

scapula, in the older this “bridge” is ossified and both bones seems to be fused to a single one; (2)
anterior part of cleithrum is ossified in both specimens; the posterior part is ossified in the older
specimen but is cartilaginous in the younger specimen. Terminal phalanges T-shaped, cross part (of

262

Fig. 9. Rainforest habitat of Oreophryne atrigularis in the Wondiwoi Mts., 550 m a.s.l.

the T) relatively short. Eight nonimbricate presacral vertebrae, the third one with broadest transverse
processes, sacral diapophyses broadly expanded, urostyle with a weakly developed bifide crista on its
dorsal anterior half, which becomes single-ridged posteriorly.

Etymology. The specific epithet refers to the colouration of the throat. Ater (atra, atrum) is a Latin adjective and
means black, and gularis is also Latin and means “belonging to throat”.

Distribution. Known only from the type locality, the slopes and valleys between 350 and 750 m a.s.l.
inthe Wondiwoi Mountains west of the coastal village of Yeretuar, base of the Wandammen Peninsula,
Irian Jaya.

Habitat and habits. O. atrigularis is one of the most abundant amphibians in the Wondiwoi Mountains.
It lives in dense closed rain forest and in more open forest and the undergrowth varies from extremely
sparse to areas with many shrubs, grasses and herbs (Fig. 9). Frogs were found on steep slopes, and
also at the bottom of valleys and gorges, often long distances from water. At night males perched on
leaves between 0.5 m and 2.5 m high. They were spaced at least several metres apart and no more than
one male occupied the same shrub. Some males started calling from beneath the leaf litter at dusk, and
then climbed on to small shrubs where they continued calling during the night. This suggests that the
litter is used as a diurnal retreat by this species. Calling activity was most intense between shortly after
dark and 9 p.m., but single call series were heard throughout the night. Calling males were found in
May, July and August; the site was not visited during any other month.

Vocalization. Advertisement calls consist of series of notes. Most series start with notes having long
and irregular inter-note intervals (“slow” notes). During the call sequence note intervals become
shorter and more regular (“fast” notes) (Fig. 10). There are calls with only a few introductory notes and
long phases of fast notes, others have equal parts of slow and fast notes, and there are also calls with
many introductory slow notes and only a few fast notes at the end. Most call series last 15 to 30 s and
are separated from each other by intervals of at least 4s (usually more). In 30 call series, the call
components with short and regular intervals contained a mean of 17.7 notes (SD 7.83, range 6-30). Call
series end abruptly during fast note sequences. Mean note length of 319 notes was 198 ms (SD 16.2),

263

5 10 15 20s

Fig. 10. Oscillogram of an advertisement call sequence of Oreophryne atrigularis. First six notes are of lower
amplitude and the intervals between them are longer and irregular. These are followed by a sequence of 24 “fast”
notes (see text). All recordings were made at temperatures between 22 and 24.5 °C.

Fig. 11. Audiospectrogram (below) and oscillogram (above) of a series of six “fast” notes of Oreophryne atrigu-
laris.

m - ——

re Te En aer ' Ta ANVER,
0,05 0,10 0,15 0,20 s

Fig. 12. Oscillogram of a single note of Oreophryne atrigularis.

264

1 15 2 25 3 Na 5 6 65 7 75 8 8598 95 10 105 khz

Fig. 13. Frequency spectrum of one note of Oreophryne atrigularis.

minimum length was 171 ms and maximum duration was 262 ms. Inter-note intervals between 290 fast
notes had a mean duration of 226 ms (SD 18.6), a minimum of 190 and a maximum of 312 ms. Mean
repetition rate of fast notes was about 3/s. Harmonics and frequency modulations are not discernible
(Fig. 11). Notes are composed of distinct pulses. Pulses are arranged into three groups which are
separated from each other by short intervals (Figs 11 and 12). The first pulse group typically consists
of 6-8 pulses, the second group contains also 6-8 pulses but of higher amplitude, and the third group
has the longest duration and most pulses (15-25). Moreover, there is an amplitude modulation in the
latter group. Fundamental frequency is about 1.5 kHz and dominant frequency is around 3 kHz (Fig.
13). All calls were recorded at temperatures between 22 and 24.5 °C.

Comparison with other species. Oreophryne atrigularis differs from all previously described Oreo-
phryne in its possession of a protruding and sharply defined snout, and in the distinctive black
coloration of the throat and lateral surfaces of the head. Described Oreophryne from New Guinea also
differ from O. atrigularis in the following features: Oreophryne anthonyi (Boulenger, 1897), ©. idenbur-
gensis Zweifel, 1956 and O. inornata Zweifel, 1956 are easily distinguished from atrigularis by their much
larger size (snout-urostyle lengths of more than 40 mm vs 26.3 mm). Oreophryne albopunctata (van
Kampen, 1909) has webbed toes and much shorter hind limbs. O. brachypus (Werner, 1898) has quite
different mating calls (Tyler 1967) and is known only from the Bismarck Archipelago. O. brevicrus
Zweifel, 1956 has much shorter hind limbs, smaller terminal discs on the fingers and toes and is a
terrestrial species. O. crucifera (van Kampen, 1913) has webbed toes, the 3rd and 5th toe are of equal
length and the procoracoid reaches the scapula. We have examined specimens of O. flava Parker, 1934
from the AMNH and this species clearly differs from O. atrigularis in having the procoracoid reaching
the scapula and the 5th toe longer than the 3rd. We have examined the type (SMF 4197) of O. geislerorum
(Boettger, 1892) and it has very short tibiae (TL/SUL of the type 0.39) and basal webbing between the
toes. We have examined specimens of O. insulana Zweifel, 1956 from the AMNH. It differs from
O. atrigularis in having a rounded canthus rostralis, snout not protruding, 3rd and 5th toe of equal
length, and basal toe webbing. In O. kampeni Parker, 1934 the procoracoid reaches the scapula and the
toes are webbed. O. parkeri has a very distinct tympanum, its toes are webbed and its 5th toe is longer
than the 3rd (Loveridge 1955). O. wolterstorffi was treated as a member of the Hylidae by Werner (1901)
and subsequently transferred to the microhylid genus Oreophryne by Tyler (1964). We have examined
one specimen (ZMB 16853) which differs from atrigularis in its distinct toe webbing, shorter tibiae, and
5th toe longer than the 3rd. The same suite of characters distinguishes ©. moluccensis (Peters & Doria

265

Fig. 14. Oreophryne wapoga, spec. nov., paratype MZB. Amph. 7359;
specimen in the type series which has a longitudinal mid-dorsal line.

nov., same specimen as in Fig. 14.

NR SSSS

Fig. 16. Dorsal view of three paratypes and the holotype (second specimen from left in the top row) of
Oreophryne wapoga, spec. nov. and of four paratypes of Oreophryne atrigularis, spec. nov. (bottom row). Visible
numbers are field numbers.

Fig. 17. Ventral view of three paratypes and the holotype (second specimen from left in the top row) of
Oreophryne wapoga, spec. nov. and of four paratypes of Oreophryne atrigularis, spec. nov. (bottom row).

267

1878), of which we examined types of its synonym O. senckenbergiana Boettger, 1895 (SMF 4203,

lectotype; SMF 4204 and 4205, paralectotypes).

The types of O. biroi (originally described as Sphenophryne biroi by Mehely 1897) were stored at the |

Museum of Natural History in Budapest and are lost. According to Häupl et al. (1994) one syntype

received in exchange from the Natural History Museum Budapest on 18.X1.1898 was deposited as
NMW 19825 in the Naturhistorisches Museum in Wien. In fact, four specimens are catalogued under |
this number (NMW 19825: 1-4). From Mehely’s papers (1897, 1901) it is clear that his original descrip-
tion was based on only two specimens with snout-vent lengths of 17mm and 8.5 mm, collected near

Friedrich-Wilhelmshafen (today Madang). In 1900 Mehely received various specimens from Sattelberg

(about 250 km east of the type locality), the largest being 25 mm long, which he ascribed to S. biroi. It '
is extremely likely that the four specimens in the Vienna museum also originated from Sattelberg.
Because Mehely himself regarded these specimens as belonging to S. biroi, they should serve as
important material in future comparative studies (although they are certainly not syntypes of Oreo-

phryne biroi and they are larger than the types from Friedrich-Wilhelmshafen). The SUL of these 4 now
bleached specimens ranges from 20.6 to 22.5mm and they are therefore similar in size to O. atrigularis.
However, all have traces of webbing between the toes, their snouts are not protruding, TL/SUL ranges
between 0.42-0.45, the 5th toe is longer than 3rd and Mehely did not mention a black loreal region and
a blackish throat in his description of freshly preserved material. We have studied a syntype of Mehelyia

affinis Wandolleck, 1911 (NMW 19826), which was regarded as a synonym of O. biroi by van Kampen

(1923) and Parker (1934). There are sufficient differences between this syntype and the four O. biroi
specimens to suggest that they represent different taxa, and the differences between this specimen and
O. atrigularis are the same as those between O. biroi and O. atrigularis. This statement applies equally
to other names regarded as synonyms of various Oreophryne species today. None of the species
descriptions within the genus Oreophryne mention a black loreal and gular region, and a projecting
snout with a well-marked canthus rostralis.

