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Dr.ssa GIANNA ARBOCCO - Zoologia

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Gli articoli pubblicati sugli Annali del Museo Civico di Storia Naturale “Gia-

como Doria” sono citati e recensiti sui principali repertori bibliografici, tra cui:

Zoological Record, BIOSIS Previews, Biological Abstracts, Abstracts of Ento-

mology, CAB Abstracts, Review of Agricultural Entomology, Review of Medi-

cal and Veterinary Entomology.

The papers published in the Annali del Museo Civico di Storia Naturale “Giaco-

mo Doria” are abstracted and indexed in the main bibliographic records, such as:

Zoological Record, BIOSIS Previews, Biological Abstracts, Abstracts of Ento-

mology, CAB Abstracts, Review of Agricultural Entomology, Review of Medi-

cal and Veterinary Entomology.

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* ATTRAE

CESARE BARONI URBANI* & MARIA L. DE ANDRADE*

THE ANT TRIBE DACETINI: LIMITS AND

CONSTITUENT GENERA, WITH DESCRIPTIONS

OF NEW SPECIES

(HYMENOPTERA, FORMICIDAE)

“As new species are discovered... the characters separat-

ing... genera are gradually losing their distinctiveness, and tit

seems likely that we may eventually see most of these genera

merged again in one large genus”.

W. L. Brown (1973a: 3)

1. INTRODUCTION

BARONI URBANI & DE ANDRADE (1994) synonymized under the

tribal name Dacetini the two tribal names Phalacromyrmecini and

Basicerotini and merged a number of genera within the tribe.

This result was promptly contrasted by BOLTON (1995: 47) who

wrote: “I consider this action too extreme by far as it is based on a

grossly inadequate character set”, and added “In consequence their

[i.e. Baroni Urbani & de Andrade’s] results are very insecure, and

their proposed synonyms are all revived from synonymy [bold-

face by Bolton] here”. As a matter of fact BARONI URBANI & DE

ANDRADE (1994) reached their conclusions after examining each and

every character used by Bolton and others to separate the genera in

question. As a consequence of this, Bolton’s statement of character

inadequacy applies much better to his generic resurrections than to

the synonymies proposed by BARONI URBANI & DE ANDRADE (I. c.).

All these generic names re-appeared hence as valid names in

BoLTon (1995) and most of them also in a number of subsequent

papers, including BoLTON (1998) and in other papers published

afterwards by uncritical entomologists.

* Institut fiir Natur-, Landschafts- und Umweltschutz, Biogeographie, Universitat

Basel, St. Johanns-Vorstadt 10, CH-4056 Basel, Switzerland.

Z | _ C. BARONI URBANI & M.L. DE ANDRADE

None of these 1995-1998 generic revivals, however, is listed

where one would expect to find them, i.e. under the “taxonomic

history” of generic names in BoLTON (2003).

The reason for this omission is that only one year after his unex-

plained revival of our synonyms, BOLTON (1999), with a dramatic

inversion of course, synonymized again all genera already synony-

mized by BARONI URBANI & DE ANDRADE (1994) under Strumigenys

under two genera, i.e. Strumigenys (including only Quadristruma as

proposed by ourselves in 1994) and the revived genus Pyramica, a

former synonym of Strumigenys now including all other genera that

we previously synonymized under Strumigenys.

Pyramica, described by ROGER (1862), was considered as a syn-

onym of Strumigenys only one year after its description by the same

ROGER (1863b) and never received generic status again until BOLTON

(1999). This last, entirely new and radically different classificatory

scheme is adopted by BOLTON (2000, 2003) and by all his uncritical

followers that rejected our synonymies only one year before.

BOLTON’s (1999-2003) classificatory system is supported by an

impressive display of morphological erudition enforcing generic

boundaries, phylogenies, and conclusions by introducing a remark-

able set of new characters. Unfortunately, many of these characters

are used for the first time with the sole support of unclear descrip-

tions, and are seldom backed by good illustrations, definitions and

unequivocal statements about their presence or absence among criti-

cal taxa.

