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Volume 78, 1950
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Notes on Staphylinidae, Chiefly from New Zealand

[Read before the Nelson Philosophical Society, May 17, 1948; received by the Editor, June 16, 1948.]

(2) A new Genus and Three new Species of Eleusiini

A Close examination of the New Zealand Eleusis virgula Fvl. indicated that this species is not congeneric with typical Eleusis spp. and belongs to an, as yet, undescribed genus, differing chiefly in the structure of the tarsi and the meso- and metasternum.

The type of Eleusis, E. tibialis Lap. was not available for study and species from the type locality (Madagascar) were taken as typical of the genus. These included E. ernestini Lap. (the type of Chasolium Lap., a synonym of Eleusis.), E. complanatus Er. and E. brachyterus Boh.

In order to ascertain whether any other species were congeneric with virgula, all available Eleusis spp. (well over 100) were examined and ten found which are closely related to the New Zealand insect, three of them being new. This examination also showed that there is a considerable variation in the form of the mandibles, gular sutures and maxillary palpi in Eleusis. In the majority of the species examined, the second segment of the maxillary palpi, though varying greatly in length, is always distinctly longer than broad. In a few species, however, of which E. humilis Er. may be taken as representative, this second segment is distinctly transverse. In none of the species examined was this segment as long as broad.

With a view to simplifying the classification of a large and polymorphic genus, humilis and its allies may well be separated off as a separate subgenus on the character of the maxillary palpi. The name Isomalus Er. is available for this subgenus, as although in his description, Erichson (1840) figures complanatus Er., which has the maxillary palpi of a typical Eleusis, Duponchel (1841) fixed humilis Er. as the genotype. The other characters mentioned above (mandibles and gular sutures) may be of value in erecting further subgenera, but this is outside the scope of the present work.

A description of the new genus (Zeoleusis) erected for E. virgula Fvl. and the like species is given below, together with a key to and descriptions of the known species of the genus and a key to the genera and subgenera of the Eleusiini.

I am greatly indebted to Dr. H. Scott and Mr. E. B. Britton for facilities for examining the material in the British Museum collection and for the loan of specimens, to Dr. R. Malaise, of the Naturhistoriska Riksmuseum, Stockholm, for the loan of a series of Z. parva (BIb.), to Dr. M. Cameron for the use of his collection and for numerous specimens, to Dr. G. C. Hale Carpenter (Oxford University Museum), Dr. E. C. Zimmermann (Bernice P. Bishop Museum. Honolulu), and

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Fig. 1—Zeoleusis virgula Fvl. (Large male) The scale = 2 mm, and is divided into ½ mm.

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Mr. E. S. Gourlay (Cawthron Institute) for the loan of specimens of Z. virgula (Fvl.), to Mr. J. R. Helfer for a series of Z. helferi n. sp., and to M. J. Jarrige for specimens of E. brachypterus Boh.

Key to the Genera and Subgenerara of Eleusiini

1. Tarsi with five movable segments −2
— Tarsi with the first three segments fused and separ ated by sutures, i.e. only three movable segments Genus Zeoleusis n.
2. Pronotum dull, very strongly and rather coarsely alutaceous, much longer than broad, the sides sub-parallel for the anterior two-thirds, thence somewhat narrowed to base. Second segment of maxillary palpi distinctly transverse Genus Eumalus Sharp
— Pronotum always at least moderately shining, never strongly alutaceous, at most slightly longer than broad, the sides sub-parallel for at most the anterior half. Second segment of maxillary palpi variable Genus Eleusis Lap.—3
3. Second segment of maxillary palpi as long as or longer than broad Subgenus Eleusis s.str.
— Second segment of maxillary palpi distinctly transverse Subgenus Isomalus Er.

Genus Zeoleusis, n.

Small, moderately convex (insularis, sororcula) or depressed species.

Head in male rather large or large, the post ocular region distinctly longer than the longitudinal diameter of the eye in some species (parva, annae, politus, semirufus), about as long as (virgula, exigua, sororcula). or distinctly shorter (banghaasi, helferi, fusciceps) than the eye in others.* Head in female smaller, the post ocular region barely half as long as the longitudinal diameter of the eye in all species examined (politus and sororcula not seen), except parva, where it is about as long as the eye. Gular sutures converging behind and meeting at about the middle of the ventral surface of the head, obliterated just in front of this point (Fig. 2). Antennae slightly shorter than head and pronotum together, all the segments sparingly setose, the fourth to eleventh also sparingly pubescent; the first segment much longer than the second and broader than the second to fifth. Labrum (Fig. 3) transverse, bearing several long setae, the anterior margin semicircularly emarginate and with a very narrow densely ciliate membranous fringe which is forwardly produced on each side. Mandibles markedly asymmetrical in males in all species (except possibly insularis and sororcula, the mandibles of which were not examined), the right sharply deflexed inwards (almost at right angles) at about the middle of its length and with a small tooth internally behind middle, the left varying according to the species. Mandibles of females varying according to the species, curved and not markedly asymmetrical except in parva, where they are similar to those of the male. Inner lobe of maxilla (lacinia) (Fig. 4)

[Footnote] * Examination of a large series of virgula shows that there is some variation in the length of the post ocular region in the male, this varying from about as long as the eye (large specimens) to about three-quarters as long as the eye (small specimens). In the small specimens. there is also a difference in the form of the mandibles. No such variation was noted in any other species of the genus, but much smaller series were available for examination.

