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(Copepoda, Poecilostomatoida, Splanchnotrophidae) Ulrike Haumayr & Michael Schrödl

Haumayr, U. & M. Schrödl (2003): Revision of the endoparasitic copepod genus Ismaila Bergh, 1867, with description of eight new species (Copepoda, Poecilosto- matoida, Splanchnotrophidae). Spixiana 26/1: 1-33

The genus Ismaila Bergh, 1867 was a poorly known group of endoparasitic copepods associated with shell-less opisthobranchiate hosts. Descriptions were limited to the gross body shape or, where given, details did not agree regarding the number and identity of cephalic and body appendages. The present study gives a complete revison of the known species of the genus Ismaila, including the descrip- tion of eight new species (I. obtusa, spec. nov., 1. jenseniana, spec. nov., I. androphila, spec. nov., 1. aliena, spec. nov., I. damnosa, spec. nov., 1. robusta, spec. nov., 1. socialis, spec. nov., I. magellanica, spec. nov.). The parasites‘ morphology is described in detail based on SEM examinations. All Ismaila species studied possess 5 pairs of highly complex cephalic appendages which are two pairs of antennae, one pair of sickle-shaped, hairy mandibles, and 2 pairs of special-shaped maxillae, all of which showing surprisingly little intraspecific and interspecific variation. Differences between the congeners are mainly related to body proportions (stocky vs. delicate body), shape and proportions of thoracopods and dorsal body processes, and to the shape of the egg-sacs. All Ismaila species appear to be host-specific, showing special biological adaptations to the host species such as different site preferences and sex ratios. While most species do not obviously damage their hosts, I. damnosa, spec. nov. usually sterilize their hosts, the aeolidoidean nudibranch Flabellina sp. 1.

The homology of Ismaila body structures, such as cephalic, thoracic and abdom- inal segments and appendages, is discussed showing that the aberrant morphology of Ismaila can be plausibly related to a general copepod bauplan. A set of potential autapomorphies, e.g. the presence of an unique unpair dorsal process, strongly suggests the monophyly of the genus Ismaila.

Ulrike Haumayr and Michael Schrödl, Zoologische Staatssammlung München, Münchhausenstr. 21, D-81247 München, Germany; e-mail: schroedl@zi.biologie.uni-muenchen.de.

Introduction

Copepods of the Poecilostomatoida Thorell, 1859, which are characterized by a sickle-shaped mandi- ble, generally parasitize a variety of marine fishes and invertebrates. The Splanchnotrophidae are high- Iy specialized and aberrantly shaped parasites of shelless opisthobranchs (see review by Jensen 1987). Most recently, the family has been critically revised

and reorganized by Huys (2001). A new family Micrallectidae Huys, 2001 was established to com- prise the genus Micrallecto Stock, 1971 and its junior synonym Nannallecto Stock, 1973, ectoparasites of pteropod Gymnosomata which previously were dis- cussed to belong to the Splanchnotrophidae (Ho 1981, Belcik 1981, Jensen 1987). Of the four endopar- asitic genera traditionally placed into the Splanch- notrophidae, Huys (2001) transferred Briarella Bergh,

1876, and, with some reservations, Chondrocarpus Basset-Smith, 1903, to the Philoblennidae, formerly only known as ectoparasites of prosobranch gastro- pods in the Far East. The genus Splanchnotrophus Hancock & Norman, 1863 was divided into Splanch- notrophus s.s. and Lomanoticola Scott & Scott, 1895. In addition, two new monotypic genera, Arthurius Huys, 2001, and Ceratosomicola Huys, 2001, were estab- lished with S. elysiae Jensen, 1990 and S. sacculatus O’Donoghue, 1924 as the type species, respectively. The placement of the genus Ismaila Bergh, 1867 with- in the Splanchnotrophidae was confirmed by Huys (2001). Based on the re-examination of the antennae and mouthparts of I. belciki Ho, 1987, Huys (2001) also briefly discussed the mandibular morphology within the Splanchnotrophidae and presented an hypothesis on the evolution and historical distribu- tion of Ismaila.

However, morphological knowledge of most Splanchnotrophus (s.l.) and Ismaila species is still lim- ited to the gross body shape from few female spec- imens examined. In particular, taxonomically im- portant structures such as antennae, mouthparts or leg structures are either unknown or may differ considerably even between congeners (e.g. Belcik 1981 vs. Ho 1981). Dwarf males have been rarely studied at all. Hence, the taxonomy of Splanchno- trophus (s.l.) and Ismaila still can be considered as being tentative with a strong need of revision on the basis of detailed structural data. Since segment lim- its are hardly recognizable in splanchnotrophids, it is also difficult to draw conclusions on the homolo- gy of certain body appendages and body portions with corresponding structures of a general copepod or crustacean bauplan.

Within the genus Ismaila, only three species were described. Ismaila monstrosa Bergh, 1867, the type species, seemed to have an extremely wide geo- graphical distribution and low host specifity. The original description was based on a single specimen (Bergh 1867) found at St. Thomas, Virgin Islands, Caribbean Sea in the aeolid nudibranch host Phidi- ana lynceus Bergh, 1967. Jensen (1987) mentioned a further specimen of I. monstrosa which was found at the same locality and in the same host species in 1897. Two parasites from the Chilean Pacific coast found in two different hosts, Archidoris incerta Bergh, 1898 (Nudibranchia: Doridoidea) and Aeolidia papil- losa serotina Bergh, 1873 (Nudibranchia: “Aeolidio- dea”), were also assigned to I. monstrosa by Bergh (1898), who, however, mentioned slight differences to the Caribbean specimens. Belcik (1981) described further specimens of Ismaila monstrosa from the Cal- ifornian Janolus fuscus O’Donoghue, 1924 (Nudi- branchia: Arminoidea), which Ho (1987a) consid- ered to be a distinct species, Ismaila belciki Ho, 1987.

Finally, Jensen (1987) assigned a few parasites found at St. Thomas, Virgin Islands in Ercolania funerea Costa, 1867 (Sacoglossa) to I. monstrosa.

In a very detailed light-microscopical study, Ho (1981) described the third species, Ismaila occulta Ho, 1981 from the Californian nudibranch Dendrono- tus iris Cooper, and was the first to add ontogenetic data from larval stages (Ho 1987b).

More recently, endoparasitic Ismaila species have been reported from a variety of Chilean opistho- branchs (Millen et. al. 1994, Schrödl 1996, 1997, 2002). These findings induced the present study, which 1) describes the parasite morphology in de- tail using SEM, 2) discusses the homology of Ismai- la body structures, 3) gives a complete revison of the known species of the genus Ismaila Bergh, 1867, including the description of eight new species.

Material and Methods

From 1991 to 1995, 42 species of nudibranch and saco- glossan Opisthobranchia have been collected along the Chilean coast (Millen et al. 1994, Schrödl 1996, 1997). Eight of these species were infected with endoparasites of the genus Ismaila (see Schrödl, 2002). Parasites of five nudibranch species, Okenia luna Millen et al., 1994; Thec- acera darwini Pruvot-Fol, 1950; Flabellina sp. 1; Phidiana lottini (Lesson, 1831); Aeolidia papillosa serotina Bergh, 1873, and the sacoglossan Elysia patagonica Muniain & Ortea, 1997 have been examined in this study. In addi- tion, with the courtesy and support of Dr. Francis Belcik, Dr. Charles Coleman, Dr. Frank Ferrari and Dr. Kathe Jensen, it was possible to re-examine specimens of T. occulta, I. belciki and I. monstrosa. The type of I. mons- trosa and Chilean material assigned to I. monstrosa by Bergh (1898) have also been examined. For the first time, specimens of the genus Ismaila have been studied with the aid of SEM. This kind of examination is suitable to identify and document even very fine structures and, thus, to critically challenge bibliographic light-micro- scopical data.

Class Copepoda H. M. Edwards, 1840 Order Poecilostomatoida Thorell, 1859 Family Splanchnotrophidae Norman & Scott, 1906

Genus Ismaila Bergh, 1867

Diagnosis (see Figs 1,2; for details see description of I. androphila, spec. nov., and I. aliena, spec. nov.: Figs 17,18).

Female

Body elongate, and either delicate or stout. Ce- phalothorax distinctly set off from trunk, consisting of five head-segments and the first thoracic seg-

Fig. 1. Morphology of Ismaila spp. by SEM (specimens critical point dried, gold coated). A. ? Ismaila androphila, spec. nov., ventral view. B. d Ismaila aliena, spec. nov., ventral view. C. Detail of mouthparts (Ismaila aliena, spec. nov.). Note the triangular labrum (1), biramous, setigerous maxillules (2) and maxilles (3), the tongue-shaped labium (4), and the tips of the large, sickle-like second antennae (5).

Fig. 2. Nudibranch hosts of Ismaila spp. A. Okenia luna (lateral and ventral view), with parasite egg-sacs (arrows) protruding under the notal rim. B. Thecacera darwini (lateral view), with parasite egg-sacs protruding behind the gill circle (arrows). C. Flabellina sp. 1 (lateral view) with parasite egg-sacs between cerata (arrow).

ment, no external segmentation detectable. Head with two ventral protrusions framing the mouth- parts. Thorax with remaining five segments (no su- perficial segmentation). Abdomen with three exter- nally detectable segments.

Cephalothorax contains five pairs of head-ex- tremities: antennule (Figs 1C, 17A): unbranched and 2-segmented, distal limb with terminal hairs. An- tenna (Figs IC, 17B): unbranched and 3-segmented, distal segment formed as large hook. Labrum (Fig. IC): triangular and scalloped. Mandible (Figs 1C, 17C): biramous, with one atrophied ramus and a second long, sickle-shaped and pointed ramus (or,

as interpreted by Huys (2001), “gnathobase bearing one stylet-like and several short teeth”). Maxillule (Figs 1C, 17D): inwards-bent lobes, terminal part biramous; rami equally short, with short hairs (="two setae” according to Huys (2001). Maxilla (Figs 1C, 17E): 3-segmented, second limb bears 1) an additional limb with long elongate outgrows, and 2) two pointed, hairy processes (= “unarmed syn- coxa with allobasis drawn into multipinnate endite with 2 accessory elements” cf. Huys, 2001). Labium (Fig. 1C): hairy, with paragnath lobes. First thoraco- pod (= maxilliped): absent.

Thorax, ventral: second thoracopod (Fig. 18A):

Fig. 3. Ismaila monstrosa Bergh, 2. A. Holotype (specimen collected in 1867), dorsal view. B. Holotype, ventral view. C. Specimen collected in 1897, dorsal view. D. Specimen collected in 1897, ventral view.

biramous; exopodit either conical or flattened, with a rounded tip, endopodit conical with a terminal claw. Third thoracopod (Fig. 18B): 3-branched; ex- opodit either conical or flattened, with a rounded tip, endopodit conical with a terminal claw, en- dopodit bears one basal process. Second and third thoracopods long, extending considerably beyond the body laterally (see also Jensen 1987). Fourth

thoracopod (Fig. 18C): rudimentary, uniramous. Fifth thoracopod (Fig. 18D): two very small, rudi- mentary and pointed processes, arise either togeth- er from one common base or are separate but close to each other. Between fourth and fifth pair of tho- racopods: sclerotized ring. Sixth thoracopod: ab- sent.

Thorax, dorsal: three pairs of long uniramous,

20 um

50 um

20 um

10 um

D

Fig. 4. Ismaila monstrosa Bergh, cephalic appendages of female collected in 1897. A. Antennule (left). B. Antenna (right). C. Mandible (right). D. Maxillule (right). E. Maxilla (left).

conical or flattened processes. One single medial dorsal process between third pair of dorsal process- es (Fig. 16B). An additional pair of dorsolateral proc- esses (Fig. 6) may be present inserting between sec- ond dorsal processes and second thoracopods.

Abdomen: first segment, externally visible: bears genital openings, mature specimens usually with a pair of straight, curved or coiled egg sacs (e.g. Fig. 16).

Third, externally visible segment: pair of caudal rami with basal hairs.

Male (dwarf)

Body pear-shaped with, depending on maturi- ty, strongly enlarged cephalothorax (head plus first and second, strongly swollen thoracic segments). Thorax with four segments (no external segmenta- tion detectable). Abdomen with three externally de- tectable segments (Fig. 1B).

Cephalothorax: indistinctly set off from trunk. Cephalic extremities as in female. First thoracopod: absent. Second thoracopod (Fig. 18E): uniramous (exopoditreduced), long, conical, with terminal claw.

Thorax, ventral: third thoracopod (Fig. 18F): biramous (exopodit reduced); endopodit long, with terminal claw; with shorter inner process with blunt

tip. Fourth thoracopod (Fig. 18G): rudimentary, unbranched. Fifth thoracopod (Fig. 18H): very small, rudimentary processes, arising either together from a common base or being separate. Sclerotized ring (Fig. 18I) located between fourth and fifth pairs of thoracopods. Sixth thoracopod (Fig. 18I): three setae on elongated genital lobes (see Ho 1981).

Thorax, dorsal: without processes.

Abdomen: first segment with genital openings. Pair of caudal rami with basal hairs (Fig. 181).

Type species: Ismaila monstrosa Bergh, 1867: 97-130, Tab. II+IV.

Ismaila monstrosa Bergh, 1867 Figs 3, 4

Ismaila monstrosa Bergh, 1867: 97-130, Tab. III +VI; Hecht 1895: 625, 630 (partim); Belcik 1981: 16-25 (partim); Ho 1981: 130-136 (partim); Ho 1987a: 67-83 (partim); Ho 1987b: 109-111 (partim); Jensen 1987: 75-84 (partim); Jensen 1990:291-296 (partim).

Types. Holotype: 2, Zoological Museum Copenhagen (ZMUC,), collected at St.Thomas, Virgin Islands, Carib-

bean Sea, in 1867. Host: Phidiana Iynceus Bergh, 1867. Examined under stereomicroscope.

Additional material: 19, collected at St.Thomas, Virgin Islands, Caribbean Sea, in 1897. Host: Phidiana Iynceus Bergh, 1867. Examined by SEM.

Description of holotype (?) (Figs 3A,B)

Elongate and delicate body, measuring 3.2 mm in length. Head severely damaged, antennule and antenna present. Two pairs of thoracopods (2"‘ and 3") macroscopically detectable. Second thoracopod: exopodits and endopodits conical, equal in length, exopodit thicker than endopodit. Third thoracopod: exopodit longer and thicker than endopodit. En- dopodit and its inner process equally long and thick.

First pair of dorsal processes damaged on both sides. Second and third pair each 1 mm long. Single medio-dorsal process damaged, giving it a bifid appearance.

Description I. monstrosa det. Bergh, 1897 (?) (Figs 3C,D, 4)

Hind body damaged. Body size approx. 4 mm. Cephalic appendages (Fig. 4) as described in the genus diagnosis. Exopodit of second thoracopod thicker and slightly longer than endopodit. Third thoracopods damaged, exopodit thicker than en- dopodit.

First and second pair of dorsal appendages still in situ, remaining part of trunk damaged.

Remarks. The original description of Ismaila monst- rosa Bergh, 1867 refers exclusively to the female holotype from 1867, since the second specimen was collected 30 years later in 1897. Bergh (1867) men- tioned the single medio-dorsal process of I. monstro- sa as being branched, respectively paired. Re-exam- ining both individuals, from 1867 and 1897, Jensen (1987) showed Bergh’s description to be erroneous since the medio-dorsal process of the holotype was damaged and that of the specimen from 1897 is unbranched. Both specimen were also re-examined in this study and each definitely have an unpaired and unbranched medio-dorsal process.

Both individuals agree regarding gross body shape and proportions. The exopodit and endopodit ofthe second thoracopod have about the same length in both specimens. The inner process of the en- dopodit of the 3" thoracopod is as long as the en- dopodit. This feature is distinctive to all other con- geners in which the inner process is shorter than the endopodit (see Tab. 1), except for I. jenseniana, spec. nov. The latter species, however, has an additional pair of dorso-lateral processes and, thus, is clearly distinct from I. monstrosa.

In absence of morphological differences it is

concluded that both individuals from St. Thomas, Virgin Islands, and from the same host species (Phidiana lynceus), belong to I. monstrosa. All other specimens formerly assigned to I. monstrosa by Bergh (1898), Belcik (1981) and Jensen (1987) clearly differ morphologically (see Tab. 1) and, thus, are regarded to be distinct species. Ismaila monstrosa occurs in tropical Caribbean waters parasitizing an aeolid nudibranch host, while I. jenseniana, spec. nov. was found at the same locality (St. Thomas) but in a sacoglossan host. All other Ismaila species occur in temperate waters, i.e. the southern and northern East-Pacific.

Ismaila obtusa, spec. nov. Fig. 5

Ismaila monstrosa Bergh, 1867: Bergh 1898: 506, Tab.. 29; Monod & Dollfus 1934: (partim); Belcik 1981: 23 (par- tim); Jensen 1987: 76 (partim).

Types. Holotype: ?, Zoologisches Museum Berlin (ZMB), No. 13512, Chile. Host: nudibranch, probably Anisodoris fontaini (D’Orbigny, 1837) (see below).

Etymology. Specific name comes from the latin obtusus (= stocky) and refers to the stocky body shape with large and broad dorsal processes and exopodits of thoraco- pods.

Description (2) (Figs 5A,B)

Body size at least 11 mm, hindmost parts of thorax and abdomen damaged. General body shape stocky. Cephalic appendages lacking, since cepha- lothorax damaged in this region.

Exopodit of second thoracopod thick and distal- ly flattened. Endopodit shorter, conical and much thinner. Exopodit of third thoracopod thicker than that of second thoracopod, distally flattened. En- dopodit shorter and much thinner than exopodit. Inner process of endopodit thinner and shorter than endopodit (ratios see Tab. 1).

Dorsal processes voluminous, relatively short, with very blunt tips. First two pairs distally flat- tened, third pair conical. Single medio-dorsal proc- ess shorter, thinner, conical.

Remarks. Bergh (1898) mentioned two parasite specimens which he assigned to I. monstrosa from two different nudibranch hosts, the doridoidean Archidoris incerta (junior synonym of the common Anisodoris fontaini D’Orbigny, 1837; see Schrödl 2000) and the aeolidoidean Aeolidia papillosa serotina Bergh, 1873. According to Bergh (1898), both parasite spec- imens were collected at the same locality, Tumbes, Chile, and during the same time period. However, the specimen from the ZMB examined in this study

Ex

ARE Eirre

>

Fig. 6. Ismaila jenseniana, spec. nov. Note the additional, dorsolateral pair of processes (dl). A. Dorsal view. B. Ventral view.

Fig. 7. Ismaila occulta Ho. A. 2, dorsal view. B. 2, ventral view. C. d, ventral view. D. d, dorsal view.

was labelled with ‘I. monstrosa, Puerto Montt’, and no host was mentioned. This specimen (Fig. 5) re- sembles the parasite from A. incerta which was sketched by Bergh (1898: pl. 29, fig. 26) as having very thick exopodits. According to Bergh (1898), the parasite specimen from A. papillosa serotina also looks like that from A. incerta. However, there is no orig- inal illustration nor any museum material remain- ing, thus, this statement cannot be verified. Para- sites found in A. papillosa serotina and described in this study (Figs 25B,C) are clearly distinct from I. obtusa since they have a grazile body shape.

In spite of considerable morphological differ- ences, Bergh (1898) assigned the parasite of A. incer-

ta to I. monstrosa and did not differentiate it from the parasite found in A. papillosa serotina. It thus seems that Bergh generally assigned all splanchnotrophids to I. monstrosa as far as they were distinguishable from Splanchnotrophus. With our present knowledge we can conclude that A. incerta is the host of the parasite specimen from the ZMB described above and that the museum label was probably wrong.

The specimen from the ZMB differs significant- ly from the holotype of I. monstrosa, as well as from all other congeners, due to its stocky body with voluminous and unique, distally flattened dorsal processes and exopodits. Thus it is described as I. obtusa, spec. nov.

9

Fig. 8. Ismaila occulta Ho, cephalicappendages. A. Antennule (left). B. Antenna (right). C.Mandible (left). D. Maxillule

(left). E. Maxilla (left).

Ismaila jenseniana, spec. nov. Fig. 6

Ismaila monstrosa Bergh, 1867: Jensen 1987: 75-84 (partim); Schrödl 1997: 45 (partim).

Types. Holotype: 2, ZMUC, collected at St. Thomas, Virgin Islands, from Ercolania funerea Costa, 1867, 28 April 1987.

Etymology. Named in honor of Dr. Kathe Jensen.

Description (Figs 6A,B)

Delicate body measuring 1.2 mm in length. Ce- phalic appendages were lost due to damage of the head.

Exopodit of second thoracopod longer and thick- er than endopodit. Exopodit of third thoracopod shorter than endopodit (artifact?). Endopodit about the same length as inner process, both equally thick.

First pair of dorsal processes damaged. Process-

10

es of second pair measuring 0.4 mm, those of third pair 0.3 mm in length, just like the medio-dorsal process. One additional pair of unbranched proc- esses is situated laterally between first and second pair of dorsal processes and second and third tho- racopods. They are called “dorso-lateral processes” in the following.

Remarks. This specimen from St.Thomas which was described by Jensen (1987) as I. monstrosa, at first glance resembles the holotype of I. monstrosa by Bergh (1867), despite of the fact that both specimens were found parasitizing two rather different host groups (sacoglossan vs. nudibranch hosts). Body proportions are just slightly different, and, in con- trast to all other congeners, these specimens have 3" thoracopods with equally long endopods and the endopods‘ inner processes (see Tab. 1). The funda- mental difference of I. monstrosa det. Jensen, 1987, however, is the unique additional pair of dorso-

200 um

20 um

50 um

G

50 um

10 um

5 pm

Fig. 9. Ismaila occulta Ho, legs. A. 2" thoracopod ($, right). B. 3°“ thoracopod (9, right). C. 4* thoracopod (9, right). D. 2” thoracopod (G, left). E. thoracopod (G, left). F. 4" thoracopod (G, left). G. 5" thoracopod (d, right).

lateral processes which is absent in I. monstrosa and in any other species of the genus Ismaila (Tab. 1). Therefore, the parasites from the sacoglossan Ercola- nia funerea described by Jensen (1987) are consid- ered to be a new species, Ismaila jenseniana, spec. nov.

Ismaila occulta Ho, 1981 Figs 7-9

Ismaila occulta Ho, 1981: 130-136; Ho 1987: 67-83; Jensen 1987: 75-83.

Types. Allotype: d, National Museum of Natural Histo- ry Washington (USNM), No. 184045; collected at Alam- itos Bay, California. Host: the nudibranch Dendronotus iris Cooper, 1863. Paratypes (322, 333): USNM No. 184046; collected together with the allotype. 1? (3.4 mm) and 1 (1.1 mm) paratype examined by SEM.

Description (2) (Figs 7A,B, 8, 9A-C) Delicate body, measuring up to 3 mm in length. Cephalic appendages resemble Ho’s (1981) original

description and are similar to cephalic appendages of other congeners which could be examined in detail by SEM (see I. androphila, spec. nov. and ge- nus diagnosis).

Exopodit of second thoracopod thicker and long- er than endopodit. Exopodit. of third thoracopod longer and thicker than endopodit. Inner process of endopodit rudimentary and very small. Two point- ed processes of fifth thoracopod arise from a com- mon base.

Three pairs of dorsal processes, medio-dorsal process between third pair; all processes conical with tip almost pointed.

Description (d) (Figs 7C,D; 9D-G)

Body size up to 1.1 mm. Cephalic appendages as in females and as in all congeners examined in detail.

Third thoracopod with a, compared to en- dopodit, very small and reduced inner process. Both processes of fifth toracopod arise from a common base.

11

A

B

Fig. 10. Ismaila belciki Ho, ? (damaged postero-dorsally). A. Dorsal view. B. Ventral view.

Remarks. The results of our re-examination using SEM correspond to a great extent with the light- microscopical original data presented by Ho (1981). The mandibles seem to be longer than illustrated by Ho (1981: fig 11). The shape of the (second) maxilla differs from the original description in that the fin- ger-shaped third segment does not bear stiff spinules but numerous long hairs. Thus, the maxilla of I. occulta closely resembles that of other congeners.

The number and morphology of thoracopods agrees with the original description of 1. occulta. In- terestingly, the fifth pair of thoracopods in 1. occulta consists of two small, pointed processes arising from a common base (Fig. 9G), while in all other Ismaila species which could be studied in detail the two processes arise separately. Due to this feature, with present knowledge, it is possible to distinguish I. occulta from I. aliena, spec. nov. (Fig. 18D,H), I. damnosa, spec. nov. (Fig. 21G) and I. belciki (Fig. 12F).

Ismaila occulta can be easily distinguished from I. belciki Ho, 1987 and from all other species of the genus Ismaila: the inner process of the endopodit of the third thoracopod is just an atrophied, rudimen- tary appendage in female I. occulta. The same is true for the endopodit of the third thoracopod of male I. occulta. In all other congeners these processes are much longer (Tab. 1).

12

Ismaila belciki Ho, 1987 Figs 10-12

Ismaila monstrosa Bergh, 1867: Belcik 1981: 16-25 (partim)); Jensen 1987: 75-84 (partim). Ismaila belciki Ho, 1987: 109-111; Huys 2001: 118-123.

Material studied. 622, 539, partly damaged (329, 338: ZSM No. 20010026); collected at Fossil Point, Empire, Oregon, 20 June 1963 (preserved 23 June 1963). 322, 18, partly damaged (222, 15: ZSM No. 20010027); collected at Charleston, Small Boat Basin, Oregon, 24 July 1963. Host: all specimens were from the nudibranch Janolus fuscus O’Donoghue, 1924. 322, 383 examined by SEM.

Description (2) (Figs 10, 11, 12A,B)

Delicate body measuring 1.8-4.4 mm. None of the specimens had egg sacs.

Cephalic appendages like in other Ismaila spe- cies (see I. androphila, spec. nov. and genus diagno- sis).

Endopodit of second thoracopod almost as long as exopodit. Exopodit of third thoracopod slightly longer and thicker than endopodit. Inner process of endopodit thinner but almost as long as endopodit. Further thoracopods not visible, because this region is covered by host tissue in all specimens examined.

Thorax dorsally with three pairs of processes and one single medio-dorsal process. All processes are conical, unbranched and relatively thin com- pared to body.

u —— 5 um @ 20 um A 20 um B D 10 um 10 um 20 um F E

Fig. 11. Ismaila belciki Ho, cephalic appendages (2). A. Antennule (right). B. Antenna (right). C. Mandible (right). D. Maxillule (left). E. Maxilla (right). F. Tip of Maxilla, inner side.

50 um B @ 200 um 500 um A

10 um

10 um

Fig. 12. Ismaila belciki Ho, legs. A. 2" thoracopod (?, right). B. 3“ thoracopod (9, left). C. 2" thoracopod (G, right). D. 3“ thoracopod (d, right). E. 4" thoracopod (d, right). F. 5" thoracopod (d, right).

Description (8) (Figs 12C-F)

Body size 1.5-2.0 mm. Cephalic appendages re- semble those of females as well as those of conge- ners.

Second to fourth and sixth thoracopods as usual (see genus diagnosis). Fifth thoracopod consisting of two very small, pointed processes arising sepa- rately.

Remarks. Belcik (1981) described one pair of anten- nules, antennae, mandibles and maxilles in his ma- terial. Additionally, a labrum and a labium with lateral appendages were mentioned. Small sclero- tized structures underneath the labium were inter- preted as being remainders of maxillipeds. Regard- ing antennulae and antennae, the results of our re-examination of I. belciki confirm the original de- scription. The sickle-like mandibles seem to be longer than illustrated by Belcik (1981: Figs 9, 10); in addi- tion, the SEM-examination showed tiny hairs, which were not detected by light microscopical examina- tion, on the mandibles. Overlooked by Belcik (1981), I. belciki possesses maxillules (Fig. 11D) which re- semble in structure those of other congeners. As also seen by Huys (2001), the maxilles of 1. belciki bear a second hind process, and thus closely resem- ble the maxilles of I. occulta. In contrast to the origi- nal description by Belcik (1981), no traces of maxil- lipeds have been detected in I. belciki, nor in any other Ismaila species. Ho (1981) assumed that there are no maxillipeds in adult I. occulta, while in larval stages (copepodit II) he found small knobs, which represent the maxillipeds; these knobs already dis- appeared in the copepodit III (Ho, 1987b).

Due to tissue covering the relevant regions, the small thoracopods of female I. belciki could not be examined by SEM. Since male 1. belciki agree with congeners regarding the number and structure of thoracopods (see genus diagnosis), this can also be assumed for females.

Ismaila belciki is distinguishable from other Isma- ila species with delicate body shape due to body proportions (Tab. 1). The inner process of the endo- podit of the third thoracopod is slightly shorter than the endopodit, while in I. monstrosa and I. jenseniana, spec. nov. they have the same length. In I. androphi- la, spec. nov, I. magellanica, spec. nov., and 1. socialis, spec. nov. the inner process reaches just half the length of the endopodit and the exopodit is obvi- ously thicker than the endopodit. In 1. belciki they have about the same diameter. In I. occulta the inner process of the endopodit of the third thoracopod is only a rudimentary, tiny appendage. The dorsal processes of I. aliena, spec. nov. are as long as the body, while they measure about half of the body size in 1. belciki.

14

Ismaila androphila, spec. nov. Figs 1A, 2A, 13-15

Ismaila sp. Schrödl, 1996: 31.

Types. Holotype: 2, ZSM No. 20010011. Allotype: ZSM No. 20010012. Paratypes: 5353 (ZSM 20010013), 522 (ZSM 20010014), 533, 62? on SEM-mounts (ZSM 20010015). All collected by M. Schrödl, Bahia de Coliumo (36°32'S, 72°57'W), Chile, 25 January 1994. Hosts: 50 Okenia luna Millen, Schrödl, Vargas & Indacochea, 1994.