One of the greatest difficulties that we faced was interpreting a

number of Bolton’s characters. Often slightly different rewordings

of the same sentence are used in different papers to define presum-

ably different synapomorphies. For instance:

BoLTON (1998: 72) states that the “labrum with an impression...

on the labral shield, distal of the basal hinge but proximal to the

labral lobes” is synapomorphic for all his “dacetonines”. But, one

year later, BOLTON (1999: 1681) defines the “labrum mediodorsally

with a very broadly and deeply concave depression in its proximal

half” as autapomorphic for the genus Epopostruma. We are unable to

understand why these two very similar definitions should be consid-

ered as two independently derived characters related to two differ-

ent clades and not as two slightly different rewordings of the same

ANT TRIBE DACETINI 3

character description. We already raised this point in a web docu-

ment (BARONI URBANI & DE ANDRADE, 2006b) and BOLTON (2006b)

replied by adding that his 1998 definition is repeated in BoLTON

(2003:54). This is perfectly true, of course, but it does not justify at

all the double role attributed to the same character in two different

positions within the same clade.

In other instances, two characters used by Bolton in the same

context but for different purposes simply contradict each other:

BoLTON (1998: 73) writes that the “propodeal spiracle low on

side of sclerite, abutting the margin of the small metapleural gland

bulla” is unique to the Basicerotini and Phalacromyrmecini, while

the “Dacetonini (sic!)... [have the] propodeal spiracle... widely sepa-

rated from metapleural gland”. But BOLTON (1999: 1649, character

22 and table 1), contradicting his own former statement, attributes

the “metapleural gland bulla abutting or very close to annulus of

propodeal spiracle” to 5 out of 9 genera of Dacetini considered...

Other characters, like the katepisternal groove, given as synapo-

morphic for the tribe Phalacromyrmecini, proved to be present in

only one species and absent even in the type genus of the tribe,

Phalacromyrmex (see later the discussion of our character # 36),

i.e. they appear to have been imagined only to support a particular

classification. BOLTON (2006a) admits obtorto collo the failure of this

character but insists that it should be re-defined and considered just

the same...

These three examples, among many more possible ones, should

be sufficient to show the need for the present study and to justify

our previous appreciation of Bolton’s work. Other examples will be

dealt with under the individual character analysis in this paper.

Another very important difference lies on the fact that in BOLTON

(1999-2000), absence or presence of a given trait in a taxon are often

declared a priori as due to secondary loss or to homoplastic gain.

These statements, however plausible they may be, are equivalent

to the assertion that the evolutionary path of the group is already

known and invalidate the meaning of any phylogenetic deduction

based on them.

We made an effort to ban potential proclamations of faith from

our analysis.

+ C. BARONI URBANI & M.L. DE ANDRADE

This was already anticipated by BARONI URBANI & DE

ANDRADE (2006b) but BOLTON (2006b) insists on the correctness

of his approach and this forces us to further explain two additional

aspects: 1. In this paper, to infer phylogeny and classification, we

shall code a trait as present or absent or polymorphic in a given

genus according to its observed record in that genus; BOLTON

(1999), on the contrary, codes it in the way that he thinks better

reflecting his idea of the phylogeny. This has nothing to do with

the plausibility of Bolton’s ideas; it is simply a matter of method.

2. Assuming that a clade can be defined by a trait presumably lost

in some of its members is unacceptable since this would poten-

tially allow creation of unlimited clades (and taxa) for any set of

species with and without the trait in question. To do so, one needs

only to suppose the secondary loss of a critical apomorphy among

the species without it.

Our working hypothesis is that the differences in method

between Bolton and us are worth consideration and should yield

significantly different results.