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FIg. 2–Z. virgula Fvl. (small male). Underside of head and thorax

narrow, the apex drawn out into one or two strong, inwardly curved spinules, the inner margin with a eiliate area towards apex, the outer (galea) broader, with a few rows of ciliae at apex, the tips of the ciliae being sharply curved. Maxillary palpi with the first segment short, transverse, the second longer, about one and one-half times as broad as long, widened apically, the third more than twice as long as the second, the sides gently rounded, the fourth slightly shorter than and about half as wide as third, parallel sided with the apex rounded. Mentum (Fig. 5) transverse, very short at sides, angularly produced in middle anteriorly. Glossae (Fig. 5) small, membranous, fused, the resultant lobe with a longitudinal median chitinized strip; on each side of this with a ciliate membranous process. Paraglossae large, distinct, extending forwards beyond the glossae. Lobes of the hypopharynx well developed, strongly ciliate internally and extending to the apex of the paraglossae. Labial palpi with the first segment about as broad as long, the second about as long as and slightly narrower than the first, the third narrower than and more than one and

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Fig. 3—Z. virgula Fvl. Labrum

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Fig. 4—Z. virgula Fvl. Maxilla

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Fig. 5—Z. virgula Fvl. Labium

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Fig. 6 Z. virgula Fvl. Intermediate tarsus

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one-half times as long as the second, parallel sided, with the apex rounded.

Pronotum with the anterior margin distinctly wider than the posterior, the sides rounded and with, at most, a trace of a small notch towards base. Pronotal epipleura narrow, the epimera represented by a slight thickening of the epipleura near base. Prosternum rather long, very short at sides, its process extending backwards between the anterior coxae for about half their length. Mesosternum short, apparently fused with the metasternum,* meso- and metasternal processes fused and forming a narrow ridge separating the intermediate coxae, this ridge in the middle not reaching the level of the sterna. Scutellum distinet, rather large. Wings well developed. Elytra as long or longer than broad, the epipleural folds well marked.

Tergite of the second abdominal segment short, chitinized, entire, the sternite entire and very lightly chitnized. Third to sixth (first to third visible) segments, parallel sided, about equal in length and width, the seventh a little longer than the sixth, slightly narrowed posteriorly, abdomen narrowed thence to apex. Tergites of the third to seventh segments with a very narrow raised margin at sides, but paratergites completely lacking. Sternite of the third segment with a well marked longitudinal median keel.

Legs rather short, the coxae small, the anterior globose, the intermediate oval, the posterior expanded above (dorsal to) the femora, all the coxae very narrowly separated. Femora rather broad. Tibiae narrow, the anterior not emarginate externally towards apex, externally with one or two setae near apex and a longer one about middle, setose internally on apical half; the intermediate with rows of spines externally on apical two-thirds and two long setae externally about middle, sparingly setose internally on apical half; the posterior with rows of spines externally on apical third and a long seta externally about middle, sparingly setose internally. All the tibiae with stout apical spurs. Tarsi (Fig. 6) with the first three segments fused and separated by sutures, so that there are only three movable segments. The first segment about as long as broad, the second extremely short, the third about as long as broad, deeply excised apically on dorsal surface, the base of the fourth (second movable) segment fitting into the excision, the fourth almost as long as the three fused segments together, deeply excised apically on dorsal surface to take the base of the fifth (third movable), which is much longer than the four preceding together.

Genotype: Eleusis virgula Fvl.

Range: India, Ceylon, Malay Peninsula, Philippine Is., East Indies, Australia. New Zealand, Rapa Is., and Juan Fernandez Is.

This genus may easily be separated from its allies by the form of the tarsi and the meso- and metasternum.

[Footnote] * Thirteen microscopical preparations of Zeoleusis spp. have been examined, including two dissections of the thoracic sterna, and in no case has it been possible to see any trace of a suture separating either the meso- and metathorax or their processes.

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With the exception of parva, the female of which is somewhat aberrant, the species form a more or less homogenous group and the females (with the exception of the above-mentioned species and possibly politus, of which no female has been seen) are very similar in facies so that identification is sometimes difficult. However, the spermathecae of five species have been examined and show differences which appear to be of significant taxonomic value.*

The males are somewhat easier to identify as there is a difference in the length of the post ocular region of the head in most of the species. The aedeagi of a few species have been examined, but do not appear to be of much value for differentiation.

Besides those mentioned above, other characters available for classification are the relative length and breadth of the elytra, the presence or absence of, and density of, ground sculpture and the form of the mandibles. The shape of the pronotum appears to be remarkably constant throughout the genus, as does the form of the antennae, politus being the only known species which can be separated using antennal characters.

The two species from Rapa Island (insularis and sororcula) differ from the others in the distinctly less depressed form, but are otherwise very similar.

Nothing appears to be known as to the habits and life-histories of Zeoleusis spp., except that they are found under the bark of decaying trees.

Key to the Known Species of Zeoleusis

1. Pronotum black or brown, elytra black or brown, abdomen black, legs pitchy −2.
— Pronotum reddish testaceous, elytra normally blackish with about basal third reddish testaceous, sometimes wholly reddish or pitchy testaceous, abdomen with at least the third to sixth (first to third visible) segments reddish or pitchy testaceous −4.
2. Size smaller (2–2 ½ mm.), moderately convex. Head, pronotum and abdomen black, elytra black with the sutural region sometimes obscurely brownish. (Rapa Island.) −3.
— Size larger (3 mm.). Head and abdomen black, pronotum and elytra brown, the latter sometimes with apical two-thirds black. Left mandible of male stout, curved, pointed apically, internally from base to in front of middle with a chitinized plate which is rounded anteriorly (Fig. 8). (Juan Fernandez Is.) 3. semirufus (Fairm. & Germ.)
3. Size larger (2 ½ mm.), elytra slightly longer than broad, about one and one-third times as long as pronotum, pronotum with two rather superficial longitudinal impressions −1. insularis (Cam.)