Additional material. 3822, 10933, collected together with the types from 50 Okenia luna. 1483 and 922 exam- ined by SEM.

Etymology. The species name refers to the several males associated with a single female parasite per host.

Description (2) (Figs 2A, 13B,C, 14, 15A-C) Delicate body measuring 1.9-4.2 mm in length. First segment of antennule (Fig. 14A) with three

short hairs on ventral side; distal portion bears two

long hairs. Base of second segment bearing two or three thin hairs ventrally, one shorter than the other(s); distal edge with usually seven long, thin hairs. First and second segments of antenna (Fig. 14B) with a stubby inner seta each. Third segment hook-shaped, bearing three setae on ventral side, of which one covers a small hole, possibly the opening of an antennal gland. Mandible (Fig. 14C) runs cross- wise under the labrum, often covered by the latter. Base of mandible thick and oval, extending to a sickle-shaped ramus sparsely covered with fine, short hairs. Maxillule (Fig. 14D) curved inwards, distal third biramous, each ramus with hairs on the edge. First segment of maxilla (Fig. 14E) trapezoi- dal. Second segment smaller, bearing third segment and an additional process. Third segment carries approximately 15 long, terminal hairs. The process of the second segment is thin, with a pointed tip, and has short hairs on one side. It is not evident from any of the individuals examined by SEM ifa second process behind the first one (as present in

I. occulta and 1. belciki) is present or absent.

Labium tongue-shaped and hairy. Distally, hairs are limited to the lateral portions of the labium. From the base of the labium arises a pair of para- gnath, hairy lobes laterally. In the centre of the labium there is a triangular area with hairs. On both sides, between the central hairy area and the hairy paragnath lobes, there is a passage without hairs (see also Fig. 1C). e

First thoracopod (maxillipeds) absent. Second thoracopod (Fig. 15A) branched into exo- and en- dopodit with about same lengths. Exopodit conical with a rounded tip, endopodid thinner, its tip bear- ing a claw. Exopodit and endopodit of third tho- racopod having same length, but endopodit thinner

In

BRNN.:

Fig. 13. Ismaila androphila, spec. nov. A. Nudibranch host specimen (Okenia luna) with egg-sacs protruding laterally under the mantle rim. B. ? parasite, ventral view. C. ? parasite, dorsal view. D. Ö parasite, ventral view.

than the exopodit and bearing a basal, small and thin process (Fig. 15B). Protopodit of fourth tho- racopod (Fig. 15C) round in cross-section, bearing a rather thin and pointed second limb. The fourth thoracopod is very small compared to second and third thoracopods. Posterior to the fourth thoraco- pod the body narrows and carries a sclerotized ring. No fifth thoracopods are visible because the area around the sclerotized ring is covered with host tissue or contracted. Behind the ring the body nar- rows further, sixth thoracopods are absent.

First visible segment of abdomen bears elongate

and straight egg sacs.

Thorax with three pairs of dorsal processes. In adult females these processes are shorter than whole body, reaching about 60 % of total body length. One single process is situated medio-dorsally between the third pair of dorsal processes.

Description (8) (Figs 13D, 15D-F)

Body length 0.6-1.6 mm, width of cephalothorax 0.3-1.5 mm.

Cephalic appendages are identical with those of the 2.

Fig. 14. Ismaila androphila, spec. nov., cephalic appendages (2). A. Antennule (right). B. Antenna (left). C. Mandible (left). D. Maxillule (left). E. Maxilla (left).

20 um

10 um

Fig. 15. Ismaila androphila, spec. nov., legs. A. 2" thoracopod (9, left). B. 3" thoracopod (9, right). C. 4" thoracopod (2, right). D. 2" thoracopod (d, left). E. thoracopod (d, right). F. 4" thoracopod (G, right).

16

Fig. 16. Ismaila aliena, spec. nov. A. ? parasite, ventral view. B. ? parasite, dorsal view. C. d parasite, ventral view.

Thorax ventral: tip of the second thoracopod (Fig. 15D) armed with a claw. Protopodit of third thoracopod (Fig. 15E) with two branches; a long, thick and pointed exopodit with a claw, and amuch shorter and thinner endopodit with a blunt tip. Fourth thoracopod resembling that of the female. Behind the fourth thoracopod there is a sclerotized ring. Males are fixed on hind body of females within a ring of (host?) tissue around the sclerotized ring. It was not visible if there is a fifth thoracopod or not because this area was strongly contracted in all indi- viduals examined. Sixth thoracic segment (segmen- tation indistinct, see discussion) is dorso-laterally drawn into a pair of elongate genital lobes. Three apical setae on each of these lobes probably repre- sent thoracopod six. Two of the setae are close to each other, the third is slightly higher and situated more laterally. The area between the genital lobes is deepened.

Genital pores presumably are situated laterally on the first abdominal segment but covered by the genital lobes. The third visible segment of the abdo- men bears, as in females, two caudal rami each with two or three hairs on the base.

Males lack any dorsal processes, but there are sac-shaped lateral protrusions on the cephalothorax. One is above the second thoracopod and a smaller one above the third thoracopod. Both protrusions are distinctly set off from each other.

Remarks. Millen et al. (1994) and Schrödl (1996) already observed parasites of the genus Ismaila in Okenia luna (see Fig. 2A). Not only these endopara- sites are confirmed as belonging to Ismaila androphi- la, spec. nov. but, obviously, also the cuticular giant “penis” of O. luna described by Munoz et al. (1996: fig. 2B) can be recognized as a somewhat abstracted female Ismaila clearly showing three pairs of dorsal

17

@)

D

Fig. 17. Ismaila aliena, spec. nov., cephalic appendages (2). A. Antennule (left). B. Antenna (right). C. Mandible (right). D. Maxillule (right). E. Maxilla (right).

20 um

Fig. 18. Ismaila aliena, spec. nov., legs. A. 2" thoracopod (9, left). B. 3' thoracopod (2, left). C. 4" thoracopod (2, right). D. 5" thoracopod ($, right). E. 2" thoracopod (dle, left). F. 3" thoracopod (d, right). G. 4" thoracopod (3, left). H. 5% thoracopod (d, right). I. Posterior body portion. Note the cuticular ring (c), genital lobes (gl) with 3 setae (= 6'" thoracopod) each, and caudal rami (cr).

18

Fig. 19. Ismaila damnosa, spec. nov. A. Nudibranch host specimen (Flabellina sp. 1) with egg-sacs protruding dorsally, posterior to the pericardium. B. ? parasite, dorsal view. C. ? parasite, ventral view. D. d parasite, dorsal view.

E. d parasite, ventral view.

processes and a single median process.

All individuals examined in this study possess very little morphological variation even regarding details like the number and position of hairs, claws and setae (see Tab. 1), therefore they all are regard- ed to belong to the same species. Ismaila androphila, spec. nov. is distinguishable from the otherwise similar Ismaila magellanica, spec. nov. and 1. socialis, spec. nov. due to slight but consistent morphologi- cal differences regarding proportions of the 3" tho- racopod, i.e. the endopod being much thinner than the exopod, while being equally thick in I. magellani- ca, spec. nov. and I. socialis, spec. nov. (see Tab. 1).

Biological information. The infection rate of O. luna with I. androphila, spec. nov. was 100 % of the exten- sive material studied herein; this is the highest in- fection rate of an opisthobranch population with splanchnotrophids ever documented. In each host there was just a single female parasite together with one to seven males. At least one male was always associated with its hind body to the hind body of the female. The hosts were never obviously dam-

aged by the parasites. In 98 % of the 50 examined hosts the female was lying crosswise in the anterior body cavity of the host, with egg-sacs protruding from the anterior left body portion of the host, most- ly from the lateral body wall just below the mantle rim (Figs 2A, 13A).

Ismaila aliena, spec. nov. Figs 1B,C, 2B, 16-18

Types. Holotype: ?, ZSM 20010001, Bahia de Coliumo, Chile, 27 February 1995. Allotype: ZSM 20010002, 8 April 1992. Paratypes: 1133 (ZSM 20010003), 6 April 1992; 5538 (ZSM 20010004), 22 April 1992; 12 (ZSM 20010005), 6 April 1992; 107%, 1 associated d (ZSM 20010006), 8 April 1992; 52? (ZSM 20010007), 22 April 1992; 1422, 6 juvenile 22, 6334 on 11 SEM-mounts (ZSM 20010000), April 1992. All types collected by M. Schrödl, Javier Sanchez and Claudio Perez, Bahia de Coliumo, Chile. Host: Thecacera darwini Pruvot-Fol, 1950.

Additional material. 30°? and 249Sparasites, collected by M. Schrödl, Javier Sanchez and Claudio Perez, Bahia

19

Fig. 20. Ismaila damnosa, spec. nov., cephalic appendages (2). A. Antennule (left). B. Antenna (right). C. Mandible (left). D. Maxillule (left). E. Maxilla (right). F. Tip of maxilla.

de Coliumo, Chile between 31 March 1992 and 27 Febru- ary 1995. 222? and 7685 examined by SEM. Etymology. The specific name is derived from the first finding of splanchnotrophids in T. darwini: while being observed under the binocular microscope, a male para- site protruded through the body wall of its host at the tip of one peribranchial process. This immediately recalled the first outbreak of an alien in the famous movie.

Description (2) (Figs 16A,B, 17, 18A-D)

Delicate body, 1.2-6.9 mm body length. Body shape, cephalic and thoracic appendages as in I. an- drophila (see also genus diagnosis). The following description refers only to characters varying be- tween different Ismaila species. Second thoracopod: exopodit longer and thicker than endopodit. Ex- opodits conical and rounded. Third thoracopod: exopodit longer and thicker than endopodit. Endo- podit with small process. Fifth thoracopod: two ru- dimentary small, pointed processes arise separately but very close to each other.

First visible abdominal segment bears coiled, white to pink egg sacs forming one whor!.

Thorax with three pairs of unbranched dorsal processes, in adult females they are as long as the whole body. The dorso-median, unpaired process is

20

relatively short and does not project behind the rear of the abdomen.

Description (8) (Figs 1B,C, 16C, 18E-I)

Body length 0.7-3.1 mm. Width of cephalotho- rax 0.2-1.8 mm. For details regarding body shape, cephalicand body appendages see I. androphila, spec. nov. and genus diagnosis. Fifth thoracopod visible in some specimens, shape as in females.

Remarks. Splanchnotrophid parasites in T. darwini (see Fig. 1B) were already mentioned by Schrödl (1996), but not further identified. They clearly be- long to the genus Ismaila (see Jensen 1987; genus diagnosis, this paper). All parasites from T. darwini examined in the present study are conspecific; they show little variation regarding number and posi- tion of hairs and setae, and proportions and shape of processes. h

The most distinctive feature of female 1. aliena, spec. nov. are the dorsal processes. They are at least as long as (or longer than) the total body, and thus much longer than in any other congener (Tab. 1). Male I. aliena, spec. nov. can reach considerable sizes being up to 2 times longer than males of any other congener known so far.

100 um 10 um 100 um - Vi 5 e

20 um

WE me 50 pm F G 10 um D E

Fig. 21. Ismaila damnosa, spec. nov., legs. A. 2" thoracopod ($, left). B. thoracopod (9, left). C. 4" thoracopod (2, left). D. 2" thoracopod (G, left). E. thoracopod (G, left). F. 4" thoracopod (g, left). G. 5" thoracopod (Z, left).

Biological notes. 84.6 % of 52 (unselectively) col- lected T. darwini (31 March-22 April 1992, Bahia de Coliumo) were infected with I. aliena, spec. nov. 15 hosts had more than one female parasite. In every host one mature female was situated laterally left in the body cavity, with egg-sacs protruding posterior to the gills (Fig. 1B). While hosts infected with a single female parasite show no obvious damage, those infected with more than a single female tend to have swollen bodies and appear disturbed. Inter- nally, no obvious organ damages caused by immo- bile mature females (beside penetrating the body wall with the abdomen) have been detected. How- ever, the numerous males migrating freely in the body cavity of their hosts may produce obvious canals damaging the connective tissue.

Ismaila damnosa, spec. nov. Figs 1C, 19-21

Ismaila sp. Schrödl, 1996: 37; 1997: 45-47.

Types. Holotype: ?, ZSM 20010016. Allotype: ZSM 20010017. Paratypes: 52? (ZSM 20010018); 533 (ZSM 20010019); 1533, 922, 3 juvenile 2? on 4 SEM-mounts (ZSM 20010020). All types collected by M. Schrödl, Bahia de Coliumo, 22 January 1994. Host: Flabellina sp. 1 (see Schrödl 1996).

Additional material. 2233, 1322, collected by M. Schrödl, Bahia de Coliumo, 22 January and 31 March 1994. Host: Flabellina sp. 1 (see Schrödl 1996). 142 and 248 parasites studied by SEM.

Etymology. The specific name of I. damnosa, spec. nov. refers to the destructive effect of these parasites to their hosts.

Description (2) (Figs 19B,C, 20, 21A-C)

Body length 0.9-3.4 mm. Body stocky. Dorsal appendages and exopodits of thoracopods volumi- nous. Cephalothorax not set off from trunk as dis- tinctively as in congeners with delicate body shape.

Second thoracopod with voluminous exopodit. Endopodit as long as exopodit or slightly longer. Third thoracopod with thick exopodit; endopodit as long as or longer than exopodit.

First visible abdominal segment bears coiled egg sacs forming one whorl.

All dorsal processes (three pairs and one medi- an process) voluminous and conical.

Description (d) (Figs 19D,E, 21D-G)

Body length 0.7-1.6 mm. Cephalic and body ap- pendages see I. androphila, spec. nov. and genus diag- nosis.

Fifth thoracopod detectable, consisting of two processes with separate origins.

500 um

Fig. 22. Ismaila robusta, spec. nov. A. Nudibranch host specimen (Phidiana lottini) with egg-sacs protruding medio- dorsally posterior to the rhinophores. B. ? parasite, dorsal view (dorsal processes damaged). C. ? parasite, ventral view. D. d parasite, dorsal view. E. d parasite, ventral view.

Remarks. Splanchnotrophids were already men- tioned from Flabellina sp. 1 (Schrödl 1996) and their harmful influences on their hosts were shown by Schrödl (1997). The present study proves the para- sites of Flabellina sp. 1 as belonging to the genus Ismaila (see Jensen 1987; genus diagnosis). All spec- imens examined are conspecific since there is very little morphological variation. Ismaila damnosa, spec. nov. shows a conspicuous, stocky and voluminous shape which is distinctive to all congeners with delicate body shape (Tab. 1). Also the proportions of dorsal processes and of thoracopod expodits are different to those of delicate species. Only two fur- ther Ismaila species have a stocky shape, I. robusta, spec. nov. and I. obtusa, spec. nov. Ismaila damnosa, spec. nov. has a less stocky body and longer dorsal processes in relation to the body length than I. ro- busta, spec. nov. With its flat dorsal processes and exopodits I. obtusa, spec. nov. clearly differs from any other Ismaila species.

22

Biological notes. The infection rate of the 33 Flabel- lina sp. 1 examined was 66.6 %. Five of the hosts were infected with more than one (up to five) fe- male parasites. Gonadal tissue of several hosts was heavily damaged or almost absent, while gonads of non-infected Flabellinasp. 1 of comparable sizes were normally developed. Strict host specifity of I. dam- nosa, spec. nov. is assumed, since a second, syntopic and common species of Flabellina (Flabellina sp. 2, see Schrödl 1996) was not at all parasitized.

Ismaila robusta, spec. nov. Figs 22-24

Types. Holotype: ?, ZSM 20010021 (on SEM-mount), col- lected 7 March 1994. - Allotype: ZSM 20010021 (on SEM- mount), collected together with holotype. Paratypes: 23d, (ZSM 20010021, on SEM mount), collected together with holotype.48d, 1? (ZSM 20010022, on SEM-mount), collected 7 March 1994; 18, 12 (ZSM 20010023, on SEM-

10 um

Fig. 23. Ismaila robusta, spec. nov., cephalic appendages (?). A. Antennule (left). B. Antenna (right). C. Maxillule. D. Maxilla.

20 um

200 um

20 um

F 10 um

D

Fig. 24. Ismaila robusta, spec. nov., legs. A. 2" thoracopod (?, right). B. 3" thoracopod (2, right). C. 4" thoracopod (2, left). D. thoracopod (d, right). E. 3" thoracopod (d, right). F. 4" thoracopod (G, left).

500 um

Fig. 25. Ismaila socialis, spec. nov. A. Nudibranch host specimen (Aeolidia papillosa serotina) with 2 pairs of egg-sacs (es) protruding latero-dorsally. B. ? parasite, dorsal view (medio-dorsal process damaged). C. ? parasite, ventral view. D. d parasite, dorsal view. E. d parasite, ventral view.

mount), collected 11 May 1996. Alltypes collected by M. Schrödl, J. Sanchez and C. Perez, Bahia de Coliumo, Chile. Host: the aeolidoidean nudibranch Phidiana lottini (Lesson, 1831).

Etymology. The specific name refers to the robust body form of the female parasites.

Description (2) (Figs 22B,C, 23, 24A-C)

Very stocky body, measuring 1.9-2.4 mm. Ce- phalothorax not distinctly set off from trunk, body processes voluminous. Cephalic appendages as in other species of Ismaila (see I. androphila, spec. nov. and genus diagnosis).

Dorsal appendages and exopodits of thoraco- pods inflated. Second thoracopod with voluminous exopodit, endopodit slightly shorter and much thin- ner. Third thoracopod with slightly less volumi- nous exopodit and endopodit, both with about the

24

same length. Fifth thoracopod not visible due to contraction.

First visible abdominal segment with pair of coiled egg sacs, forming double whorls.

Description (8) (Figs 22D,E, 24D-F)

Body length 1.9-2.4 mm. Cephalic appendages, thoracopods and other body processes as described for I. aliena, spec. nov. (see also genus diagnosis). Fifth thoracopod not visible.

Remarks. All parasites found in Phidiana lottini be- long to the genus Ismaila (see Jensen 1987; genus diagnosis in this paper). Females have almost no morphological variation. Ismaila robusta, spec. nov. differs from I. damnosa, spec. nov. due to its even more voluminous shape, more inflated dorsal proc- esses, due to 2" thoracopods with exopodits being

20 um

10 um

Fig. 26. Ismaila socialis, spec. nov., cephalic appendages (?). A. Antennule (right). B. Antenna (right). C. Maxilla

(right).

slightly longer than endopodits (vs. having the same length), and due to egg sacs coiled to a double whorl (vs. single whorl) (see Tab. 1). The third known congener with stout body shape, I. obtusa, spec. nov., differs due to its uniquely flattened dorsal processes and exopodits.

Biological information. Of more than 200 P. lottini examined, just three were infected by single ? par- asites with 1-44 associated to their abdomens. Egg sacs protruded dorsally between the hosts’s cerata (Fig. 22 A).

Ismaila socialis, spec. nov. Figs 25-27

Types. Holotype: ?, ZSM 20010008. Allotype: ZSM 20010009. Paratypes: 52? (ZSM 20010010, on SEM- mounts). All types collected by S. Millen, S. Gigglinger, J. Sanchez, C. Perez, Bahia de Coliumo, 17 December 1994, from a single Aeolidia papillosa serotina Bergh, 1873 (Fig. 25A). 52? examined by SEM, 559 were lost during critical point drying.

Etymology. The specific name refers to six ? and six d parasites within a single host.

Description (2) (Figs 25B,C, 26, 27)

Delicate body, measuring 2.6-3.6 mm. Antennules, antennes and maxilles as in other congeners (see I. androphila, spec. nov.; genus diagnosis). Mandibles and maxillules were not visible in the specimens studied since they were covered with host tissue.

Second and 3" thoracopods with endopodits being shorter and thinner than exopodits. Third thoracopod with small process of endopodit, meas- uring about one third of the endopodit. Fifth tho-

racopod not visible. First abdominal segment with pair of sausage- shaped and relatively short egg sacs.

Description (8) (Figs 25D,E) Body length 1.0-1.6 mm.

Remarks. There is almost no morphological varia- tion among the female parasites from A. papillosa serotina. Ismaila socialis, spec. nov. has to be compared with other Chilean congeners with delicate body shape (see Tab. 1): Ismaila aliena, spec. nov. has much longer dorsal processes relative to the body length than 1. socialis, spec. nov. In I. androphila, spec. nov. the exopodit and endopodit of the 2nd thoracopod have the same length, while in I. socialis, spec. nov. the endopodit is shorter than the exopodit. Ismaila magellanica, spec. nov. morphologically is very sim- ilar to 1. socialis, spec. nov.. However, Ismaila magel- lanica, spec. nov. infects a sacoglossan host in the Magellan Strait area, while 1. socialis, spec. nov. was found in A. papillosa serotina in central Chile.

Ismaila magellanica, spec. nov. Figs 28-30

Types. Holotype: 2, ZSM 20010024 (on SEM-mount). Allotype: ZSM 20010024, on SEM-mount). Paratypes: 222 (ZSM 20010024, on SEM-mount); 12, 2355 (ZSM 20010025, on SEM-mount). All types collected by S. Mil- len, Fuerte Bulnes (53°39'S, 70°56'W), Magellan Strait, 5 January 1995. Host: Elysia patagonica Muniain & Ortea, 1997 (Figs 28A,B).

Additional material. 322, collected together with the types, all together from 4 host specimens. All parasite specimens examined by SEM.

m O1

| 10 __10um 50 __50um 50 __50um |

Fig. 27. Ismaila socialis, spec. nov., legs (2). A. 2" thoracopod (right). B. 3 thoracopod (left). C. 4* thoracopod (right).

500 um

500 um

Fig. 28. Ismaila magellanica, spec. nov. A. Sacoglossan host specimen (Elysia patagonica) with egg-sacs protruding dorsally. B. Host with abdomina of parasites protruding the body laterally, ventral view. C. ? parasite, ventral view. D. 2 parasite, dorsal view. E. & parasite, ventral view. F. ö parasite, dorsal view.

26

C 10 um

de I a:

10 um

D

Fig. 29. Ismaila magellanica, spec. nov., cephalic appendages (?). A. Antennule (right). B. Antenna (left). C. Maxillule

(right). D. Maxilla (left).

Etymology. The specific name refers to the Magellan Strait, the type locality.

Description (2) (Figs 28C,D, 29, 30A,B)

Delicate body measuring 1.1-2.5 mm. Cephalic appendages as in other congeners (see I. androphila, spec. nov. and genus diagnosis).

Exopodit and endopodit of second thoracopod with almost equal length, but endopodit thinner. Fifth thoracopod not visible.

First abdominal segment with pair of short, straight and relatively thick egg sacs.

Description (8) (Figs 28E,F, 30C-E) Body length 0.8-1.1 mm. Fifth thoracopod not visible.

Remarks. The endoparasitic copepods of E. pata- gonica belong to the genus Ismaila (see Jensen 1987; genus diagnosis in this paper), there is hardly any morphological variability detectable. Ismaila magel- lanica, spec. nov. comes closest to Ismaila socialis, spec. nov. (see Tab. 1), but due to different hosts and considerable geographical distance (and hydro- graphical differences; see Brattstöom & Johanssen 1983) of the collecting localities they are considered to be different species.

Biological notes. There are only few records of endoparasitic copepods in Sacoglossa. Ismaila jense- niana, spec. nov. was reported to parasitize Ercolania funerea (see Jensen 1987), Monod & Dollfus (1934) described a member of Splanchnotrophus from Elysia ornata Pease, and Jensen (1990) found Arthurius ely- siae (Jensen, 1990) in Elysia australis (Quoy & Gaim- ard, 1832). All host species appear to be just sporad-

ically infected. No parasites were mentioned from Argentinian Elysia patagonica by Muniain & Ortea (1997). In contrast, the population of Elysia pata- gonica from the Magellan Strait showed a high infec- tion rate (90 %; see Schrödl 2002) with Ismaila magel- lanica, spec. nov. Two of the four host specimens examined had single female parasites, one host had two and one had even three female parasites, thus also the infestation of Chilean E. patagonica may be much higher than previously known from any sacoglossan hosts.

Discussion

Cephalic appendages. There was little and contra- dictory information on the morphology of cephalic appendages of Splanchnotrophidae. According to Gotto (1993), there are first and second antennae, sickle-shaped mandibles and second maxilles. How- ever, this referred to data on only one genus, Splanch- notrophus Hancock & Norman, 1863, of which only two species had been studied in more detail (see Laubier 1964), with contradictory results on number and identity of cephalic appendages. Most recently, Huys (2001) made an end to that long lasting debate showing S$. gracilis Hancock & Norman, 1863, the type species, and $. angulatus Hecht, 1893, to pos- sess a pair of 2-segmented antennules, 3-segmented antennae, one bilobate labrum, a pair of blade-like mandibles with denticulate tips, a pair of maxillules with an apical seta, and a pair of 2-segmented max- illae. Maxillipeds are definitely absent in adults. Descriptions of head appendages of Ismaila spe-

27

200 um

E 20 um

50 um

50 um

Fig. 30. Ismaila magellanica, spec. nov., legs. A. 2" thoracopod (2, right). B. 3’ thoracopod (2, left). C. 2"! thoracopod (3, right). D. thoracopod (d, right). E. 4" thoracopod (G, right).

cies also were highly contradictory: Ismaila monstro- sa was mentioned to have two pairs of antennae, a labrum, and one pair of mandibles and maxillae by Bergh (1867). The cephalic appendages of 1. belciki and I. occulta were described in detail (Belcik 1981, Ho 1981) but show significant differences concern- ing the number (4 vs. 5) and morphology of head appendages. Thus, one major aim of this study was to find out, with aid of the SEM, which kind of cephalic appendages are present in the different species of Ismaila, whether their structure is really that different, and to evaluate their taxonomic val- ue.

All Ismaila specimens examined herein have an- tennules and antennae. Also the presence of sickle- shaped mandibles could be confirmed. Under the SEM, the mandibles show a broad base and a long, elongate blade with fine hairs (Fig. 8C) which had not yet been reported for any Ismaila species. In accordance with Ho (1981), all Ismaila species are shown to possess two pairs of maxillae. Maxillules similar in shape to those of other congeners are definitely present in Ismaila belciki, obviously they had been overlooked by Belcik (1981). The second segment of the maxillae bears three hairy processes in I. occulta (see Ho 1981), I. belciki, and, possibly, also in other congeners, but the third process was difficult to detect by SEM examination due to its

28

posterior position (see Fig. 8E). In contrast to Belcik (1981) who mentioned maxillipeds from 1. belciki, no traces of maxillipeds could be found in any adult Ismaila specimens examined. Thus, there is no vari- ation regarding the number of cephalic appendages between the different Ismaila species. Also the shape of cephalic appendages of all species examined is similar even regarding details. Slight differences concerning the number of hairs or thorns of cephalic appendages, especially of the terminal hairs of an- tennules and maxilles, indicate that these may not be suitable features for taxonomic distinction: hairs may easily been torn off or may be hidden depend- ing on the angle of view. The SEM results suggest that all Ismaila species have complex, special shaped antennae and mouthparts which differ clearly from those of other splanchnotrophids (see Huys 2001), and thus appear to be diagnostic for Ismaila. Future cladistic analyses may also confirm such special structures as autapomorphies for Ismaila as assumed by Huys (2001).

Our results on head appendages of Ismaila agree well with the general assumption that heads of cope- pods consist of five segments, each bearing one pair of extremities, i.e. two pairs of antennae, mandibles and two pairs of maxilles (e.g. McLaughlin 1980). Additionally,.there is a labrum and a so-called la- bium with paragnath lobes also present in Ismaila.

Body segmentation and appendages. Several at- tempts have been made to homologize splanchno- trophid body portions, segments, legs and body outgrows, i.e. by Ho (1981), Jensen (1987), and Huys (2001) who, regarding Ismaila, largely followed Ho’s interpretation. The following interpretation is de- rived from results on different Ismaila species stud- ied herein. In both sexes, the five head segments of Ismaila are characterized by the presence of one pair of extremities each. In females one thoracic segment is aggregated to the head (Jensen 1987). It lacks extremities in adult specimens, but one pair of max- illipeds was shown to be present in the copepodit II of I. occulta by Ho (1987b). In female Ismaila, the second thoracic segment bears a pair of large, bi- ramous (second) thoracopods, the third thoracic seg- ment has triramous (third) thoracopods (“legs 1 and 2” by Ho 1981). The first pair of dorsal appendages is situated slightly anterior to the second thoraco- pods and is supposed to belong to the second seg- ment. The second pair of dorsal appendages lies in between the second and third thoracopods, and the third pair of dorsal appendages, together with the medio-dorsal process, is situated posterior to the third thoracopods. In contrast to Ho (1981) who assumed that the third pair of dorsal appendages belongs to the fourth thoracic segment, we think both the second and third dorsal appendages may more likely belong to a long third thoracic segment since Ho (1987b) showed copepodits IV to possess a long third but a short fourth thoracic segment. Ac- cording to our interpretation it is the fourth thoracic segment that bears a vestigial, unbranched, 2-seg- mented process, the fourth thoracopod (“leg 3” by Ho 1981). The anterior portion of the fifth thoracic segment forms a strong cuticular ring, posteriorly it bears a vestigial (fifth) thoracopod, a small process tipped with two setae in I. occulta (“leg 5” by Ho 1981) or two separate setae in other congeners. We have not found further thoracopods, thus the sixth thoracopods appear to be completely reduced in female Ismaila. Ho (1981) interpreted small elements inserting in the area of egg-sac attachment of female I. occulta as being an additional “leg 6”. However, no such structures have been detected in the present study. If present, such structures may also be ab- dominal outgrows rather than thoracopods.