In spite of its pertinence to the subject, we shall pay less atten-

tion to a recent contribution by DieTz (2004). The reasons for this

are twofold: 1) Dietz’s analysis is largely inspired by and, in its

motivations, differs insignificantly from the ones of BOLTON (1998-

2000), and 2) Dietz’s data differ considerably from those of Bolton

and also from Dietz’s own character descriptions. We tentatively

explain this anomaly by a number of typing mistakes in Dietz’s

published data matrix, mistakes presumably absent from or unlike

those of the matrix that Dietz used for calculations. Our hypothesis

is supported by the fact that searching for the shortest tree(s) for

DieTtz's (2004) published data by means of PAUP 4.0b (see later,

the methods chapter) yields results radically different from those of

Dietz (2004).

These discrepancies between character descriptions and char-

acter coding render difficult any attempt to evaluate Dietz’s argu-

ments and conclusions and, for this reason, in this paper we will

discuss only the most controversial of them.

In this paper, on the contrary, we shall discuss all the charac-

ters used by Bolton on the base of the factual evidence available

tor us:

ANT TRIBE DACETINI 5

2. MATERIAL AND METHODS

We were able to study material deposited in the institutions

listed below.

ANIC. Australian National Insect Collection, C.S.I.R.O., Can-

berra, Australia.

BMNH. The Natural History Museum, London, UK.

FMNH. Field Museum of Natural History, Division of Insects,

Chicago, U.S.A.

GOPC. The amber collection George O. Poinar, Jr. maintained

at the Oregon State University, Corvallis, Oregon.

MCZC. Museum of Comparative Zoology, Harvard University,

Cambridge, Mass.

MHNG. Muséum d’Histoire Naturelle, Genève, Switzerland.

MIZA. Museo del Instituto de Zoologia Agricola “Francisco

Fernandez Yépes”, Maracay, Venezuela.

MSNG. Museo Civico di Storia Naturale “Giacomo Doria”,

Genoa, Italy.

MZSP. Museu de Zoologia, Universidade de Sao Paulo, Brazil.

NHMB. Naturhistorisches Museum, Basel, Switzerland.

PUCE. Museo de Zoologia QCAZ, Escuela de Biologia, Pontifi-

cia Universidad Catélica del Ecuador, Quito, Ecuador.

In addition, we were able to study two Baltic amber specimens

(a worker and a gyne) from the collection of Mr Jorg Wunderlich,

Straubenhardt (Baden-Wirttemberg, Germany) that we identified

as Agroecomyrmex duisburgi (Mayr) or a species very close to it. In

spite of its relevance for the present study, the genus Agroecomyrmex

was not explicitly included in our analysis. Examination of the two

specimens from the collection Wunderlich did not allow assessment

of some characters needing proper mounting or dissection and did

not permit morphological inferences much better than those already

possible from the figures of WHEELER (1915a).

To assess optimal rank of and phylogenetic relationships between

taxa, we performed a cladistic analysis including all genera consid-

ered as valid by BOLTON (1998-2000) and as many as possible of the

characters used by BOLTON (1999) and elsewhere.

6 _ _ C. BARONI URBANI & M.L. DE ANDRADE

2.1. OUTGROUP COMPARISON

BARONI URBANI & DE ANDRADE (1994), for their cladistic analy-

sis, choose as outgroup the Attini and two genera with presumably

superficial similarities with the Dacetini, i.e. Stegomyrmex Emery

and Calyptomyrmex Emery. BOLTON (1999) used only Myrmica

Latreille as outgroup. Bolton’s choice was not explained but it can

be probably justified since it includes an apparently unspecialized

myrmicine. We are still convinced that our earlier choice has better

chances to include a near relative of the Dacetini than the one of

Bolton, but pro bono pacis, for this study we consider most of our

former outgroups plus Myrmica as Bolton did. On the other hand,

the Attini were not considered any more in the present analysis. The

reasons for this are that, for a better comparison with our ingroup

taxa, all Attini genera should have been individually considered for

the analysis and this would have raised the total number of taxa

under study to a figure preventing any form of exact search for the

shortest tree(s).