[Footnote] * In all cases where the spermatheca has been examined, at least two specimens of each species have been used. The organ appears to be constant in form for any one species.

[Footnote] † For this and the following species (sororcula), Dr Cameron's measurements for the length of the elytra are given and these were, presumably, taken from the apex to the extreme base. In all the other species dealt with here, the elytra are measured from the humeral angle to the apex.

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— Size smaller (2 mm.), elytra much longer than broad, nearly twice as long as the pronotum, which is without longitudinal impressions 2. sororcula (Cam.)
4. Elytra about one and one-fifth times as long as broad and at least one and one-half times as long as pronotum. Loft mandible of male normally strongly emargiutate internally just before the apex, which is pointed (Figs. 20, 24). Head with the post ocular region varying in length according to the species, very short in females, at most half as long as the eye (except possibly in politus, of which no female has been seen) −5.
— Elytra not or scarcely (one and one-eighth times) longer than broad, at most one and one-third times as long as pronotum. Left mandible of male variable. Head with the post ocular region varying in length in both sexes, according to the species −9.
5. Head and pronotum very shining, at most with traces of ground sculpture at extreme front and sides. Head in male with a well marked ridge from inner border of each eye almost to the base 6.
— Head and pronotum less shining, with a distinct though sometimes diffuse ground sculpture over the whole surface. Head in male without longi tudinal ridges −7.
6. Head brown, pronotum reddish testaceous, abdomen reddish testaceous, with the seventh and eighth (fifth and sixth visible) segments brown. Head with the post ocular region slightly longer than the longitudinal diameter of the eye (Fig. 19), from inner margin of each eye almost to base of head, with a longitudinal sulcus which is bounded externally by a ridge. Antennae with the second segment slightly longer than the third. Length 2 ½ mm. (Ceylon) 6. politus n.sp. ♂
— Head and pronotum reddish testaceous, abdomen reddish testaceous sometimes slightly infuseate apically. Head with post ocular region about one and one-half times as long as the longitudinal diameter of the eye (Fig. 14), the longitudinal ridges not bounding sulci. Antennae with the second segment slightly shorter than the third. Length 2 ½ mm. (Australia) 5. annac n.sp. ♂
7. Head with the post ocular region at most half as long as the longitudinal diameter of the eye (Figs. 15, 23) −8.
— Head with the post ocular region at least threequarters as long as the longitudinal diameter of the eye. Length 2–2i mm. (New Zealand) 7. virgula (Fvl.) ♂
8. Ground sculpture close and very distinct. Mandibles curved, the left (Fig. 27) bifid at apex and without internal teeth, the right (Fig. 28) pointed apically and with an indistinct tooth internally in front of middle. Spermathecae characteristic. Length 2 mm. (New Zealand) 7. virgula (Fvl.) ♀
— Ground sculpture less close and more superficial. Mandibles curved, the left (Fig. 16) pointed apically and edentate internally, the right (Fig. 17) pointed and with a well-marked tooth in front of middle internally. Spermathecae charac teristic (Fig. 18). Length 2 mm. (Australia) 5. annac ii.sp. ♀
9. Size larger (2–2J mm.). Ground sculpture absent mi head and pronotum except for alight traces towards sides −10.
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— Size smaller (1 ½ mm.). Ground sculpture on head and pronotum distinct, though sometimes diffuse, covering the whole surface −11.
10. Head with the post ocular region almost twice as long as the longitudinal diameter of the eye (Fig. 9). Left mandible (as Fig. 20) stout, semicircularly emarginate just before the apex, which is pointed. Length 2–2½ mm. (Australia) 4. parva (Bib.) ♂
— Head with the post ocular region about as long as the longitudinal diameter of the eye (Fig. 10). Right mandible similar to that of male, the left narrower, pointed apically and with a well marked pointed tooth internally in front of middle (Fig. 11). Length 2 mm. (Australia) 4. parra (Blb.) ♀
11. Elytra as long as broad. Head in male at most as long as broad (Fig. 30) −12.
— Elytra slightly but distinctly (about one and one-eighth times) longer than broad. Head in male as long as or longer than broad (Figs. 34, 35) −13.
12. Head reddish testaceous, antennae testaceous, slightly infuscate apically, abdomen reddish testaceous. Ground sculpture on head and pronotum not very close and sometimes rather indistinct. Length 1 ⅓–1 ½ mm. (Philippine Islands) 9. banghaasi (Bernh.)
— Head black, antennae black with first three segments testaceous, abdomen pitchy testaceous. Ground sculpture on head and pronotum close and very distinct. Length 1 ¼–1 ½ mm. (New Guinea) 8. helferi n.sp.
13. Ground sculpture on head and pronotum not very close and sometimes indistinct. Head of male with the post ocular region slightly longer than the longitudinal diameter of the eye (Fig. 34). Length 1 ½–2 mm. (Malay Peninsula, Philippine Islands) 10. exigua (Bernh.) [= decipiens (Cam.) n.syn.]
— Ground sculpture on head and pronotum close and distinct. Head of male with the post ocular region about three-quarters as long as the longitudinal diameter of the eye (Fig. 35). Length 1 ¼–1 ½ mm. (India, Ceylon, Malay Peniusula, Philippine Islands) 11. fusciceps (Kr.)

The ground sculpture on the head, pronotum and elytra (when present) consists, unless otherwise stated, of fine wavy lines of which most, but not all, may be said to be longitudinal in direction. That on the abdomen is similar, though somewhat finer and transverse in direction.