In male Ismaila two laterally swollen thoracic segments are aggregated to the head (see Jensen 1987). In adult males the first thoracic segment lacks legs (i.e. maxillipeds), the second thoracic segment bears a pair of large uniramous (second) thoraco- pods. The third thoracic segment is distinctly set off from the cephalothorax; it bears a pair of large bi- ramous (third) thoracopods. The thoracopods of the fourth and fifth segments resemble those of the

females. In contrast to the females, the sixth thoracic segment of males bears a pair of (sixth) thoracopods which are modified into plate-like, triangular lobes with three distal setae which, as usual in in gymno- plean copepods, may serve as copulatory organs. The gonopores open at the following, first abdomi- nal segment.

Although the abdomen of I. occulta was inter- preted to be 1-segmented by Ho (1981), at least three segments may be distinguished in all Ismaila species examined. The first abdominal segment bears the genital openings in both sexes. The second visible segment does not bear any outgrows. The posterior segment is characterized by a pair of caudal rami.

Revision of the genus. Previously, the genus Isma- ila was represented by three species. The type spe- cies is Ismaila monstrosa Bergh, 1867 with only four female museum specimens available for re-exami- nation. Two of them, I. obtusa, spec. nov. (formerly I. monstrosa det. Bergh, 1898) and I. jenseniana, spec. nov. (formerly I. monstrosa det. Jensen, 1987), are shown herein to significantly differ from the holo- type of I. monstrosa described by Bergh (1867). This confirms Monod & Dollfus‘ (1934) and Belcik‘s (1981) doubts that the Chilean specimen illustrated by Bergh (1898) belong to I. monstrosa.

Californian specimens originally assigned to I. monstrosa by Belcik (1981) already were shown to be distinct from I. monstrosa and another Califor- nian species, Ismaila occulta Ho, 1981, by Ho (1987a) and described as I. belciki Ho, 1987. The main distin- guishing criteria were related to the cephalic ap- pendages and the thoracopods, i.e. the proportions of exopodits relative to endopodits of the 3" tho- racopods of females. Since Jensen (1987) showed the bilobed dorso-median process of I. monstrosa to be an artifact, this feature could no longer be used as a distinguishing feature to other congeners (see Ho 1981, Belcik 1981).

In the framework of this study specimens of the three currently known Ismaila species were re-ex- amined critically. In addition, parasites of six differ- ent Chilean host species were described. All para- sites out of one host species (if there were more than one parasite specimen to examine) are highly simi- lar to each other and, thus, are certainly conspecific. On the other hand, except for I. socialis, spec. nov. and I. magellanica, spec. nov., they are distinguisha- ble morphologically from the parasites of the other host species (Tab. 1).

Other than expected from the literature (see Ho 1981, Belcik 1981), all species of Ismaila show nearly identical cephalic appendages. Those of I. jensenia- na, spec. nov. and I. obtusa, spec. nov. could not be examined because the head region was damaged in

29

Tab. 1. Comparision of 11 known Ismaila species (literature data in quotation marks)

I. monstrosa I. obtusa, I. jenseniana, I. occulta 1. belciki Bergh, 1867 spec. nov. spec. novV. Ho, 1981 Ho, 1987 Host Phidiana lynceus Anisodoris fontaini Ercolania funerea Dendronotus iris Janolus fuscus (Nudibranchia: (Nudibranchia: (Sacoglossa) (Nudibranchia: (Nudibranchia: Aeolidoidea) Doridoidea) Dendronotoidea) “Arminoidea”) Locality St. Thomas, Tumbes, St. Thomas, Alamitos Bay, Oregon Virgin Islands, central Chile Virgin Islands, California Caribbean Sea Caribbean Sea No. of females examined in detail herein 2 1 1 6 & Body length of females (mm) (without egg-sacs) 3.2-4 >11 1.2 up to 3 1.8-4.4 Body shape (females) delicate very stocky, delicate, delicate delicate unique flattened additional pair of dorsal processes dorso-lateral processes Shape of egg-sacs coiled ? “elongate, “rather short, “short, (one whorl) straight” straight” . straight” Ratio length of first dorsal process/body length damaged 0.4 0.3 0.2-0.3 0.5-0.6 2"d thoracopods females:- relative length of exo-/endopodit exo = endo exo > endo exo > endo exo > endo exo = endo 2"d thoracopods: relative diameter of exo-/endopodit exo > endo exo > endo exo > endo exo > endo exo = endo 34 thoracopods females: relative length of exo-/endopodit exo > endo exo > endo ? exo > endo exo (>) endo 3” thoracopods females: relative diameter of exo-/endopodit exo > endo exo > endo 8 exo > endo exo (>) endo 3“ thoracopods females: ratio inner process/endopodit ji 0.3 1 0.1 0.8 No. of males examined 0 0 0 6 6 Body length males (mm) ? ® ? up to 1.1 1.5-2.0 34 thoracopods males: ratio inner process/endopodit ? 0.7 0.05 0.4 Setae of 5th thoracopod females ? % common base separate

both specimens. Differences between the species refer to general body shape, number, proportions and structure of body appendages and the shape of egg sacs.

As in Splanchnotrophus, which was subdivided into subgenera Lomanoticola (stocky body shape) and Splanchnotrophus (delicate body shape) by Dela- mare Deboutteville (1950), the Ismaila species can be

easily divided according to their stocky vs. delicate body shape (Tab. 1). Ismaila robusta, spec. nov., I. damnosa, spec. nov. and I. obtusa, spec. nov. have a conspicuous robust and stocky body. Within this group, species are distinguishable from another by the length and diameter of dorsal processes and proportions of exopodits to endopodits of the fe- males‘ 2" thoracopods (Tab. 1). Ismaila obtusa, spec.

I. androphila, I. aliena, I. damnosa, spec. nov. spec. noV. spec. noVv. Okenia luna Thecacera darwini Flabellina sp. 1

(Nudibranchia: (Nudibranchia: (Nudibranchia:

Doridoidea) Doridoidea) Aeolidoidea)

Bahia de Coliumo, Bahia de Coliumo, Bahia de Coliumo,

central Chile

central Chile

central Chile

15 15 15 1.9-4.2 1.2-6.9 0.9-3.4 delicate delicate, stocky very long dorsal processes elongate, coiled coiled straight (one whorl) (one whorl) 0.4-0.7 0.7-1.0 0.4-0.7 exo = endo exo = endo exo = endo exo > endo exo > endo exo > endo exo = endo exo > endo exo < endo exo > endo exo > endo exo = endo 0.5 0.2 0.4 10 10 10 0.6-1.6 0.7-3.1 0.7-1.6 0.3 0.5 0.3 % separate separate

I. robusta, I. socialis, I. magellanica, spec. nov. spec. nov. spec. nov. Phidiana lottini Aeolidia papillosa _Elysia patagonica (Nudibranchia: serotina (Sacoglossa) Aeolidoidea) (Nudibranchia: Aeolidoidea) Bahia de Coliumo, Bahia de Coliumo, Magellan Strait

central Chile

central Chile

4 4 7/ 1.9-2.4 2.6-3.6 1.1-2.5 very stocky delicate delicate coiled elongate, short, (two whorls) banana-like straight 0.3-0.4 0.3-0.4 0.4-0.6 exo > endo exo > endo exo > endo exo >> endo exo > endo exo > endo exo = endo exo > endo exo > endo exo > endo exo > endo exo > endo 0.2 0.3 0.3 7, 4 4 1.0-1.2 1.0-1.6 0.8-1.1 0.3 ? 0.2

nov. with its unique, large and flattened dorsal proc- esses and exopodits is clearly distinct to the delicate shaped I. monstrosa (as it was identified by Bergh 1898) and to any other species of Ismaila.

All other Ismaila species show the delicate body type. The single known female of I. jenseniana, spec. nov. has an unique additional pair of dorsolateral processes and, thus, is clearly distinct from all con-

geners. With exceptional long dorsal processes, I. aliena, spec. nov. is easily distinguishable from any other Ismaila species (see Tab. 1). In contrast to all other delicate species, females of I. aliena, spec. nov. have coiled egg sacs. Both individuals of I. monstrosa have third thoracopods with equally long endopodits and inner processes, while in all other Ismaila species (except for I. jenseniana, spec.

nov.) endopodits are longer than the inner process- es (Tab. 1). Ismaila occulta and 1. belciki are distin- guishable from each other and from all remaining gracile species of Ismaila by the short vs. long inner process of the endopodit of the third thoracopod (Ho 1987b; Tab. 1). Ismaila occulta additionally may be well characterized with 5" thoracopods consist- ing of two setae arising from a common base while other congeners in which 5" thoracopods could be examined have setae arising separately. Ismaila an- drophila, spec. nov. differs slightly but consistently regarding the proportions of the rami of the thoracopod from the otherwise similar Ismaila magel- lanica, spec. nov. and 1. socialis, spec. nov. (Tab. 1). The latter two species closely resemble each other regarding the shape of egg sacs, dorsal processes and proportions of endopodits and exopodits. How- ever, 1. socialis, spec. nov. parasitizes an aeolidoide- an nudibranch and is so far only known from cen- tral Chile, while Ismaila magellanica, spec. nov. in- fects the sacoglossan Elysia patagonica which occurs in the Magellan Strait area. Thus, as a result of this study, 11 different species of the genus Ismaila are regarded to be valid, showing more or less distinc- tive morphological features (Tab. 1). The general body shape including the structure of antennae, mouthparts, legs, and body outgrows is consistent within the different Ismaila species showing surpris- ingly little variability. This is in clear contrast to Splanchnotrophus where the prosomal region and its lateral outgrows may show substantial intraspecific variability (Huys 2001).

An alternative hypothesis would be to regard at least some delicate Chilean Ismaila specimens as belonging to one single species; on one hand imply- ing certain intraspecific varibility, and on the other hand quite rigid adaptations to certain host species causing slight but consistent morphological differ- ences. However, biological data argue against this one-species-hypothesis. The parasites of Thecacera darwini and Okenia luna, two Chilean phanerobranch doridoidean nudibranchs, are highly specific regard- ing their position in the host (this study). Also the number of females in each host differs, in Okenia luna there was always just one single female para- site together with several males, while in Thecacera darwini (and other hosts) there may be more than one female. While the populations of O. luna and T. darwini in the Bahia de Coliumo were highly in- fected with Ismaila, another syntopic and common Chilean phanerobranch nudibranch, Holoplcamus papposus Odhner, 1926 was not infected at all. An- other, morphologically well-characterized Ismaila species, I. damnosa, spec. nov., also infects just a single host species (Flabellina sp. 1) while other syn- topic aeolids and even a very common and external-

32

ly similar congener, Flabellina sp. 2, were not infect- ed at all. In conclusion, it seems that the species of Ismaila are highly host specific.

Host specifity may be quite common in certain copepod groups. For example, Ho (1994) found a strong trend towards host specifity for the Chondr- acanthidae (parasites of marine fishes). Humes (1974) considered Anthessius dolabellae Humes & Ho, 1965, Doridicola audens Humes, 1959 and Metaxymolgus commodus Humes, 1964, three exoparasitic copepods which are associated with Opisthobranchia, as host specific. Over a geographical distance from Madagas- car to New Caledonia, they occur in the same host species. However, in the future, it will be interesting to use molecular techniques to test such taxonomic hypotheses which until now are solely based on quite ambiguous morphological or biological data.

With the results of this study it is now possible to give a detailed diagnosis of the genus Ismaila and to identify allknown members (Tab. 1). Attempts to define the genus had already been made by Belcik (1981), Jensen (1987), and Huys (2001). Belcik con- sidered the maxilla of 1. belciki as being characteris- tic for the genus. The present study shows a special type of mandible, maxillule and maxilla to be present in all Ismaila species and to be different in structure from such mouthparts of other splanchnotrophids studied by Huys (2001). These features, together with the sexual dimorphism of the 2" and 3" tho- racopods (“legs 1 and 2”, see Huys 2001) are certain- ly diagnostic for Ismaila and may also be autapo- morphies. Jensen’s (1987) observation that the fe- males of Ismaila, in contrast to other genera of the Splanchnotrophidae, have always long and slender (anterior) thoracopods extending beyond the body laterally, could be verified in this study. A single medio-dorsal process is present in all Ismaila species which appears to be an unique autapomorphy of the group. Due to the absence of a comprehensive phylogenetic analysis it is still problematic to judge on the significance of characters and polarity of character states. However, all these potential aut- apomorphies coincide with each other clearly sug- gesting that the genus Ismaila is monophyletic. Three pairs of large dorsal processes, which are not ho- mologous to thoracopods but formed de novo (Lau- bier 1966 fide Huys 2001), are common to Ismaila, Splanchnotrophus, Lomanoticola, and Ceratosomicola) and may indicate a common ancestry. The genus Arthurius is morphologically aberrant and, accord- ing to Huys (2001), shows a mix of highly derived and plesiomorphic characters which additionally differ between both sexes. However, as long as there isno sound hypothesis on the phylogeny of Splanch- notrophidae, either on their systematic position, it is difficult to draw reliable evolutionary conclusions.

Acknowledgements

We would like to thank Dr. Francis Belcik, Dr. Niels Bruce (ZMUC), Dr. Charles Coleman (ZMB), Dr. Frank Ferrari (USNM), and Dr. Kathe Jensen (ZMUC) for giving us museum material for examination. Sebastian Gigglin- ger (Munich), Sandra Millen (Vancouver), Claudio Perez and Javier Sanchez (Concepciön, Chile) helped collect- ing specimens in the field. Prof. Horst Bohn and Prof. Gerhard Haszprunar are thanked for laboratory facili- ties in Munich. Dr. Ju-shey Ho (Long Beach) made com- ments on the manuscript, Teresa Saks kindly corrected the language.

References

Belcik, F. P. 1981. The male of Ismaila monstrosa Bergh, 1867 (Copepoda, Splanchnotrophidae). Crusta- ceana 40: 16-25

Bergh, L. S. R. 1867. Phidiana lynceus og Ismaila monstrosa. Vidensk. Medd. Dan. Naturh. For. (1866): 97-130, pls. 3-4

-- 1898. Die Opisthobranchier der Sammlung Plate. Zool. Jb. Suppl. 4: 481-582

Brattström, H. & A. Johanssen 1983. Ecological and re- gional zoogeography of the marine benthic fauna of Chile. Report no. 49 of the Lund University Chile Expedition 1948-49. Sarsia 68: 289-339

Delamare Deboutteville, C. 1950. Contribution a la co- naissance des Copepodes genre Splanchnotrophus Hancock & Norman parasites de mollusques. Vie Milieu 1: 74-80

-- 1951. Description du male du genre Splanchno- trophus Hancock & Norman (Crust. Copepoda). Vie Milieu 2: 366-370

Gotto, R. V. 1993. Commensal and parasitic copepods associated with marine invertebrates (and whales). In: Synopses of the British Fauna, No. 46. D. M. Kermack, R. S. K. Barnes & J. H. Crothers (eds). Universal Book Services, Dr. Backhuys

Hancock, A. & A.M. Norman 1863. On Splanchnotrophus, an undescribed Genus of Crustacea, parasitic in Nudibranchiate Mollusca. Trans. Linn. Soc. Lon- don 24: 49-60, pls. XV-XVI

Hecht, E. 1893. Note sur un nouveau Copepode parasite des Nudibranches. - Arch. Zool. Exp. Gen. 3: 13-16

-- 1895. Contributions a l’etude des Nudibranches. Mem. Soc. Zool. Fr. 8: 539-711

Ho, J.-S. 1981. Ismaila occulta, a new species of poecilosto- matoid copepod parasitic in a dendronotid nudi- branch from California. J. Crust. Biol. 1: 130-136

-- 1987a. Reconsideration of Ismaila monstrosa Bergh, 1867 (Copepoda, Splanchnotrophidae) from Ore- gon. Crustaceana 52: 109-111

-- 1987b. Larval stages of Ismaila occulta Ho, 1981 and the affinity of Splanchnotrophidae (Copepoda: Poe- cilostomatoida). Research. Crust. 16: 67-83

-- 1994. Chondracanthid copepods (Poecilostomato- ida) parasitic on Japanese dep-sea fishes, with a key to the genera of the Chondracanthidae. - ]. Nat. Hist. 28: 505-517

Humes, A. G. 1974. Cyclopoid Copepods associated with Opisthobranch Mollusks in New Caledonia. Crustaceana 26: 233-238

Huys, R. 2001. Splanchnotrophid systematics: a case of polyphyly and taxonomic myopia. J. Crust. Biol. 21: 106-156

Jensen, K. R. 1987. Ismaila monstrosa Bergh (Copepoda: Splanchnotrophidae) found parasitizing in Ercola- nia funerea (Costa) (Gastropoda: Ascoglossa). - Ophelia 28: 75-84

-- 1990. Splanchnotrophus elysiae n. sp. (Copepoda; Splanchnotrophidae) found parasitizing in the sacoglossan opisthobranch Elysia australis (Quoy and Gaymard, 1832). Proceedings of the Third Interna- tional Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia. Western Australian Museum, Perth 1: 291-296

Laubier, L. 1964. La morphologie des pieces buccales chez Splanchnotrophidae (copepodes parasites de mollusques). Crustaceana 7: 167-174

McLaughlin, P. A. 1980. Comparative Morphology of Recent Crustacea. Freeman, San Francisco

Millen, S. V., Schrödl, M., Vargas, N. & A. Indacochea 1994. A new species of Okenia (Nudibranchia: Dori- dacea) from the Peruvian Faunal Province. Veli- ger 37: 312-318

Monod, T. & R.-P. Dollfus 1934. Des copepodes parasites de mollusques. —- Ann. Parasit. 12: 309-321

Munoz, M. A., Valdes, A. & J. Ortea 1996. The genus Okenia Menke, 1830 (Nudibranchia: Goniodoridi- dae) in Chile. -— Haliotis 25: 101-106

Muniain, C. & J. Ortea. 1997. First record of a sacoglos- san (= ascoglossan, Opisthobranchia) from Patago- nia (Argentina): Description of a new species of genus Elysia Risso, 1818. Veliger 40: 29-37

Schrödl, M. 1996. Nudibranchia y Sacoglossa de Chile: Morfologia exterior y distribuciön. Gayana Zool. 60: 17-62

-- 1997. Aspects of Chilean nudibranch biology: Ef- fects of splanchnotrophid copepod parasitism on Flabellina sp. 1. - Opisthobr. Newsl. 23: 45-47

-- 2000. Taxonomic revision of the common South American nudibranch Anisodoris fontaini (D’Or- bigny, 1837) (Gastropoda: Opisthobranchia), with discussion of its systematic placement. J. Moll. Stud. 66: 49-61

-- 2002. Heavy infestation by endoparasitic copepod crustaceans (Poecilostomatoidea: Splanchnothro- phidae) in Chilean opisthobranch gastropods, with aspects of splanchnothrophid evolution. Org. Divers. Evol. 2: 19-26. Plus Electronic Supplement (Parts 1 and 2) at http:/ /www.senckenberg.de/ odes/02-03.htm

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Buchbesprechungen

1. Schmidt, G.: Giftige und gefährliche Spinnentiere. Die Neue Brehm-Bücherei Bd. 608, 2. überarb. u. erweit. Auflage. - Westarp Wissenschaften Verlags- ges. m.b.H., Hohenwarsleben, 2000. 215 S., mit 2 Abb. ISBN 3-89432-405-8.

. von den etwa 66.000 Arten der Spinnentiere können nur wenige dem Menschen wirklich gefährlich werden. Nur von ihnen handelt dieser Band” schrieb der Autor am Ende seines Vorwortes in der ersten Auflage und daran hält er sich auch in der zweiten aktualisierten Auflage. Die Hauptkapitel widmen sich den Skorpio- nen, Milben und Spinnen, daneben werden noch die Geißelskorpione und Walzenspinnen erwähnt. Nicht zu Unrecht ist es dem Autor wichtig, am Ende des ersten Kapitels (Gifte und ihre Wirkungen. Unfälle durch Gift- tiere weltweit.) darauf hinzuweisen, “... daß Unfälle, auch tödliche, durch Schlangen, Bienen und Wespen, diejenigen, welche durch Spinnentiere verursacht wer- den, bei weitem in ihrer Häufigkeit übertreffen.” Dies ist aber nicht als allgemeine Verharmlosung gedacht. So sind die für den Menschen gefährlichen Arten und deren Verbreitung, ihre medizinische Bedeutung, ihre Toxizi- tät und die Wirkungen der Gifte eingehend beschrieben. Besonders wichtig scheinen mir die Hinweise auf die Prophylaxe und das Kapitel: “Was tun bei Unfällen”.

Durch die Reisefreudigkeit in ferne Länder kommt es immer wieder zur unabsichtlichen Einschleppung von giftigen Spinnentieren im Reisegepäck, sowie bei der Einfuhr von tropischen Früchten. So ist denn auch der Arzt bei uns bei Vergiftungen gefragt. Er sollte daher dieses Werk in seine Handbibliothek aufnehmen, aber auch der Reisende selbst sollte das Buch vor seiner Reise zur Hand nehmen, einerseits um unnötige Ängste abzu- bauen, andererseits um sich durch Kenntnis und Pro- phylaxe vor Unfällen zu schützen. Dem Buch ist eine große Verbreitung zu wünschen. L. Tiefenbacher

2. Brechtel, F. & H. Kostenbader (Hrsg.): Die Pracht- und Hirschkäfer Baden-Württembergs. Verlag Eu- gen Ulmer, Stuttgart 2002. 632 S., 306 Farbfotos, 180 Diagramme und Zeichnungen, 86 Velzelunglar ten, 30 Tabellen. ISBN 3-8001-3526-4.

Auf diesen voluminösen, prachtvoll und reichhaltig be- bilderten Band haben die daran Beteiligten und Interes- sierten lange gewartet. Sie haben jedoch nicht mit einem derartig ausführlichen und gut ausgestatteten Werk ge- rechnet, das in der Tat fast keine Frage offenläßt. Ob man etwas über die Verbreitung, über spezielle Fragen der Biologie oder Ökologie, die forstwirtschaftliche Bedeu- tung, oder über Gefährdung und Schutz der zwei behan-

34

delten Käferfamilien wissen möchte, man findet es. Ei- gentlich istes sogar erstaunlich, daß man über die Pracht- käfer, eine bei uns nun wirklich nicht artenreiche Käfer- gruppe, deren eigentliche Heimat die Tropen, bzw. in Europa der Mittelmeerraum ist, so viel schreiben kann. Das heißt zugleich, daß wir auch erstaunlich viel über diese Käfer wissen. Dieses Wissen haben die beiden Herausgeber, die zugleich die Hauptautoren sind, enorm gründlich zusammengetragen, allerdings mit Hilfe einer Reihe von im Impressum genannen Mitautoren, sowie zahlreicher weiterer Helfer, die Funddaten oder andere Informationen beigetragen haben.

Eine allgemeine historische Einführung wird gefolgt von einer ausführlichen Darstellung der Morphologie und der Systematik der beiden Käferfamilien, sowie ei- ner Checkliste. Darauf folgt eine sehr ausführliche Dar- stellung der Naturräume Baden-Württembergs und der für die Käfer wichtigen Lebensräume. Ebenso ausführli- che Darstellungen der Biologie und Ökologie, der Wirts- pflanzen, der Bedeutung dieser Käfer für die Land- und Forstwirtschaft, sowie der Gefährdung und der Schutz- bemühungen schließen den insgesamt 200 Seiten umfas- senden allgemeinen Teil ab.

Der spezielle Teil behandelt die Käfer in alphabeti- scher Reihenfolge der Gattungen und innerhalb dieser der Arten, und man findet darin erneut ausführliche Angaben zu Biologie und Ökologie, Schädlichkeit, Ge- fährdung und meistens auch eine Verbreitungskarte der jeweiligen Art für Baden-Württemberg, sowie bei selte- nen Arten auch genaue Verbreitungsangaben. Naturge- mäß unterscheiden sich die Kapitel der einzelnen Arten beträchtlich in Länge und Informationsgehalt. Das liegt daran, daß eine bedeutende Anzahl der Prachtkäfer bei uns am äußersten Nordrand ihres meist mediterranen Verbreitungsgebietes leben oder aus anderen Gründen selten oder nur sporadisch verbreitet sind. Mit wenigen Ausnahmen sind die einzelnen Kapitel mit in der Regel vorzüglichen Farbfotos der in der Tat prächtigen Käfer versehen, sowie häufig noch mit Fotos von Fraßbildern oder ihrer Lebensräume.

Ein fast 30-seitiges Literaturverzeichnis sowie Regi- ster der wissenschaftlichen und der deutschen Namen beschließen diesen Prachtkäferband, der zugleich auch ein Prachtband ist und in die Bibliothek eines jeden Käferfreundes gehört, auch wenn er nicht in Baden- Württemberg beheimatet ist. Er ist in der Tat ein Nach- schlagewerk und ist daher auch ein Muß für Forstämter sowie für alle Ämter und Einzelpersonen, die in irgend- einer Weise mit dem Naturschutz zu tun haben - zumal der Preis für ein derartig voluminöses und umfassendes Buch wirklich nicht hoch ist. M. Baehr

SPIXIANA München, 01. März 2003 ISSN 0341-8391

Die Milben in der Zoologischen Staatssammlung München Teil 2. Familie Spinturnicidae

(Acari, Gamasida)

Hanna Ferenc, Czeslaw Blaszak & Rainer Ehrnsberger

Ferenc, H., Blaszak, C. & R. Ehrnsberger (2003): Die Milben in der Zoologischen Staatssammlung München. Teil 2. Familie Spinturnicidae (Acari, Gamasida). -

Spixiana 26/1: 35-41

The second part of the revision of the mites stored in Zoologische Staatssa- mmlung München deals with the family Spinturnicidae. Mites from the samples of Willmann, Vitzhum, Kneissl, Hirschmann and Zumpt are recognised. The species are listed and the condition of the slides is described.

Mgr. Hanna Ferenc, Lehrstuhl für Tiermorphologie Adam Mickiewicz Universi- tät, 28 czerwca 1956 Nr. 198, 61-485 Poznan, Polen; e-mail: hania@man.poznan.pl

Prof. Dr. Czeslaw Blaszak, Lehrstuhl für Tiermorphologie Adam Mickiewicz Universität, 28 czerwca 1956 Nr. 198, 61-485 Poznan, Polen; e-mail: blaszak@main.amu.edu.pl

Prof. Dr. Rainer Ehrnsberger, Hochschule Vechta, Institut für Naturschutz und Umweltbildung, D-49364 Vechta, Bundesrepublik Deutschland; e-mail: rainer.ehrnsberger@uni-vechta.de

Einleitung

Die Milben aus der Familie Spinturnicidae sind ob- ligatorische Ektoparasiten, die ausschließlich auf Fledermäusen (Chiroptera) leben. Sie sehen bräun- lich aus, haben eine Grösse von 400-1700 um und ernähren sich vom Blut ihrer Wirte. Die meisten Arten kommen auf den Flügelhäuten vor (beson- ders auf Plagiopatagium, Propatagium und Chiro- patagium, selten auf Uropatagium); einzelne Arten wurden auch auf dem Augenlid und am Anus ge- funden.

Die außergewöhnlichen Lebensbedingungen führen zu einer speziellen Morphologie und Biolo- gie dieser Milben. Sie sind dorsoventral abgeflacht. Zur Anhaftung an die Wirte dienen die kräftigen Beine mit Ambulacren und grosser Kralle und die häutigen Pullvillen. Die Milben dieser Familie sind sehr stark sklerotisiert und haben dicke, steife Bor- sten. Bei den ? einiger Gattungen (Paraperiglischrus,

Mesoperiglischrus, Periglischrus, Cameronieta) ist das Opisthosoma verbreitet und dient als Saugnapf. Die Spinturnicidae sind lebendgebärend.

Die Dorsalseite der Milben der Unterfamilie Spinturnicinae ist mit einem Schild bedeckt, bei der Unterfamilie Periglischrinae sind es zwei Schilde. Bei den d befinden sich auf dem Sternogenitalschild 4 Paar Borsten. Bei den ? sind Sternalschild und Genitalschild getrennt. Das Sternalschild trägt beim 2 3 Paar Borsten, das Genitalschild 1 Paar Borsten. Die Peritremen befinden sich auf der Dorsalseite, die Stigmen liegen zwischen dem 3. und 4. Bein. Sie verlaufen nach vorne bis zum 3. Beinpaar (Gattun- gen: Paraperiglischrus Rudnick, 1960 und Meso- periglischrus Dusbabek, 1968) oder sogar bis zum 2. Beinpaar (Gattungen: Ancystropus Kolenati, 1856, Meristaspis Kolenati, 1857, Oncoscelus Delfinado & Baker, 1963, Eyndhovenia Rudnick, 1960, Spinturnix von Heyden, 1826, Emballonuria Uchikawa, 1994, Paraspinturnix Rudnick, 1960, Periglischrus Kolenati,

35

1857, Cameronieta Machado-Alison, 1965). Bei eini- gen Gattungen (Spinturnix, Emballonuria, Paraspin- turnix) ziehen die Peritremen auf der Ventralseite bis zwischen das 2. und 4. Beinpaar. Das Tectum ist kegelförmig (konisch) oder rundlich. 1 Paar Gnathosomalborsten und 3 Paar Hypostomalbor- sten. Cheliceren der d mit Spermatodaktylus.

In dieser Arbeit benutzen wir die Systematik nach Rudnick (1960).

In der Zoologischen Staatssammlung München befinden sich Präparate von Spinturnicidae aus fol- genden Kollektionen: Kneissl, Vitzthum, Willmann, Hirschmann, Zumpt (Südafrika) und Ferenc (Lehr- stuhl für Tiermorphologie Poznan/Polen Abk. ZMZ).

Bei der Beschreibung wird auch der aktuelle Zustand der einzelnen Präparate angegeben. Es werden vier Erhaltungszustände unterschieden: A: Zustand sehr gut, B: Zustand gut, C: Präparate beschädigt, D: Präparat zerstört (kann nicht mehr restauriert werden).