In addition to our 1994 outgroups, we included in the analy-

sis also a representative of the genus Tatuidris Brown & Kempf, a

purportedly distantly related genus belonging to a different subfam-

ily according to BOLTON (2003) but possessing a mandibular kinetic

similar to the one apomorphic for the Dacetini according to BARONI

URBANI & DE ANDRADE (1994: 10) and BOLTON (1998: 68-70). We

regard this state of affairs as an excellent reason to include Tatuidris

in our analysis. To avoid that our search might force Tatuidris to

appear too close to the dacetine genera (e. g. by choosing Myrmuica

as sole outgroup and Tatuidris + the dacetines as ingroup) or unnat-

urally too far from them (e. g. by artificially transferring Tatuidris

among the outgroups), we included in our study two genera clearly

belonging to two separate but related subfamilies, i.e. Pseudomyrmex

Lund and Myrmecia Fabricius. In our parsimony analyses we con-

sidered Pseudomyrmex and Myrmecia as the sole outgroups. Pseudo-

myrmex is meant to represent the subfamily Pseudomyrmicinae, 1.e.

the sister-subfamily of all other ants considered in this paper and

Myrmecia is the sole genus constituting the subfamily Myrmeciinae,

also close to the Myrmicinae (BARONI URBANI et al. 1992; BARONI

URBANI 2005).

Study of the possible phylogenetic relations of Tatuidris by

means of Pseudomyrmex and Myrmecia as outgroups will not ben-

ANT TRIBE DACETINI 7

efit from consideration of their respective autapomorphies. On the

contrary, we considered for our analysis the three synapomorphies

of the Myrmicinae as given by BOLTON (2003: 52). These should

exclude Tatuidris from the Myrmicinae. But we thought it fair con-

sidering also two potential Myrmicinae synapomorphic characters

described by BARONI URBANI et al. (1992) and shared by all classic

Myrmicinae genera, including Tatuidris.

To assess presence or absence of some internal structures among

the taxa under consideration we dissected the following species:

Bastceros disciger (Mayr).

Calyptomyrmex sp.

Colobostruma sisypha Shattuck.

Eurhopalothrix bruchi (Santschi), platisquama Taylor, procera

(Emery).

Mesostruma turner (Forel).

Myrmecia pyriformis F. Smith.

Myrmica scabrinodis Nylander.

Octostruma sp., O. stenognatha Brown & Kempf.

Orectognathus sp.

Protalaridris sp.

Pseudomyrmex sp. 1 & sp. 2.

Pyramica alberti (Forel), argiola (Emery), decipula (Bolton),

denticulata (Mayr), eggersi (Emery), jacobsoni (Menozzi), kichijo

(Terayama, Lin & Wu), membranifera (Emery), minima (Bolton),

myllorhapha (Brown), nannosobek Bolton, nepalensis (de Andrade),

semicompta (Brown), subedentata (Mayr), tenuissima (Brown), urrho-

bia Bolton, zeteki (Brown).

Rhopalothrix ciliata Mayr.

Stegomyrmex vizottor Diniz.

Strumigenys chapmani Brown, doriae Emery, ekasura Bolton,

elongata Roger, emmae (Emery), exilirhina Bolton, godmani Forel,

grandidieri Forel, harpyia Bolton, koningsbergert Forel, lanuginosa

Wheeler, lyroessa (Roger), micretes Brown, paranetes Brown, rogeri

Emery, saliens Mayr.

Tatuidris tatusia Brown & Kempt.

8 _ €. BARONI URBANI & M.L. DE ANDRADE

If, on one hand, this list may be shorter than what we wished,

on the other the number of structures apparently escaped to the

attention of former students and discovered by us in the species

listed above is surprising. Moreover, the monograph by BoLTON

(2000) alone revealed a precious, nearly exhaustive source of reliable

information contrasting with the information contained in BOLTON’s

(1998 & 1999) previous character descriptions and generic boundaries.