1. Zeoleusis insularis (Cameron)

Eleusis insularis Cameron (1936)

Moderately shining, rather convex. Head, pronotum and abdomen black, the elytra black with the sutural region sometimes obscurely brownish. Antennae with the first segment black, the second and third testaceous, the remainder black. Mouth parts testaceous. Legs pitchy. Length 2 ½ mm.

♂ Head moderate, about as long as broad, the post ocular region slightly longer than the longitudinal diameter of the eye. Dorsal surface with a distinct ground sculpture and a few scattered fine punctures. Antennae with the third segment slightly longer than second.

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the fourth subquadrate, the fifth and sixth very slightly transverse, the seventh to tenth more transverse, but not strongly so, the eleventh longer than broad, bluntly pointed, nearly as long as the two preceding together.

Pronotum about as long as broad, the sides gently rounded and slightly constricted just before base; on each side of middle with a rather broad, superficial, longitudinal impression; surface with a few scattered fine punctures and a strong ground sculpture which is mainly coriaceous.

Elytra slightly longer than broad, about one and one-third times as long as pronotum, rather diffusely punctured and with a distinct ground sculpture.

Tergites of abdomen with a few scattered setiferous punctures and a distinct ground sculpture which is coriaceous in places.

Type locality: Rapa Island.

Type in the Bernice P. Bishop Museum collection.

Records: Rapa Island, Mount Vaitau, 1,100 feet (E. C. Zimmermann), under dead puru bark.

Specimens examined: One paratype in Dr Cameron's collection.

2. Zeoleusis sororcula (Cameron)

Eleusis sororcula Cameron (1936)

Moderately shining, rather convex. Head, pronotum and abdomen black, the elytra black with the sutural region sometimes brownish. Antennae with the first segment black, the second and third testaceous, the remainder black. Mouth parts testaceous. Legs pitchy. Length 2 mm.

♂ Head moderate, about as long as broad, the post ocular region distinctly shorter than the longitudinal diameter of the eye. Dorsal surface with a distinct ground sculpture and a few scattered fine punctures. Antennae with the third segment slightly longer than the second, the fourth subquadrate, the fifth and sixth slightly transverse, the seventh to tenth more transverse, the eleventh longer than broad, bluntly pointed, nearly as long as the preceding two together.

Pronotum about as long as broad, the sides gently rounded and without constriction near base; on each side of middle with a setiferous puncture; surface with a few scattered fine punctures and a distinct ground sculpture of the normal type.

Elytra distinctly longer than broad, nearly twice as long as pronotum, very finely and diffusely punctured, and with distinct ground sculpture.

Tergites of abdomen with a few scattered setiferous punctures and a distinct ground sculpture of the normal type.

Type locality: Rapa Island.

Type in the Bernice P. Bishop Museum collection.

Records: Rapa Island, Mount Vaitau, 1,100 feet (E. C. Zimmermann), under dead puru bark.

Specimens examined: One paratype in Dr Cameron's collection.

This species is closely allied to Z. insularis, from which it differs in the shape and sculpture of the pronotum and the length of the

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elytra. The specimen of sororcula examined is a female and that of insularis is most probably a male, but this is not certain. From the description, the types are of the same sexes as the specimens seen, and as all the specimens were taken in the same locality, it is possible that they may represent the sexes of a single species.

3. Zeoleusis semirufus (Fairmaire and Germain) (Figs. 7–8)

Eleusis semirufus Fairmaire and Germain (1861)

Somewhat dull, depressed. Head and abdomen black, pronotum brown, elytra brown, sometimes with the apical two-thirds black. Antennae black with the first three segments reddish testaceous. Mouth parts reddish testaceous, legs pitchy. Length 3 mm.

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Fig. 7—Z. semirufus F. and G. Head of male. Scale = ½ mm.

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Fig. 8 Z. semirufus F. & G. Left mandible. Male

♂ Head rather large, longer than broad, the post ocular region about one and one-third times as long as the longitudinal diameter of the eye. Dorsal surface with a distinct, close ground sculpture of the normal type and a few scattered fine punctures. Antennae with the second segment about as long as third, the fourth subquadrate, the fifth slightly transverse, the sixth to tenth gradually becoming more transverse, the tenth being almost twice as broad as long, the eleventh longer than broad, bluntly pointed, about as long as the preceding two together. Left mandible (Fig. 8) stout, curved, pointed apically; internally, from base to in front of middle with a chitinized plate which is rounded in front.

Pronotum about as long as broad, the sides rounded. Surface with a few scattered setiferous punctures towards sides and a rather sparse, longitudinally vermiculate ground sculpture.

Elytra about one and one-fifth times as long as broad, about one and one-half times as long as pronotum. Practically impunctate, and with a few longitudinal vermiculations.

Tergites of abdomen with a few scattered setiferous punctures and a distinct ground sculpture which is coriaceous in places.

♀ Head about as long as broad, the post ocular region barely half as long as the longitudinal diameter of the eye. Mandibles curved, the left bifid at apex and without internal teeth, the right pointed and

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with an indistinct tooth internally in front of middle. (As virgula, Figs. 27–28.) Ground sculpture of pronotum more diffuse than in male. Ground sculpture of abdominal tergites consisting of rather diffuse transverse vermiculations.

In other respects similar to male.

Type locality: Juan Fernandez Islands.

Type in the British Museum collection.

Records: Type locality only.

Specimens examined: The type and four specimens from the type locality collected by J. J. Walker and in the British Museum collection.

This species is the largest yet known in the genus and is quite distinct on account of its coloration.