Gattung Spinturnix von Heyden, 1826 Spinturnix myoti (Kolenati, 1856)

Pteroptus murinus Walckenaer & Gervais, 1847 Spinturnix murinus Oudemans, 1910 Pteroptus myoti Kolenati, 1856 Diplostaspis myoti Kolenati, 1857 Pteroptus cappaccinii Kolenati, 1856 Diplostaspis nattererii Kolenati, 1857 Diplostaspis nattererii Koch, 1865 Heterostspis octastigma Kolenati, 1859 Heterostspis octastigma Vitzthum, 1932 Diplostaspis stellata Kolenati, 1859 Spinturnix mystacinus Oudemans, 1902 Pteroptus grossus Banks, 1910 Spinturnix.vespertilionis Hirst, 1942

Diagnose

2: Länge des Idiosoma 1,19-1,72 mm, Breite 0,94- 1,33 mm; 3 Paar Borsten auf Sternalschild, das eine Länge von 213-270 um und eine Breite von 217-239 um hat. Vorderer Rand des Sternalschildes gerun- det; Anzahl der Opisthodorsalborsten 90-130. Scut- um bedeckt mit Schüppchen mit gezähntem Rand.

d: Länge des Idiosoma 1,09-1,16 mm, Breite 0,84- 0,91 mm; 4 Paar Borsten auf Sternalschild, das eine Länge von 371-413 um und eine Breite von 257- 304 um hat; Anzahl der Opisthodorsalborsten 33- 46.

Wirt: Myotis myotis

36

Präparate aus der Sammlung Willmann

1. [Präparat Nr.: 63/4] Spinturnix vespertilionis 16, leg. Leruth, det. Willmann [B], det. H. Ferenc/ 2000

2. [Präparat Nr.: 63/5] - Spinturnix sp. 16, det. Will- mann [A], det. H. Ferenc/ 2000

3. [Präparat Nr.: 63/9] - Spinturnix sp. - 15, det. Will- mann [B], det. H. Ferenc/2000

4. [Präparat Nr.: 63/10] Spinturnix sp. -— Hermanns Höhle, leg. Vornatscher, 12, det. Willmann [C], det. H. Ferenc/ 2000

5. [Präparat Nr.: 63/11] - Spinturnix sp. 42/83, Stum- nier, det. Willmann, 12 [C], det. H. Ferenc/ 2000

6. [Präparat Nr.: 63/12] - Spinturnix sp. - B. 5., leg. L. Leruth, det. Willmann, 12 [C], det. H. Ferenc/2000

7. [Präparat Nr.: 63/13] - Spinturnix sp. - B. 5., leg. L. Leruth, det. Willmann 14 [B], det. H. Ferenc/2000

8. [Präparat Nr.:63/ 14/1] - Spinturnix sp. - (Ni.?) Höh- le, leg. Vornatscher, det. Willmann, 12 [C], det. H. Ferenc/ 2000

9. [Präparat Nr.: 63/ 14/2] - Spinturnix sp. - (Ni.?) Höh- le, leg. Vornatscher, det. Willmann, 13 [C], det. H. Ferenc / 2000

10. [Präparat Nr.: 63/ 14/3] - Spinturnix sp. - (Ni.?) Höh- le, leg. Vornatscher, det. Willmann, 12 [C], det. H. Ferenc/ 2000

11. [Präparat Nr.: 63/ 14/5] - Spinturnix sp. - (Ni.?) Höh- le, leg. Vornatscher, det. Willmann, 18 [C], det. H. Ferenc/ 2000

Präparate aus der Sammlung Vitzthum

1. [Präparat Nr.: V 1134] 18.04.1904 Spinturnix myoti Kolenati - 1? [A], Vespertilio murinus, det. Furman/ 1968, det. H. Ferenc/ 2000

2. [Präparat Nr.: V 1135] 18.04.1909 Pteroptus vesperti- lionis 1? [C], von Vespertilio murinus, det. Furman/ 1968, det. H. Ferenc/ 2000

3. [Präparat Nr.: V 1150] 15.03. 1932 Spinturnix muri- nus (Walkenaer, 1847) 288 [A], det. C. Blaszak/ 2001

4. [Präparat Nr.: V 1157] 09.04.1937 Spinturnix muri- nus (Walkenaer, 1847) - 1? [B], det. H. Ferenc/2000

5. [Präparat Nr.: V 1158/1] 18.05.1933 Spinturnix murinus (Walkenaer, 1847) - 12 [A], det. H. Ferenc/ 2000

6. [Präparat Nr.: V 1158/2] 18.05.1933 Spinturnix murinus (Walkenaer, 1847) - 18 [A], det. H. Ferenc/ 2000

7. [Präparat Nr.: V 1164] 17.04.1927 Spinturnix sp. 12 [A], det. H. Ferenc/2000

8. [Präparat Nr.: V 1166] 31.01.1936 Spinturnix sp. 12 [A], det. H. Ferenc/ 2000

9. [Präparat Nr.: V 1168] 28.01.1936 Spinturnix sp. 1? [A], det. H. Ferenc/2000.

10. [Präparat Nr.: V 1189/1] 15.11.1928 Spinturnix sp. 13 [A], det. H. Ferenc/2000

Präparate aus der Sammlung Hirschmann

1. [Präparat Nr.: W 63/1] - Spinturnix myoti 2, 8, 27 [B], det. Hirschmann

2. [Präparat Nr.: W 63/2] - Spinturnix myoti 2, 28 [B], det. Hirschmann

Spinturnix andegavinus Deunff, 1977

?Diplostaspis daubentonii Kolenati, 1857 ?Diplostaspis stellata Kolenati, 1859 ?Spinturnix daubentonii Neumann, 1942 Spinturnix myoti Rudnick, 1960

Diagnose

2: Länge des Idiosoma 0,91-1,36 mm, Breite 0,75- 0,97 mm; 3 Paar Borsten auf Sternalschild, das eine Länge von 160-180 um und eine Breite von 130-152 um hat. Vorderer Rand des Sternalschildes zuge- spitzt; Anzahl der Opisthodorsalborsten 70-80. Die Borsten liegen in 4 oder mehr Reihen. Scutum mit Schüppchen mit gezähntem Rand.

d: Länge des Idiosoma 0,80-0,95 mm, Breite 0,64- 0,78 mm; 4 Paar Borsten auf Sternalschild, das eine Länge von 331-370 um und eine Breite von 241-273 um hat; Anzahl der Opisthodorsalborsten 33-46.

Wirt: Myotis daubentoniü

Präparate aus der Sammlung Vitzthum

1. [Präparat Nr.: V 1147] 11.10.1910 - Spinturnix mysta- cinus (Kolenati) 1? [A], det. H. Ferenc / 2000

2. [Präparat Nr.: V 1148] 11.10.1910 - Spinturnix mysta- cinus (Kolenati) 18 [A], det. H. Ferenc / 2000

3. [Präparat Nr.: V 1163] 11.10.1910 - Spinturnix mysta- cinus (Kolenati) - 12 [A], det. H. Ferenc / 2000

4. [Präparat Nr.: V 1167] 16.01.1936 Spinturnix sp. 12 [A], det. H. Ferenc/2000

5. [Präparat Nr.: V 1182] 13.11.1928 Spinturnix sp. 12 [A], det. H. Ferenc/2000

6. [Präparat Nr.: V 1191] 25.10.1933 Spinturnix sp. 13 [A], det. H. Ferenc/2000

7. [Präparat Nr.: V 1181] 26.3.1929 Spinturnix sp. 2 [A], det. C. Blaszak / 2001

Spinturnix americanus (Banks, 1902)

Pteroptus americanus Banks, 1902

Spinturnix americanus Banks, 1915

Spinturnix americana Rysgaard, 1942

Pteroptus echinipes Banks, 1910

Spinturnix araguensis Vitzthum, 1932

Spinturnix iowae Keegan, 1943

Spinturnix sp. Dymond, 1938

Spinturnix sp. undet., Holdenried, Evans & Longanek- ker, 1951

Diagnose

2: Länge des Idiosoma 1,04-1,24 mm, Breite 0,76- 0,93 mm; auf der Dorsalseite in Höhe des Femur II liegen 2 Mikrochaeten und 2 Makrochaeten; Anzahl der Opisthosomalborsten 10 oder 12 (4 liegen in der Mitte, 6 oder 8 liegen terminal).

d: Länge des Idiosoma 0,83-1,01 mm, Breite 0,66- 0,80 mm; auf der Dorsalseite in Höhe des Femur II liegen 2 Mikrochaeten und 2 Makrochaeten; Anzahl der Opisthosomalborsten 4.

Wirt: Myotis lucifugus

Präparate aus der Sammlung Vitzthum

1. [Präparat Nr.: V 1130] Spinturnix araguensis Vitzt- hum, 1931, Hacienda Puerto de la Cruz, westl. La Guaira, Venezuela, 27.12.1930, leg. Hase, det. Vitzt- hum (05.1930) - 1? [A] von Myotis nigricans SYNTY- PUS, det. H. Ferenc/ 2000

Spinturnix kolenatii Oudemans, 1910

Pteroptus arcuatus Kolenati, 1856 Pteroptus carnifex Kolenati, 1856 Diplostaspis carnifex Kolenati, 1857 Spinturnix carnifex Stiles & Nolan, 1931 Spinturnix vespertilionis Oudemans, 1903 Spinturnix kolenatii Oudemans,1910 Spinturnix kolenatii Neumann, 1942 Spinturnix sp. Thompson, 1935 Spinturnix murinus Allen, 1940

Diagnose

?: Länge des Idiosoma 0,75-0,79 mm, Breite 0,60- 0,62 mm; auf der Dorsalseite in Höhe des Femur II liegen 2 Mikrochaeten und 2 Makrochaeten; Geni- talschild liegt sehr dicht beim Sternalschild; Anzahl der Opisthodorsalborsten 8 (4 Mikrochaeten und 4 normale Borsten).

d: Länge des Idiosoma 0,65-0,68 mm, Breite 0,55- 0,56 mm; auf der Dorsalseite in Höhe des Femur II liegen 2 Mikrochaeten und 2 Makrochaeten; Anzahl der Opisthodorsalborsten 4 Mikrochaeten.

Wirt: Eptesicus serotinus

Präparate aus der Sammlung Vitzthum

1. [Präparat Nr.: V 1178] 13.11.1928 Spinturnix sp. 13 [B], det. H. Ferenc/ 2000

2. [Präparat Nr.: V 1188] 13.11.1928 Spinturnix sp. - 15 [B], det. H. Ferenc/ 2000

3. [Präparat Nr.: V 1179] 13.11.1928 Spinturnix sp. - 18 [A], det. H. Ferenc/ 2000

4. [Präparat Nr.: V 3300] 04.05.1909 - Spinturnix vesperti- lionis - 12, det. Furman/ 1968, [A], det. H. Ferenc/ 2000

Spinturnix punctata (Sundevall, 1833)

Pteroptus punctatus Sundevall, 1833 Spinturnix punctatus van Eyndhoven, 1943 Pteroptus barbastelli Kolenati, 1856 Diplostaspis barbastelli Kolenati, 1957 Spinturnix sp. van Eyndhoven, 1950

Diagnose

2: Länge des Idiosoma 1,11-1,15-mm, Breite 0,78-0,90 mm; von den 3 Paar Sternalborsten liegen 1 oder 2 Paar außerhalb des Sternalschildes; Anzahl der Opisthodorsalborsten 36-41; Länge der Podoso- malborsten 81-128 um. Scutum bedeckt mit Schüpp- chen.

d: Länge des Idiosoma 0,87-0,92 mm, Breite 0,70- 0,72 mm; Anzahl der Opisthodorsalborsten 18-21; Dorsalschild glatt.

Wirt: Barbastella barbastellus

Präparate aus der Sammlung Kneissl

1. [Präparat Nr.: K 289] 28.05.1910 Pteroptus sp. - 1? von Pipistrellus sp., Zwergfledermaus, Oberalting, leg. Kneissl, det. Kneissl, [B], det. H. Ferenc / 2000

Präparate aus der Sammlung Vitzthum 1. [Präparat Nr.: V 1149] 09.04.1937 Spinturnix noctu- lae Oudemans, 1902 - 12 [A], det. H. Ferenc/ 2000

Spinturnix acuminatus (Koch, 1836)

Pteroptus acuminatus Koch, 1836 Spinturnix acuminatus Hirst, 1927 Pteroptus abominabilis Koch, 1836 Pteroptus vespertilionis Koch, 1839 Pteroptus abdominalis Walckenaer & Gervais, 1847 Pteroptus arcuatus Kolenati, 1857 Monostaspis arcuatus Kolenati, 1857 Diplostaspis arcuata Kolenati,1859 Monostaspis pipistrelli Kolenati,1859 Spinturnix carnifex Oudemans, 1902 Spinturnix noctulae Oudemans,1910 Spinturnix javensis Oudemans, 1914 Spinturnix sp. Haarlov, 1943

Diagnose

2: Länge des Idiosoma 1,01-1,22 mm, Breite 0,83- 0,98 mm; von den 3 Paar Sternalborsten liegt min- destens 1 Paar außerhalb des Sternalschildes; An- zahl der Opisthodorsalborsten 24-46; Länge der Podosomalborsten 54-68 um. Scutum bedeckt mit Schüppchen.

d: Länge des Idiosoma 0,94-0,99 mm, Breite 0,80- 0,81mm; Anzahl der Opisthodorsalborsten 18-21. Dorsalschild mit punktförmiger Skulptur.

Wirt: Nyctalus noctula

Präparate aus der Sammlung Vitzthum 1. [Präparat Nr.: V 1151] 02.11.1932 Spinturnix muri- nus (Walkenaer, 1847) - 18 [A], det. H. Ferenc/ 2000

Spinturnix psi (Kolenati, 1856)

Pteroptus psi Kolenati, 1856 Diplostaspis psi Kolenati,1857 Spinturnix psi Hirst, 1927

Spinturnix amboinensis Oudemans,1925

Diagnose

2: Länge des Idiosoma 0,79-1,06 mm, Breite 0,63- 0,82 mm; von den 3 Paar Sternalborsten liegt min- destens 1 Paar außerhalb des Sternalschildes; An- zahl der Opisthodorsalborsten 37-47. Scutum be- deckt mit quadratischen oder viereckigen Plättchen.

38

d: Länge des Idiosoma 0,62-0,75 mm, Breite 0,50- 0,61 mm; Peritremalschild dreieckig, die ganze Flä- che zwischen Coxa II und Coxa III bedeckend.

Wirt: Miniopterus schreibersi

Präparate aus der Sammlung Willmann

1. [Präparat Nr.: 63/3/1] - Spinturnix psi Klt., 39/75, det. Willmann - 18 [A], det. H. Ferenc/2000

2. [Präparat Nr.: 63/3/2] - Spinturnix psi Klt., 39/75, det. Willmann - 18 [A], det. H. Ferenc/ 2000

3. [Präparat Nr.: 63/3 /3] - Spinturnix psi Klt., 39/75, det. Willmann - 12 [A], det. H. Ferenc/2000

4. [Präparat Nr.: 63/3/4] - Spinturnix psi Klt., 39/75, det. Willmann - 12 [A], det. H. Ferenc/ 2000

Präparate aus der Sammlung Vitzthum 1. [Präparat Nr.: V1133] 29.09.31 - Spinturnix amboinen- sis Ouds., 1927 2 [A], det. C. Blaszak /2001

Spinturnix semilunaris De Meillon & Lavoipierre, 1944

Diagnose

Q: Länge des Idiosoma 0,84-0,90 mm, Breite 0,67- 0,70 mm; Sternalschild mit 3 Paar Borsten. Scutum bedeckt mit dreieckigen Schüppchen.

d: Länge des Idiosoma 0,64-0,74 mm, Breite 0,52- 0,56 mm; Peritremalschild dreieckig, die ganze Flä- che zwischen Coxa II und Coxa III bedeckend, Ster- nalschild endet mit einem deutlichen Fortsatz.

Wirt: Miniopterus schreibersi natalensis

Präparate aus der Sammlung Zumpt

1. [Präparat Nr.: III/96/2] 26.12. 60 - Spinturnix semilu- naris De Meillon & Lavoipierre, 1944 ? [B] von Miniopterus schreibersi, Rooiberg near Thabazinbi, det. Zumpt?

2. [Präparat Nr.: II/96/3] 26.12. 60 - Spinturnix semilu- naris De Meillon & Lavoipierre, 1944 2 [B] von Miniopterus schreibersi, Rooiberg near Thabazinbi, det. Zumpt?

Spinturnix plecotinus (Koch, 1839)

Pteroptus plecotinus Koch, 1839 Spinturnix plecotinus Oudemans, 1910 Pteroptus plecutinus Koch, 1839 Pteroptus transversus Kolenati, 1856 Diplostaspis transversus Kolenati, 1857 Diplostaspis transversa Kolenati, 1859 Spinturnix plecoti OQudemans, 1902

Diagnose

2: Länge des Idiosoma 0,97-0,98 mm, Breite 0,72- 0,77 mm; Anzahl der Podosomalborsten 5 Paar.

d: Länge des Idiosoma 0,85-0,93 mm, Breite 0,69- 0,73 mm; Anzahl der Podosomalborsten 5 Paar.

Wirt: Plecotus auritus

Präparate von H. Ferenc, Sammlung Lehrstuhl für Tier-

morphologie der Universität Posen (ZMZ)

1. [Präparat Nr.: ZMZ/F 272) 22.09.1976 Miedzy- chöd, Polen, leg. A. Nowosad - 1? [A] von Plecotus auritus, det. H. Ferenc/ 1999

2. [Präparat Nr.: ZMZ/F 253) 22.09.1976 Miedzy- chöd, Polen, leg. A. Nowosad - 14 [A] von Plecotus auritus, det. H. Ferenc/ 1999

Spinturnix mystacinus (Kolenati, 1857)

Diplostaspis mystacinus Kolenati, 1857 Spinturnix mystacinus Oudemans,1910 Diplostaspis mystacina Koleanti, 1859 Spinturnix mystacina Stile & Nolan, 1931 Spinturnix murinus Thompson, 1935

Diagnose

2: Länge des Idiosoma 0,93-1,06 mm, Breite 0,78- 0,94 mm; 3 Paar Borsten auf Sternalschild, das eine Länge von 155-167 um und eine Breite von 148-159 um hat. Vorderer Rand des Sternalschildes zuge- spitzt; Opisthodorsalborsten in der Anzahl von 70- 80 liegen maximal in drei Reihen. Scutum bedeckt mit feiner quadratischer oder viereckiger Skulptur.

d: Länge des Idiosoma 1,00-1,10 mm, Breite 0,83- 0,90 mm; drei Paar Borsten auf Sternalschild; vier- tes Paar liegt außerhalb des Sternalschildes, das eine Länge von 277-289 uım und eine Breite von 200- 215 um hat.

Wirt: Myotis mystacinus

Präparate von H. Ferenc (Sammlung ZMZ Posen)

1. [Präparat Nr.: ZMZ/F 804] 18.09.1999 Jaskinia Wielka Litworowa, Tatra Gebirge, Polen, leg. K. Pik- sa - 1? [A] von Myotis mystacinus, det. H. Ferenc/ 1999

2. [Präparat Nr.: ZMZ/F 88] 22.09.1976 - Miedzychöd, Polen, leg. K. Piksa - 13 [A] von Myotis mystacinus, det. H. Ferenc/ 1999 _

Gattung: Paraperiglischrus Rudnick, 1960 Paraperiglischrus rhinolophinus (Koch, 1841)

Pteroptus rhinolophinus Koch, 1841 Periglischrus rhinolophinus Murray, 1877 Periglischrus rhinolophus Walckenaer & Gervais, 1847 Pteroptus euryalis Berlese, 1889

Pteroptus interruptus Kolenati, 1856 Periglischrus interruptus Kolenati, 1857 Pteroptus hipposideros Kolenati, 1856 Periglischrus hipposideros Kolenati, 1857 Periglischrus asema Kolenati, 1857

Periglischrus glutinimargo Kolenati, 1857 Periglischrus africanus Zumpt, 1950 Periglischrus africana Hiregaudar & Bal, 1956 Periglischrus rhinolophi Hiregaudar & Bal, 1956

Diagnose

2: Länge des Idiosoma 0,52-0,76 mm, Breite 0,28- 0,43 mm; kurze Peritremen, ganz auf der Dorsalsei- te liegend, 2 Dorsalschilde.

d: Länge des Idiosoma 0,33-0,43 mm, Breite 0,25- 0,35 mm; kurze Peritremen, ganz auf der Dorsalsei- te liegend, 2 Dorsalschilde.

Wirt: Rhinolophus hipposideros

Präparate aus der Sammlung Vitzthum 1. [Präparat Nr.: V 1192] 09.04.1937 Spinturnix sp. 13 [B], det. H. Ferenc / 2000

Präparate aus der Sammlung Willmann

1. [Präparat Nr.: W 58/1] 16.03.40 - Periglischrus inter- ruptus, det. C. Willmann, Fl. 12/42/44, (Kiechen- weg?) Höhle, Streitberg. Kl. Hufeisennase 2, det. Willmann, [B], det. C. Blaszak /2001

Präparate aus der Sammlung Zumpt

1. [Präparat Nr.: III/96/6] 08.06.50 - Periglischrus afri- canus Zpt. -— 2 [A] von Rhinolophus geoffroyi, Sterk- fontein Caves, Transval, det. Zumpt?, det. C. Blas- zak/2001

2. [Präparat Nr.: III/96/7] - Periglischrus africanus Zpt. - 2 [A] von Rhinolophus geoffroyi, Sterkfontein Caves, Transval, det. Zumpt?, det. C. Blaszak/ 2001

3. [Präparat Nr.: III/96/8] 15.06.58 Periglischrus afri- canus Zpt. - 22? [A], 2 DN von Rhinolophus blasii empusa, Rooiberg, Transval, det. Zumpt?, det. C. Blas- zak/2001

4. [Präparat Nr.: P517/1] 15.07.64 Paraperiglischrus africanus Zpt. - 12 [A] von Rhinolophus blasii, Rooi- berg, Transval, S. Africa, det. Zumpt (E. Kat. 573), det. C. Blaszak/2001

Paraperiglischrus moucheti (Till, 1958)

Diagnose

2: Länge des Idiosoma 0,67-0,73 mm, Breite 0,43- 0,48 mm; Peritremen kurz, ganz auf Dorsalseite lie- gend, zwei Dorsalschildchen; alle Sternalborsten außerhalb des Mediodorsalschildes; Borsten auf Femur I länger als Breite des Femurs.

d: Länge des Idiosoma 0,39-0,43 mm, Breite 0,35- 0,40 mm; Peritremen kurz, ganz auf der Dorsalseite liegend, 2 Dorsalschildchen; alle Sternalborsten au- ßerhalb des Mediodorsalschildes; Borsten auf Fe- mur I länger als Breite des Femurs.

Wirt: Hipposideros caffer guineensis

Präparate aus der Sammlung Zumpt

1. [Präparat Nr.: III/96/4] 12.04.55 Periglischrus moucheti Till-422 [A] von Hipposideros cafer, Yooun- de, Cameroons, det. Zumpt?

2. [Präparat Nr.: III/96/5] 12.04.55 Periglischrus moucheti Till - d [A] von Hipposideros cafer, Yoounde, Camero- ons, det. Zumpt?

39

Gattung Eyndhovenia Rudnick, 1960 Eyndhovenia euryalis (Canestrini, 1884)

Pteroptus euryalis Canestrini, 1884

Spinturnix euryalis Oudemans, 1902

Periglischurus interruptus Oudemans, 1910 Spinturnix oudemansi van Eyndhoven, 1941 Spinturnix omahonyi Turk, 1945

Spinturnix omahonyi Turk, 1953

Spinturnix viduus Zumpt, 1950

Spinturnix euryalis orientalis Hiregaudar & Bal, 1955

Diagnose

2: Länge des Idiosoma 0,36-0,66 mm, Breite 0,35- 0,50 mm; Peritremen ganz der auf Dorsalseite, Bei- ne I mit Pulvillen. 1 Dorsalschild, 6 Paar Podoso- malborsten, die den Dorsalschild umgeben.

d: Länge des Idiosoma 0,36-0,56 mm, Breite 0,36- 0,50 mm; Peritremen ganz auf der Dorsalseite, Bei- ne I mit Pulvillen. 1 Dorsalschild, 6 Paar Podoso- malborsten, die den Dorsalschild umgeben.

Wirt: Rhinolophus ferrumequinum

Präparate aus der Sammlung Willmann 1. [Präparat Nr.: 62/10] - Spinturnix euryalis Can. - 19, 39/77 [A], det. H. Ferenc/2000

Präparate aus der Zumpt Sammlung

2. [Präparat Nr.: III/96/1] 01.06.50 - Spinturnix viduus Zpt. - [A] von Rhinolophus geoffroyi, Sterkfontein Caves, Transval, d, det. Zumpt?, det. C. Blaszak /2001

Unbestimmte Präparate

Einige Präparate sind nicht bestimmbar, weil:

- für Jugendstadien keine Beschreibungen und kein Bestimmungsschlüssel vorliegen (])

- sie bereits völlig zerstört sind (Z)

- wegen zu großer Variabilität eine Artabgren- zung nicht möglich ist (V)

- wichtige taxonomische Merkmale nicht erkenn- bar sind (T)

Präparate aus der Sammlung Willmann

1. [Präparat Nr.: W 62/11] Sp. - 509 Spinturnix murinus —- 1? [B], det. C. Willmann, [V], [T]

2. [Präparat Nr.: W 62/12] - Spinturnix murinus Walk. 2 [A], det. C. Willmann, [V], [T]

3. [Präparat Nr.: W 62/13] - Spinturnix murinus Walk., Segeberg, Höhle - 233 [A] - 22? [A] von Myotis myotis, det. C. Willmann, [V], [T]

4. [Präparat Nr.: W 63/6/1]- Spinturnix sp. - 12 Deuto- nymphe [B], det. C. Willmann, [J]

5. [Präparat Nr.: W 63/6/2]-Spinturnix sp.- 18 Deuto-

nymphe [B], det. C. Willmann, [J]

[Präparat Nr.: W 63/7] - Spinturnix sp. 1 Proto-

nymphe, 40/49, [Z], det. C. Willmann, [J]

en

40

7. [Präparat Nr.: W 63/8] - Spinturnix sp. 1 Proto- nymphe, 42/83 [A], det. Willmann, [J]

8. [Präparat Nr.: W 63/ 14/4] - Spinturnix sp. - 1 Proto- nymphe [Z], leg. Vornatscher, det. C. Willmann, [J]

Präparate aus der Sammlung Vitzthum

9. [Präparat Nr.: V 1131] 27.12.1930 - Spinturnix araguen- sis Vitzthum, 1931 - 12 [A], von Myotis 'nigricans, Hacienda Puerto de la Cruz, westl. La Guaira, Vene- zuela, leg. Hase, det. Vitzthum -— SYNTYPUS, [T]

10. [Präparat Nr.: V 1132] 09.06.1920 Spinturnix javen- sis Ouds.,1914, Buitenzorg/Java, Fledermaus, leg. Sieber, det. Vitzthum. 05.08.1923, 222 [das ist keine Spinturnix acuminatus (Koch, 1836) nach Rudnick 1960 (H. Ferenc/2000)]

11. [Präparat Nr.: V 1155] Spinturnix mystacinus Kole- nati, 11.10.1910 - 1 Protonymphe [B]

12. [Präparat Nr.: V 1152] 11.10.1910 Spinturnix mysta- cinus (Kolenati) Ny., []]

13. [Präparat Nr.: V 1153] 13.10.1910 - Spinturnix mysta- cinus (Kolenati) 1 Ny., []]

14. [Präparat Nr.: V 1154] 13.10.1910 - Spinturnix mysta- cinus (Kolenati) Ny., J]

15. [Präparat Nr.: V 1156] 02.11.1932 Spinturnix muri- nus (Walkenaer, 1847) -— DN-Weib,, []]

16. [Präparat Nr.: V 1159] 04.10.1934 Spinturnix muri- nus (Walkenaer, 1847) - 2 Ny., []]

17. [Präparat Nr.: V 1160] 22.09.1931 - Spinturnix javen- sis Oudemans, 1914-12 [B], [das ist keine Spinturnix acuminatus (Koch, 1836) nach Rudnick 1960 (H. Fe- renc/2000)]

18. [Präparat Nr.: Zool. V 1161] 22.09.1931 Spinturnix javensis Oudemans, 1914 13 [A], [das ist keine Spinturnix acuminatus (Koch, 1836) nach Rudnick 1960 (H. Ferenc/2000)]

19. [Präparat Nr.: V 1162] 22.09.1931 Spinturnix javen-

sis Oudemans, 1914-12 [Z], [das ist keine Spinturnix

acuminatus (Koch, 1836) nach Rudnick 1960 (H. Fe- renc/2000)]

Präparat Nr.: V 1165] 17.04. 1927 Spinturnix sp. 6, VJ IT]

Präparat Nr.: V 1169] 28.01. 1936 Spinturnix sp. 9, vJ [7]

20. [ [ l l 22. [Präparat Nr.: V 1184] 09.11.1928 Spinturnix sp. 9, | | [

al

VJ IT]

Präparat Nr.: V 1185] 13.11.1928 Spinturnix sp. ®,

VJ[T]

24. [Präparat Nr.: V 1186] 28.03.1929 Spinturnix sp. 6, [VL IT]

25. [Präparat Nr.: V 1187] 13.11.1928 - Spinturnix sp. ®, [VL IT]

26. [Präparat Nr.: V 1190] 04.06.1934 Spinturnix sp. ®, [VL IT]

27. [Präparat Nr.: V 1193] 17.04. 1927 - Spinturnix sp. ®, [VL IT]

28. [Präparat Nr.: V 3301] 05.05.1909 - Spinturnix vesperti- lionis (L.), 1 Protonymphe [A], det. Furman/ 1968, [J]

28:

Präparate aus der Sammlung Kneissl 1. [Präparat Nr.: K 288] 28.05.1910 - Pteropus sp. Zwerg- fledermaus, Oberalting, leg. Kneissl, det. Kneissl

Literatur

Deunff, J. 1977. Observations sur les Spinturnicidae de la Region Palearctique occidentale (Acarina: Meso- stigmata): Specificite, repartition et morphologie. Acarologia 18(4): 602-616

-- ‚Keller, A. & V. Aellen 1997. Redescription of Spin- turnix punctata (Sundevall, 1833) a specific parasite of Barbastella barbastellus (Chiroptera Vespertilioni- dae). Nat. Hist. Mus. Geneva 1: 199-206

Dusbabek, F. 1964. Parasitische Fledermausmilben der Tschechoslowakei. II. Familie Dermanyssidae Kol., 1859 (Acarina: Gamasides). Cs. Parazitol. 11: 77-125

Evans, G. ©. 1968. The external morphology of post- embryonic developmental stages of Spinturnix myo-

ti Kol. (Acari: Mesostigmata). Acarologia 10(4): 589-608

Estrada-Pena, A. & C. Sanchez 1989. Redescription of Spinturnix dasycnemi (Kolenati) (Acarina, Spintur- nicidae). —- Acarologia 30: 107-110

Rudnick, A. 1960. A Revision of Mites of the Family Spinturnicidae (Acarina ). - Univ. Calif. Publ. En- tom. 17: 157-284

Uchikawa, K. 1979. Studies on mesostigmatic mites pa- rasitic on mammals and birds in Japan. VII. Bat mites of the genus Paraperiglischrus Rudnick, 1960 with descriptions of the males of Paraperiglischrus sternalis Petrova & Taskaaeva, 1975, and Paraperig- lischrus hipposideros Baker & Delfinado. Acarolo- gia 21: 9-17

41

Buchbesprechungen

3. Simovich, M. A., C. Sassaman & D. Belk (eds.): Stu- dies on Large Branchipod Biology and Conservati- on. In: Dumont, H. ]. (ed.): Developments in Hydro- biology 125. (Reprinted from Hydrobiologia, Vol. 359). - Kluwer Academic Publishers, 1997. pp. XI + 245. ISBN 0-7923-4966-0.