Oreophryne wapoga, spec. nov.
Figs 14-20

Types. Holotype: MZB Amph. 7358, adult male, collected by S.J. Richards and D. Iskandar on 11.1V.1998 at an |

altitude of 1070 m asl at Wapoga Alpha Exploration Camp (136°34'423"E, 3°08'687"S), Wapoga River headwa-

ters, Irian Jaya, Indonesia. — Paratypes: Four adult males with inventory numbers ZMB 63435, SAMA R55923
and MZB Amph. 7359-60. All data as for holotype except that ZMB 63435 collected on 12.IV.1998, and SAMA

R55923 collected on 16.1V.1998 by M. Moore.

Diagnosis. Oreophryne wapoga was not studied osteologically because of scarcity of material. However, |
superficial dissection of the pectoral girdle revealed the presence of reduced clavicles, demonstrating |

that this species belongs to the genus Oreophryne. Based on external morphology (Figs 14, 15) it appears

to be closely related to O. atrigularis. Diagnostically important characters are the same as outlined for |
that species. From O. atrigularis it can be distinguished by a dark brown (not black) throat, a more

spotted pigmentation, concealed parts of hind limbs are yellowish not red, a shorter head (HL/SUL
0.339-0.357), bigger eyes (ED/SUL 0.122-0.131), greater body size (males 21.8-23.3 mm SUL) and by
different advertisement calls.

Description of the holotype

Measurements and ratios of body proportions are given in tables 2 and 3. General body shape same
as for preceding species. Ground colour dorsally pale grey in fixative and beige in life, venter cream

coloured. A solid black band extends from insertion of upper arm to tip of snout, bordered behind eye |

by a whitish stripe. Dorsum and head with irregular brownish marbling extending to tip of snout.
Shape of the pupil oval, reddish-brown in life. Darker pigments concentrated dorsolaterally extend
from eyes to middle of iliae. Flanks and dorsal surfaces of limbs covered with diffuse (not clearly
marked and differing in size) brown spots. Posterior sides of thighs dusted with many very minute

spots. All fingers and toes dark brownish above with exception of a whitish spot near base of terminal |
disk. Throat densely spotted, with irregular dark brown spots merging into each other. More isolated |
dark spots on chest and ventral sides of extremities, posterior of belly unspotted. Palms and soles |

greyish-brown, ventral side of fingers and toes inconspicuously spotted.

268

. Fig. 18. Ventral view of right hand (left) and of right foot (right)

of Oreophryne wapoga, spec. nov. (ZMB 63435).

Variation in the type series. Measurements and body proportions of the type series are given in table
2 and 3 and variation of coloration in comparison to that of O. atrigularis is shown on Figs 16 and 17.
Coloration of preserved animals is very similar to that of living ones. One specimen MZB. Amph. 7359

Tab. 2. Measurements (in mm) of the type series

(5 males) of Oreophryne wapoga.

Number

SUL
TL
Tal
L4T
L3T
T4D
F3D
HL
HW
LIT
LMT
END
IND
ED
TyD
FD

MZB. MZB. MZB
Amph. Amph. Amph. 63435
7358 7359 7360
23.0 23.3 22.8
11.8 11.9 1167
7.8 7.7 TR.
11.0 10.9 10.6
8.1 8.0 7.8
1.3 1.3 1.1
1.4 1.3 1.3
7.6 HR 7.6
7.8 79 7.8
1.8 24 "9
1.3 1.2 1.1
2.0 2.4 2.4
2.2: 2.6 2.5
2.8 3.0 2.8
0.8 0.8 0.7
Ze N.2 7.0

ZMB SAMA
R55923

Tab.3. Body proportions of the type series (n=5 males)

of Oreophryne wapoga.

Ratio Mean
TL/SUL 0.51
HL/SUL 0.334
HW/SUL 0.345
HL/HW 0.969
ED/SUL 0.126
END/SUL 0.097
F3D/SUL 0.0556

0.014

0.0045
0.0076
0.012

0.0039
0.0072
0.0038

Range

0.49-0.53
0.330-0.339
0.339-0.357
0.948-0.974
0.122-0.131
0.087-0.105
0.0505-0.0608

269

— —

T si SE
15 20s

DE]
5 10

Fig. 19. Oscillogram of an advertisement call of Oreophryne wapoga. Notes at the beginning and at the end ofthe

series are produced at a slower rate than the “fast” notes in the middle of the series.

Me. un, De Bu En Po u DE Bu 2

0.5

Fig. 20. Oscillogram (above) and audiospectrogram (below) of a sequence of nine “fast” notes of Oreophryne

wapoga. Air temperature 20.6 °C.

has a relatively broad (in comparison to O. atrigularis) mid-dorsal line, bordered by irregular dark '

brown stripes. Remaining specimens lack a mid-dorsal line. Ground colour of dorsum grey in four

specimens and brown in one specimen. Light dorsum bordered by an irregular dark longitudinal stripe

dorsolaterally is typically for all specimens. Dorsal surface of head, body and limbs more or less spotted

in all specimens, concealed parts of hind limbs yellowish and dusted with minute flecks. Flanks in four
specimens lightly marbled below and more strongly marbled above; one specimen shows nearly

uniform brown flanks. Small whitish spots or strokes are conspicuous on almost all penultimate
phalanges near base of terminal disks. One specimen has a uni-coloured dark brown throat, others have

some lighter regions within dark coloration. Chest and anterior of belly with more isolated brown |
spots, posterior of belly without or with only a few spots. Ventral surfaces of limbs more or less |

unspotted. One specimen with a cross-like figure on the chest. An inconspicuous w-shaped mark in the

scapular region occurs in all specimens. Length of fingers and toes and size of terminal discs varies only |

a little, a “typical” state is shown on Fig. 18.

Etymology. Named for the Wapoga River headwaters where the type series was collected. Wapoga is consid-
ered as an invariable noun in apposition to the generic name.

Distribution. Known from the type locality in the Wapoga River headwaters, Irian Jaya, Indonesia. |
Based on morphology and structural features of vocalizations a population of frogs on Yapen Island N
appears to be closely allied to Oreophryne wapoga from the Wapoga River headwaters. We tentatively

recognise this population as O. wapoga pending further studies (see below).

270

Fig. 21. Male of Oreophryne cf. wapoga from Yapen Island (ZMB 62625).

Habitat and habits. Male Oreophryne wapoga called from heights of between 1.0 and 2.5 m on leaves
of understorey plants in lower-montane rainforest. All specimens were collected from a relatively dry
ridge, and none were heard calling in the extremely moist gullies at the Wapoga site. Males called
sporadically on clear, dry nights, and calling intensity increased during rain.

Vocalization. The advertisement call of O. wapoga has a similar basic structure to that of O. atrigularis;
a long series of notes in which the initial notes of a call series have relatively long inter-note intervals
followed by “fast” notes with much higher repetition rates. Note repetition rate of a call sequence may
decrease again at the end of the call sequence (Fig. 19). Notes consist of 3-7 pulses, and terminal pulses
are uttered at a higher rate than those at the beginning of the note (Fig. 20). 221 notes of two specimens
were recorded at an air temperature of 20.6 °C. Their mean length is 76 ms (SD 11.9), minimum length
49 ms and maximum length 104 ms. 135 intervals between “fast” notes have a mean duration of 179 ms
(SD 33.7), range 102-241 ms. Total call length varied from 10 to 31 s. The number of “fast” notes within
4 calls varies from 20-64. Fundamental frequency is around 1.5 kHz and dominant frequency is
between 2.5 and 3 kHz. Repetition rate in sequences of “fast” notes was between 4 and 6 notes/s.

Comparison with other species. O. wapoga is morphologically similar to O. atrigularis and differs from
other species of the genus in the same characters described in the “Comparison with other species”
section for that taxon. The morphological similarity of the Wondiwoi and Wapoga populations initially
led us to suspect that they might represent a single variable taxon. However, a number of consistent
differences in morphology and call structure lead us to recognise the two populations as distinct
species. The two taxa are of similar size although the SUL of O. wapoga is slightly and significantly
greater (mean 22.6 mm) than that that of O. atrigularis (mean 21.5 mm; t=3.5, P=0.00081). The head of
O. wapoga is longer than that of O. atrigularis; HL/SUL of the former 0.330-0.339, of the latter 0.278-
0.321, t=6.25, P=<0.0001; HL/HW for O. wapoga 0.948-0.974, for O. atrigularis 0.735-0.888, t=9.08,
P<0.0001; HW/SUL for O. wapoga 0.339-0.357, for O. atrigularis 0.333-0.404, t=3.96, P=0.00024; ED/SUL
for O. wapoga 0.122-0.131, for O. atrigularis 0.102-0.120, t=6.30, P<0.0001. Tibia length of ©. atrigularis
is slightly higher than that of O. wapoga; t=2.87, P=0.0038.