4. Zeoleusis parva (Blackburn) (Figs. 9–13)

Elcusis parva Blackburn (1892)

Shining, depressed. Head, pronotum and abdomen reddish testaceous, elytra blackish with about basal third reddish testaceous. Antennae reddish testaceous, slightly infuscate apically. Mouth parts testaceous. Legs reddish testaceous. Length 2–2 ½ mm.

♂ Head large, distinctly longer than broad, post ocular region practically twice as long as the longitudinal diameter of the eye, the posterior angles rounded, not very distinct. From inner margin of each eye almost to base, with traces of an indistinct longitudinal ridge. Dorsal surface practically impunctate and without ground sculpture

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Fig. 9—Z. parra Blb. Head of male Scale = ½ mm.

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Fig. 10—Z. parra Blb. Head of female

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Fig. 11—Z. parra Blb. Left mandible Female

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Fig. 12—Z. parra Blb. Right mandible Female

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Fig. 13—Z. parra Blb. Spermatheea

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except for very few short vermiculations just internal to the eyes and posterior angles. Antennae with the second segment slightly shorter than the third, the fourth subquadrate, the fifth slightly transverse, the sixth to tenth gradually becoming more transverse, the eleventh longer than broad, bluntly pointed, about as long as the two preceding together. Left mandible broad, semi-circularly emarginate internally just before the apex which is pointed (similar to politus. Fig. 20).

Pronotum about as long as broad, the sides rounded. Surface with a few scattered setiferous punctures towards sides and at most with faint traces of ground sculpture towards extreme sides.

Elytra about one and one-eighth times as long as broad, about one and one-third times as long as pronotum. Practically impunctate and without visible ground sculpture.

Tergites of abdomen with a few scattered setiferous punctures and traces of ground sculpture at extreme sides.

♀ Head rather large, longer than broad, the post ocular region about as long as the longitudinal diameter of the eye. Right mandible (Fig. 12) similar to that of male, the left (Fig. 11) curved, pointed apically, with a well-marked pointed tooth internally in front of middle. Spermatheca characteristic (Fig. 13).

In other respects similar to male.

Type locality: New South Wales (Blue Mountains).

Type in the British Museum collection.

Records: New South Wales—Blue Mountains, under eucalyptus bark; Queensland—Blackal Range (Mjoberg), Atherton (Mjoberg).

Specimens examined: The type, eight specimens from the Naturhistoriska Riksmuseum, Stockholm, and one specimen in Dr Cameron's collection.

This species is noteworthy in that it is the only one so far known in which the shape of the head in the female shows a departure from the normal type. It is also the only species in which the right mandible of the female is similar to that of the male.

Bernhauer (1920) recorded specimens of this insect from Blackal Range, Queensland, as Eleusis apicipennis Fairm., to the female of which it bears a very superficial resemblance. A specimen from Atherton in Dr Cameron's collection also bears Bernhauer's label “apicipennis.”

5. Zeoleusis annae n.sp. (Figs. 14–18)

Shining, depressed. Head, pronotum and abdomen reddish testaceous, the head sometimes slightly darker, elytra blackish with about basal third reddish testaceous or (?immature specimens) entirely testaceous. Antennae reddish testaceous, slightly infuscate towards apex. Month parts testaceous, legs reddish testaceous. Length 2–2 ½ mm.

♂ Head rather large, longer than broad, post ocular region rather more than one and one-half times as long as the longitudinal diameter of the eye, the posterior angles rounded, distinct; from inner margin of each eye almost to base, with a distinct ridge. Dorsal surface practically impunctate and without ground sculpture except for very few short vermiculations just internal to the eyes and posterior angles. Antennae with the second segment slightly shorter than the

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third, the fourth subquadrate, the fifth slightly transverse, the sixth to tenth gradually becoming more transverse, the eleventh longer than broad, bluntly pointed, about as long as the preceding two together. Left mandible broad, semicircularly emarginate internally just before the apex which is pointed (similar to politus, Fig. 20).

Pronotum about as long as broad, the sides rounded. Surface with a few scattered setiferous punctures towards sides and at most with faint traces of ground sculpture at extreme sides.

Elytra about one and one-fifth times as long as broad, just over one and one-half times as long as pronotum. Practically impunctate and without visible ground sculpture.

Tergites of abdomen with a few scattered setiferous punctures and an indistinct ground sculpture of the normal type at extreme sides.

♂ Head rather small, scarcely as long as broad, the post ocular region barely half as long as the longitudinal diameter of the eye. Dorsal surface practically impunctate and with a distinct though diffuse ground sculpture of the normal type. Left mandible (Fig. 16) curved, pointed at apex and edentate internally, the right (Fig. 17) curved, pointed apically, and with a small tooth internally in front of middle. Pronotum and elytra with a distinct though not very close

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Fig. 14—Z. annae n.sp. Head of male Scale = ½ mm.

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Fig. 15—Z. annae n.sp. Head of female

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Fig. 16—Z. annae n.sp. Left mandible Female

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Fig. 17—Z. annae n.sp. Right mandible Female

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Fig. 18—Z. annae n.sp. Spermatheca

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ground sculpture of the normal type. Spermatheca characteristic (Fig. 18).

In other respects similar to male.

Type locality: New South Wales—Sydney.

Type (♂) and one damaged ♀ in my collection, two paratypes, (♂,♀) in the British Museum collection.

Records: New South Wales—Sydney, Richmond River.

Specimens examined: The three types and the damaged ♂ only.

The male of this species is very similar to Z. parva (Blb.), but may be distinguished by the longer elytra, more marked posterior angles of the head and the longitudinal ridges on the head. The female is abundantly distinct by reason of the much smaller head and different mandibles.

The aedeagi of this species and Z. parva have been examined, but do not appear to be of much value as a taxonomic character.