Der vorliegende Band umfaßt 28 Beiträge, die sich mit den großen branchiopoden Crustaceen (Anostraca, No- tostraca und Conchostraca) beschäftigen und die auf dem Third International Large Branchiopod Symposi- um (ILBS - 3) vom 15.-18.Juli 1996 an der University of San Diego, CA, USA vorgestellt wurden. 9 Beiträge zur Regionalfauna von Südafrika, Mexico, den USA, der Region an der Barentssee, Österreich und dem Balkan leiten ein. Daran schließen sich 4 Beiträge zur Taxono- mie und Systematik. 5 Beiträge zur Genetik und Evolu- tion, 6 Beiträge zur Ökologie und 3 Beiträge zur Aqua- kultur zeigen die Vielfalt der Forschungsvorhaben, die auf diese interessante Tiergruppe eingehen. Für Wissen- schaftler und Studierende, die sich der Erforschung der Branchiopoda widmen, ist dieser Symposiumsband un- verzichtbar. L. Tiefenbacher

4. Nietzke, G. (unter Mitarbeit von P. M. Kornacker): Die Terrarientiere 3. Krokodile und Schlangen (4. neu bearbeitete und neugestaltete Auflage). - Eugen Ul- mer Verlag, Stuttgart, 2002. 375 S., 129 Farbphotos, 35 Zeichnungen und Verbreitungskarten. ISBN 3-8001-7459-6.

Der vorliegende dritte und letzte Band der “Terrarientie- re” schließt die schon 1989 begonnene vierte Auflage dieses Klassikers ab. Aufbau und Gliederung des Schlan- genbandes folgen dem schon in den ersten beiden Teilen (Amphibien; Echsen, Schildkröten) bewährten Muster: Nach einführenden Kapiteln - hier u.a. zu rechtlichen Problemen (der Bedeutung entsprechend nicht als klei- ner Appendix irgendwo versteckt, sondern gleich auf den ersten. Seiten), Entwicklungsgeschichte, Anatomie, Morphologie, Verhalten, Fortpflanzung und Haltung folgen Artkapitel, die neben Beschreibungen und Anga- ben zur Biologie ausführliche terraristische Informatio- nen enthalten. Abgeschlossen werden die Artkapitel je- weils von einem eigenen, kleinen Literaturverzeichnis, das die Übersichtlichkeit wesentlich erhöht. Schließlich folgen Kapitel über Krankheiten, Präparation, Literatur- beschaffung, ein Fachbuchverzeichnis, eine Übersicht über herpetologisch-terraristische Gesellschaften und Vereine und deren Publikationsorgane, ein Glossar so- wie Register.

Die Texte des Schlangen-Haupteils aus der Feder von Paul Kornacker sind durchweg von hoher Qualität

42

und, trotz der langen Vorlaufzeit bis zum Erscheinen des Buches, in den allermeisten Fällen auf dem neuesten Stand. Allein bei der Durchsicht der kurzen Literatur- verzeichnisse der Artbearbeitungen fällt auf, welche immense Informationsfülle hier berücksichtigt wurde. Aufgeführt werden nämlich nicht nur terraristische Ar- beiten, sondern Schlüsselzitate aus dem gesamten Spek- trum der betreffenden wissenschaftlichen Literatur. Dem- entsprechend lassen die Artkapitel meist keine Wünsche offen: Die terraristisch relevanten Informationen umfas- sen die Aspekte Terrariengröße- und Ausstattung, Hei- zung, Licht und Luftfeuchtigkeit, Futter, Nach- und Aufzucht sowie Verträglichkeit.

Man mag an der Ausgabe kritisieren, daß die An- zahl der Bilder noch reichhaltiger hätte sein können. Tatsächlich findet sich lange nicht auf jeder Seite eine Photographie, und oft sind die Abbildungen dann eher klein geraten. Ich sehe in der etwas zurückhaltenden Bildauswahl jedoch durchaus eine der großen Stärken des Werkes. Terraristische “Bilderbücher” sind derzeit zuhauf auf dem Markt und in den meisten dieser Fälle geht eine üppige Bildpräsentation auf Kosten der Texte, die oft mit heißer Nadel gestrickt, beliebig und belanglos daherkommen. Der “Nietzke” folgt dagegen seiner Tra- dition als terraristisches Standardwerk und setzt hier eindeutig den Schwerpunkt auf Information. Genau aus diesem Grund wird das Buch auch noch in vielen Jahren in Gebrauch sein, während sich andere Terrarienbücher als Eintagsfliegen entpuppen werden.

Naturgemäß kann an der Auswahl der vorgestellten Arten immer kritisiert werden, da man es hier, je nach Sichtweise und persönlichen Präferenzen, kaum jemals allen wird recht machen können. Ich möchte aber trotz- dem folgendes zu bedenken geben: Bei einigen Arten hat man den Eindruck, daß sie seit der ersten Auflage 1972 nur noch aus reiner Tradition mitgeschleppt werden. Ein solcher Fall ist zum Beispiel die Kreuzotter. Die Art spielt heute in der Terraristik praktisch keine Rolle mehr, während der betreffende Text im Buch aber mit etwa vier Druckseiten zu den längeren Artbearbeitungen ge- hört. Auch muß an dieser Stelle erlaubt sein, darüber nachzudenken, ob es zu rechtfertigen ist, ineinem Grund- lagenwerk mit extremer Breitenwirkung für solche Ar- ten zu “werben”, die nur ein sehr kleines Areal aufwei- sen und schon seit Jahren in der Natur einem hohen Sammeldruck ausgesetzt sind (wie z.B. die Kaukasusot- ter, V. kaznakovi).

Diese kleineren Kritikpunkte sollten aber nieman- den ernsthaft terraristisch Interessierten von der An- schaffung des Buches abhalten." Nach wie vor gibt es kein umfassenderes und genaueres Terrarienbuch als “den Nietzke”. M. Franzen

SPIXIANA

München, 01. März 2003

ISSN 0341-8391

Polythore spaeteri, spec. nov. from the Peruvian tropical rainforest (Panguana), with remarks on its ecology

(Odonata, Zygoptera, Polythoridae)

Ernst-Gerhard Burmeister & Läszlo Börzsöny

Burmeister, E. G. & L. Börsöny (2003): Polythore spaeteri, spec. nov. from the Peruvian tropical rainforest (Panguana), with remarks on its ecology (Odonata, Zygoptera, Polythoridae). - Spixiana 26/1: 43-48

A new species of Polythore from the area of Panguana (Prov. Huanuco, Peru) is described and compared with species of the groups of Polythore (Bick & Bick 1985, 1986, 1990a, 1990b). The coloration of wings in males and females and the structure of the penis differ from all other species. The observation of males and females in tandem or copula can open some aspects into the aquatic biotopes of the larvae. Larvae of Polythore have not been described.

Dr. Ernst-Gerhard Burmeister, Zoologische Staatssammlung München, Münch- hausenstraße 21, D-81247 München, Germany

Läszlö Börzöny, Cserenyhegyi U. 30, H-8233 Balatonszölös, Hungary

Introduction

Montgomery (1967) presented a list of Polythoridae species, an Odonata family with an exclusively Ne- otropical distribution, including 16 species of the genus Polythore. Later, a revision of the picta-group and the other species of this genus followed (Bick & Bick 1985, 1986, 1990a, 1990b) that revealed two additional species. Thus, the picta-group now com- prises six species, including one new description, the major group of the remaining ones comprises ten species split up in five groups. The descriptions of Polythore neopicta (Bick & Bick, 1990) from the area of Tingo Maria (Peru) at the foot of the Andes, and Polythore manua (Bick & Bick, 1990) from Manu National Park (Dept. Madre de Dios, Peru) show, that the recording of species is still by far not nearly finished, and not only as far as that group is con- cerned.

Five species of the genus Polythore (neopicta, lamerceda, ornata, victoria, manua) are only known from Peru. These and further undescribed species

have been found in material of the museum in Lima (Peru) by L. Börzsöny, but until now, only other specimens collected by E.-G. Burmeister in 2000 could be worked. Field observations and the catch- ing of a copulating pair has proved especially use- ful. Females proved to be relatively easily distin- guishable from other species, a remarkable fact, because otherwise females are notoriously difficult to identify in this genus. The new species of the genus Polythore, recorded from the mouth area of the Rio Llullapichis and the Pachitea, a tributary of the river Ucayalli near the foot of the Andes (Peru) and described here, certainly is not closely related to the other species, because it shows some special characteristics.

The conspicuous larvae of this dragonfly family are not yet described from anyone of the species of the genus Polythore. Only larvae of the genus Cora in the family Polythoridae were described. The mating places and the timing of distribution on the waters, as well as emergence patterns, possibly indicate habitat and larval ecology.

43

Fig. 1. Polythore spaeteri, spec. nov., allotype female (top), holotype male (bottom) (photo M. Müller, ZSM).

Polythore spaeteri, spec. nov. Figs 1-5

Types. Holotype: d, Peru, Panguana, Rio Llullapichis (Yuyapichis), rechter Nebenfluß des Rio Pachitea, 37'5/74°56'W, 28.09.-06.10.2000, leg. Burmeister, ZSM (Zoologische Staatssammlung München). Allotype: 9, same data (ZSM, d holotype and 2 allotype collected in copula). Paratypes 8d, 32, same data: 68, 22 in ZSM (13 and 1% preserved in alkohol), 18, 1? in MSM (Museo de Historia Natural, Lima, Peru), 18, 12 in col. L. Börzsöny; 13 Panguana, Rio Llullapichis, 220 m, Nov.- Dec. 1987, leg. K. Voß (MSM).

Etymology. Named to honour a person for his sponsor- ship of biological investigations at the biological obser- vation center of Panguana (Prov. Huanuco, Peru).

44

Description

Male (holotype).

Length of abdomen, excluding appendages, 41 mm, hind wing length 38 mm, maximal width 11.3 mm.

Head. Labrum yellowish to orange narrowly bordered by black, anterior border and midline black; postelypeus black with a minute pale spot on either side, distal membrane dark brown; vertex with 4 yellow to orange spots, which are rectangularly ar- ranged, front edges also yellowish; Labium pale yellowish, mouthparts shiny black.

Prothorax. Dorsally and laterally largely black, shading into pale beneath; broad transverse orange to yellowish band on the middle lobe, medially interrupted.

Fig. 2. Polythore spaeteri, spec. nov., female (paratype) (photo M. Müller, ZSM).

Pterothorax. Black with 5 orange to yellowish stripes arranged in the typical manner described by Bick & Bick (1985) in Polythore lamerceda. Legs black, inner side of femora pale.

Abdomen. Black with lateral yellow markings on each side as described by Bick & Bick (1990) in Polythore neopicta. Ventral area of pterothorax and abdomen (basal segments) covered with bluish pat- terns like hoarfrost.

Appendages (Fig. 4). Black, length 2.05 mm, with a mid-length ventromedial process.

Penis (Fig. 3). Terminal segment (horns) length c. 0.16 mm, weakly divergent, flagella not totally visible, in paratypes long and 2-segmented.

Wings. Length of hind wing (hw) 38 mm, fore wing (fw) 42 mm, base to nodus fw 15.5/hw 14 mm, nodus to pterostigma along costal margin fw 19.5/ hw 18.5 mm, Petiole fw 2.6/hw 3.0 mm. Pterostig- ma dark brown, fw 5.5/hw 4.8 mm along posterior border, surmounting left fw 20+(right fw 19)/left hw. 16+(right hw 16) cells. Antenodals fw 54 (54)/ hw 37, the fw 16. (18.)/hw 12. (15.) thickened; post- nodals fw 69 (70)/hw 67. Basal space crossed by fw 16 (17)/hw 15 crossveins, the quadrangular by fw 8 (8)/hw 11 (11) crossveins. Membrane transparent amber, nearly uniform throughout somewhat light-

ened at base, wings at apex (ca % ofthe winglength) very diffusely darkened to brownish.

Female (allotype).

Length of abdomen, excluding appendages, 33 mm, hind wing 34 mm, fore wing 36 mm, maxi- mal width 10.5. Head, prothorax, pterothorax col- our patterns as in male holotype. Abdomen black with yellow-orange marks on each side. The yellow

Fig. 3. Terminal segment of penis of Polythore spaeteri, spec. nov. under medially fused genital lobes, ventral view: holotype (left); paratype (right), scale bar = 0.1 mm.

45

Fig. 4. Polythore spaeteri, spec. nov., last segments of abdomen: a. male (lateral view); b. male (dorsal view); c. female (lateral view) (figured by R. Kühbandner, ZSM).

marks all darker than in holotype, segment I with yellow area also more ventrally, segment V with spot and long thin yellow stripe extending over nearly the whole length.

Abdomen (Fig. 4.). Apendages black, gonapo-

physis distally brownish transparent. Wings. Both wings basally and apically light trans- parent brown with a greenish-bronze metallic shine. In the area of the last third of wings (postnodal area about fw 21.0-26.5/hw 22.0-28.0 mm from base of wing) a band of dark brown is spread across the whole width (Fig. 2). This pattern is homogenous in colour and greenish metallic shine - especially from the underside and in living specimens (Fig. 4.), there is no shadow of this marking distally. In front of the dark band there is a small transparent area. Ptero- stigma chocolate brown, cells under Piseeukyne fw 15+/hw12+ (Fig. 2).

Variation. Body markings in all males and fe- males generally similar. Essential differences only in the lateral pointed and stretched markings of abdominal segments I-V, of lateral stripes of ptero- thorax, and in basal yellowish area of femora. The wing bands in females are mostly located in the same area (venation: 26-28/44-47 postnodal veins). Penis horns variable in length by 0.15-0.16 mm. Wings of females differ in the width of the pale area proximal of the dark band in the distal area between nodus and pterostigma (Fig. 5). Immature colour patterns or different morphs especially in females are unknown.

46

Diagnosis and discussion

The characteristics of the species of the genus Poly- thore have been explained by Fraser (1946, 1957) and Montgomery (1967). Most of them are characters of wing colouration and male secondary genitalia. The width, length ratio, position of'the first visible thick- ened antenodals and the number of veins are very similar and often “obviously not useful in separat- ing the species” (Bick & Bick 1985).

Species of Polythore are characterized by:

1. the overall black body colour with pale mark- ings in both sexes;

2. the absence of inferior abdominal appendages in males;

3. males with uniformly black superior appendag- es each with a conspicuous process extending ventro-medially at mid length;

4. males with lateral flagella and terminal horns on the terminal segment of the penis;

5. females with dark, elongate, triangular mesostig- mal lamina, pointed ventrally and rimmed on all sides by a different elevation.

Diagnosis of Polythore spaeteri, spec. nov. in compar- ison:

There are only two species with uniformly col- oured wings in the male, P. williamsoni and P. concin- na.

Male wings of Polythore spaeteri nearly uniform transparent amber, with the apical area diffusely

darkened to brown area. Apart from this darkening there are no markings such as lunules, etc. The base of the wings with lighter coloration like in other species. In P. concinna (McLachlan, 1881) there are amber wings and absolutely no other pattern, nota- bly no apical darkening, in P. williamsoni (Foerster, 1903) the fore and hind wings are smoky gold with a very obscure pale lunule and a transparent basic area (Bick & Bick 1986).

Penis horns of Polythore spaeteri are nearly straight, divergent but less so than in P. concinna. In contrast to P. concinna the tips are rounded more like in P. aurora (Selys, 1879) or P. boliviana (McLach- lan, 1878). The flagellum of P. spaeteri is long and roundly bent and clearly 2-segmented in contrast to P. concinna.

Polythore concinna is common in Ecuador, but three specimens are recorded from Peru, one male without locality date deposited in Paris, one male from Pasco: Chucharas leg. P. Martin in British Museum, another single male from Junin: Camino del Pichis, leg. W. T. M. Forbes, VII.1920 in Michi- gan Museum. Because in the subadult males of the new species the apical darkening may not have developed, it seems possible that these three Peru- vian records belong to this new species. One male is recorded from Pasco, Obda. Castilla, NW de Iscoz- acin, 345m, 10°10'S, 75°15'W, 3.X1.86, P. Lozada (MSM), which belongs to the new species. There is a considerable gap between the distribution areas of P. concinna and P. spaeteri.

Female fore and hind wings are transparent light amber with a broad, dark brown transversal band midway between nodus and pterostigma. There is no shadow or lunule of other colour distal- ly of this band. This clearly differs from all other species of the genus Polythore especially in compar- ison with the position of the wing band in P. concin- na and P. williamsoni. ö

Habitat and distribution of Polythore spaeteri, spec. nov.

The new species of Polythore is recorded from the area of Panguana in the rainforest of Peru (Amazo- nia) near the basis of the Andes mountains at 250 m (Koepcke 1987). Directly north of the building at Panguana there is the Rio Llullapichis (Yuyapichis), flowing into the Rio Pachitea. In the northeast the Sira-Mountains rising up to 3000 m are a centre of endemic species. The area of Panguana is covered with primary rain forest typically with light under- growth, in the south there exists a path through this protected area of 2 square kilometers. Imagines of Polythore spaeteri, spec. nov. have been collected near

small dry brooks (quebradas, flowing only during the wet season) or furrows without water. Only very small puddles existed in the dry season, which were muddy and warm, and at a level of 8-10 me- ters above the Rio Llullapichis. There is no direct drainage into the river. No specimens of P. spaeteri could be observed at the banks of the perennial small brooks of this area. With the beginning of a short rainy period on 5. October 2000, individuals of P. spaeteri could be observed in tandems and copula positions (Fig. 4). It seems that copulation and ovi- position take place after a longer raining period when the brooks have been filled with water. This area has two rainy seasons per year with variable onset. No specimen of Polythore could be found during a visit in 1982 (6.-20. July) at the same local- ities. No other species of Polythoridae is reported from Panguana. The revisions and descriptions of species of Polythoridae by Bick & Bick (1985, 1986, 1990a, 1990b) include no information about copula- tion or oviposition. The larvae of Polythore species are unknown. Most records of imagines exist from short-term observations, only few were carried out all over the year, and the distribution areas reach from 100 to 2800 m (P. concinna). Aspects of territo- rial and reproductive behaviour of Cora, a genus related to Polythore, are given by Fraser & Herman (1993).

The larvae of Polythoridae are marked by ven- tral gills at the abdomen as in Euphaeidae. This may be a plesiomorphic character of the ancestral Odo- nata. This character allows the larvae to exist in aquatic biota with changing oxygen levels, especial- ly due to alternating rain and dry seasons.

Polythore spaeteri, spec. nov. is recorded from the type locality and also known by one male from Pasco, Obda. Castilla, NW de Iscozacin, 345 m, 1010 S, 7515 W, 3. XI. 86, leg. P. Lozada (MSM). This location is near Rio Palcazu which unites with Rio Pichis and both together form the Rio Pachitea. This single specimen is not included in the type series; it is much smaller, but otherwise indistinguishable from the same species.

Resumen

Se describe una nueva especie de Polythore del area de Panguana (Depto. de Huänuco, Peru), la cual se com- para con los otros grupos de Polythore (Bick & Bick 1985, 1986, 1990a, 1990b). La coloraciön de las alas de machos y hembras y la estructura del pene se distinguen de todas las otras especies. Se pudo observar machos y hembras en tändem o cöpula, lo que permite cierta infor- maciön sobre algunos aspectos de los biotopos acuaticos colonizados por las larvas. Las larvas del genero Poly- thore aün no han sido descritas hasta la fecha.

Fig. 5. Polythore spaeteri, spec. nov., male (holotype) and female (allotype) in copulation (photo E.-G. Burmeister).

Acknowledgements

Our thanks are due to Dr. J. Hoffmann (MSM) for the kind loan of the Polythore specimens. We are very grate-

Fig. 6. Locality of Polythore spaeteri, spec. nov. (copula) in the rain forest of Panguana after first rain (photo E.-G. Burmeister).

ful to Mr. Martin Spiess for reading and correcting the English version of the manuscript. Special thanks are due to Dr. Juliane Diller who kindly translated the ab- stract into Spanish.

References

Bick, G.H. & J. C Bick 1985. A revision of the picta group of Polythore, with a description of a new species, P. lamerceda, spec. nov., from Peru (Zygoptera: Polythoridae). - Odonatologica 14: 1-28

-- 1986. The genus Polythore exclusive of the picta group (Zygoptera: Polythoridae). - Odonatologica 15(3): 245-273

-- 1990a. Polythore neopicta, spec. nov. from Peru (Odonata: Polythoridae). - Opusc. zool. flumin. 49: 1-7

-- 1990b. Polythore manua, spec. nov. from southern Peru (Zygoptera: Polythoridae). -— Odonatologica 19(4): 367-373

Fraser, F. C. 1946. Notes on Amazonian Odonata in the Leeds Museum. Trans. R. ent. Soc. Lond. 96: 11-46

-- 1957. A reclassification of the order Odonata. - R. zool. Soc. N.S.W., Sydney

-- &T.B. Herman 1993. Territorial and reproductive behavior in a sympatric species complex of the neotropical damselfly Cora Selys (Zygoptera: Poly- thoridae). - Odonatologica 22(4): 411-429

Koepcke, H.-W. 1987. Gründungsgeschichte und Bes- timmung des Studiengebietes Panguana. - ÖGH- Nachrichten 12/13: 1-24

Montgomery, B. E. 1967. Studies in the Polythoridae. Acta biol. venez. 5: 123-158

Verhaagh, M. 1986. Panguana Wald und Wasser im tropischen Südamerika. Führer zu Ausstellun- gen, 8; Museum am Friedrichsplatz Karlsruhe

Zöckler, C. 1995. Panguana Naturkundliches Tage- buch aus dem peruanischen Regenwald. Bremen

oma 1]

49-55 München, 01. März 2003

ISSN 0341-8391

Abstracts and brief versions of some talks held at the Workshop: “Systematics and Biogeography of Tenebrionoidea” at Zoologische Staatssammlung München, 14.-15.3.2002

The Palorus Group - a new subfamily of Tenebrionidae

(Insecta, Coleoptera)

Eric G. Matthews

Matthews, E. G. (2003): The Palorus Group - a new subfamily of Tenebrionidae (Insecta, Coleoptera). Spixiana 26/1: 49-50

Evidence is provided for the Palorus-group of genera to represent a separate subfamily of Tenebrionidae.

Dr. Eric G. Matthews, The South Australian Museum, Adelaide, S. A., 5000,

Australia

The higher classification of the Tenebrionidae is moving towards a consensus although there are still some points of disagreement. With regard to the Australian fauna I recognise three major divisions or branches with their included subfamilies as fol- lows: the lagrioid branch (Lagriinae, Phrenapatinae, Palorinae), the pimelioid branch (Pimeliinae, Zolo- dininae), and the tenebrionoid branch (Tenebrion- inae, Opatrinae, Alleculinae, Diaperinae and Coelo- metopinae, with Toxicini and Bolitophagini possi- bly being subfamilies).

Halstead (1967) recognised and revised an in- formal grouping which he called the Palorus genus group comprising seven genera characterised by 1) eyes entire, 2) antennae with only a poorly dif- ferentiated club, 3) scutellum transverse, 4) meten- dosternite without lamellae, 5) males with deep internal ventral abdominal pits, 6) elytra striate- punctate, and 7) wing venation with reduced anal area. He believed that the group was otherwise most closely related to Tribolium Macleay and its relatives, consequently the Palorus group has been placed within the tribe Triboliini, subfamily Tene- brioninae.

With our present better understanding of the characters on which tenebrionid higher classifica- tion is based, we can discern that the Palorus group has the following additional key character states:

8) asubquadrate labrum, 9) simple antennal sensil- la, 10) absent tentorial bridge, 11) unstriated man- dibular mola, 12) unarmed lacinia, 13) sometimes ten elytral striae, 14) an elytro-abdominal interlock- ing groove along the edges of the last three ven- trites, 15) absent aedeagal alae, 16) inverted aedea- gus, 17) very short basal piece of aedeagus, and 18) spermatheca not derived from bursa copulatrix. The Palorus group differs from Triboliini in all the characters listed except 6 and 9. Character states 8, 10, and 13-18 are unknown in the Tenebrioninae and rare in the tenebrionoid branch (14 is entirely unknown there). Conversely, character states 8-15 are common in the lagrioid branch.

It is clear that the Palorus group cannot be asso- ciated with Triboliini, Tenebrioninae, or even the tenebrionoid branch. It falls within the lagrioid branch, but the question then is should it be a tribe of Lagriinae or a subfamily in its own right? This is a difficult question because the lagrioid branch is probably a polyphyletic assemblage of taxa display- ing mainly primitive characteristics. The subfamily Lagriinae itself is defined mainly on a larval apo- morphy: pubescent two-segmented antennae, but the larvae of the lagriine Cossyphini and Chaerodi- ni are unknown. Palorine larvae have three-seg- mented antennae.

On the whole, it may be best to consider the

49

Fig. 1. Eutermiticola sculpticollis Lea, 1916. Habitus (left), underside of prothorax (right). Scale bar 1 mm.

Palorus group to be a subfamily, possibly the sister group of Phrenapatinae with which it shares char- acter states 1, 4, 7, 8,9, 12, 14, 15, 17 and 18. Phrena- patinae differs from Palorinae mainly in having a tentorial bridge, an antennal club, a striated molar surface, and absent defensive glands. Subfamily sta- tus for Palorinae is further supported by two ex- traordinary autapomorphies: the abdominal pits (character 5, not always present) and the inverted aedeagus (character 16). The former have not been observed in any other tenebrionids, and the latter is almost unknown outside the pimelioid branch. In fact, the aedeagus of Palorinae is identical to that of some pimeliines in both shape and orientation, but the presence of defensive glands makes it impossi-

50

ble to place the group in the pimelioid branch.

Palorinae have endemic genera in Madagascar, the Oriental Region and northern Australia, while Palorus is more widespread in Africa, Eurasia and the Pacific. The group does not occur naturally in the New World. The original seven genera included by Halstead are Palorus Mulsant, Coelopalorus Blair (shown by Scupola [2002] to be asynonym of Ulomi- na Baudi di Selve), Palorinus Blair, Prolabrus Fair- maire, Astalbus Fairmaire, Austropalorus Halstead and Pseudeba Blackburn. Subsequently Doyen et al. (1990) added Platycotylus Olliff, and I am adding Eutermicola Lea here (Fig. 1). In the checklist of Doy- en et al. (1990) Eutermicola was left in an uncertain position because at that time the characteristics of the Palorinae were not understood. In fact Eutermi- cola agrees with all the character states of the sub- family as listed above but it has some striking aut- apomorphies as well. It is a monotypic genus repre- sented by just three males collected in a termite nest prior to 1916 in north-western Australia (Fig. 1, up- per right), and never seen again. Pseudeba is also termitophilous and may be the most closely related genus.

References

Doyen, J. T., Matthews, E. G. & J. F. Lawrence. 1990. Classification and annotated checklist of the Aus- tralian genera of Tenebrionidae (Coleoptera). In- vert. Taxon. 3 (1989): 229-260

Halstead, D. G. H. 1967. A revision of the genus Palorus (sens. lat.) (Coleoptera: Tenebrionidae). Bull. Brit- ish Mus. Nat. Hist., Ent. 19: 61-148

Scupola, A. 2002. A proposito di Ulomina carinata Baudi di Selve, 1876 (Coleoptera, Tenebrionidae). Boll. Mus. reg. Sci. nat. Torino 19: 185-189

The systematic position of the ignote tribe Lachnogyini Reitter, 1904, with comments on the evolution of the aedeagus in the subfamilies Pimeliinae and Opatrinae

(Insecta, Coleoptera, Tenebrionidae)

Julio Ferrer

Ferrer, J. (2003): The systematic position of the ignote tribe Lachnogyini Reitter, 1904, with comments on the evolution of the aedeagus in the subfamilies Pimeliinae and Opatrinae (Insecta, Coleoptera, Tenebrionidae). Spixiana 26/1: 51-53

This paper deals with the systematic position of the Central Asiatic tribe Lachno- gyini, that was previously placed in the subfamily Tenebrioninae near the tribe Opatrini. A revision of available types belonging to the genera of this tribe indicates that the position of the Lachnogyini is within the subfamily Pimeliinae. The system- atic position of the genus Lachnodactylus Reitter, 1904, is in the subfamily Diaper- inae, tribe Trachyscelini. The evolution of the aedeagus in the subfamilies Pimelii- nae and Opatrinae is discussed and illustrated.