There are consistent differences in coloration between the two species. Throat colour of
O. atrigularis is uni-coloured black while that of O. wapoga is dark brown and speckled. Concealed parts
of the hind limbs are red in O. atrigularis, without minute dark spots; those of ©. wapoga are yellowish
and dusted with fine specks. Notes within advertisement calls of O. atrigularis are much longer (171-

271

262 ms vs 49-104 ms in O. wapoga), have a complicated substructure showing 3 different pulse groups,

and have a higher number of pulses (more than 20 vs 3-7 in O. wapoga). Internote intervals within fast
note sequences have a duration of 190-312 ms in O. atrigularis and of 102-241 ms in O. wapoga (t=18.6).
As a result mean note repetition rate in the very fast note sequences was about 3/s in O. atrigularis and
about 6/s in O. wapoga at a slightly lower temperature. The opposite trend would be expected if
temperature was solely responsible for differences in note repetition rate. Sequences of fast notes in
about 50 call series from O. atrigularis consisted nearly exclusively of less than 30 notes whereas 3 out
of 4 series of fast notes in O. wapoga contained more than 30 notes, and two of these had more than 60
notes.

Oreophryne cf. wapoga on Yapen Island

Eight frogs collected by the senior author about 15km north-east of Serui on Yapen Island closely

resemble O. wapoga and we tentatively assign them to this species. However we exclude them from the

type series because slight but consistent morphological differences raise some doubts about the

relationships of the two populations. Two males (ZMB 62159 and 62160) were collected on 10 Septem- |
ber 1999 and five males (ZMB 62622 and 62624-27) and one female (ZMB 62623) were collected on 18

and 19 May 2000 at an altidude of 610 to 630 m a.s.l. Selected body proportions are presented in table
4. Statistically the Yapen series differs significantly from Wapoga animals in the following characters:
mean snout-urostyle length in O. wapoga males 22.6 mm, in Yapen males 20.9 mm (t=3.61, P=0.0034);
mean ratio of head length /snout-urostyle length in ©. wapoga 0.334, in frogs from Yapen 0.310 (t=3.77,
P=0.0030); and mean ratio of head length/head width in O. wapoga 0.969 and in Yapen specimens 0.862
(t=4.09, P=0.0017). The only female from Yapen has a SUL of 22.5mm and has large whitish and
possibly immature ovarian oocytes that measure about 2 mm in diameter.

There are also differences in coloration between O. wapoga and wapoga-like frogs from Yapen. The |
blackish colour laterally on the head and on the throat is less intensive in most specimens from the
Yapen population, and in two specimens (ZMB 62159 and 62160) is missing entirely. However the |
coloration of some frogs (for example ZMB 62622 and 62627) is remarkably similar to that of a paratype |

(ZMB 63435) of O. wapoga. None of the eight Yapen frogs has a whitish mid-dorsal longitudinal line

(Fig. 21).

The advertisement calls of Yapen specimens consist of single creaks, small groups of creaks with |

comparatively long and often irregular inter-note intervals, and longer series of creaks. Notes consist

of 4 to 5 clearly defined pulses (Fig 22). Longer call series often start with notes having long internote- |
intervals, and in the course of the series intervals become shorter and more regular. Longer series have
a very similar structure and number of notes/s to those documented for O. wapoga. Some males called

from under leaf litter and others called from shrubs at heights between 0.30 and 1.50 m.

Both colouration and advertisement call structure distinguish the Yapen Island population from |
O. atrigularis. Although differences in morphology and possibly in calling behaviour between
O. wapoga from the type locality and wapoga-like frogs from Yapen Island doubtless exist, we believe

that the data currently available do not support recognition of the Yapen population as a distinct
species.

Tab. 4. Body proportions of a series of seven males and one female of Oreophryne cf. wapoga from Yapen Island. |

Ratio Mean SD Range
TE/SUE 0.53 0.019 0.48-0.53
HL/SUL 0.31 0.013 0.29-0.33
HW/SUL 0.36 0.018 0.34-0.39
HL/HW 0.86 0.057 0.74-0.93
ED/SUL 0.124 0.0052 0.116-0.132
END/SUL 0.095 0.0018 0.092-0.098
F3D/SUL 0.055 0.0049 0.047-0.060

DD,

ih Il) 11 ee T] ee) | Be) 7) ee en

0.5 1 S

Fig. 22. Oscillogram (above) and audiospectrogram (below) of a sequence of 11 “fast” notes of Oreophryne cf.
wapoga from Yapen Island. Air temperature during recording 22.5 °C.

Acknowledgements

RG would like to thank Mr Suyomo, head of Departemen Kehutanan, Kantor Wilayah Propinsi Irian Jaya for
permission to do field work and to collect some voucher specimens in the district of Nabire. Furthermore he is
very grateful to Dr. L. Ford (AMNH, American Museum of Natural History, New York), Dr. G. Köhler (SMF,
Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt/Main) and Dr. F. Tiedemann (NMW, Naturhis-
torisches Museum Wien) for lending material under their care. Financial support by the Deutsche Forschungs-
gemeinschaft is also appreciated. Thanks are due to M. Kapisa, G. Mareku, S. Marani, E. Konyorah and I. Tetzlaff
for their help during field work and V. Heinrich for preparing final copies of most figures.

SJR and DI are extremely grateful to Dr Arie Budiman of Puslitbang Zoology, and the head of Museum
Zoologicum Bogoriense — LIPI, Ibu Liliek Prijono, for their assistance in Jakarta and Bogor. This research was
part of Conservation International’s Rapid Assessment Program, and we greatly appreciate their assistance.
Funding support to SJR and DI was provided by the CI-USAID Cooperative Agreement # PCE-5554-A-00-4028-
00. We are especially grateful to Andy Mack, Yatna Supriatna, and Burke Burnett of CI for their support and
assistance during the survey, and to Mike Moore who helped to collect specimens. P.T. Freeport Indonesia
provided invaluable logistical support for which we are most grateful.

The authors are grateful also to Ibu Mumpun at MZB for her assistance.

References

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History) 208pp.

nella sotto-regione Austro-Malese. — Ann. Mis. Civ. Stor. Nat. Genova, ser. 1, 13: 323-450

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Jaya, Indonesia. — Raffles Bull. Zool. 48(2): 257-262

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65-132. Columbia, Missouri; Univ. Missouri Press

Tyler, M. J. 1964. Systematic position of the New Guinea frog Hylella wolterstorffi Werner. — Rec. South Aust. Mus.
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Wandolleck, B. 1911. Die Amphibien und Reptilien der papuanischen Ausbeute Dr. Schlaginhaufens. - Abh. Ber.

Königl. Zool. Anthropol.-Ethnogr. Mus. Dresden XIIK(6): 1-15

Werner, F. 1898. Vorläufige Mittheilung über die von Herrn Prof. F. Dahl im Bismarckarchipel gesammelten |

Reptilien und Batrachier. - Zool. Anz. 21: 552-556

-- 1901. Über Reptilien und Batrachier aus Ecuador und Neu-Guinea. II. Reptilien und Batrachier aus Deutsch-
Neu-Guinea. — Verh. Zool. bot. Ges. Wien 51: 602-614

Zweifel, R. G. 1956. Results of the Archbold Expeditions. No. 72. Microhylid Frogs from New Guinea, with
Descriptions of New Species. - Amer. Mus. Nov. 1766: 1-49

-- &M.]. Tyler 1982. Amphibia of New Guinea. - In Gressitt, J. L. (ed); Biogeography and ecology of New
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| SPIKIANA 275-286 ManenenolNevember2001 ISSN 0341-8391

A new monitor lizard from Panay Island, Philippines

(Reptilia, Sauria, Varanidae)*

Maren Gaulke & Eberhard Curio

Gaulke, M. & E. Curio (2001): A new monitor lizard from Panay Island, Philip-
pines (Reptilia, Sauria, Varanidae). — Spixiana 24/3: 275-286

Varanus mabitang, spec. nov. is described from Antique Province, Northwest
Panay, Philippines. The new species is closely related to V. olivaceus from South-
Luzon, Catanduanes, and Polillo Islands, Philippines, with which it shares the
general morphological appearance, the blunt teeth, a large caecum, and several
aspects of its highly specialized feeding habits. It differs from V. olivaceus by its
almost uniform black colouration, the different head shape with a slightly domed
snout and a strongly bulging nasal and temporal region, the finer scalation and
consequently higher standard scale counts, the tail with a triangular cross section
and a well developed double keeled scale row on its crest, the strongly keeled
ventrals, and an exclusively vegetarian diet at least in the holotype.