For our phylogenetic analysis the evolution of all characters was

considered as unordered. BOLTON (1999) states that only three of

the characters considered by him were treated as unordered without

specifying how the remaining characters were considered. By ana-

lysing Bolton’s data and considering all characters as unordered, we

obtain the same results as Bolton (I. c.). We explain this coincidence

by supposing that Bolton coded most of his characters as ordered

but, as a matter of fact, since most of his characters are binary, there

is no difference between ordered and unordered binary characters.

Differences arise only when computing with multi-state characters.

In the following we give a list of all the characters that we

retained as potentially phylogenetically significant and of their

coding. Differences with BOLTON (1999), if any, will be explained

and justified in each character description. We felt compelled to

exclude from computation some characters resulting from the lit-

erature. A list of these excluded characters and the justification for

their exclusion will be given after the list of the characters used.

A parsimony analysis of all genera treated as valid in the most

recent literature was performed by PAUP 4.0b10 (SworrorD 2002).

Search for the shortest tree(s) was done by means of the mathemati-

cally exact algorithm “Branch-and-Bound”.

Some graphic display and character tracing was obtained by

MacClade 4.05 (Mappison & Mappison 2002).

To enable a simple judgement on valid or invalid genera, when-

ever possible, we tried to include in our data at least one known and

undoubted autapomorphy for each genus. Genera that, at the end

of our analysis, resulted destitute of any kind of apomorphies (i.e.

genera without valid apomorphies already known from the literature

and without apomorphies representing secondary gains or losses of

other characters appearing as a result of our character optimization)

will be considered as synonyms of their closest related genus.

ANT TRIBE DACETINI 9

While discussing the results of the phylogenetic analyses and

in order to give a possible evaluation term of the characters on

which our classification is based, all critical apomorphies implying

a nomenclatorial decision are reported with their individual Con-

sistency Index and Retention Index as computed by PAUP 4.0b10

and MacClade 4.05 on the Strict Consensus Tree of Figs. 33 and

34. Determining the number and quality of the synapomorphies at

critical branches was performed by means of MacClade.

Measurements and indices used in the text for species descrip-

tions are the following:

P.I

TEL

MT I

Head Length: the maximum measurable distance between

the medial margin of the vertexal angles and the antero-

medial margin of the clypeus with the head in full frontal

view.

Head Width: maximum measurable head width with the

head in full frontal view.

Mandible Length: maximum length of the mandible

between the antero-medial margin of the clypeus (concave

margin excluded when present) and the mandibular apex.

Eye Length: maximum length of the eye.

Scape Length: length of scape shaft, excluding the basal

condyle.

Weber’s Length: diagonal length of mesosoma from the

anterior pronotal border (excluding neck) to the distal

edge of the propodeal lobe.

Total Length: combined head length in full-face view

(closed mandibles included), Weber’s length of mesosoma,

petiole and postpetiole lengths (in profile) and length of

gaster (in profile).

= Cephalic Index: (HW/HL)x100

= Mandibular Index: (ML/HL)x100

= Scape Index: (SL/HL)x100

Mandibular- Torular Index: (Distance between the inter-

sections of the external margins of the closed mandibles

with the anterior clypeal border/Distance between the

outermost points of lower margins of toruli)x100. BOLTON

(1998) proposes an operational “Mandibular-Torular

10 | CC. BARONI URBANI & M.L. DE ANDRADE

Index (MTI)” to quantitatively express the old qualitative

dacetine character “pear-shaped head”. However, accord-

ing to the anthropological and biometric practice as already

codified in myrmecology by RAIGNER & van Boven (1955)