Dr. Julio Ferrer, Stora hundensgata 631, S-13664 Haninge, Sweden.

Introduction

The higher classsification of the Coleoptera Tene- brionidae has been discussed recently by Iwan (2001), who reestablished the validity of the subfamily Opatrinae sensu Mevdeved (1968) that was degrad- ed to tribal level by Doyen & Tschinkel (1982).

The tribe Lachnogyini was created by Reitter (1904) to receive two monotypic, central asiatic gen- era and species: Lachnogya squamosa Menetries, 1849, from Turkestan and Afghanistan and Netuschilia hauseri (Reitter, 1897) described under the preoccu- pied name Lachnopus, from Buchara. The tribe was posteriorly increased with another three genera: Kleweria colydiiformis Reitter, 1910 from Transcaspia and Lachnodactylus digitatus (Seidlitz, 1898) from Turkmenistan, previouly placed by Reitter (1904) in the Trachyscelini, and a synonym of the common African opatriine Phitammus sericans Fairmaire, 1870, redescribed as Canariella arenapta by Uyttenbogaart (1929) from the Canary Islands.

The representatives of this tribe are extremely rare in all collections and confined to the arid steppes of the Central Asian region from Transcaucasia, Turkmenistan, Afghanistan to the desert regions of China.

During a recent French expedition to a remote locality in the cold Desert of Taklamakan, in the Xing Kiang Province, China, a French archeologist, Mr. Sebastian Lepetz, found under an excavation a tenebrionid beetle which was sent to me for study by Mr. Jean Herve Yvinec, a French archeologist and practising entomologist.

I was puzzled searching for a systematic posi- tion of this hitherto unknown insect. The only pos- sible option, according to the available literature, was to place this species in the tribe Lachnogyini, which finally was revised under this research. The specimen from the French expedition to Xing Kiang was compared with type material of all previous known genera of the Lachnogyini and found to be a new representative of the tribe. However, this spe- cies, new for science, has to be placed in a new genus, that was created to receive it and will be described in a separate paper in preparation, to- gether with Mr. Jean Herve Yvinec.

In the first Catalogue of Tenebrionidae (Gebien 1910) the tribe Lachnogyiini was placed after Epi- tragini, between other tribes of the subfamily Ten- tyriinae, but, after several changes, all genera were placed by Reitter and Gebien in an own tribe, after the tribe Opatrini sensu Gebien (1939), and were placed at this time in the subfamily Tenebrioninae. Gebien (loc. cit.) noted that Lachnogyiini was con- sidered in his Catalogue a provisional group, com- posed by “heterogenere Gattungen”, which succes- sively were transferred by Reitter to very different tribes, e.g. Leptodini, Trachyscelini and Blaptini.

Several tribes from Gebien’s Catalogue (1939), were transferred by later authors to an own sub- family, the Opatrinae sensu Medvedev (1968). This classification was rejected by Doyen & Tschinkel (1982), was but finally adopted by Iwan (2001), who redefined the subfamily Opatrinae auct. based on the special configuration and position of the aedeagus.

51

Fig. 1. Derivative, apomorphic position of the male cop- ulatory organ in Pimeliinae. Aedeagus of Moluris muata Harold, from Namibia. Scale bar: 1.3 mm.

Fig. 2. Primitive, normal position of the male copulatory organ in Opatriinae. Aedeagus of Opatrum sabulosum Linne, from Italy. Scale bar: 0.7 mm.

Fig. 3. Derivative, apomorphic position of the male cop- ulatory organ in Lachnogyini. Aedeagus of Lachnogya squamosa Menetries. Scale bar: 0.6 mm.

Results

Examination of the aedeagus of the new genus and species from Taklamakan and of the genotypic spe- cies Lachnogya squamosa Menetries reveal that the choice of Reitter (1904) and Gebien (1939), who trans- ferred the genera Lachnogya, Netuschilia, and Klewe- ria to the subfamily Opatrinae, was inadequate, be- cause of the ventral exposition in dorsal view, of the male copulatory organ, the aedeagus. The initial choice of Reitter (1904), however, who placing Lach- nodactylus within the tribe Trachyscelini, was cor- rect.

In a cladistic analysis towards a new higher classification of Tenebrionidae, Doyen & Tschinkel (1982) concluded that the dorsal position, i.e. the ventral side of the aedeagus being contiguous to the abdominal sternites, is the normal, primitive posi- tion in Coleoptera, the plesiomorphic status of the male copulatory organ. In Opatrinae, however, the

52

aedeagus exhibits a “rotated” position, in which the dorsal face is visible from dorsally, and the ventral side is contiguous to the inner surface of the abdom- inal sternites (Fig. 2). This position is considered a derived, apomorphic status.

In fact, the contrary is true: the normal position of an organ - be it a hand or an aedeagus - is with the dorsal face exposed dorsally. The rotation of a member exhibiting the ventral face dorsally (Figs 1,3), is a very unusual, evolved, apomorphic state. The goal of this evolution is probably a configura- tion of the genitalia, in which the disposable space of the abdominal cavity is increased, to dispose the aedeagus completely contiguous to the surface of the abdomen, to reduce the empty space, to permit a dilatation of the ventral sack to accumulate food and to increase humidity, by means of reduction of evaporation under the elytra. These adaptative apo- morphic states are characteristic for such a xerophi- lous subfamily as Pimeliinae, that generally live in extreme arid, desert habitats, e.g. the tribes Molurini, Tentyrini, Adesmiini, Stenosini, Asidiini, Pimeliini, etc. from African dry savannas and deserts, and likewise the Central Asiatic Lachnogyini.

The new representative of Lachnogyini, pos- sesses an aedeagus that is similar to those of several genera of Pimeliinae (Figs 1, 3), which indicate that the Lachnogyini, hitherto placed within Opatrinae, have to be transferred. This is the consequence of several features that are absent in the genera of the subfamily Opatrinae: absence of a clypeo-genal in- cision; the habitual aspect of a relative of Ulomini; the median and posterior legs finely spinose and elongate, as in the genera of Zoophosini and Cryp- ticini; and of course, the rotated aedeagus. Special characters of Lachnogyiini resulting from conver- gent evolution, are the long pubescence covering the sides of the body and the very dilated protibiae, which are similar to some psammophile genera of Opatrini.

Examination of Lachnodactylus digitatus Seidlitz indicates that this genus has to be retransferred to the tribe Trachyscelini, today placed in the subfami- ly Diaperinae, because a combination of various different characters, as the club of antennae, the short, massive legs, well adapted to digging, but unable to run, and the simple position of the aedea- gus with the dorsal surface being exposed dorsally.

Acknowledgements

I am deeply indebted to Mr Jean Herve Yvinec, Centre de Recherche Archeologique de la Vallee de l’Oise, Compiegne, to Dr. Ottö Merkl, Hungarian Museum of Natural History Budapest, for communicating material

for this study, and to Dr. Martin Baehr, Zoologische Staatssammlung, München, for valiable comments about this note.

References

Doyen, T. J. &W.R. Tschinkel 1982. Phenetic and cladis- tic relationship among tenebrionid beetles. Syst. Ent. 7: 127-183

Gebien, H. 1910. Katalog der Coleoptera Tenebrionidae. - W. Junk., Coleopterorum Catalogus, Berlin.

-- 1939. Katalog der Tenebrioniden. Teil 2. - Mitt. Münchner Ent. Ges. 28(3): 444-463

Iwan, D. 2001. A comparative study of male genitalia in Opatrinae sensu Medvedev, 1968 (Coleoptera, Ten- ebrionidae), with notes on the tribal classification. Part 1. - Ann. Zool. Warszawa 51(3): 351-390

Medvedev, G. S. 1968. Coleoptera, Darkling Beetles. Tenebrionidae, Subfamily Opatrinae: Platynotini, Dendarini, Pedinini, Dissonomini, Pachypterini, Opatrini (part), Heterotarsini. - Fauna of URSS 19, 2, 1-288. Acad. Sci. Leningrad (in russian)

Reitter, E. 1904. Bestimmungs-Tabelle der Tenebrioni- den. Unterfamilien: Lachnogyini, Akidiini, Pedini- ni, Opatrini und Trachyscelini aus Europa und der angrenzenden Ländern. Verh. Natur.-forsch. Ver. Brünn 42: 25-129

The genus Scaurus: Biogeography and Ecology

(Insecta, Coleoptera, Tenebrionidae)

Harold Labrique

Labrique, H. (2003): The genus Scaurus: Biogeography and Ecology (Insecta, Coleoptera, Tenebrionidae). Spixiana 26/1: 53-54

Ecological and biogeographical parameters of the mediterranean genus Scaurus are discussed. The high grade of endemism is tressed and it is shown that most taxa live at low altitudes and preferably in regions with a mediterranean bioclimate.

Dr. Harold Labrique, Centre de Conservation et d’Etude des Collections (Mu- seum de Lyon) 13A, rue Bancel, F-69007 Lyon; e-mail: harold.labrique@rhone.fr

The genus Scaurus Fabricius actually contains 30 species and 8 subspecies. This genus ranges around the Mediterranean basin with the exception of the Balkans and Asia minor where it is replaced by the genus Cephalostenus Solier. Like many other tenebri- onid beetles living around the Mediterranean Sea, species of Scaurus are wingless. Most species are twilight or nocturnal beetles and, during the day, many are hidden under stones.

The genus Scaurus presents different types of distribution: betico-rifenian (such as S. tingitanus Peyerimhoff (f. typ.), maghrebian (such as 5. sancti- amandi Solier), north-west mediterranean (such as S. striatus Fabricius), saharo-mediterranean (such as S. aegyptiacus Solier), turanian (such as S. puncticollis ssp. macricollis Allard), etc.

There are two interesting regions where the di- versity is remarkable. The first one is the tripoli- tano-cyrenaic region with 5-6 occurring species. In this region, Scaurus sancti-amandi Solier reaches his eastern limit of distribution and S. carinatus ssp. vicinoides Schuster reaches his western limit. The second region includes north-eastern Marocco and west-eastern Algeria with altogether 11-12 species occurring. In this region, some species reach their

western limit of distribution (S. varvasi Solier, 5. an- gustus Reiche, S. atratus Fabricius and S. dubius So- lier) and others their eastern limit (S. vicinus Solier and S. mesatlanticus Peyerimhoff). One species is endemic in this region: S. camelus Kocher. General- ly, the orano-maroccan block has a high diversity with several endemic species.

Concerning endemism, this is very important in the genus. Among the 38 known taxa, 23 could be considered as endemic, that is about 60.5 %. For example, in Marocco live 22 taxa and among these 11 are endemic, that is 50 % of all taxa living in this country and about 50% of all endemic taxa. This richness of the maroccan fauna could be explained by the high diversity of habitats in this country.

Concerning the ecology of the species of the genus Scaurus, it can be stated that most taxa (66 %) inhabit areas with mediterranean bioclimate and only 4 taxa strictly inhabit the desert region. A mor- phological and phylogenetic study showed that the genus Scaurus can be divided into three lineages: the Scaurus angustus lineage (1 species), the Scaurus tristis lineage (24 taxa) and the Scaurus punctatus lineage (13 taxa). If we remove Scaurus angustus which is a very particular species, we could argue

53

that in the Scaurus tristis lineage 11 taxa among 24 (46 %) strictly inhabit subdesertic and desertic are- as, whereas in the Scaurus punctatus lineage only 2 species among 13 (16 %) inhabit areas with compa- rable bioclimate. On the contrary, in the Scaurus punctatus lineage 10 taxa among 13 (76 %) strictly inhabit areas with mediterranean bioclimate, where- as in the Scaurus tristis lineage 10 taxa among 24 (41 %) inhabit similar areas.

Concerning the altitudinal distribution of the taxa of the genus Scaurus, only 3 among 38 taxa can be found at high altitude (> 2000 m): namely Scaurus mairei Peyerimhoff (Hoggar), 5. alticola Escalera and S. kocheri Peyerimhoff (High-Atlas, Marocco). Thus, the large majority of taxa (33=87 %) live at low altitude from 0-1000 m. If considering the two dif-

ferent lineages defined above we find in the Scaurus uncinus lineage 7 species among the 11 ranging from 0-1000 m can be found at an upper altitude which is 63.6 %. In the Scaurus tristis lineage the situation is quite different: only 4 species among 21 ranging from 0-1000 m can be found at an upper altitude which is only 19 %.

In conclusion we can state that in the genus Scaurus the number of endemic taxa is very high (60 %), but very few species (6) possess a very large distribution. Most taxa live at low or medium alti- tudes in areas of mediterranean bioclimate. Very few species (3) only have been found at high altitude.

This work has been done at the Laboratoire de Zoogeographie, Universite Montpellier III, BP 5043, F-34032 Montpellier cedex 1.

A tenebrionid society in southeastern Kazakhstan: composition, zoogeography and ecology

(Insecta, Coleoptera, Tenebrionidae)

Wolfgang Schawaller

Schawaller, W. (2003): A tenebrionid society in southeastern Kazakhstan: com- position, zoogeography and ecology (Insecta, Coleoptera, Tenebrionidae). Spixi-

ana 26/1: 54-55

Species composition of Tenebrionidae at a river bed with dense gallery forest in Kazakhstan is enumerated and the ecological and biogeographical conditions are discussed. No tenebrionid was found in the forest which is ascribed to rather recent origin of the forest which prevented forest-dwelling species to immigrate.

Dr. Wolfgang Schawaller, Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, Gemany; e-mail: schawaller.smns@naturkundemuseum-bw.de

Locality of study

Southeastern Kazakhstan, Charyn Canyon W Chun- dzha, 650 m, 43°37'N, 79°21'E, 10.-13.V1.1993 & 29.- 31.V.2001. The Charyn river as a tributary of the river Ili comes from the Tien Shan mountains and has washed out an up to 20 metres deep canyon in the adjacent flat semidesert area. A sharp ecological contrast exists here: a quite humid forest as a river gallery and a quite dry belt of loamy soil. The level of the ground water is high, so the soil within this canyon is partly salty. The dense forest consists mainly of Fraxinus, Populus and Salix trees, and is often flooded, in daily as well as in seasonal inter- vals, depending from the weather and snow condi- tions in the upperstream mountains. In contary, the loam belt is quite open, only a few bushes, mainly tamarisks and Haloxylon are growing here.

54

Composition of fauna

27 species of Tenebrionidae have been recorded (all leg. and det. W. Schawaller, material in Museum Stuttgart). The faunal composition concerning sub- families is: Lagriinae (7 %), Pimeliinae (45 %), Tene- brioninae (37 %), Alleculinae (7 %), Diaperinae (4 %), Coelometopinae (0 %).

Belopini: Belopus calcaroides Reitter,1920, Belopus fili- formis (Motschulsky, 1872);

Tentyriini: Anatolica lata (Steven, 1829), Anatolica subquadrata Tauscher, 1812, Microdera iliensis Sko- pin, 1961, Microdera tscharynensis Kaszab, 1966, Scythis affinis Ballion, 1878, Tentyria acuticollis Reitter, 1900;

Adesmiini: Adesmia panderi Fischer von Waldheim, 1835;

Stenosini: Dichillus pusillus (Menetries, 1849);

Akidini: Cyphogenia gibba (Fischer von Waldheim, 1821);

Pimeliini: Ocnera lepidacantha Fischer von Waldheim, 1830, Pimelia cephalotes Pallas, 1781;

Boromorphini: Boromorphus opaculus Reitter, 1887;

Blaptini: Blaps deplanata Menetries, 1832, Blaps granu- lata Gebler, 1825, unidentified Blaps species, Prosodes rugulosa Gebler, 1841;

Platyscelidini: Oodescelis clavatipes Kaszab, 1938;

Opatrini: Cheirodes dentipes (Ballion, 1878), Dilamus gnom Skopin, 1961, Lobodera rufescens (Mulsant & Rey, 1859), Pachypterus serrulatus (Reitter, 1904);

Helopini: Unknown species of unknown genus;

Alleculinae (2 tribes): Unidentified species of Omo- phlus, Steneryx dejeani Faldermann, 1836;

Crypticini: Crypticus latiusculus (Menetries, 1849);

Coelometopinae (2 tribes): No record.

Zoogeography

The species have been sorted in 5 distributional groups, from a wider transpalaeartic distribution (Europe, all Siberia, West Caspian and Central Asia) up to endemic species in southeastern Kazakhstan. Within these distributional groups, the tenebrionid tribes are represented quite differently. In the group II with a wider distributional pattern (West Cas- pian, South Russia, Central Asia, Iran, Afghanistan) species of Opatrini represent 49% and of Tentyriini 0% of the fauna, in the group IV with smaller areas (Central Asia, Mongolia, Xinjiang) species of Opatrini represent only 10% and of Tentyriini 40%; in the group V (endemics) Tentyriini represent 100 %. In other words: Tentyriini obviously show a higher rate of speciation in the investigated area than the other recorded tribes of Tenebrionidae.

Ecology

Shifting sand and consequently psammophilous species are missing in the Charyn canyon, whereas only 50km away in the sand dunes of the Ili river sand-adapted species (Trichosphaena, Lachnodacty- lus, Leichenum, Psammestus) are not rare. The soil of the canyon is loamy and quite compact, at least in dry condition, thus the sand digging species with a smaller body size are unable to exist. Also missing in the Charyn canyon are quite common species from adjacent Artemisia steppe habitats (Podhomala, Pterocoma, Trigonoscelis, Earophanta, Platyscelis). Prob- ably these species need a denser vegetation occur- ing in the Artemisia steppe but not at the Charyn. It is striking, that these species mostly from the sub- family Pimeliinae have a bigger body size, probably the food resources are too low in the open Charyn habitats for additional big species. All 27 tenebrion- id species occur in the open loamy belt and not a single tenebrionid species (for example from the genera Diaclina, Eledona, Uloma, Platydema, Corti- ceus, Scaphidema) has been found in the dense forest, although old and dead trees and a lot of fungi as usual biotopes for tenebrionids occur. Probably we face here old trees, but not an old forest with an age of only 500-1000 years. Before that time, the canyon was filled by water up to the slopes. Then, in con- nection with the known desertification in historical times the water sunk and the forest developed by seeds from the upperstream mountains. But the forest fauna from the mountains could not reach this place because of its absolute isolation by huge semidesert surroundings.

Acknowledgements

The field work was supported by D. Bartsch (Stuttgart), M. Hauser (Urbana/Illinois) and Dr. S. Golovatch (Mos- cow). J. Reibnitz (Stuttgart) helped with the preparation of figures and diagrams.

Q1

Buchbesprechungen

5. Samper, J. R.: Iconografia del genero Ibeodorcadion (Coleoptera, Cermabycidae). Argania editio, Bar- celona, 2002. 197S, zahlreiche Farbabb. ISBN 84- 931847-2-1

Die flugunfähigen Bockkäfer der Großgattung Dorcadi- on, die allerdings von fleißigen Systematikern inzwi- schen in mehrere Gattungen aufgespalten worden ist, besitzen ein verschworene Gemeinde, die mit großer Energie diese “Erdböcke” genannten Käfer sammelt und bearbeitet. Man kann die Wertschätzung dieser Käfer durch die Sammler durchaus verstehen, denn sie sind einerseits sehr ansprechend gefärbte und gezeichnete Tiere das zeigen die Abbildungen in diesem Band sehr eindrucksvoll-andererseits neigen sie infolge ihrer Flug- unfähigkeit zur Bildung von zahlreichen Rassen und Farb- und Zeichnungsvarietäten. Das läßt das Herz der Sammler höher schlagen, haben sie doch so die Möglich- keit, neue Rassen oder Varietäten zu finden und zu benennen.

Der vorliegende Band richtet sich denn auch vor- nehmlich an die Sammler und besteht vor allem aus Farbabbildungen der Tiere, wobei insbesondere auch die Farbvariationen berücksichtigt werden. Erfreulicher- weise verzichtet der Autor auf die unsinnige Benennung einerjeden Variation. Gegenüber den sehr schönen, hand- gemalten Abbildungen sind die Beschreibungen und sonstigen Angaben sehr kurz gehalten. Die für die Art- differenzierung wichtigen männlichen Genitalorgane werden gar nicht erwähnt. Offenbar wurden und wer- den die Arten ausschließlich nach äußeren Merkmalen der Form, Farbe, Zeichnung und Oberflächenstruktur unterschieden. Inwieweit diese Art der Taxonomie den tatsächlichen Verhältnissen gerecht wird, entzieht sich der Beurteilung durch den Rezensenten.

Allerdings ist es offenbar das Ziel dieser “Ikonogra- phie”, dem Sammler die Möglichkeit zu geben, seine Tiere nach der Färbung und Zeichnung zu bestimmen, und dieses Ziel wird sicherlich erreicht, denn zum ersten Mal überhaupt wird eine Gesamtübersicht über die Erd- böcke der Iberischen Halbinsel gegeben. Selbst den kriti- schen Benutzer werden aber die schönen Bilder beein- drucken. Es ist also kein eigentlich wissenschaftliches Werk, aber dennoch ein nützliches und schönes’ Buch. Dem Sammler und Dorcadienfreund sei es deshalb emp- fohlen, auch wenn der Preis recht hoch erscheint.

M. Baehr

6. Lawrence, J. F., A. M. Hastings, M. J. Dallwitz, T. A. Paine & E. J. Zurcher: Beetles of the World. A Key and Information System for Families and Subfami- lies. Version 1.0 for Microsoft Windows. CSIRO Publishing, Collingwood 2000. CD and User Manu- al. ISBN 0-643-06557-1

56

This is the first of a scheduled series of CD’s containing interactive identification keys for various large animal groups. The keys were generated using the DELTA data- base program that has been developed by some of the authors. So, in particular the first author is responsible for the scientific contents, the others for the development of the databases and for the presentation.

Although such interactive keys request some trai- ning for the quick use, the pathways to find the required information are rather easily tracked and largely self- evident. Therefore, the user rather soon may be ac- quainted with the program and after few attempts he will be able to find his way to the taxon he is looking for. Once having tracked the family or subfamily, the user will find very fine and informative pictures of represen- tatives of the taxon he wants. Although the interactive key in some parts requires quite a lot of morphological knowledge, it is furnished with numerous, highly infor- mative figures of morphological details that are very helpful. A glossary of morphological terms further faci- litates the use of the keys, and a quite extensive general bibliography allows further reading about the subject.

The classification follows the recent work of Law- rence & Newton (1995) which since has become the basis for any modern beetle classifications. Nothing more should be said about contents than that it is the best available at present.

The unusual approach may not only be used for pure identification, but it likewise much facilitates cross checks, for example to see in which families a special morphological character state is present, and many others that have to be detected while using. So, the user will easily find a lot of information that would be more difficult to gather using printed, non-computerized sources.

As a conclusion: this is an extremely useful aid for anybody not only seeking for identification of suprage- neric beetle taxa, but also searching for information about beetle morphology, systematics, and even purely pic- tures of beetles and their lifestyle types. Certainly, many sorts of information are more easily gathered than from books, but tracking can be quite time consuming, partly because the user is easily deluded into searching for additional information that is unimportant for the im- mediate question. But this is a general problem for users of computers and their sometimes confusingly numerous applications. Also use of the program while examinating a specimen under a microscope probably will be more circumstantial and time consuming. However, these are minor complaints that by no means impair the high value of this CD that clearly belongs in the library of everybody seeking for general information about syste- matics and morphology of beetles. M. Baehr

SPIXIANA ur 5 57-63 München, 01. März 2003 ISSN 0341-8391

New taxa and new records of Odacanthinae from Sulawesi

(Insecta, Coleoptera, Carabidae)

Martin Baehr

Baehr, M. (2003): New taxa and new records of Odacanthinae from Sulawesi (Insecta, Coleoptera, Carabidae). Spixiana 26/1: 57-63

Ophionea nigrofasciata fortestriata, subspec. nov. is described from central Sulawe- si. The species was not yet recorded from this island and apparently, there a distinct subspecies occurs. Some records of additional odacanthine species from Sulawesi are dealt with, including a specimen of Archicolliuris bimaculata Redtenbacher hav- ing completely black pronotum, though the single available specimen from central Sulawesi is not regarded more than a colour variant and thus, it is not given a nomenclatorially valid name.

Considerations about the odacanthine fauna of Sulawesi suggest rather an Orien- tal than Australian origin of the fauna. For odacanthines at least, Wallace’s line may not have acted as a past or present faunal border.

Dr. Martin Baehr, Zoologische Staatssammlung, Münchhausenstr. 21, D-81247 München, Germany, e-mail: martin.baehr@zsm.mwn.de.

Introduction

Samples of carabid beetles recently received from Sulawesi include a number of odacanthine species some of which are interesting in the light of the very limited knowledge we still possess about the ground beetle fauna of that island. Especially the odacanth- ine fauna never was systematically sampled and most records are based on rather casual collections or samples made at light. Hence, neither the classi- cal works on the Oriental fauna, nor even more recent papers about this fasciating group - fascinat- ing in view of both, external structure and ecology contain much information on the odacanthine fau- na of Sulawesi.

Previous authors, e.g. Andrewes (many papers during the 20thies and 30ties of last Century), Lieb- ke (1938), Jedlicka (1963), had little access to materi- al from Sulawesi, and, although since the eighties of 20'" Century Sulawesi is being much better explored, there is still much more information available con- cerning the faunas of neighbouring areas than Su- lawesi itself: e.g. Andrewes for Buru (1930), Sumat- ra (1933), and Bali (1937); Habu (1962, 1967) for

Japan; Darlington (1968) for New Guinea; Stork (1986) for Borneo; and Baehr for New Guinea (1995, 1996b, 1997a, 1998), Australia (1986, 1996c, 1999a, in press), the Philippines (1996d), Java (1997b), and the New Guinean- West Pacific region (1996a, in prep.).

Only Baehr (1996a) described a new odacanth- ine species from Sulawesi and gave some additional records of species occurring on that island. Howev- er, the odacanthine fauna of the Moluccas still is far less well known than the faunas of the surrounding areas. Therefore, this paper, likewise far from being an exhausting treatment, could add to the knowl- edge and also could encourage future collectors and workers to direct their attention to the fauna of this island.

Format and style of the description, as well as measurements and ratios follows those used in my revision of the genus Casnoidea (Baehr 1996a). The full list of synonymies of the described species of Ophionea (=Casnoidea) may also be taken from this paper.

The specimens recorded are stored either in Zoo- logische Staatssammlung, München (ZSM), or in the working collection of the author at Zoologische

Staatssammlung (CBM), or in Staatliches Museum für Naturkunde, Stuttgart (SMNS).

Eucolliuris fuscipennis (Chaudoir)

Chaudoir, 1850: 26 (Casnonia); Andrewes 1927: 106 (Oda- cantha); Csiki 1932: 1527 (Colliuris); Liebke 1938: 65 (Colliuris); Jedlicka 1963: 494 (Colliuris); Habu 1967: 17; Darlington 1968: 205 (Colliuris); Lorenz 1998: 417.

The nominate form occurs through the whole of southern Asia including the Greater and Lesser Sun- da Islands, the Philippine Islands, and according to Darlington (1968) even New Guinea, and it also occurs on Sulawesi. The available Sulawesi speci- mens would rather correspond to the description of punctata Nietner, if this taxon at all could be main- tained as a separate one. According to Andrewes (1927), Darlington (1968), and Lorenz (1998), how- ever, the individual variability within C. fuscipennis is so great as to prevent maintenance of any subspe- cies. After examination of large samples of speci- mens of C. fuscipennis from India, Thailand, Sumat- ra, and Sulawesi I fully agree with this opinion, because even specimens collected at the same local- ity and within the same sample in some instances differ more substantially in shape and colouration than specimens from areas far away.

New records: Ugung Pandang (S-Sulawesi), 17.12. 1994, leg. Hiermeier (CBM); Sengkang, 10.4.1995, leg. Gerst- meier (CBM).

Eucolliuris celebensis (Gestro)

Gestro, 1875: 854 (Casnonia); Csiki 1932: 1527 (Colliuris); Liebke 1938: 65 (Colliuris); Lorenz 1998: 417.

Liebke (1938, p. 65), in his “Key to the indo-malayan species of the subgenus Eucolliuris” noted this spe- cies as a subspecies of E. fuscipennis, while Jedlicka (1963) did not mention the species at all. The taxo- nomic status certainly is disputable, though Lorenz (1998) mentioned it as a separate species. The series of specimens at hand corresponds well with the description of Gestro (1875). From my view, the specimens are so different from all E. fuscipennis I have seen that their specific status should not be questionable.

New records: 20 km ne. Bantmurung (S-Sulawesi), 30. 1II.1999, leg. Becvar & Zabransky (CBM).

58

Archicolliuris bimaculata (Kollar & Redtenbacher)

Kollar & Redtenbacher, 1842: 498 (Odacantha); Csiki 1932: 1525 (Colliuris); Liebke 1938: 61 (Colliuris); Jedlicka 1963: 497 (Colliuris); Habu 1967: 26; Lorenz 1998: 418.