6. Zeoleusis politus n.sp. (Figs. 19–20)

Shining, depressed. Head brown, pronotum reddish testaceous, elytra brownish with the basal third lighter, abdomen with the third to sixth (first to third visible) segments reddish testaceous, the seventh and eighth (fifth and sixth visible) brown. Antennae, mouth parts and legs reddish testaceous. Length 2 ½ mm.

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Fig. 19—Z. politus n.sp. Head of male. Scale = ½ mm.

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Fig. 20 Z. politus n.sp. Left mandible of male

♂ Head moderate, about as broad as long, post ocular region about as long as the longitudinal diameter of the eye. From inner margin of eye almost to base with a narrow longitudinal sulcus which is bounded internally by a ridge. Dorsal surface with a few scattered punctures and without ground sculpture except for traces at extreme sides of neck. Antennae with the second segment slightly longer than third, the fourth and fifth slightly longer than broad, the sixth subquadrate, the seventh to tenth slightly transverse, the eleventh longer than broad, bluntly pointed, nearly as long as the two preceding together. Left mandible (Fig. 20) stout, angularly emarginate internally just before the apex, which is pointed.

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Pronotum about as long as broad, the sides rounded. Surface practically impunctate and without visible ground sculpture.

Elytra just over one and one-fifth times as long as broad, about one and one-half times as long as pronotum. Practically impunctate and without ground sculpture.

Tergites of abdomen with a few scattered setiferous punctures and towards sides and apex with traces of a coriaceous ground sculpture.

Type locality: Ceylon.

Type (♂) in my collection.

Records: Type locality only.

Specimens examined: The unique type only.

This species, though allied to Z. parva and annae, may be easily distinguished by the colour, longer second segment of the antennae, and the well-marked longitudinal sulci on the head.

7. Zeoleusis virgula (Fauvel) (Figs. 1–6, 21–29)

Eleusis virgula Fauvel (1889)

Moderately shining, depressed. Head, pronotum and abdomen reddish testaceous, the head sometimes darker, elytra blackish with about basal third testaceous. Antennae reddish testaceous, somewhat infuseate apically. Mouth parts and legs reddish testaceous. Length 2–2 ½ mm.

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Fig. 21—Z. virgula Fvl. Head of large male. Scale = ½ min.

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Fig. 22—Z. virgula Fvl. Head of small male

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Fig. 23—Z. virgula Fvl. Head of female

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Fig. 24—V. virgula Fvl. Left mandible (large male)

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Fig. 25—Z. virgula Fvl. Right mandible (male)

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Fig. 26—Z. virgula Fvl. Left mandible (small male)

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Fig. 27—Z. virgula Fvl. Left mandible Female

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Fig. 28—Z. virgula Fvl. Right mandible Female

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Fig. 29—Z. virgula Fvl. Spermatheca

♂ Head moderate, about as broad as long, post ocular region about three-quarters as long as, to as long as, longitudinal diameter of eye. Dorsal surface with a few scattered fine punctures and a distinct close ground sculpture of the normal form. Antennae with the second segment slightly shorter than the third, the fourth subquadrate, the fifth to tenth gradually becoming more transverse, the eleventh longer than broad bluntly pointed, about as long as the two preceding together. Left mandible variable, in large specimens (those with the post ocular region about as long as the eye) broad, semicircularly emarginate internally just before the apex, which is pointed, the inner margin sometimes slightly angulate about middle (Fig. 24), in small specimens (those with the post ocular region distinctly shorter than the eye) pointed and with the inner margin straight and without emargination (Fig. 26).

Pronotum about as long as broad, the sides rounded. Surface with a few scattered setiferous punctures at sides and a distinct close ground sculpture of the normal type.

Elytra slightly over one and one-fifth times as long as broad, nearly one and two-thirds times as long as pronotum. Practically impunctate and with a ground sculpture similar to that of pronotum, but less close.

Tergites of abdomen with a few scattered setiferous punctures and a distinct, close ground sculpture of the normal type.

♀ Head rather small, scarcely as long as broad, post ocular region from distinctly less than half to about half as long as the longitudinal diameter of the eye. Mandibles symmetrical, curved, the left (Fig. 27) bifid at apex and edentate internally, the right (Fig. 28) pointed apically and with an indistinct tooth in front of middle internally. Ground sculpture as for male. Spermatheca characteristic (Fig. 29).

In other respects similar to male.

Type locality: New Zealand.

Type presumably in the collection of Blanche Rancin, Caen, France.

Records: New Zealand, North Island—Tairua [Auckland] (Broun), Parna [Whangarei] (Broun), Kerikeri (Brown), Wellington (J. J. Walker), South Island—Nelson (E. S. Gourlay), Mount Robert (Broun), Picton (Helms), Greymouth (Helms).

Specimens examined: 58 in the British Museum collection, 7 from the Broun collection, 7 from the Bernice P. Bishop Museum collection, 3 from the Hope Department, Oxford University Museum, 4 from the collection of Mr E. S. Gourlay, and 8 in my own collection.

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The male of this species is quite distinct, due to its size and sculpture. The female is very similar to that of Z. parva Bib., but may be distinguished by the closer ground sculpture and the form of the mandibles and spermatheca.

8. Zeoleusis helferi n.sp. (Figs. 30–33)

Rather dull, depressed. Head black, pronotum reddish testaceous, elytra black, with about basal third lighter, abdomen pitchy testaceous. Antennae black, with the first three segments testaceous. Mouth parts and legs testaceous. Length 1¼–1½ mm.

♂ Head moderate, about as long as broad, post ocular region distinctly shorter than the longitudinal diameter of the eye. Dorsal

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Fig. 30–Z. helferi n.sp. Head of male Scale = ¼ mm.