Maren Gaulke, PESCP, Bodenseestr. 300, D-81249 München, Germany

Eberhard Curio, PESCP, Ruhr-University Bochum, Faculty for Biology, Conser-
vation Biology Unit, D-44780 Bochum, Germany

Introduction

At present, the number of monitor lizard species occurring on the Philippine Islands is somewhat
unclear. Well known is the allopatric distribution of three different forms of the Varanus salvator group,
V. s. marmoratus, V. s. cumingi, and V. s. nuchalis, throughout the Philippines, and the occurrence of
V. olivaceus, an endemic species of the Luzon region. The type locality of V. rudicollis is given by Gray
(1845) with “Philippines”. The specimen was collected by H. Cuming, supposedly at Borongan on
Samar Island. However, as is known in the meantime, several of Cuming’s distribution records are
erroneous. No conclusive evidence that this species occurs on the Philippines has been forthcoming
since then, and it is widely assumed that the type locality given for V. rudicollis is erroneous (e.g. Taylor
1922, Bennett 1998). Other authors (Mertens 1959, Auffenberg 1976) list it as part of the Philippine
varanid fauna. Auffenberg (1988) mentions that it may be part of the Philippine fauna in spite of
statements to the contrary, referring to a preserved adult specimen (FSM) supposedly coming from the
Philippines. Auffenberg (1976) also lists V. dumerili for the Philippines, but corrects this as an erroneous
report later on (Auffenberg 1988). All monitor lizards known from the Philippines so far belong to the
large sized group, with a total length of 150 cm or above.

The recent discovery of another large monitor lizard (a specimen with a total length of 175 cm still
was considered as modestly large by hunters) on one of the biggest Philippine islands suggests that this
country may still hold some more monitors to be discovered in the future. In view of the highly
secretive life habits of V. mabitang, it is not too surprising that it escaped scientific discovery until

* Publication No. 41 of the Philippine Endemic Species Conservation Project (PESCP) of the Frankfurt Zoolog-
ical Society.

275

recently. To our present knowledge, this arboreal lizard is confined to some of the remaining rainforest
patches on Panay, where it obviously spends most of the time hiding in tree holes or on branches of
high forest trees. Its completely dark colouration further helps to disguise the shape of this animal
within its natural, shady and densely vegetated surrounding. The newly discovered species is most
closely related to V. olivaceus, with which it shares the general morphological appearance. Like
V. olivaceus, V. mabitang has slit-like nostrils, blunt teeth, very large feet, enlarged scales on the head,
and a large caecum.

Material and methods

About four years ago (in 1996), during the process of setting up a base for the work of the Philippine Endemic
Species Conservation Project (PESCP) on Panay, E. C. was informed by different hunters from the NW Panay
area about the existence of a black, very large, arboreal monitor lizard. This lizard was said to be very rare
compared to the West Visayan form of the water monitor (V. salvator nuchalis), which is widespread and common
all over Panay. All informations pertaining to this “mystery lizard” were collected. However, it took more than
three years until the first specimen of this lizard could be examined by project members. Since the project did
not hold a collecting permit for reptiles during that time, the animal was released after a non invasive
examination and picture taking.

To obtain data on the habitat, in March 2001 M. G. made a field trip to the area where the animal had been
caught. The trip was guided by the project member N. Paulino, a former hunter with a profound knowledge of
the flora and fauna within his range, the person whom we owe most of the information regarding the biology
of the new species.

Only when we were in the possession of a permit including reptiles (Gratuitous Permit No. 93), N. Paulino
was asked to secure a second specimen. It was finally caught on May 19%, 2001, and brought to the PESCP
research station at Sibaliw in the West of the NW Panay Peninsula. The animal was kept alive in a large outdoor
enclosure, until M. G. and the cooperating DENR member of PESCP, Mr. Nilo Subong from Kalibo, Aklan
Province, Panay, were able to go there. For observations, picture taking, and trials on its food choice, the animal
was kept inside the enclosure for another week (May 29% to June 4" 2001).

Description of pholidosis (using a magnifying lense) and colouration, and biometric measurements were
done immediately after death. For the first two days the animal was preserved in 7 % formalin, afterwards
transferred to 70 % alcohol. On June 23th, 2001 the specimen was exported to Germany (CITES Export Permit
No. 5571, issued by the Department of Environment and Natural Resources, Protected Areas and Wildlife
Bureau, Quezon City, and CITES Import Permit No. E: 1884/01, issued by the “Bundesamt für Naturschutz”,
Bonn) for further examination and description. The body cavity was opened to determine the reproductive
status, the development of the gastrointestinal tract, and its contents. For a closer examination of its dentition
the head was x-rayed with a Faxitron 805 (Field Emission Corp. Ore., USA) in ZSM, and to obtain cross-sections
without cutting, parts of the tail were moulded using “Palgat Plus” (ESPE Dental-Medizin GmbH & Co. KG,
Seefeld), and the form later on filled with silica rubber, to obtain an easy to cut positive. Isotopic analyses were
performed by the GeoBio-Center, Munich, with a Finnigan MAT Delta S. A Carlo Erba (EA 1108) elementar
analyser was coupled to the MAT by a Finnigan Conflo-Interface. ’N/“N ratios are given by elevational delta
notation (ö”N) versus air N, as relative standard.

For comparison, four V. olivaceus from ZFMK, Bonn, and two from PNM, Manila, were investigated. Not all
scale counts and measurements could be taken from all individuals, as some had scars covering large parts of
their bodies. For further comparison, data from Auffenberg (1988) were taken. He gives some scale counts and
measurements for more than 100 specimens of V. olivaceus, and very comprehensive data on the feeding habits
and biology of this monitor lizard.

Museum abbreviations: FSM: Florida State Museum, Gainesville; PNM: Philippine National Museum,
Manila; ZFMK: Zoologisches Forschungsinstitut und Museum A. Koenig, Bonn; ZSM: Zoologische Staats-
sammlung München.

276

Results and Discussion
Varanus mabitang, spec. nov.

Type. Holotype: PNM 7272, female, caught in the South Pandan Forest, ca. 250 m a. s. 1., Municipality of Pandan,
Antique Province, NW Panay Island, Philippines, on 19 May 2001 by Narciso Paulino. - Due to the scarcity of
this animal we collected no paratypes and we strongly advise to refrain from collecting further specimens at least
as long as its population status must be considered as critically endangered. However, we possess some
measurements, observations, and pictures from a second specimen, which actually was caught prior to the
holotype, end of October 2000 (same person and same locality as holotype), and released unharmed after
examination.

Diagnosis. V. mabitang can be distinguished from Varanus olivaceus Hallowell, 1856, as follows:

- Deorsal side black with scattering of tiny yellow dots on the posterior end of some scales of neck,
back, and extremities (vs. greenish gray with darker transverse bands across neck, back, and tail,
and extremities irregularly mottled yellowish-olive and gray);

- ventral surface of head, neck, tail, extremities, and belly dark gray to blackish (vs. grayish, grayish
green, or yellow-gray with 3-4 longitudinal brownish black to black stripes on throat);

- nuchal scales adjoining head scales smaller than these (vs. same size);

- extremely small scales on neck, body, and tail, and consequently very high standard scale counts:
scales from rictus to rictus 70 (vs. a maximum of 61, average 58,4), transverse rows of ventral scales
from gular fold to a theoretical line connecting the insertion of hindlegs ventrally 124 (vs. a
maximum of 121, average 109), transverse rows of dorsals from gular fold to a theoretical line
connecting the insertion of hindlimbs dorsally 138 (vs. a maximum of 122, average 112), Tab. 1;

- tail triangular in cross section, upper scale crest with a well defined, double, longitudinal keel (vs.
irregular oval in cross section, low double keel on tail hardly discernible), Fig. 1;

— head elongate, snout region slightly domed (vs. more massive with sloping snout region), Fig. 2;

- cranial table with well developed bulges above temporal regions (vs. flat);

- ventrals strongly keeled (vs. smooth or feebly keeled);

- scales of tail strongly keeled throughout entire length (vs. tail scales close to vent smooth), Fig. 3;

- exclusively vegetarian diet at least in the holotype (vs. a balanced molluscivorous-frugivorous diet).