This species is very widely distributed through the whole of south Asia from northwestern India in the west to Japan in the east, and through the Philippine and Indonesian Archipelagos including Sulawesi. From the latter island I have seen no material apart from a single female specimen that is distinguished by completely black instead of red prothorax. In other features, e. g. elytral pattern and striation, density and distinctness of microreticulation, shape and structure of prothorax, and shape of head, it is within the variation limits of A. bimaculata. Hence, the specimen most probably does not merit a no- menclatorial valid name, but at most it represents an infrasubspecific variety that may be called bimacu- lata s. str var. nigricollis. It is not known, however, whether this colour variety is accidental and indi- vidual, whether it has a wider distribution and might be characteristic even for alocal population. Itshould be mentioned, however, that A. bimaculata appar- ently is a quite variable species with respect to col- ouration and pattern. Indeed, I have seen some specimens from different provenance completely lacking the white spots on the elytra.

New record: 20 km ne. Bantmurung (S-Sulawesi), 30.IH. 1999, leg. Becvar & Zabransky (CBM).

Ophionea indica (Thunberg)

Thunberg, 1784: 68 (Attelabus); see Baehr (1996a: 1055, as Casnoidea) for further citations; Lorenz 1998: 417.

A species with very wide range that includes almost all of South Asia including the Philippine and Indo- nesian Archipelagos. It was also recorded from Su- lawesi, but neither from New Guinea nor from Aus- tralia, although it was accidentally introduced into Western Australia by ship (Baehr 1996a). Available specimens from Sulawesi in general structure are similar to those from other areas, though they slightly differ in their less distinct microreticulation of the elytra which gives the elytra a glossier appearence. However, these slight differences that, moreover, should be corroborated by additional material, prob- ably do not merit any taxonomic differentiation of the Sulawesi specimens.

New records: 20 km ne. Bantmurung (S-Sulawesi), 30. 11.1999, leg. Becvar & Zabransky (CBM, ZSM).

Ophionea celebensis (Baehr)

Baehr, 1996a: 1060 (Casnoidea); Lorenz 1998: 417.

Species closely related to O. interstitialis Schmidt- Göbel and its allies that so far was only recorded from Sulawesi. Apparently it is quite common there, as I have lots of specimens collected at light in different parts of Sulawesi. It seems that this species is the most common Ophionea on this island.

New records: 20 km ne. Bantmurung (S-Sulawesi), 30.11.1999, leg. Becvar & Zabransky (CBM, ZSM); 8 km W Mamasa (S-Sulawesi), 950 m, 18.-21.VIl.1999, leg. Bolm (SMNS); 20 km NE Sabbang (S-Sulawesi), 5.-7.VII. 1999, leg. Bolm (SMNS).

Ophionea nigrofasciata Schmidt-Göbel

Schmidt-Göbel, 1846: 21; see Baehr (1996a: 1052, as Cas- noidea) for further citations; Lorenz 1998: 417.

A widespread species that is, inter alia, character- ized by the red base, black fascia, and presence of two small, circular, white spots on the elytra; and by absence of any lateral seta on the prothorax. The nominate form is also characterized by the striation of the elytra becoming very weak in the apical half. The species is distributed through continental South- east Asia, the Greater Sunda Islands, and Buru in the Moluccas, though was not yet known from Su- lawesi nor from New Guinea (Baehr 1996a). The specimens mentioned below, therefore, represent the first record of this species for Sulawesi. The available specimens from this island, however, dif- fer rather characteristically from those of other are- as (India, Thailand, Sumatra), so that their descrip- tion as a separate subspecies is justified.

Ophionea nigrofasciata fortestriata, subspec. nov. Figs 1,2

Types. Holotype: d, INDONESIA - C-SULAWESI W coast of lake POSO TAIPA env. 10.-11.IV.1999 BECVÄR & ZABRANSKY leg. (CBM-ZSM). Paratype: 12, same data (CBM).

Diagnosis. Distinguished from nominate subspe- cies by the following character states: posterior half of head slightly shorter, prothorax considerably shorter and wider, puncturation of elytra distinct and coarse also in posterior half, and apex of aedea- gus even shorter and more asymmetrically turned to right side.

Fig. 1. Ophionea nigrofasciata fortestriata, subspec. nov. Aedagus, parameres, and genital ring. Scale: 0.5 mm.

Description

Measurements. Length: 6.8-7.1 mm; width: 1.70- 1.75 mm. Ratios. Length/width of prothorax: 2.05- 2.08; length/width of elytra: d 1.86, ? 1.84.

Colour (Fig. 2). Elytra quadrimaculate and with wide black band as in nominate subspecies, though posterior white spot even more enclosed in black fascia. Head, prothorax, antennae, and legs of same colour as nominate subspecies.

Head. Much alike nominate subspecies, though posterior part of head slightly less elongate, head, therefore, of slightly wider appearance.

Pronotum. Of same shape and surface struc- ture as in nominate subspecies, but distinctly short- er and wider (l/w: 2.05-2.08, vs. 2.23-2.33 in nomi- nate subspecies).

Elytra. In size and shape similar to nominate subspecies, though puncturation of striae not or barely finer towards apex, and intervals still dis- tinctly convex in apical half.

Lower surface. Similar to nominate subspecies.

Legs. Similar to nominate subspecies.

d genitalia (Fig. 1). Generally similar to those of nominate subspecies, though genital ring (in the single available male!) wider. Aedeagus as short and compact, but apex even shorter and more turned to right side. Parameres of similar size as in nomi- nate subspecies, but left paramere bearing a more acute apex.

? genitalia. Similar to nominate subspecies.

Variation. Due to limited material very little variation noted.

ll)

ll ll NT ORARUR

Fig. 2. Ophionea nigrofasciata fortestriata, subspec. nov. Habitus. Length of specimen: 7.1 mm.

Distribution. Central Sulawesi. Known only from type locality.

Collecting circumstances and habits. Largely un- known, both available specimens collected at light. Lives probably in and on reeds and grass near water or in swamps, like its nominate subspecies.

Etymology. The name is an adjective and refers to the coarse striation of the elytra.

Note. Although degree of variation in shape and structure of male genitalia is unknown due to the very limited material, the differences in length of pronotum and surface structure of the elytra seem to be constant. Hence description of the Sulawesi specimens as a separate subspecies is reliable.

Recognition. For recognition of the new subspecies the key to the nigrofasciata-group of the genus Ophio- nea (= Casnoidea) in Baehr (1997b) has to be altered in some ways. Therefore, the whole key to this species- group is repeated below.

60

Key to the Oriental-Australian species of the nigrofasciata-group of the genus Ophionea Castelnau

For identification of the species included, follow up to couplet 13. in the key in Baehr (1996a).

13. Prothorax coarsely punctate and with a row of lateral setae; only one large white spot in poste- rior half of each elytron. Northern and north- eastern Australia, New Guinea .......eeeeeeeseeeeeneeen

EEE OR re puncticollis (Sloane)

- Prothorax not coarsely punctate, without lateral setae or with only one seta; two white spots in anterior third and near apex of each elytron. Distribution@different. 2. ne 14.

14. Dark parts of upper surface distinctly blue. 15. - Dark parts of upper surface black ................. 16.

15. Elytra with anterior white spot situated outside dark fascia, posterior spot completeley included in fascia; head longer, posteriorly slightly con- vex; aedeagus unknown. Burma ......eeeeneeen

a bhamoensis (Bates)

- Elytra with both white spots at least partly in- cluded in dark fascia; head shorter, posteriorly rather straight; aedeagus with hook-shaped apex (Baehr 1996a, fig. 12). Philippines .....n.. nun...

ER bocebonda bakeri (Dupuis)

16. Prothorax without distinct transverse strioles; elytra with barely indicated microreticulation, surface highly glossy; both white elytral spots large and elongate, anterior spot completely outside of dark fascia, posterior spot clearly sur- passing posterior margin of fascia; posterior part of head rather short (Baehr 1997b, fig. 1). Java

ee bonn insignis (Baehr)

- Prothorax with distinct transverse strioles; elytra with distinct microreticulation, surface dull; ei- ther both white elytral spots small and circular, or only posterior spot large, in latter case both spots completely included in dark elytral fascia; posterior part of head longer ...........n. 17.

17. Prothorax without lateral seta, narrower and more elongate, surface regularly transversely striolate and distinctly microreticulate; posteri- or white elytral spot circular and rather small, clearly surpassing posterior margin of elytral fascia; head anteriorly longer, shape of head regularly rhomboidal with eyes situated about in middle (Fig. 2); microreticulation of elytra distinct, entire surface dull ..........neee- 18.

- Prothorax with a pair of lateral setae, wider and shorter, surface less regularly transversely stri- olate, rather superficially microreticulate; pos- terior white elytral spot elongate and very large, completely included in elytral fascia; head ante- riorly shorter, shape of head irregularly rhom- boidal with eyes clearly situated in front of mid- dle (Baehr 1996a, fig. 20); microreticulation of elytra less distinct, surface more glossy. North- Erlhatland ....-.....2oatssuaunsaraasass malickyi (Baehr)

18. Prothorax longer, about 24x as long as wide; elytral striae distinct and coarsely punctate only in basal half; posterior part of head slightly long- er; apex of aedeagus longer (Baehr 1996a, fig. 10). Southern Asia including the Greater Sunda IElands except Sulawesi...r...u.uusurneesneonneneassenencenene

Re nigrofasciata nigrofasciata Schmidt-Göbel

- Prothorax shorter, just twice as long as wide; elytral striae distinct and coarsely punctate even in apical half (Fig. 2); posterior part of head slightly shorter; apex of aedeagus shorter (Fig. 1). EIFEIESENESTRRER 222.0 022 000204 »002500205555 02080520 nn unensatsasatnege

Remarks

The odacanthine species mentioned from Sulawesi in this paper demonstrate quite different states of endemism. Whereas some species, e.g. Eucolliuris fuscipennis, Ophionea indica, and perhaps also Archicol- liuris bimaculata, do not differ perceptibly from their extra-Sulawesian forms, taxa like Eucolliuris celeben- sis and Ophionea celebensis apparently are specifical- ly distinct, and Ophionea nigrofasciata occurs in a separate subspecies on this island (O. nigrofasciata fortestriata).

In this context it should be mentioned, however, that recognition of subspecific rather than specific differentiation of insect taxa is rather a matter of opinion, as long as we do not know the extent to which the gene flow between taxa actually is inter- rupted. In particular in insular populations such decisions are extremely difficult, the more when the respective taxa either are able and willing to fly over considerable distances, or are easily transported by natural or human carriers. Classifications, there- fore, are rather based on characters and, at least for tropical insects like those mentioned in the present paper, usually they make advantage of more or less extensive similarity of morphological character states. In other words: without possessing much more information about genetics, in particular pop- ulation genetics, but also about way of life and behaviour of the respective taxa, we have little

chance to draw any final decision about their taxo- nomic status; and so we call ‘subspecies’ those taxa that exhibit little morphological differences and, at the same time, are allopatric, and ‘species’ those taxa that are more strikingly different and/or occur sympatrically. For the further discussion, therefore, one should bear in mind that more or less extensive similarities, as a rule, indicate more or less close relationships of taxa, though without adhering too much to terms like ‘species’ or ‘subspecies’.

Certainly, it is premature to draw any biogeo- graphical conclusions about the odacanthine fauna of Sulawesi, because our knowledge about the actu- al number of species occurring is still quite unsatis- factory. Nevertheless, we could argue that most of the odacanthine species yet mentioned from Su- lawesi have their nearest relatives in southern Asia, or even belong to species being widely distributed there. The odacanthine fauna of New Guinea and Australia, on the other hand, is rather different and includes several genera and species groups that do not reach further west than New Guinea or in few instances, the Moluccas (Darlington 1968, 1971, Baehr 1986, 1996a, 1996c, 1999a, 2000a, in press, in prep.). One recorded exception from this rule is the occur- rence of Dicraspeda brunnea Chaudoir on Sulawesi. This species belongs to a genus that has its centre of specific and morphological diversity in New Guin- ea and Australia, although a single species (D. brun- nea) occurs on Sulawesi, the Moluccas, the Philip- pines, and the Greater Sunda Islands (Baehr in prep.). As a conclusion one could argue, then, that - at the present state of knowledge - the odacanthine fauna of Sulawesi generally is an Oriental one.

The insular belt that runs from the Greater Sun- da Islands in the north and northwest, to the Moluc- cas and New Guinea in the south and southeast, since a long time has been noted as a major area of faunal transition, where Oriental and Papuan-Aus- tralian faunal elements are intermixed to a remark- able extent. To clarify the difficult situation, some ‘lines’ have been drawn by early scientiststhatshould depict certain faunal boundaries, or better, lines of faunal balance or of a certain procentual degree of preponderance of the Australian faunal elements over the Oriental ones, or vice versa (Fig. 3). The most familiar lines are “Wallace’s line’ that runs between Borneo and Sulawesi, and Bali and Lombok, and depicts an equilibrium of elements of both faunal provinces; “Weber’s line’ that runs east of Timor and Sulawesi but west of the Moluccas, and depicts a more than 75% advantage of Papuan-Australian faunal elements and also marks the western bound- ary of some Papuan-Australian elements; and ‘Lydekker’s line’ that divides New Guinea and some nearby islands from the Moluccas and marks the

61

Fig. 3. The Oriental and Papuan-Australian area of faunal transition and the important zoogeographic lines.

1: Wallace’s line; 2: Weber’s line; 3: Lydekker’s line.

Papuan (and Australian) faunal province(s) proper.

For the subfamily Odacanthinae, at least, Wal- lace’s line apparently has not been of any biogeo- graphic significance in the past, but, for Odacanthi- nae, the faunal border is running further east, pre- sumably just west of the Moluccas proper (Weber’s line). Unfortunately, very little is known about the odacanthine faunas of the Moluccas, especially of Seram, as well as about the faunas of Timor and the neighbouring Lesser Sunda Islands and of the is- land groups east of Timor, so, a more exact defini- tion cannot been given at present.

Asa conclusion, as far as Odacanthinae are con- sidered, Sulawesi, although showing a certain de- gree of endemism, clearly belongs to the Oriental faunal region. This situation, at least at the present state of knowledge, apparently deviates from re- sults gathered from some other carabid genera, for example in the subgenus Coeloprosopus Chaudoir of the Oriental-Papuan genus Pericalus W.S. Macleay (Baehr 1994), though apparently not in the nomi- nate subgenus Pericalus s. str. (Baehr 2000b). Similar differences are presumably seen in the large genus Dolichoctis Schmidt-Göbel. Here, only a single spe- cies from the decidedly Papuan subgenus Spinidoli- choctis Baehr (D. aculeata Chaudoir) occurs on Su- lawesi, though many species of the predominantly Oriental nominate subgenus Dolichoctis s. str. occur, of which probably no one is conspecific with those -

62

very few species - that occur in New Guinea and/or Australia (Baehr 1999b, in prep.).

Already these apparent differences within sing- le genera demonstrate the complexity of the bioge- ographic situation in this mentioned area of faunal transition. They also suggest that, in some instanc- es, Sulawesi apparently was re-colonized by single species of Papuan-Australian origin which, never- theless, belong to genera that originally came from the Oriental region. It seems, thus, that not only general trends of a northwest to southeast range extension and of a similar migration in the reverse direction, respectively, took place, or even still takes place, in this region, but moreover, that even within restricted groups of taxa to and fro range extensions in both directions occurred which, therefore, neces- sitates very scrutinized examinations of the biogeo- graphic history even on the species-group level.

References

Andrewes, H. E. 1927. Papers on Oriental Carabidae XIX..- Ann. Mag. Nat. Hist. (9) 19: 97-111

-- 1930. Fauna Buruana. Coleoptera, Fam. Carabidae. - Treubia 7, Suppl.: 331-347

-- 1933. A Catalogue of the Carabidae of Sumatra. Tijdschr. Ent. 76: 318-382

-- 1937. On a collection of Carabidae from the island of Bali. - Bull. Ann. Soc. ent. Belg. 77: 37-46

Baehr, M. 1986. Revision of the Australian ground-beetle genus Porocara Sloane (Coleoptera: Carabidae: Odacanthinae). Aust. J. Zool. 34: 717-731

-- 1994. Revision of the subgenus Coeloprosopus Chau- doir of the ground beetle genus Pericalus MacLeay. Taxonomy, phylogeny, zoogeography (Coleoptera, Carabidae, Lebiinae). Mitt. Münch. ent. Ges. 84: 13-58

-- 1995. A new genus of Odacanthinae from New Guinea (Insecta, Coleoptera, Carabidae). Spixiana 18: 45-48

-- 1996a. The ground beetle genus Casnoidea Castel- nau. Taxonomy, phylogeny, zoogeography (Insec- ta, Coleoptera, Carabidae, Odacanthinae). Inver- tebr. Taxon. 10: 1041-1084

-- 1996b. Three new species of the genus Dicraspeda Chaudoir from New Guinea (Insecta, Coleoptera, Carabidae, Odacanthinae). Spixiana 19: 137-146

-- 1996c. The Australian ground beetle genus Porocara Sloane. Second revision (Insecta, Coleoptera, Cara- bidae, Odacanthinae). - Spixiana 19: 253-265

-- 1996d. Two new species of the genus Lachnothorax Motschoulsky from the Philippines (Insecta, Cole- optera, Carabidae, Odacanthinae). Stuttg. Beitr. Zool. A, 539: 1-8

-- 1997a. Three further new species of the genus Di- craspeda CHAUDOIR from New Guinea (Coleoptera, Carabidae, Odacanthinae). - Mitt. Münch. Ent. Ges. 8729-37

-- 1997b. A new species of the genus Casnoidea Castel- nau from Java (Coleoptera, Carabidae, Odacanthi- nae). Entomofauna 18: 385-389

-- 1998. Two further new species of the genus Dicras- peda Chaudoir from New Guinea and the New He- brides (Insecta, Coleoptera, Carabidae, Odacanth- inae). - Entomofauna 19: 173-184

-- 1999a. A new genus of Odacanthinae from north- ern central Australia (Insecta, Coleoptera, Carabi- dae). - Coleoptera 2 (1998): 115-119

-- 1999b. The genus Dolichoctis Schmidt-Göbel, 1846 in New Guinea and surrounding areas (Carabidae: Lebiinae). - Coleoptera 2 (1998): 121-169

-- 2000a. Some genera and species of ground beetles new to Australia (Coleoptera: Carabidae). -— Mem. Qld. Mus. 46: 9-14

-- 2000b. Review of the Pericalus guttatus-complex (In- secta, Coleoptera, Carabidae, Lebiinae). Spixiana 23: 33-39

-- in press. A peculiar new genus of Odacanthinae from northern Australia (Insecta, Coleoptera, Cara- bidae). Boll. Mus. Sci. nat. Torino

Chaudoir, M. de 1850. Memoire sur la famille des Cara- biques. 2. Bull. Soc. Nat. Moscou 23: 3-85

Csiki, E. 1932. Coleopterorum Catalogus, pars 124, Har- palinae VII: 1279-1598. - W. Junk, Berlin

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

-- 1971. The Carabid beetles of New Guinea. Part IV. General considerations, analysis and history of the fauna, taxonomic supplement. Bull. Mus. Comp. Zool. 142: 129-337

Gestro, R. 1875. Nota sopra alcuni Carabici. - Ann. Mus. Civ. Stor. Nat. Genova 7: 840-894

Habu, A. 1962. Odacanthini of Japan (Coleoptera, Cara- bidae). Bull. Nat. Inst. Agric. Sci. (C) 13: 91-126

-- 1967. Fauna Japonica: Carabidae. Truncatipennes group (Insecta: Coleoptera). Biogeogr. Soc. Jap.: 1-338 + -XXVII

Jedlicka, A. 1963. Monographie der Truncatipennen aus Ostasien. Lebiinae - Odacanthinae Brachyninae (Coleoptera, Carabidae). Ent. Abh. Ber. Staatl. Mus. Tierkde Dresden 28: 269-579

Kollar, V. & L. Redtenbacher 1842. Aufzählung und Beschreibung der von Freiherrn Carl v. Hügel auf seiner Reise durch Kaschmir und das Himalayage- birge gesammelten Insekten.

Liebke, M. 1938. Denkschrift über die Carabiden-Tribus Colliurini. In: Festschr. Embrik Strand 4: 37-141

Lorenz, W. 1998. Systematic List of extant Ground Bee- tles of the World (Insecta Coleoptera “Geade- phaga”: Trachpachidae and Carabidae incl. Paussi- nae, Cicindelinae. Rhysodidae). Tutzing, printed by the author. 502 pp.

Schmidt-Göbel, H. 1846. Faunula Coleopterorum Birma- nae: 94 pp. Prague

Thunberg, C. G. 1784. Novas Insectorum Species III: 53- 68

Buchbesprechungen

7. Alonso-Zarazaga, M. A. &C.H.C. Lyal: A world catalogue of families and genera of Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Pla- typodidae). - Entomopraxis S.C.P., Barcelona, 1999. 315 pp. ISBN 84-605-9994-9

Curculionoidea certainly represent the most speciose beetle group and also a group of remarkable economic importance. The more regrettable was the confusion with regard to systematics of suprageneric taxa within this large assemblage. This confusion was mainly due to the fact that, apart from few modern regional treatments, the most recent catalogue of the whole group dates back more than 40 years, in large parts even 60 years. In the meantime not only many thousands of new species and hundreds of new genera have been described, but the suprageneric system of the weevils and their kind was changed again and again. Thus, a new, modern basis was absolutely necessary. The authors took over this labour and have presented a new system that apparently takes an intermediate position between the extreme po- sitions of splitting and of lumping of suprageneric taxa.

It is easily understood why such new approach was necessary, simply by consideration of the many new names, new synonyms, resurrected names, changes of status, and transfers between families and subfamilies that are presented in the introductory section of the catalogue. This part is followed by the synopsis of the system of families and subfamilies chosen by the au- thors, and by the catalogue that lists the subfamilies, tribes, and genera in alphabetical order. In the generic list, synonyms are included and the genotypic species is mentioned. An extensive bibliography of more than 50 pages, lists of genus-group names not included and of those included in Nomenclator Zoologicus, of the species names mentioned in the catalogue, and of family- and genus-group names conclude the volume.

Certainly this is a very valuable basis for any further work on this important beetle family that is most useful, or even absolutely necessary, for anybody interested in the higher classification of weevils, but certainly is also much appraised by collection managers who for a long time have lost the overview over this extremely speciose beetle group. M. Baehr

64

8. Bittmann, W. & B. Fugger: Reiseführer Natur. Au- stralien: Queensland. -BLV Verlagsgesellschaft mbH, München, Wien, Zürich, 1999. 129 S., zahlr. Fabfotos u. Karten. ISBN 3-405-15522-3

Australien wird seit einigen Jahren als Reiseland für deutsche Touristen immer beliebter, weil es mit dem Flair des Abenteuers und der Weite lockt. In der Tat ist es ein abwechslungsreiches, zwar sehr weitläufiges, aber gut zu bereisendes und sehr sicheres Reiseland, in dem vor allem Besucher auf ihre Kosten kommen, die eher ursprüngliche Natur als Ferienrummel erleben möch- ten. Und da kommen sie besonders im nordöstlichen Bundesstaat Queensland voll auf ihre Kosten. Was es dort an geologischen und biologischen Naturschönhei- ten zu sehen gibt, ist nicht nur großartig, sondern in vielen Fällen inzwischen auch problemlos zu erreichen.

Der Reiseführer vermittelt einen sehr guten Ein- druck dieser touristischen Highlights und geht insbe- sondere auch auf die reichhaltige Pflanzen- und Tierwelt ein. Natürlich ist er in erster Linie ein Führer durch die sehr zahlreichen Nationalparks, was aber darin begrün- det ist, daß in den letzten Dekaden erfreulicherweise viele Nationalparks neu gegründet wurden, so daß ein Großteil der besonders interessanten Gebiete heutzuta- ge als Nationalpark Schutz genießt. Es sollte jedoch nicht verschwiegen werden, daß auch in Queensland ausge- dehnte Gebiete entweder recht eintönige Trockengebie- te darstellen, die nicht dem normalen Touristen, sondern höchstens dem Geologen oder Biologen Interessantes bieten, oder aber durch Überweidung, Weizen- oder Zuckerrohranbau, und auch durch Anpflanzung gebiets- fremder Pflanzen zerstört oder jedenfalls nicht mehr in ihrer natürlichen Form erhalten sind. Dies wird aber nur dem Touristen auffallen, der auch weitere Strecken mit dem Auto zurücklegt.

Der normale Tourist wird Mühe haben, aus den sehr zahlreichen Sehenswürdigkeiten diejenigen herauszu- picken, die er in seiner beschränkten Zeit erreichen kann. Denn wenn auch viele sehenswerte Nationalparks ent- lang der Ostküste und zum Teil auch recht nah beieinan- der liegen, sollte man nicht vergessen, wir gewaltig die Entfernungen immer noch sind. Der vorliegende Reise- führer kann bei dieser Auswahl dank seiner gründlichen Information, der zahlreichen Karten und der schönen und instruktiven Bebilderung sehr gute Hilfe leisten. In der Tat machen die vielen schönen Farbfotos auch dem Rezensenten den Mund wässrig, einmal wieder in dieses prächtige Reiseland aufzubrechen, und den übrigen Le- sern wird es ähnlich ergehen. Ein guter, vor allem für naturverbundene Reisende sehr informativer Reisefüh- rer, der durchweg empfohlen werden kann.

M. Baehr

München, 01. März 2003

ISSN 0341-8391

Neue Arten aus der Gruppe der Amiserica insperata (Brenske, 1898)

(Coleoptera, Scarabaeoidea, Sericini)

Dirk Ahrens

Ahrens, D. (2003): New species of the Amiserica insperata (Brenske, 1898) species group (Coleoptera, Scarabaeoidea, Sericini). Spixiana 26/1: 65-81

Nine new species of the Amiserica insperata (Brenske) species group are described from Meghalaya, Indochina and Sichuan: Amiserica chiangdaoensis, spec. nov., A. flavolucida, spec. nov., A. langbianensis, spec. nov., A. lii, spec. nov., A. malickyi, spec. nov., A. nanensis, spec. nov., A. nokrekensis, spec. nov., A. omeiensis, spec. nov., and A. rejseki, spec. nov. From the studies one new combination results: Amiserica pardalis (Arrow), comb. nov. Male genitalia are figured. Further distribution records and a key to the species are given.

Dirk Ahrens, Deutsches Entomologisches Institut, Schicklerstr. 5, D-16225 Ebers- walde, Germany, e-mail: dahrens@zalf.de

Einleitung

Im Rahmen der Revision der indochinesischen Seri- cini wurden weitere, bisher nicht bekannte Arten aus der Verwandtschaft der Amiserica insperata (Bren- ske) aufgefunden, die in der vorliegenden Arbeit beschrieben werden. Unter den Arten der Gattung Amiserica Nomura, 1974 können diejenigen des öst- lichen Himalaya und Indochinas aufgrund der dor- salen Hell-Dunkel-Zeichnung des Körpers, den vorhandenen Schuppenhaaren auf Pronotum und Elytren sowie der charakteristischen Ausbildung der Phallobasis des Aedoeagus (dorsomedian tief und breit ausgerandet) deutlich von den übrigen Arten unterschieden werden. Sie werden daher als gesonderte Artengruppe behandelt. Habituell äh- neln sie stark den Vertretern von Lasioserica Brens- ke, 1896.

Abkürzungen BMNH The Natural History Museum, London BPBM Bernice P. Bishop Museum, Honolulu CA coll. Dirk Ahrens, Eberswalde CK coll. D. Kräl, Praha

CP coll. P. Pacholätko, Brno

MEMU Mississippi State University

MHNP Museum National d’Histoire naturelle, Paris MMBC Moravske Muzeum, Brno

NHMW Naturhistorisches Museum, Wien

SMNS _ Staatliches Museum für Naturkunde, Stuttgart SMTD Staatliches Museum für Tierkunde, Dresden ZSM Zoologische Staatssammlung, München

Bemerkungen zur Systematik von Amiserica Nomura, 1974

Gattungstypus: Amiserica rufidula Nomura, 1974 (durch Monotypie).

Nachdem Nomura (1974) die Gattung Amiserica für Arten aus Taiwan mit krenulierter Leiste an der Metatibia und Metafemur sowie 9- oder 10-gliedri- ger Antenne mit dreigliedriger Fühlerkeule (3) auf- gestellt hat, wurden weitere Arten von dort durch Kobayashi (1980, 1988, 1991) beschrieben. Später stellte Ahrens (1996, 1999) auch Arten vom asiati- schen Festland dazu, doch ist nach den bisherigen Untersuchungen des Autors die Monophylie der Gruppe keineswegs sicher begründet. Diese Ver- mutung begründet sich auf folgende Tatsachen:

ON 01

1) bei der krenulierten Leiste des Metafemur von Amiserica handelt es sich mit großer Wahrschein- lichkeit um eine Plesiomorphie, denn das Merkmal tritt in der gleichen Ausprägung auch bei Lasioserica Brenske, 1896 auf (ob es sich bei den sehr ähnlichen Zuständen bei Eumaladera Nomura, 1967 und Tai- wanoserica Nomura, 1974 um Analogien handelt, bedarf noch einer Klärung, da hier die gesamte äußere Morphologie stark abweicht); 2) bei der krenulierten Leiste der Metatibia existieren verschie- dene Merkmalszustände, wobei sie bei der Typus- art (Amiserica rufidula Nomura) vor der letzten Au- ßendorngruppe der Metatibia deutlich unterbro- chen, bei den Arten der A. insperata-Gruppe aber durchgehend ist; 3) die Lesrichtung des Merkma- les der Anzahl der Antennomeren in der Fühlerkeu- le bedarf einer eindeutigen Klärung, dazu sind wei- tere Studien der ostasiatischen Fauna notwendig. Die Zuordnung der hier behandelten und neu be- schriebenen Arten zu Amiserica, die auf die bisher lediglich diagnostischen Merkmale Nomura’s (1974) beruht, ist daher als vorläufig anzusehen.