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Fig. 31—Left mandible (male)

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Fig. 32—Right mandible (male)

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Fig. 33—Z. helferi n.sp. Spermatheen

surface with a few scattered fine punctures and a close, fine ground sculpture. Antennae with the second and third joints subequal in length, the fourth subquadrate, the fifth slightly transverse, the sixth to tenth gradually becoming more transverse, eleventh longer than broad, bluntly pointed, about as long as the two preceding together. Left. mandible (Fig. 31) broad, bluntly pointed, the internal margin rounded.

Pronotum about as long as broad, the sides rounded. Surface with a few fine setiferous punctures towards sides and a close, distinct ground sculpture as on head.

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Elytra about as long as broad, about one and one-third times as long as pronotum. Practically impunctate, and with a ground sculpture similar to that of head and pronotum, but somewhat less close.

Tergites of abdomen with a few scattered setiferous punctures and a fine ground sculpture of the normal type.

♀ Head hardly as long as broad, post ocular region less than half as long as the longitudinal diameter of the eye. Mandibles symmetrical (as fusciceps, Figs. 37–38), the left bifid apically and edentate internally, the right pointed apically and with a small tooth internaly in front of middle. Ground sculpture as for male. Spermatheca characteristic (Fig. 33).

In other respects similar to male.

Type locality: Bosnek, Biak Island (New Guinea).

Holotype (♂), allotype (♀) and five paratypes in my collection, two further paratypes have been deposited in Dr Cameron's collection, and one more in the British Museum collection.

Records: Bosnek, Biak Island (New Guinea), in moist situations under the bark of dead trees, 30/7–6/8/44 (J. R. Helfer).

Specimens examined: The 10 types and 3 damaged females in my collection.

I have much pleasure in naming this insect after my friend, Mr -I. R. Helfer, of Mendocino, California, U.S.A., in gratitude for many Staphylinidae collected for me during his service in the Pacific area with the United States Army.

This species closely resembles Z. banghaasi (Bernh.) in facies and Z. fusciceps (Kr.) in sculpture. It is, however, easily separated from both by the colour and from banghaasi by the close ground sculpture and from fusciceps by the shorter elytra.

9. Zeoleusis banghaasi (Bernhauer)

Elcusis banghaasi Bernahauer (1927)

Rather shining, depressed. Head, pronotum and abdomen reddish testaceous, elytra blackish with about basal third testaceous. Antennae reddish testaceous, somewhat infuscate apically, mouth parts and legs testaceous. Length 11/4–11/2 mm.

♂ Head moderate, about as long as broad, post ocular region about three-quarters as long as the longitudinal diameter of the eye. Dorsal surface with a few scattered fine punctures and an indistinct, fine, not very close ground sculpture of the normal type. Antennae with the second and third segment subequal in length, the fourth subquadrate, the fifth slightly transverse, the sixth to tenth gradually becoming more transverse, eleventh longer than broad, bluntly pointed, about as long as the two preceding together. Left mandible short, bluntly pointed, the inner margin almost straight (as fusciceps, Fig. 36).

Pronotum about as long as broad, the sides rounded. Surface with a few setiferous punctures at sides and ground sculpture as on head.

Elytra about as long as broad, about one and one-third times as long as pronotum. Practically impunctate and with a ground sculpture similar to that on head and pronotum, but less close. Tergites of abdomen with a few scattered setiferous punctures, and a fine close ground sculpture of the normal type.

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♀ Head scarcely as long as broad, the post ocular region less than half as long as the longitudinal diameter of the eye. Mandibles symmetrical, curved (as fusciceps, Figs. 37–38), the left bifid at apex and edentate internally, the right pointed and with a small tooth internally in front of middle. Ground sculpture as for male, but somewhat closer.

In other respects similar to male.

Type locality: Philippine Islands.

Type in the collection of the late Dr M. Bernhauer.

Records: Philippine Islands—Siargao: Dapa; Palawan: Binaluan; Luzon: Provinz Mountain, Balbalasang; Mindanao: Provinz Lanae, Mumungan; Masbate: Arory. All the above records ex Bernhauer.

Specimens examined: One specimen from the type series in Dr Cameron's collection.

The above description has been drawn up from the specimen seen and from the original description.

The species may be identified by the length of the elytra and the not very close ground seulpture on the head and pronotum.

10. Zeoleusis exigua (Bernhauer) (Fig. 34)

Eleusis banghaasi var. exigua Bernhauer (1926)

Eleusis decipiens Cameron (1932) n.syn.

Rather shining, depressed. Head, pronotum, and abdomen reddish testaceous, the head sometimes darker, elytra reddish or pitchy testaceous, with about the basal third sometimes lighter. Antennae with the first three segments testaceous, the rest darker. Mouth parts and legs reddish testaceous. Length 11/2–2 mm.

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Fig. 34–Z. exigua Bnh. Head of male. Scale = ¼ mm.

♂ Head moderate, as long as broad, the post ocular region slightly longer than the longitudinal diameter of the eye. Dorsal surface with a few scattered fine punctures and a fine, distinct but not very close ground sculpture. Antennae with the second and third segments subequal in length, the fourth subquadrate, the fifth slightly transverse, the sixth to tenth gradually becoming more transverse, the eleventh longer than broad, bluntly pointed, about as long as the two preceding together. Left mandible stout, bluntly pointed, the inner margin almost straight (as fusciceps. Fig. 36).

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Pronotum about as long as broad, the sides rounded. Surface with a few setiferous punctures at sides and a ground sculpture similar to that of head.