Description of holotype (Figs. 4-7)

Habitus slender. Head forming a small, elongate triangle from above, with pointed and slightly
domed snout. Nostril closer to tip of snout than to eye. Narial opening slit-like, angled upward
posteriorly. Canthal ridge well developed between eye and nostril. Nasal region swollen, with a

Tab. 1. Scalation differences between V. mabitang and Varanus olivaceus. If no n is given, based on one individu-
um only. XVII. Scales from rictus to rictus in a straight line; XIX. Scales around midbody; XX. Transverse rows
of ventral scales from gular fold to a theoretical line connecting the insertion of hindlegs ventrally; XXI.
Transverse rows of dorsal scales from hind margin of head to gular fold; XXI. Transverse rows of dorsal scales
from gular fold to a theoretical line connecting the insertion of hindlegs dorsally.

characteristics Varanus mabitang, Varanus olivaceus, Auffenberg (1976) Auffenberg (1988)

holotype counts of specimens

in ZFMK & PNM

xVI 70 (n=2) 56.5 (n=-4) 51-61 48 58.4 (n=106) 50-61
XIX 212 179.4 (n=5) 165-200 186 186.1 (n=106) 169-214
xx 124 104 (n=5) 95-107 111 109 (n=106) 101-121
xxl 53 42.4 (n-5) 40-45
XxXH 138 112.2 (n=5) 105-122
Ventrals strongly keeled smooth (n=6) not mentioned feebly keeled (n=106)
Double crest high, well low, hardly discernible not mentioned low
on tail developed

Fig. 1. Cross sections of the tail of V. mabitang and V. olivaceus (both females of almost same size), showing shape
differences at one, two, and three head length behind vent (from left to right). The tail of V. mabitang (below)
has a triangular shape in cross section, while the tail of V. olivaceus (above) is irregular oval.

median, longitudinal concave groove. Parietal region with prominent bulges above temporal regions.
Two enlarged supraoculars left and right, distinctly longer than broad. Scale covering the pineal organ
slightly enlarged, roundish, black with a yellowish-whitish centre. Scales on dorsal surface of head flat,
relatively large, polygonal, largest in the intraorbital and parietal area. Each scale with six to eight
pustules (plaques sensu Auffenberg 1994). Scales on sides of head in temporal region very small, oval.
Scales on sides of head between eye and snout and below eyes enlarged, roundish to polygonal, with
several pustules on each.

There are 10 maxillary teeth in one half of the jaw, they are roundish in transverse section, blunt
posteriorly and conical anteriorly. The last two are very short, the following four are the longest, and
the first four again small. There are five roundish, conical premaxillary teeth. The thirteen dentary teeth
are round and blunt, gradually increasing in size posteriorly. Both in the alive and freshly dead lizard,
the dentary teeth do not extend beyond the gum, being visible only as almost translucent, flat and
round ovals within the gum.

Neck long and slender. Dorsal neck scales anteriorly roundish to ovally broadened, smaller than
adjoining head scales. Posterior nuchal scales elongate, very small, and high domed; surrounded by
wide interspaces with minute intercalary granules; each nuchal scale bearing a pore at its posterior
edge. Dorsal scales longish oval, surrounded by minute intercalary granules; smallest between front
limbs but not smaller than posterior nuchal scales, gradually increasing in size towards tail; keeled,
with keels becoming much more pronounced posteriorly. Scales on sides of dorsum smaller than along
the middle. Scales on neck and dorsum arranged in relatively regular transverse rows.

Scales below head enlarged at snout region, otherwise small; gular scales longish oval, arranged in
more or less regular transverse rows.

Ventral scales larger than dorsals, longish rectangular, strongly keeled, with a well defined pit
(external opening of a scale pore) on posterior part on each. On the border between ventrals and
dorsals, each ventral scale row divides into two dorsal scale rows.

Dorsal scales of limbs longish, partly rectangular, high domed on front limbs and keeled on hind
limbs, keels more pronounced on lower limb. Scales on undersurface of front- and hindlimbs roundish
and flat. Very large, long fore- and hindfeet, with long and slender toes and very long, strong, curved |
claws (Fig. 4).

Scales on tail rectangular, strongly keeled, arranged in irregular whorls; scales larger on ventral
than on dorsal side, the anterior ventral scale rows divide into two dorsal scale rows in irregular

Fig. 2. Profiles of V. mabitang (PNM 7272) and V. olivaceus (ZFMK 57589), showing the more pointed and slightly
upward turned snout of V. mabitang (above) and the sloping snout region of V. olivaceus (below). Both animals
are females of almost same size.

distances at the tail base, after 12 scale rows every second ventral row divides, and after about one third
of the tail, each ventral row divides into two dorsal rows. Upper crest of tail with prominent,
longitudinal double keeled scale row. Tail slender, triangular in cross section, with a sharp upper edge
defined by the double keel (Figs. 1 and 3).

Measurements. I. Total length: 1268mm; II. Snout vent length: 527 mm; III. Tail length from
cloaca to tip of tail: 741 mm, with some centimetres of tip missing; IV. Length of hindlimb from inner
insertion of hindlimb to end of longest toe without claw: 227 mm; V. Length of forelimb from inner
insertion of forelimb to end of longest toe without claw: 188 mm; VI. Length of head from tip of snout
to anterior margin of tympanum: 89.9mm; VII. Head width (maximum width between eyes and
tympanum): 43.85 mm; VIlla/VIIIb. Head height (above eye/maximum height between eyes and
tympanum): 32.90 mm/ 35.00 mm; IX. Distance from anterior margin of eye to middle of nostril:
27.20 mm; X. Distance from middle of nostril to tip of snout: 21.85 mm; XI. Distance from anterior

279

Er ch = y Fl

Fig. 3. Tail scalation of V. mabitang (above) and V. olivaceus (below) at one head length behind vent. Same
animals as Figs 1 and 2.

margin of tympanum to anterior margin of eye: 42.30 mm.

Proportion indices. XII. Relative tail length (III:II): 1.41 (several cm of tail are missing!); XII.
Position of nostril between tip of snout and eye ((IX:X): 1.25; XIV. Position of nostril to snout tip ([VI-
X1]:DX): 1,75; XV. Relative head length in relation to head width (VI:VII): 2.05; XVI. Relative head
length in relation to maximum head height (VI:VIIb): 2.57.

Scale counts. XVII. Scales from rictus to rictus in a straight line: 70; XVIII. Scales around tail base:
113; XIX. Scales around midbody: 212; XX. Transverse rows of ventral scales from gular fold to a
theoretical line connecting the insertion of hindlegs ventrally: 124; XXI. Transverse rows of dorsal
scales from hind margin of head to gular fold: 53; XXII. Transverse rows of dorsal scales from gular
fold to a theoretical line connecting the insertion of hindlegs dorsally: 138; XII. Scales around neck
before gular fold: 160; XXIV. Ventrals from tip of snout to gular fold: 117; XXV. Supralabials: 35.

Weight. 1850 g.

280

Colouration (in life). Dorsal and lateral sides of head black. Nuchal and dorsal region black, skin
between scales and some of the intercalary granules, especially on neck und anterior dorsum, partly
yellowish. Dorsal side of extremities black, with a tiny yellow dot at posterior end of scales (Fig. 5),
yellow dots most prominent on hindlimbs. Dorsal side of tail black. Ventral side of head, neck, body,
extremities, and tail anthrazite. The yellow colouration is only visible at a short distance, from a
distance of 2 m or more, the animal looks uniform black (Fig. 6). Claws dark grey; eyes reddish brown;
tongue pink.

Special anatomical features. V. mabitang possesses a large caecum, such as V. olivaceus. PNM 7272
contains ovarian follicles of a size between 5 and 7 mm, showing that it is mature.

Additional material. End of October 2000 a much larger specimen of V. mabitang was caught by Narciso Paulino
in the same area as PNM 7272 in the South Pandan Forest. This specimen was brought to our field station at
Sibaliw, NW-Panay Peninsula, for examination and picture taking, and released afterwards. Total length:
1750 mm; snout vent length: 640 mm; tail length: 1110 mm; weight: 5750 g; XII. 1.73; XVII. 70 (scale count taken
from pictures); no enlarged supraoculars.

The overall appearance is very similar to PNM 7272, but the habitus is more massive, and the bulges above the
temporal region much more pronounced (Fig. 7), with only a small longitudinal groove between both bulges on
the parietal region. The colour is black, with only very faint, tiny yellow dots on scales of front feet, and no
yellowish colouration on neck and back. Scales on head largest in parietal region. The body scales are very small,
with the anterior nuchal scales smaller than the posterior head scales, and the posterior nuchals same size as
smallest dorsals.

Etymology. The name mabitang is used for this species since generations within its range by the local people.
The meaning is somewhat like big monitor lizard (in Kinarayan dialect). The name is used as invariable noun
in apposition to the generic name.

Distribution. So far, V. mabitang is only known from forested areas of the NW-Panay Peninsula and
the Western Panay Mountain Range. The area belongs to the West Visayan region, one of the well
distinguished Philippine faunal regions (Heaney & Regalado 1998, de Jong & Treadaway 1993, Leviton
1963, and others), which are characterized by a high level of endemism. It just now starts to show that
Panay is an endemism centre on its own within this region. Recent faunal investigations revealed
several species new to science, which to present knowledge are endemic to Panay (e.g. Brown et al.
1997, Brown et al. 1999, Brown et al. in press, Ferner et al. 1997, Gaulke in press, Gonzales & Kennedy
1996). The closest relative of V. mabitang, V. olivaceus, is distributed over southeastern Luzon, Catan-
duanes and Polillo islands. The area belongs to another faunal region, the Luzon region.