Hilfsschlüssel zur Determination der Arten der Amiserica-insperata-(Brenske)-Gruppe

B. Eühlerld-shednie®...n. Ar... ern 2, , Fühler 9-chedrig................ 0 @22seeneereretennen 7.

2. Körperoberfläche matt. Oberseite einzeln, mä- Big lang und wenig abstehend behaart. .......... 3:

- Körperoberfläche irisierend glänzend. Obersei- te einzeln, lang abstehend behaart. .................... BEE A LAN 1 ce A. flavolucida, spec. nov.

3. Metatibia lang und schmal, Verhältnis Breite / Länge: 1/4,5. Phallobasis ventral mit zwei gro- ßen, lamellenartigen Längshöcker. ..................

ayhesashegenhishhentgereeege A. semipunctata Ahrens, 1999

= Men lang und mäßig breit, Verhältnis Brei- te/Ianger> 149,9. 0.0.02. 2m enennenassnneen een 4.

4. Laterale Ventralkante der Phallobasis allmäh- lich apikad längs erhöht. Intervalle der Elytren mit einzelnen weißen schuppenförmigen aber senmalensHliaaten wre 2 er 5,

- Laterale Ventralkante der Phallobasis weder all- mählich apikad längs erhöht noch mit deutlich abgesetzten großen Längshöckern. ................. 6.

5. Linke Paramere deutlich breiter als die rechte (Dorsalansicht). Fühlerkeule 1,3mal so lang wie die übrigen Glieder zusammen. ........ueseseseeeee SELL EURE NENNE A. nokrekensis, spec. nov.

66

10.

ill,

12:

119),

Beide Parameren etwa gleich breit (Dorsalan- sicht). Fühlerkeule 1,5mal so lang wie die übri- gen Glieder zusammen... Ga nstgerengen A. mawphlangensis Ahrens, 1999

Phallobasis ventral mit zwei sehr kleinen Hök-

| A. insperata (Brenske, 1898) Phallobasis ventral vollkommen ohne Erhaben- eiten A. manipurensis Ahrens, 1999

Fühlerkeule beim d doppelt so lang die übrigen Glieder zusammen und kräftig nach außen ge- bogen. Phallobasis relativ kurz und robust .......

BE RR ES). LET Bi A. nanensis, spec. NOV.

Fühlerkeule beim d maximal 1,5mal so lang die übrigen Glieder zusammen... re er 8.

Fühlerkeule beim d nur etwas länger als die übrigen Glieder zusammen. Phallobasis apikal starksverlangenty nn A. rejseki, spec. nov.

Fühlerkeule beim d 1,5mal so lang die übrigen Glieder zusammen... ee 2)

Phallobasis apikal mäßig verlängert. Parameren apikal stark hakenförmig ventrad gebogen. Phallobasis an der Spitze rechts länger als links.

ren nee A. sparsesetosa Ahrens, 1999

Phallobasis apikal lang ausgezogen. Parameren apikal nur schwach ventrad gebogen. ........... 10.

Pronotum in der Mitte am breitesten. Phalloba- sis im apikalen Drittel nur schwach gebogen und einfach zugespitzt (Lateralansicht). Basal- lappen der linken Paramere schmal und im rech- ten Winkel zur Aedoeaguslängsachse................

RE, A. malickyi, spec. nov.

Pronotum an der Basis am breitesten. Phalloba- sis im apikalen Drittel kräftig ventrad gebogen, präapikal dorsoventral verbreitert (Lateralan- sicht), an der Spitze breit abgerundet. .......... Al;

Phallobasis mesoventral mit einer deutlich ab- gesetzten und kräftig erhöhten Längslamelle. Linke Paramere an der Außenseite mit einem

serastentenaReldem ee A. lii, spec. nov. Phallobasis ventral ohne deutlich erhöhte Längs- lamelle....:.....2.0.222.2: 22222 22ER 112.

Basallappen der linken Paramere im Querschnitt rundlich oder oval. ..................... 2 ee 13:

Basallappen der linken Paramere stark abge- plattet und mehr oder weniger konkav gebo- SEM. ee LEE 14.

Basallappen der linken Paramere schmal und distal mit einigen feinen Zähnchen. ............. ne RR A. chiangdaoensis, spec. nov.

—- Basallappen der linken Paramere breit, fast recht- winklig zur Aedoeaguslängsachse, distal breit MEILEN

ESS IEE IR En SE UERERIRERN." A. pardalis (Arrow, 1946)

14. Rechte Paramere apikal sehr spitz. .................... ken hauader A. langbianensis, spec. nov.

Rechte Paramere apikal schräg abgeschnitten und außen leicht abgerundet... EN A. omeiensis, spec. nov.

Amiserica flavolucida, spec. nov. Abb. 1-3

Typen. Holotypus: d, NE India Meghalaya state West Garo Hills Nokrek Nat. Park 9.-17.V.1996 alt. 1100+150 m GPS N25°29,6', E 90°19,5' (WGS 84) E. Jendek & O. Sausa leg. (MMBC via CP). Paratypen: 1833, 162%, gleiche Angaben wie der Holotypus (CP, CA); 6039, 8922, NE India Meghalaya 1400 m, Nokrek n.p., 3km S Daribokgiri 25°27'N 90°19'E, 26.iv.1999 Debicky & Pa- cholatko leg. (CP, CA); 384, 12, NE India; Meghalaya; 1999; 3 km E Tura 25°30'N 90°14'E; 1150 m; 4.v.; Dembik- ky & Pacholätko leg. (CP, CA); 12, Thai, Chiang Mai prov., 18°49'N 98°54'E, 1600 m Doi Pui mt. 2.-6.v. (CP).

Beschreibung

Länge: 5,8-6,7 mm, Elytrenlänge: 4,4-4,7 mm, Breite: 3,5-3,7 mm.

Gestalt länglich oval; dunkel gelbbraun, Elytren wie auch Pronotumseiten und Labroclypeus heller und mit kleinen dunklen Flecken, gesamte Oberflä- che irisierend glänzend; Oberseite einzeln, lang ab- stehend behaart.

Labroclypeus relativ klein, deutlich breiter als lang, Seitenränder in der basalen Hälfte gerade und schwach verengt, vorn kräftig nach innen gebogen, Vorderecken breit abgerundet, Ränder leicht aufge- bogen und vorn median deutlich ausgerandet, Ober- fläche in der Mitte sehr schwach konvex, fein und dicht punktiert, der Abstand der Punkte ist kleiner als ihr Durchmesser, mit wenigen abstehenden Bor- sten, deren Punkte etwas größer als die übrigen sind; Frontoclypealnaht sehr fein eingeritzt und schwach gebogen; Ocularcanthus mäßig lang und sehr schmal, sehr fein und zerstreut punktiert, mit einer einzelnen Terminalborste, Augenspiegel 2,5 mal so breit wie lang. Punktur der Stirn fein und mäßig dicht bis locker, neben den Augen jeweils zwei bis drei einzelne Haare. Augen groß, Verhält- nis Augendurchmesser/-abstand: 0,9 (3), 0,64 (?). Fühler 10-gliedrig, gelb, Keule 3-gliedrig, beim d 1,5 mal so lang wie die übrigen Glieder zusammen und schwach nach außen gebogen, beim 2 Fühler- keule etwas kürzer als die übrigen Glieder zusam- men. Mentum erhaben, vorn abgeplattet.

Pronotum mäßig breit, an der Basis am breite- sten, Seitenränder von der Basis zur Mitte nur sehr schwach geradlinig verengt, im vorderen Drittel schwach nach innen gebogen, Vorderwinkel spitz und deutlich vorgezogen, Seitenrand neben den Vorderwinkeln schwach konvex, Hinterwinkel stumpf, in der Spitze kaum verrundet, Vorderrand fein gerandet und in der Mitte leicht konvex vorge- zogen, wie die Seitenränder lang einzeln bewim- pert; Oberfläche fein und dicht punktiert, in den Punkten mikroskopisch fein behaart, auf jeder Seite mit einem einzelnen langen, abstehenden Haar. Scu- tellum mäßig breit, dreieckig, dicht und fein punk- tiert, bisweilen median an der Basis punktfrei, wie das Pronotum mikroskopisch fein behaart.

Elytren kurz hinter der Mitte am breitesten, äu- ßerer Spitzenwinkel nur mäßig verrundet, Streifen wenig deutlich eingedrückt und fein punktiert, In- tervalle flach, fein und ungleichmäßig dicht punk- tiert, die lockere Punktur spart auf den Intervallen kleine Bereiche aus, die teilweise eine fleckenförmi- ge Form haben, ungeradzahlige Intervalle mit ein- zelnen langen, abstehenden Haaren, in den Punk- ten sonst nur mikroskopisch behaart; Epipleural- kante kräftig, endet am äußeren Spitzenwinkel, Epipleuren locker und lang behaart, Elytrenspitzen mit breitem Saum aus feinen weißen Mikrohärchen.

Unterseite einschließlich Metasternum nur fein und spärlich behaart, Metacoxalplatten bis auf die lateralen Borsten nur mikroskopisch fein behaart. Abdominalsternite bis auf je eine Querborstenreihe nur mikroskopisch fein behaart, letztes Sternit zu- sätzlich mit einzelnen langen Haaren, vorletztes Ster- nit beim d mit undeutlicher Querschwiele. Meso- sternum zwischen den Mesocoxae so breit wie der schmale Metafemur. Verhältnis Metepisternen-/ Me- tacoxalänge: 1/1,9. Pygidium kräftig gewölbt, kräf- tig und dicht punktiert, längs der Mitte breit glatt und schwach kielartig erhöht, mit mäßig dichter, kurzer und langer, gelber Behaarung, Pygidium beim ? weniger stark gewölbt.

Meso- wie Metafemur schmal und glänzend, Metafemur mit durchgehend krenulierter Kante am Vorderrand, dicht und kräftig punktiert und dicht, mäßig lang behaart, die Längshaarpunktreihen zwi- schen der übrigen Behaarung wenig deutlich, Hin- terrand fast gerade, unbehaart, Hinterrandkante ventral und apikal auch dorsal fein krenuliert. Me- tatibien mäßig schlank, in der Mitte am breitesten, basal stark verschmälert, Verhältnis: Breite/ Länge der Metatibien: 1/3,6 (4), 1/3,0 (?); dorsal sehr scharf gekantet, mit krenulierter Längsleiste paral- lel zur der Oberkante, mit dieser konvergent ab der schwach angedeuteten, distalen Querleiste bei % der Metatibienlänge, neben der krenulierten Leiste einzelne kräftige Haare; außen schwach längs ge-

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1

Abb. 1-3. Amiserica flavolucida, spec. nov. 1, 3. Aedoea- gus lateral. 2. Parameren dorsal. Maßstab 1 mm.

wölbt, sehr dicht und kräftig punktiert, dicht und mäßig lang behaart; Ventralkante mit drei kräfti- gen, leicht gehöckert stehenden Borsten in etwa gleichen gleichem Abstand zueinander, Innenseite fein, locker punktiert, apikal in Nähe der Tarsalein- lenkung nur wenig tief und etwas konkav ausge- schnitten. Tarsen ventral fein und mäßig dicht be- haart, Meso- und Metatarsen dorsal dicht punktiert und seitlich ungekantet, Metatarsen lateral gefurcht, ventral neben der krenulierten Leiste eine kräftige Längskante, Metatarsomer 1 deutlich kürzer als die zwei folgenden Glieder zusammen und % länger als der obere Enddorn der Metatibien. Protibia mäßig lang, 2-zähnig, Protarsomere dorsal unpunktiert, Glieder 1-4 nur wenig breiter als lang, alle Klauen einfach ausgebildet und symmetrisch. Aedoeagus: Abb. 1-3.

Differentialdiagnose. Amiserica flavolucida, spec. nov. unterscheidet sich deutlich von den übrigen Amiserica-Arten mit 10-gliedriger Antenne durch die stark irisierend glänzende Körperoberfläche.

Amiserica mawphlangensis Ahrens

Ahrens, 1999: 235.

Untersuchtes Material: 10 ex., India 26.VI.95 Chera- punjee Meghalaya Werner leg. (CA).

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Amiserica manipurensis Ahrens

Ahrens, 1999: 236.

Untersuchtes Material: 55 ex., NE India, Meghalaya state Jaitia Hills, reg. Jowai 6.-8.V1.1996 alt. 1350+100 m GPS N 25°27', E 92°12' (WGS 84) E. Jendek & O Sausa leg./ IS 64 (CP); 1 ex., NE India, Meghalaya state Khasi Hills, Shillong paek 4.-5. VI.1996 alt. 1850+50 m GPS N 25°32', E 91°52,5' (WGS 84) E. Jendek & O. Sausa leg. (CA).

Amiserica nokrekensis, spec. nov. Abb. 4-6

Typen. Holotypus: d, NE India Meghalaya state West Garo Hills Nokrek Nat. Park 9.-17.V.1996 alt. 1100+150 m GPS N25°29,6', E 90°19,5' (WGS 84) E. Jendek & O. Sausa leg./ IS7 (MMBEC via CP). - Paratypen: 18, gleiche Angaben wie der Holotypus (CA); 12, Khasis Nat. Coll./ Coll. C. Felsche Kauf 20, 1918" (SMTD); 13, Mu- seum Paris Haut Tonkin Laokay (Vitalis de Salvaza) M. A. Vuillet 1920 (MHNP).

Beschreibung

Länge: 5,6-6,8 mm, Elytrenlänge: 3,7-5,6 mm, Breite: 3,3-4,3 mm.

Gestalt länglich oval; braun, teilweise mit grün- lichem Schimmer, Labroclypeus rotbraun, Prono- tum und Elytren mit mehr oder weniger ausge- dehnten, hellen Flecken in den Punkten, Elytren fast vollkommen hell gelbbraun, nur vor der Spitze eine große dunkle Makel, gesamte Oberfläche bis auf den schwach glänzenden Vorderkopf matt; Ober- seite einzeln, mäßig lang, abstehend behaart.

Labroclypeus mäßig breit, deutlich breiter als lang, Seitenränder in der basalen Hälfte gerade und schwach verengt, vorn kräftig nach innen gebogen, Vorderecken mäßig abgerundet, Ränder leicht auf- gebogen und vorn median deutlich ausgerandet; Oberfiäche in der Mitte schwach konvex, fein und dicht punktiert, der Abstand der Punkte ist kleiner als ihr Durchmesser, mit zahlreichen abstehenden Borsten, deren Punkte etwas größer als die übrigen sind; Frontoclypealnaht sehr fein eingeritzt und schwach gebogen; Ocularcanthus mäßig lang und sehr schmal, sehr fein und zerstreut punktiert, mit einer einzelnen Terminalborste, Augenspiegel 2,5 mal so breit wie lang. Stirn irisierend glänzend, caudal leicht matt, Punktur fein und locker, neben den Augen und auf der Stirnmitte wenige einzelne lange Haare, im hinteren Viertel kurz und dicht, abstehend behaart. Augen mäßig groß, Verhältnis Augendurchmesser/-abstand: 0,74 (5), 0,57 (2). Fühler 10-gliedrig, gelb, Keule 3-gliedrig, beim Ö etwa 1,3 mal so lang die übrigen Glieder zusammen

und kaum gebogen, beim ? Keule so lang wie die übrigen Glieder zusammen. Mentum erhaben, vorn abgeplattet.

Pronotum breit, an der Basis am breitesten, Seiten- ränder in der basalen Hälfte subparallel bis schwach craniad verengt und wenig gebogen, in der vorderen Hälfte mäßig stark nach innen gebogen, Vorderwin- kel mäßig spitz und deutlich vorgezogen, Seitenrand neben den Vorderwinkeln konvex, Hinterwinkel mäßig abgerundet, Vorderrand sehr fein gerandet und in der Mitte leicht konvex vorgezogen, wie die Seitenränder lang einzeln bewimpert; Oberfläche kräftig und dicht, teilweise unregelmäßig punktiert, weißlich, fein schuppenartig, nach hinten gebogen behaart, dazwischen mikroskopisch fein behaart. Scutellum mäßig breit, dreieckig, dicht und fein entlang der Seitenränder punktiert, median an der Basis eine dreieckige punktfreie Fläche, wie das Pronotum mikroskopisch fein behaart.

Elytren kurz hinter der Mitte am breitesten, äu- ßerer Spitzenwinkel breit verrundet, Streifen fein eingedrückt und dicht, fein punktiert, Intervalle flach, fein und ungleichmäßig dicht punktiert, an der Basis mit kurzer, weißer, feiner, anliegender Behaarung in den Punkten, alle Intervalle mit ein- zelnen weißen schuppenförmigen aber schmalen Haaren, deren Punkte doppelt so groß wie die übri- gen sind, in den feinen Punkten nur mikroskopisch behaart; Epipleuralkante kräftig, endet am äußeren Spitzenwinkel, Epipleuren locker und lang behaart, Elytrenspitzen mit einem schmalen Saum aus fei- nen weißen Mikrohärchen.

Unterseite einschließlich Metasternum nur fein und spärlich behaart, Metacoxalplatten bis auf die lateralen Borsten nur mikroskopisch fein behaart. Abdominalsternite bis auf je eine Querborstenreihe nur mikroskopisch fein behaart, letztes Sternit zu- sätzlich mit einzelnen langen Haaren, vorletztes Ster- nit ohne Erhabenheiten. Mesosternum zwischen den Mesocoxae so breit wie der schmale Metafemur. Verhältnis Metepisternen-/Metacoxalänge: 1/1,85. Pygidium kräftig gewölbt, fein und dicht punktiert, längs der Mitte schmal glatt, mit mäßig dichter, kurzer und langer, gelber Behaarung.

Meso- wie Metafemur schmal und matt, Meta- femur mit durchgehend krenulierter Kante am Vor- derrand, fein und spärlich punktiert und kahl, Längs- haarpunktreihen deutlich, Hinterrand fast gerade, unbehaart, Hinterrandkante ventral und apikal auch dorsal fein krenuliert. Metatibien mäßig schlank, in der Mitte am breitesten, basal und apikal leicht verschmälert, Verhältnis Breite/Länge der Metati- bien: 1/3,3; dorsal deutlich gekantet, mit krenulier- ter Längsleiste parallel zur der Oberkante, mit die- ser konvergent ab der schwach angedeuteten, dista- len Querleiste bei % der Metatibienlänge, neben der

4 6

Abb. 4-6. Amiserica nokrekensis, spec. nov. 4, 6. Aedoea- gus lateral. 5. Parameren dorsal. Maßstab 1 mm.

krenulierten Leiste einzelne kräftige Haare; außen schwach längs gewölbt, fein und mäßig dicht punk- tiert, im basalen Drittel in den Punkten kurz be- haart, sonst nur mit mikroskopischen Härchen in den Punkten; Ventralkante mit drei kräftigen, leicht gehöckert stehenden Borsten in etwa gleichen glei- chem Abstand zueinander, Innenseite fein, locker punktiert, apikal in Nähe der Tarsaleinlenkung nur wenig tief und etwas konkav ausgeschnitten. Tar- sen ventral fein und mäßig dicht behaart, Meso- und Metatarsen dorsal locker punktiert und seitlich ungekantet, Metatarsen lateral schwach längs ge- furcht, ventral neben der krenulierten Leiste eine kräftige Längskante, Metatarsomer 1-4 ventral kahl, Metatarsomer 1 etwas kürzer als die zwei folgen- den Glieder zusammen und % länger als der obere Enddorn der Metatibien. Protibia mäßig lang, 2-zäh- nig, distaler Zahn fast rechtwinklig nach außen ge- richtet. Protarsomere dorsal unpunktiert, Glieder 1-4 nur wenig breiter als lang, alle Klauen einfach ausgebildet und symmetrisch. Aedoeagus: Abb. 4-6.

Differentialdiagnose. A. nokrekensis, spec. nov. unterscheidet sich von A. mawphlangensis Ahrens durch die etwas kürzere Fühlerkeule beim d sowie vor allem durch die Form der Parameren.

Amiserica pardalis (Arrow), comb. nov. Abb. 7-10

Serica pardalis Arrow, 1946: 8.

Typen. Syntypen: 13, Cotype/ N. E. Burma Kambaiti 7000 ft. 13/5.1934/ N. E. Burma R. Malaise B. M. 1945- 71/ Serica pardalis, spec. nov. Arrow (BMNH); 15, 3/ Co- Type/ N. E. Burma Kambaiti 7000 ft. 3-7.5.1934/ N. E. Burma R. Malaise B. M. 1945-71/ Serica pardalis co-type

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Abb.7-9. Amiserica pardalis (Arrow) (Yunnan, Gaoligong Mts.). 7, 9. Aedoeagus lateral. 8. Parameren dorsal. Maßstab 1 mm.

Arrow (BMNH); 12, N. E. Burma Kambaiti 7000 ft. 22/ 6 1934 R. Malaise/ Cotype/?/ N. E. Burma R. Malaise B. M. 1945-71 (BMNH); 12, N. E. Burma Kambaiti 6000 ft. 10/5/1934 R. Malaise/ N. E. BurmaR. Malaise B. M. 1945-71/ Serica pardalis co-type Arrow (BMNH); 12, N. E. Burma Kambaiti 6000 ft. 2/6/1934 R. Malaise/ N. E. Burma RR. Malaise B. M. 1945-71 (BMNH).

Weiteres Material: 13, 12, Yunnan 1500-2500 m 25.22N 98.49E 17-24/5 Gaoligong mts. Vit Kubän leg. 1995 (CP).

Beschreibung

Länge: 6,3-7,0 mm, Elytrenlänge: 4,3-4,6 mm, Breite: 3,4-3,7 mm.

Gestalt länglich oval; dunkel rotbraun, teilweise mit grünlichem Schimmer, Pronotum und Elytren mit mehr oder weniger ausgedehnten, hellen Flek- ken, gesamte Oberfläche bis auf den schwach glän- zenden Läbroclypeus matt; Oberseite einzeln, kurz abstehend behaart.

Labroclypeus mäßig breit, deutlich breiter als lang, Seitenränder in der basalen Hälfte gerade und schwach verengt, vorn kräftig nach innen gebogen, Vorderecken mäßig abgerundet, Ränder leicht auf- gebogen und vorn median deutlich ausgerandet; Oberfläche in der Mitte schwach konvex, fein und dicht punktiert, der Abstand der Punkte ist kleiner als ihr Durchmesser, mit zahlreichen abstehenden Borsten, deren Punkte etwas größer als die übrigen sind; Frontoclypealnaht sehr fein eingeritzt und schwach gebogen; Ocularcanthus mäßig lang und sehr schmal, sehr fein und zerstreut punktiert, mit einer einzelnen Terminalborste, Augenspiegel 2,5 mal so breit wie lang. Punktur der Stirn fein und mäßig dicht, neben den Augen und auf der Stirn-

70

mitte wenige einzelne Haare. Augen mäßig groß, Verhältnis Augendurchmesser/-abstand: 0,66 (4), 0,62 (?). Fühler 9-gliedrig, gelb, Keule 3-gliedrig, beim d 1,5 mal so lang die übrigen Glieder zusam- men und kaum gebogen, beim 2 Fühlerkeule etwas kürzer als die übrigen Glieder zusammen. Mentum erhaben, vorn abgeplattet.

Pronotum breit, an der Basis am breitesten, Sei- tenränder in der basalen Hälfte subparallel und nur sehr schwach gebogen, in der vorderen Hälfte kräf- tig nach innen gebogen, Vorderwinkel mäßig spitz und mäßig vorgezogen, Seitenrand neben den Vor- derwinkeln konvex, Hinterwinkel mäßig abgerun- det, Vorderrand sehr fein gerandet und in der Mitte leicht konvex vorgezogen, wie die Seitenränder lang einzeln bewimpert; Oberfläche kräftig und mäßig dicht teilweise unregelmäßig punktiert, weißlich, fein schuppenartig, nach hinten gebogen behaart, dazwischen mikroskopisch fein behaart. Scutellum mäßig breit, dreieckig, dicht und fein punktiert, median an der Basis ausgedehnt punktfrei, wie das Pronotum mikroskopisch fein behaart.

Elytren kurz hinter der Mitte am breitesten, äu- ßerer Spitzenwinkel breit verrundet, Streifen fein eingedrückt und dicht, fein punktiert, Intervalle flach, fein und ungleichmäßig dicht punktiert, an der Basis mit kurzer, weißer, feiner, anliegender Behaarung in den Punkten, alle Intervalle mit ein- zelnen weißen schuppenförmigen aber schmalen Haaren, deren Punkte doppelt so groß wie die übri- gen sind, in den feinen Punkten nur mikroskopisch behaart; Epipleuralkante kräftig, endet am äußeren Spitzenwinkel, Epipleuren locker und lang behaart, Elytrenspitzen mit einem schmalen Saum aus fei- nen weißen Mikrohärchen.

Unterseite einschließlich Metasternum nur fein und spärlich behaart, Metacoxalplatten bis auf die lateralen Borsten nur mikroskopisch fein behaart. Abdominalsternite bis auf je eine Querborstenreihe nur mikroskopisch fein behaart, letztes Sternit zu- sätzlich mit einzelnen langen Haaren, vorletztes Ster- nit ohne Erhabenheiten. Mesosternum zwischen den Mesocoxae so breit wie der schmale Metafemur. Verhältnis Metepisternen-/Metacoxalänge: 1/1,5. Pygidium kräftig gewölbt, fein und dicht punktiert, längs der Mitte breit glatt, mit mäßig dichter, kurzer und langer, gelber Behaarung, Pygidium beim ? fast flach.

Meso- wie Metafemur schmal und matt, Meta- femur mit durchgehend krenulierter Kante am Vor- derrand, fein und spärlich punktiert und kahl, Längs- haarpunktreihen deutlich, Hinterrand fast gerade, unbehaart, Hinterrandkante ventral und apikal auch dorsal fein krenuliert. Metatibien mäßig schlank, im hinteren Drittel am breitesten, basal leicht verschmä- lert, Verhältnis Breite/Länge der Metatibien: 1/3,3

(3), 1/3,0 (2); dorsal deutlich gekantet, mit krenu- lierter Längsleiste parallel zur der Oberkante, mit dieser konvergent ab der schwach angedeuteten, distalen Querleiste bei % der Metatibienlänge, ne- ben der krenulierten Leiste einzelne kräftige Haare; außen schwach längs gewölbt, locker und mäßig kräftig punktiert, im basalen Drittel in den Punkten kurz behaart, sonst nur mit mikroskopischen Här- chen in den Punkten; Ventralkante mit drei kräfti- gen, leicht gehöckert stehenden Borsten in etwa gleichen gleichem Abstand zueinander, Innenseite fein, locker punktiert, apikal in Nähe der Tarsalein- lenkung nur wenig tief und etwas konkav ausge- schnitten. Tarsen ventral fein und mäßig dicht be- haart, Meso- und Metatarsen dorsal dicht punktiert und seitlich ungekantet, Metatarsen lateral schwach längs gefurcht, ventral neben der krenulierten Lei- ste eine kräftige Längskante, Metatarsomer 1-4 ven- tral kahl, Metatarsomer 1 fast so lang wie die zwei folgenden Glieder zusammen und % länger als der obere Enddorn der Metatibien. Protibia mäßig lang, 2-zähnig, Protarsomere dorsal unpunktiert, Glieder 1-4 nur wenig breiter als lang, alle Klauen einfach ausgebildet und symmetrisch. Aedoeagus: Abb. 7-10.

Amiserica omeiensis, spec. noV. Abb. 11, 12

Typen. Holotypus: d, Mt. Omei Szechuan, China VII- 16-32 Franck, Coll./ F. C. Hadden Collection” (BPBM). Paratypen: 1d, 12, gleiche Angaben wie der Holotypus (BPBM, CA); 14, 222, China: Sichuan 27.V1.-2.VD. Ji- ulonggou near Dayi 1995 (= Chongging Jiulong Valley) ca 60 km W of Chengdu 31°00'N 103°30'E M. Tryzna et O. Safränek Igt. (CP, CA).

Beschreibung

Länge: 6,8-7,6 mm, Elytrenlänge: 5,3-5,4 mm, Breite: 4,3-4,4 mm.

Gestalt länglich oval; dunkel rotbraun, teilweise mit grünlichem Schimmer, Pronotum und Elytren mit mehr oder weniger ausgedehnten, hellen Flek- ken, gesamte Oberfläche bis auf den schwach glän- zenden Vorderkopf matt; Oberseite einzeln, kurz abstehend behaart.

Labroclypeus mäßig breit, deutlich breiter als lang, Seitenränder in der basalen Hälfte gerade und schwach verengt, vorn kräftig nach innen gebogen, Vorderecken mäßig abgerundet, Ränder leicht auf- gebogen und vorn median deutlich ausgerandet; Oberfläche in der Mitte schwach konvex, fein und dicht punktiert, der Abstand der Punkte ist kleiner als ihr Durchmesser, mit zahlreichen abstehenden Borsten, deren Punkte etwas größer als die übrigen

sind; Frontoclypealnaht sehr fein eingeritzt und schwach gebogen; Ocularcanthus mäßig lang und sehr schmal, sehr fein und zerstreut punktiert, mit einer einzelnen Terminalborste, Augenspiegel 2,5 mal so breit wie lang. Punktur der Stirn fein und mäßig dicht, neben den Augen und auf der Stirn- mitte wenige einzelne Haare. Augen mäßig groß, Verhältnis Augendurchmesser/-abstand: 0,55 (d), 0,46 (2). Fühler 9-gliedrig, gelb, Keule 3-gliedrig, beim d 1,5 mal so lang die übrigen Glieder zusam- men und kaum gebogen, beim ? Fühlerkeule etwas kürzer als die übrigen Glieder zusammen. Mentum erhaben, vorn abgeplattet.

Pronotum breit, an