Elytra about one and one-eighth times as long as broad, about one and one-third times as long as pronotum. Practically impunctate and with a ground sculpture similar to that on head and pronotum, but rather less close.

Tergites of abdomen with a few scattered setiferous punctures and a fine, rather close, ground sculpture of the normal type.

♀ Head scarcely as long as broad, the post ocular region less than half as long as the longitudinal diameter of the eye. Mandibles symmetrical, curved (as fusciceps, Figs. 37–38), the left bifid at apex and edentate internally, the right pointed and with a small tooth internally in front of middle. Ground sculpture as for male, but somewhat closer.

In other respects similar to male.

Type locality: exigua (Bernh.), Philippine Islands; decipiens (Cam.), Pahang.

Type of exigua (Bernh.) in the collection of the late Dr M. Bernhauor; of decipiens (Cam.), in Dr Cameron's collection.

Records: Philippine Islands (ex Bernhauer); Pahang: Frasers Hill, 4,000 ft.; Perak; Selangor.

Specimens examined: The type of decipiens and two other specimens in Dr Cameron's collection, one specimen in the British Museum collection, one specimen in my own collection.

The specimen in my collection (from Perak) is a male and was identified by Bernihauer as banghaasi. It does not, however, agree with the description of this species, but agrees well with that of the variety exigua. The type of decipiens Cam. was compared with this specimen and no significant difference could be found; the two species are therefore synonymised.

There is a marked difference in the shape of the head of the male and the length of the elytra between banghaasi and the var. exigua, and it does not seem likely that they represent only one species. The variety is therefore raised to specific rank.

Exigua is closely allied to fusciceps, from which the males may be separated by the length of the post ocular region of the head. The females, however, are very similar and the only means of separating them seems to be the density of the ground sculpture. This is distinctly closer in fusciceps and when the two species are seen together, the difference is easily seen. It has, unfortunately, not been possible, as yet, to examine the spermatheca of exigua, but this would be expected to be characteristic and should serve as a useful determinative feature.

11. Zeoleusis fusciceps (Kraatz) (Figs. 35–39)

Eleusis fusciceps Kraatz (1859)

Rather dull, depressed. Pronotum and abdomen reddish-testa-cous, the head darker, elytra lighter or darker testaceous, sometimes with about basal third lighter. Antennae with the first three segments testaceous, the rest darker. Mouthparts and legs testaceous. Length 11/4–11/2 mm.

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♂ Head moderate, broader than long, post ocular region about three-quarters as long as the longitudinal diameter of the eye. Dorsal surface with a few scattered fine punctures and a fine, close ground sculpture of the normal type. Antennae with the second and third segments subequal in length, the fourth subquadrate, the fifth slightly transverse, the sixth to tenth gradually becoming more transverse, eleventh longer than broad, bluntly pointed, about as long as the two

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Fig. 35—Z. fusciceps Kr. Head of male Scale = ¼ mm.

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Fig. 36 Z. fusciceps Kr. Left mandible (male)

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Fig. 37—Z. fusciecps Kr. Left mandible (female)

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Fig. 38—Z. fuseiceps Kr. Right mandible (female)

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Fig. 39—Z. fusciceps Kr. Spermatheca

preceding together. Left mandible stout, bluntly pointed, the inner margin almost straight. (Fig. 36).

Pronotum about as long as broad, the sides rounded. Surface with scattered setiferous punctures at sides and a close, distinct ground sculpture as on head.

Elytra about one and one-eighth times as long as broad, about one and one-third times as long as pronotum. Practically impunctate and with a distinct, rather close ground sculpture similar to that on head and pronotum.

Tergites of abdomen with a few scattered setiferous punctures, and a fine close ground sculpture of the normal type.

♀ Head scarcely as long as broad, the post ocular region less than half as long as the longitudinal diameter of the eye. Mandibles symmerical, the left (Fig. 37) bifid at the apex and edentate internally, the right (Fig. 38) pointed and with a small tooth internally

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in front of middle. Spermatheca characteristic (Fig. 39). Ground seulpture as for male.

In other respects similar to male.

Type locality: Ceylon.

Type: Presumably in the Deutsche Entomologische Institut, Berlin.

Records—India: Nilgiri Hills. Ceylon. Pahang: Fraser's Hill, 4,000 ft. Singapore: Bukit Panjang. Sumatra: Fort de Kock. Philippine Islands: Leyte (ex Bernhauer); Palawan, Binaluan (ex Bernhauer); Siargao (ex Bernhauer); Masbate Aroroy (ex Bernhauer).

Specimens examined: About 20 in the British Museum and Dr. Cameron's collection. Three in my own collection.

Specimens, apparently of this species, from India have been seen which have the elytra distinctly longer than normal. However, it has not been possible to examine any of these critically, so they are not dealt with here.

This species appears to be the most widely distributed of the genus.

References

Bernhauer, M., 1920. Ark. Zool., 13, no. 8.

— 1920. Philipp. J. Sci., 31, no. 2, 245–263.

Blackburn, T., 1892. Trans. Roy. Soc. S. Aust., 15.

Cameron, M., 1932. Bull. Raffles Mus., no. 7, 124–142.

— 1930. Occ. Pap. Bishop Mus., Honolulu, 12, no. 14, 3–9.

Duponchel, P. A. J., 1841. Rev. Zool. Soc. Cuvier.

Erichson, W. F., 1839. Gen. et Spec. Staph. Berlin.

Fairmaire, L., and Germain, P., 1861. Ann. Soc. Ent. Fr., ser. 4, 1.

Fauvel, A., 1880. Rev. Ent., Caen, 8, 243.

Kbaatz, G., 1859. Arch. Naturgesch., 25.