The vertical distribution of V. mabitang within the Panay mountain range is still unknown. We
know from sightings that it occurs at least up to a height of 450m on the NW Panay peninsula.
Generally we assume that its vertical distribution is restricted by similar factors as in V. olivaceus. This
lizard occurs in heights up to about 400 m. Important temperature and moisture changes between 300
and 500 m are probably limiting many of the food plants of V. olivaceus to lower elevations, and
therefore limit its upper distribution (Auffenberg 1988).

Habitat and life habits. To our present knowledge, V. mabitang is a highly arboreal, secretive lizard
of the lowland rainforest.

Both specimens defecated seeds of the fruit of screw palms after capture. In captivity, different
kinds of forest tree fruit and land snails were offered to PNM 7272. While it fed on the fruit of two
species of screw palms (Pandanus spp.) and on the fruit of a fig tree (Ficus minahassae), it showed no
interest in the snails. Even when crawling directly in front of its snout, the sole reaction was short
tongue flicking. According to N. Paulino, V. mabitang does not eat any carnivorous food, only fruit and
leaves. He witnessed them eating fruit of different screw palms (Pandanus spp.), of a palm tree (Pinanga
sp.), of a fig tree (Ficus minahassae), and leaves from screw palms and a kind of shrub (local name is
“topsi”). When opening the stomachs of some individuals previously, he found seeds and leaf remains.

These observations are surprising, because no exclusively vegetarian monitor lizard is known so
far, and no monitor lizard feeding on leaves. To verify the observations, the stable isotope compositions
of nitrogen of PNM 7272 was analysed for scaling its diet and trophic level (see Schoeninger et al. 1997
for methodology). We analysed fruit and leaves of screw palms, fruit of fig trees, and also claw
material, different soft tissues, and contents of the gastrointestinal tract . The plants range at -0.1 to
1% 8"N, and the biomass of PNM 7272 is shifted by around 1.5-3.1%o 8"°N. This coincides with only

281

Ye,
DE

Fig. 6. V. mabitang looks uniformly black when seen from some distance.

one single step in a food chain (Fry 1988). The food ingested by this lizard has to be expected at near
0-1%o 8°N. Significant amounts of carnivorous diets can be excluded, because this should shift the
nitrogen isotope composition towards values near or above 6% ö"’N. A publication of the nitrogen
isotopic composition of V. mabitang (near 3%o ö8"N) and V. olivaceus (near 9%o ö”N) is in preparation
at the GeoBio-Center Munich.

The exclusively vegetarian diet is one of the most unusual features of PNM 7272. So far, V. olivaceus
was the only known varanid with frugivorous feeding habits. Auffenberg (1988) proposed for
V. olivaceus an evolutionary dietary shift from insectivory or faunivory to frugivory, which neccesitated
special adaptations in its gastrointestinal tract like the large caecum. So far V. olivaceus did not complete
this evolutionary shift, it is a molluscivorous-frugivorous animal, with both food components being of
equal importance for its nutrition. As main reason for its still partially faunivorous diet, Auffenberg
(1988) mentioned that fruit is an adequate source of carbohydrate, but is generally inadequate in

Fig. 7. Profile and hand of the first caught, large V. mabitang (175 cm), drawing by Helga Schulze.

protein. By feeding on land snails, in much less important percentages also on other animals like
insects, arachnids, and vertebrates, V. olivaceus compensates this lack in protein, and at the same time
adds calcium to its diet. Auffenberg (1988) refuted that V. olivaceus ever feeds on leaves as stated by
some of the local inhabitants. While dissecting more than 100 specimens he never found evidence of
leaves, nor did they take leaves in captivity or were observed feeding on leaves in the wild by any of
the project members.

From the isotopic measures we conclude that PNM 7272 is unique among all other varanids. The
nitrogen isotope measures inevitably fit with the herbivorous diets discussed. The general appearance
of PNM 7272, the well developed ovarian follicles, and the well developed fat bodies give no indication
that this specimen was suffering from malnourishment. Nitrogen isotope analyses of claw material and
muscle tissue of the folivorous-frugivorous sailfin lizard (Hydrosaurus pustulatus) from the same habitat
range at 1.5 to 1.9%o "N, and this is nearly identical to the ö"N level observed in V. mabitang. For the
mabitang one may presume a continuous evolutionary line, shifting from a mixed faunivorous-
frugivorous diet to a frugivorous-folivorous diet.

So far, we have data for one animal only. Future investigations will show whether the trend
towards an exclusively vegetarian diet has been reached in V. mabitang in general.

Both animals behaved very timid after capture. They never tried to defend themselves or showed
any sign of threatening behaviour typical for monitor lizards, like tail coiling and uncoiling, gular
extension, or hissing. Being held, they let extremities, tail, and head hang down. Most remarkable were
the extended periods of time, which both animals spend in complete tanatosis (no movement could be
observed for two consecutive days in PNM 7272). Letisimulation was never observed in V. olivaceus by
Auffenberg (1988), but is described for V. exanthematicus, an African monitor lizard, by Barbour (1926).
According to N. Paulino, feigning death is a common behaviour of the mabitang after capture.

Systematic relationships. V. mabitang is closely related to V. olivaceus, whose phylogenetic relations to
other varanids still is subject to discussion, in spite of all studies done on the latter (e.g. Auffenberg
1976, 1978, 1979a, 1979b, 1988). In 1962 the monotypic subgenus Philippinosaurus was erected for
V. grayi (now V. olivaceus) by Mertens, based on different skull features and the dentition. Auffenberg

284

‚ (1976) accepted the subgenus Philippinosaurus in his redescription of V. grayi, but later on (Auffenberg

1988) classified this species among the “slit-nosed” monitors, and as being closest to V. bengalensis, a
, member of the subgenus Empagusia. In a pers. comm. (cited in Böhme 1991) he later on stated that a
relation to V. salvator is also feasible. Based on the investigation of its hemipenial morphology, Böhme
‚ (1991) supported the idea of a rather isolated position (see also Ziegler & Böhme 1997). Based on the
investigation of DNA sequences, Fuller et al. (1998) place V. olivaceus in close relationship to V. prasinus,
a much smaller, arboreal monitor lizard from New Guinea, which was classified within the Australian
subgenus Odatria by Mertens (1942), but according to its hemipenial morphology belongs in the Asian
subgenus Euprepiosaurus (Böhme 1988). From this short overview, it is obvious that we still are far from
a satisfying resolution of the phylogenetic relationship of V. olivaceus, and therefore also of its close
relative V. mabitang. The results depend highly on the method used.

Acknowledgements

The work of the Philippine Endemic Species Conservation Project of the Frankfurt Zoological Society is
formalized under the aegis of a Memorandum of Agreement with the Department of Environment and Natural
Resources (Quezon City, Philippines), and the help of the Protected Areas and Wildlife Bureau (Director R.C.
Bayabos: Collecting Gratuitous Permit No. 93) and RED J. Amador (DENR Region VI) is gratefully acknowl-
edged. The project is sponsored by the Zoological Society Frankfurt, further by the ABC (Advocates of Bird
Conservation), AZ, B. Bacsal, Daimler-Chrysler Foundation, A. de Dios, Prof. Dr. Dr. mult. h. c. E. Mayr, Pentax,
H. Kessler v. Sprengeisen, Swiss Society for Bird Protection, and Vitakraft-Werke. Furthermore, M. Gaulke is
indebted to the Zoological Society Frankfurt for the funding of a first distributional survey on V. mabitang. Many
project members and people from the NW Panay area have part in the discovery of the new monitor lizard,
and/or in the investigation of the first, later on released animal. Foremost Mr. Narciso “Narsing” Paulino should
be mentioned, to whom we owe most of the information on the biology of this lizard, and who caught both
specimens. We acknowledge gratefully the help of Fel C. Cadiz, Arnold Demegillo, Tai Felimon, Eric Garrett,
Lucia Lastimoza, Stefan Luft, Enrique Sanchez, Benjamin “June” Tacud, Henry Urbina, and Vicente “Manong
Viseng” Hironimo, who in 1996 gave the first hints to the existence of the animal. Hans-Georg Horn helped
E. Curio in putting together all topically relevant monitor lizard information for a manual on the “mystery
lizard” reinforcing the research. We are grateful to Mr. Nilo Subong, CENRO Kalibo, who took interested part
in our investigations in the Philippines during all stages, and gave us the permit to preserve the holotype.

Thanks are due also to Wolfgang Böhme, ZFMK, Frank Glaw, ZSM, and to Rogelio Sison, PNM, for the loan
of specimens and/or the permission to examine specimens under their care. The x-ray was done by Marianne
Müller in the ZSM. Ulrich Struck and Alexander Altenbach, GeoBio-Center Munich, suggested and conducted
all isotope analyses in the laboratory of the Bavarian State Collection for Paleontology and Geology. Finally we
wish to thank Wolfgang Böhme, who critically revised the manuscript.

References

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-- 1979a. A monitor lizard in the Philippines. -— Oryx 15(1): 38-46

-- 1979b. The butaan: World’s rarest lizard? — Topic Jan.-Feb.: 14-16

-- 1988. Gray’s monitor lizard. — University Press of Florida, Gainesville, 419 pp.

-- 1994. The Bengal monitor. — University Press of Florida, Gainesville, 560 pp.

Barbour, T. 1926. Reptiles and amphibians. Their habits and adaptations. - Houghton-Mifflin and Co., Boston,
125 pp.

Bennett, D. 1998. Monitor lizards — Natural history, biology & husbandry. — Edition Chimaira, Frankfurt, 352 pp.

Böhme, W. 1988. Zur Genitalmorphologie der Sauria: funktionelle und stammesgeschichtliche Aspekte. - Bonn.
zool. Monogr. 27: 1-176

-- 1991. New findings on the hemipenial morphology of monitor lizards and their systematic implications. —
Mertensiella 2: 42-49

Brown, R. M., Leviton, A. E. & R. V. Sison 1999. Description of a new species of Pseudorabdion (Serpentes:
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7-12

-- ,--,J. W. Ferner & R. V. Sison (in press). A new snake of the genus Hologerrhum Günther (Reptilia;
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Brown, W. C., Brown, R. M. & A.C. Alcala 1997. Species of the hazelae group of Platymantis (Amphibia: Ranidae)

from the Philippines, with descriptions of two new species. — Proc. Cal. Acad. Sci. 49(11): 405-421

Ferner, J. W., Brown, R. M. & A. E. Greer 1997. A new genus and species of moist closed canopy forest skinks '

from the Philippines. - J. Herpetol. 31(2): 187-192

Fry, B. 1988. Food web structure on Georges Bank from stable C, N, and S isotopic compositions. — Limnol.
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Fuller, S., Baverstock, P. & D. King 1998. Biogeographic origins of Goannas (Varanidae): a molecular perspec-
tive. - Molecular Phylogenetics Evolution 9(2): 294-307

Gaulke, M. (in press). A new species of Lycodon from Panay Island, Philippines (Reptilia, Serpentes, Colubr-
idae). - Spixiana 25

Gonzales, P. C. & R. S. Kennedy 1996. A new species of Crateromys (Rodentia: Muridae) from Panay, Philip- |

pines. — J. Mammalogy 77(1): 25-40

Gray, J. 1845. Cat. Liz. Brit. Mus.: 10.

Heaney, L. R. & J. C. Regalado 1998. Vanishing treasures of the Philippine rain forest. - The Field Museum,
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de Jong, R. & C. G. Treadaway 1993. The Hesperiidae (Lepidoptera) of the Philippines. - Zool. Verhand., Leiden
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Leviton, A. E. 1963. Remarks on the zoogeography of Philippine terrestrial snakes. - Proc. Cal. Acad. Sci. 4" ser.
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Taylor, E. H. 1922. The lizards of the Philippine Islands. - Phil. J. Sci. Monogr. 17: 1-269

Ziegler, T. & W. Böhme 1997. Genitalstrukturen und Paarungsbiologie bei squamaten Reptilien, speziell den
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286

‚ Buchbesprechungen

37. Mebs, D.: Gifttiere - Ein Handbuch für Biologen, Toxikologen, Ärzte und Apotheker. - Wissenschaftliche
Verlagsgesellschaft GmbH, Stuttgart, 2000, 350 S., 2. neu bearbeitete Auflage mit 320 meist vierfarbigen Abb.
ISBN 3-8047-1639-3.

Nach dem Erscheinen der ersten Auflage dieses Buches, das allgemein großes Interesse fand, hat sich das Heer
der “Gifttiere” erweitert. So wurden vom Autor bis dahin unbekannte oder unbeachtete Tiergruppen, wie
Giftvögel oder unter den Säugetieren die ursprünglichen Insektenfresser, ebenso neu aufgenommen, wie die
Vergiftungserscheinungen durch den Verzehr von Haifleisch, besonders deren Leber. Letztere Vergiftungen
stehen der vielfach als Fischvergiftung bezeichneten Ciguatera sehr nahe, die durch mikroskopisch kleine,
einzellige Geißeltierchen (Dinoflagellaten) hervorgerufen wird. Unter den passiven Vergiftungen, die durch den
Verzehr von Meerestieren hervorgerufen werden (z.B. Muschelvergiftungen) spielen die Toxine dieser winzigen
Meeresbewohner eine herausragende Rolle. Außerdem werden in dem durch zahlreiche charakterisierende
Farbabbildungen der entsprechenden Gifttiere sehr anschaulichen Buch die aktiven Giftwirkungen der Meeres-
und Landtiere vorgestellt. Dabei wird jede Tiergruppe, von den Schwämmen bis zu den Giftschlangen, ausführ-
lich behandelt, wobei jedoch einschränkend hinzugefügt werden muß, daß nicht jedes der Tiere besonders
erwähnt werden kann. Neben der Kurzcharakteristik der Merkmale, der Verbreitung und des Lebensraumes
bzw. der Lebensweise der Einzelarten wird die Tiergruppe als Einheit vorgestellt. Ebenso werden die Vergif-
tungsumstände selbst, die Vorsichtsmaßnahmen, der Giftapparat, der Vergiftungsverlauf und die möglichen
Erste-Hilfe-Maßnahmen beschrieben. Diesen vielfach fachlich medizinischen Dokumentationen werden oft
Fallbeispiele beigegeben, in denen sowohl überlebte wie auch tödlich verlaufende Vergiftungen detailliert
beschrieben werden. Jedem der Tiergruppenkapitel ist ein umfangreiches Literaturverzeichnis angehängt. Eine
Zusammenfassung der ‘Grundlagen und Hinweise’ wird der Zusammenschau der Tiergruppen vorangestellt,
in der die Gifttiere allgemein beschrieben werden, ebenso wie die Toxine, die Bedeutung dieser für die
Pharmazie. Besonders eindrücklich sind die Beschreibungen, wie es zum Kontakt mit Gifttieren kommt, und die
Ratgeber zu Maßnahmen bei einer Vergiftung. Auf drastische Abbildungen von Körperreaktionen auf Biß- oder
Stichverletzungen wurde verzichtet. Viele altertümliche und traditionelle Behandlungsmaßnahmen, wie etwa
das Aufschneiden von Schlangenbissen oder das Aussaugen der Bißwunden, werden kritisch unter die Lupe
genommen. Dieses sehr informative Buch mit seiner Bebilderung spricht die im Titel aufgeführte Interessenten-
gruppe sicher besonders an. E. G. Burmeister

38. Sternberg, K. & R. Buchwald (Hrsg.): Die Libellen Baden-Württembergs; Band 1: Allgemeiner Teil, Kleinli-
bellen (Zygoptera). - Eugen Ulmer Verlag Stuttgart, 1999, 712 S., 241 Farbfotos, 49 Diagramme und Zeich-
nungen, 29 Verbreitungskarten und 21 Tab. ISBN 3-8001-3514-0.

Libellen gehören neben Schmetterlingen und Käfern zu den auffälligsten auch allgemein bekannten Insekten,
die auf Grund ihrer geringen Artenzahl in Deutschland auch wissenschaftlich besonders gut bearbeitet sind.
Diesem Umstand verdankt das vorliegende Buch seine herausragende Dokumentationsweise. In einzigartiger
Weise haben die beiden Herausgeber, vorzügliche Kenner der europäischen Libellenfauna, mit zahlreichen
Kollegen alles bisher Bekannte zu dieser Insektengruppe zusammengetragen und ein Standardwerk geschaffen,
das nicht nur den Raum Baden-Württemberg erfaßt, sondern allgemein die Biologie, Faunistik und Ökologie
dieser faszinierenden Fluginsekten mit ihren bizarren aquatischen Larven präsentiert. Neben den verschieden-
sten Aspekten, die auch die Namensgebung, die Systematik und Evolution und das Verhältnis Libelle - Mensch
beleuchten, werden die 26 Kleinlibellenarten der insgesamt 75 Arten der Fauna Baden-Württembergs vorge-
stellt. Diese artlichen Dokumentationen enthalten neben den Verbreitungsangaben die Phänologie, Lebens-
raumansprüche, Larval- und Imaginalhabitate sowie die Biologie der beiden Stadien, Parasiten und, was
besonders wichtig erscheint, die Einnischungsstrategien der nächstverwandten Arten. Diese Kapitel zu den
einzelnen Arten schließen Anmerkungen zur Gefährdung und zum Schutz ein, wobei auch die Pflege und der
Schutz der Lebensräume besonders Erwähnung finden. Bei der Darstellung der Verbreitung ist die ausschließ-
liche Verwendung der Rasterpunktkarten antiquiert, da sie keine Aussagen zur lokalen Verbreitungsmustern
zulassen. Hier scheint die Erfassung der bayerischen Libellen einen besseren Weg gegangen zu sein. Dennoch
zeugt dieses Buch vom umfangreichen Wissen der Bearbeiter, das auch im Text umgesetzt wurde. Auf Grund
fehlgeleiteter Naturschutzaktivitäten scheint der zweite Band, der die Großlibellen (Anisoptera) dokumentieren
sollte, in Frage zu stehen, was ganz besonders bedauerlich wäre, zumal das Bundesland Baden-Württemberg
sich durch die Unterstützung dieser besonders gelungenen Reihe zur Fauna auszeichnete.

E.-G. Burmeister

287

Buchbesprechungen

39. Thenius, E.: Lebende Fossilien - Oldtimer der Tier und Pflanzenwelt, Zeugen der Vorzeit. — Verlag Dr.
Friedrich Pfeil, 2000, 228 S. ISBN 3-931516-70-9.

Heute lebende Organismen, die aus einer fernen Zeit zu kommen scheinen und auch in täuschend ähnlicher
Form als Fossilien heute auf Steinplatten freipräpariert zu bestaunen sind, haben in allen Zeiten nicht nur
Wissenschaftler fasziniert, sondern auch das Interesse eines breiten Publikums geweckt und die Phantasie
angeregt. Wie konnten sich über Jahrmillionen gegenüber einer sich beständig ändernden Umwelt diese Formen
erhalten? Hatten sie eine sogenannte Nische gefunden, in der sie sich behaupten konnten im Gegensatz zu ihren
nächsten Verwandten? Sicher sind diese Pflanzen und Tiere meist sehr selten und nur auf kleine Regionen
beschränkt, aber ihr Überleben dokumentiert eine erfolgreiche Strategie, über die vielfach noch sehr wenig
bekannt ist. Meist sind sie selten, sieht man etwa vom Spinnentier “Pfeilschwanzkrebs” Limulus polyphemus ab,
der kaum verändert im Vergleich zum Mesolimulus walchi der Solnhofener Plattenkalke erscheint, und auch
heute noch zur Laichzeit an der nordamerikanischen Ostküste in großer Zahl zusammengeschaufelt und als
Hühnerzusatzfutter verwendet wird. Der aus China stammende Ginkgo-Baum wird inzwischen in Fußgänger-
zonen der Städte betonumsäumt gepflanzt. Schicksal Lebender Fossilien? Sicher Ausnahmen, denn wir betrach-
ten mit Erfurcht die Zeugen der Vergangenheit.

Das vorliegende Buch des bekannten Palaeontologen Erich Thenius vermindert nicht diese Ehrfurcht, macht
aber die zahlreichen behandelten Organismen zugänglich, d.h. ihr Werdegang in der Stammesgeschichte wird
beleuchtet und anhand anschaulicher Grafiken präsentiert. Nach einer Begriffsbestimmung “Lebender Fossili-
en”, die nicht einheitlich ausfallen kann, werden von den Archaebakterien bis zu den Säugetieren Vertreter
vorgestellt, die als Zeitzeugen fungieren, aber auch solche, die als einzige Überlebende einer großen Gruppe
noch existieren, aber stark abgeleitet sind. Hier stößt der Begriff “Lebendes Fossil” an seine Grenzen. Die
abschließende Übersicht zum System der Organismen mit den behandelten Aspiranten für den Titel Lebendes
Fossil, aber auch mit ausgestorbenen Vertretern, erleichtert die Zuordnung. Das umfangreiche Literaturver-
zeichnis ist vorzüglich geeignet, die Erfahrung mit diesen außergewöhnlichen Objekten der Botanik wie Zoo-
logie fortzusetzen. Neben dem Autor gilt auch dem Verlag besonderer Dank für diese Zusammenfassung, die
nicht nur Wissenschaftlern der Biologie und Palaeontologie, sondern auch einer breiten interessierten Öffent-
lichkeit einen Einblick in die Vielfalt des Lebens und Überlebens eröffnet. E.-G. Burmeister

40. Gorissen, I.: Die großen Hochmoore und Heidelandschaften in Mitteleuropa - Natur — Landschaft -
Naturschutz. - Selbstverlag Igmar Gorissen, Siegburg, 1998, 190 S., 141 Abb., 7 Tab. ISBN 3-00-003890-6.

Natürliche offene Landschaften wie Hochmoore und Heiden sind inzwischen sehr selten geworden und
verdienen besonderen Schutz. Die in diesem Buch beschriebenen 79 Areale des mitteleuropäischen Raumes, der
den verschiedenen biogeographischen Einflußzonen von borealen bis zu mediterranen Elementen unterliegt,
umfassen vielfach Großlebensräume, die besonders gefährdet sind. Ein großer Teil dieser Gebiete war durch
militärische Nutzungen “geschützt”, deren Wegfall sich nachhaltig in Flora und Fauna wiederspiegelt. Der
Autor hat von Belgien bis Ostpolen Informationen zusammengetragen und zu jedem der Gebiete eine Struktur-
analyse, die Bedeutung für die Vogelwelt im europäischen Rahmen, die jeweilige Größe, die Entwicklung und
Nutzung sowie die Probleme des Gebietes zusammengetragen.

Einige der Moore und Heideflächen hätten vermutlich eine detailliertere floristisch und faunistische Be-
handlung verdient, zumal hierzu eine Fülle von Literatur vorliegt, die jedoch nicht berücksichtigt wurde.
Bedauerlicherweise werden auch einige Arten angegeben, ohne zu prüfen, ob es sich dabei um Invasionisten
oder um Besiedler von Rückzugsarealen handelt. Neben den Gebietscharakteristiken werden auch Areale
außerhalb Mitteleuropas, wie in der Ukraine, in ihrer Bedeutung vorgestellt. Insgesamt ist diese Zusammenstel-
lung eine Informationsquelle für Landschaftspfleger, wobei hier deutlich wird, daß diese Lebensräume Unikate
sind und nach einer Degradierung unwiederbringlich verloren sind. Eine Ersatzlebensraumschaffung wird
dadurch ad absurdum geführt. Darum sollten auch die im behördlichen Naturschutz tätigen Personen die in
ihrem Zuständigkeitsbereich liegenden Moore und Heiden mit ihrer Sukzession, die in diesem Buch dokumen-
tiert sind, ganz besonders im Auge behalten, was aber nicht zu einer Naturschutzkäseglocke über den Gebieten
führen darf. E.-G. Burmeister

288

SPIXIANA - Zeitschrift für Zoologie SPIXIANA - Journal of Zoology
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SPIXIANA 193-288 München, 01. November 2001 ISSN 0341-8391

INHALT - CONTENTS

Hausmann, A. & R. Trusch (ed.): Proceedings of tne FORUM HERBULOT 2001 Neotrop-
ical Geometridae: Approaches to a Modern Concept of the Geo-
metrid System on Genus and Tribe Level (8.3.-9.3.2001) ..............

Giachino, P. M.: New data on Anillina of the Oriental Region (Insecta, Coleoptera,
Garabidae, ‚BembidiiNi)i.....::. a...

Horstmann, K.: Revision der bisher zu /Iselix Förster gestellten westpaläarktischen
Arten von Phygadeuon Gravenhorst (Insecta, Hymenoptera, Ichneu-
monidae,. Enyptinae)inaze-sceseeareeesnrenssregene nee en

Leistikow, A.: Araucoscia Verhoeff, 1939 is a juniour synonym of Pseudophiloscia
Budde-Lund, 1904 (Crustacea, Isopoda, Oniscidea) ......................-

Weigmann, G.: Contribution to the taxonomy of European Poronota I. Oribatella and
Anachipteria (Acari, Oribatida) ........................... en.

Hausmann, A. & M. Sommerer: Oenospila kopperi, spec. nov., eine neue grüne Geome-
tride aus Sumatra (Insecta, Lepidoptera, Geometridae, Geomettri-

Kreipl, K.& A. Al: A new Oocorys from Western Australia (Mollusca, Gastropoda,
ERIC) er ER passbancbenocacgene

Glaw, F., M. Vences & K. Schmidt: A new species of Paroedura Günther from northern
Madagascar (Reptilia, Squamata, Gekkonidae) ...............................

Günther, R., S. J. Richards & D. Iskandar: Two new species of the genus Oreophryne from
Irran Jaya, Indonesia (Amphibia, Anura, Microhylidae) ....................

Gaulke, M. & E. Curio: A new monitor lizard from Panay Island, Philippines (Reptilia,
Sauna, Varanidae)e... sn ann... oalen Bellere

Seite

193-202

203-206

207-229

231-233

235-240

241-244

245-247

249-256

Büichbesprechumngem en en... m eaken...... ..aeeer MeenneeBennun een reeReRRer. 230, 234, 248, 287-288

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