and current use in ant literature including other Bolton’s

printed papers, Bolton’s MTI is a ratio (and also Bolton

calls it so) and not an index. An index should be the ratio

between two measures where the presumably smallest one,

multiplied by 100, is divided by the larger one. The ant

literature already offers several examples of such indices,

like the Cephalic Index (CI), the Scape Index (SI), Peti-

olar Index (PI), etc. Bolton’s quotient, hence, should be

called “Mandibular-Torular Ratio” or, to maintain the

name index and the acronym MTI as we did in the pres-

ent paper for uniformity with the other indices, Bolton’s

M'TI should be multiplied by 100. Furthermore, percent

expressions like CI = 125, SI = 56, for example, offer

small diction and mnemonic facilities as compared with

rough ratios like CI = 1.25 and SI = 0.56.

Defenders of the deregulation may object that, in this same

paper, we also use the name index for ratios like the Consistency

Index and the Retention Index for characters and for phylogenetic

trees. ‘hese terms, however, are already established in the cladistic

literature and the linguistic consistencies or homoplasies of cladism

are far beyond the scope of the present work.

3. CHARACTER CHOICE AND TAXONOMIC RANK

There are no general rules but only intuitive guesses drawn on

taxonomic experience on which characters could be “good” or “bad”

in defining genera or tribes.

Though the temptation was strong, we will not discard any of

Bolton’s generic characters as BOLTON (1987: 285) did for the genus

Diplorhoptrum by means of vague statements like “I am unable to

regard... [this morphology of the male volsellae] as being significant

above the species-group level”, even if we believe that this same

statement would better fit any of Bolton’s Dacetini genus-level char-

acters rather than Diplorhoptrum. As a matter of fact, Diplorhoptrum

ANT TRIBE DACETINI 11

appears as the perfectly sound and well-defined sister-genus genus

of Solenopsis as shown by BARONI URBANI (1995), a citation omit-

ted by BOLTON (2003) in his synopsis of ant classification. Irreduc-

ible sceptics, in addition, may have a glance to the Diplorhoptrum

male volsellae by means of a SEM: they will discover a previously

unknown, new structure in ant morphology. We consider this, still

unpublished, ultrastructural particularity as a redundant, impressive

argument in favour of Diplorhoptrum’s generic validity.

The current situation of the Dacetini classification forces us

to acknowledge the obvious: ‘good’ supraspecific taxa like genera

and tribes, in ants as in other organisms, must be characterized by

at least one unequivocal synapomorphy holding for all the species

involved. Homoplasy occurs among distantly related taxa. When the

same trait appears occasionally in closely related clades, this is more

likely to be due to common ancestry and, until proof of the contrary,

this trait cannot be used to separate monophyletic genera or tribes.

Stated otherwise, all members of a taxon (species, genus, tribe, or

subfamily), and not just the majority of them, should share at least

one, clear synapomorphy characterizing that taxon. This is not the

case of the Dacetini genera as they are currently defined. In this

respect, the present study should be considered as a considerable

but still far from optimal improvement of the existing classification.

The numerical and morphological diversity of the Dacetini is

still much inferior e.g. to the one known for the ant genus Cam-

ponotus (see e. g. EMERY 1925). On the other hand, some of the

repeatedly blamed Camponotus subgenera are better defined and

biogeographically more meaningful than some currently accepted

Dacetini genera.

In our research we entirely concur with BOLTON (1999:1640) in

adopting “an opening hypothesis that every genus-group name listed

be regarded as invalid until proved otherwise by the establishment of

apomorphic characters”. But, contrarily to BOLTON (I. c.), decisions

about individual character apomorphy in this study will be taken

only by means of standard algorithms like PAUP 4.0b10 (SwoFFoRD,

2002) and MacClade 4.05 (Mappison & Mappison, 2002) applied

to observed character distributions and not on personal opinions.

We are aware that a species-level cladistic analysis of the ca. 900

known Dacetini species might suggest additional branches supported

12 | _ C. BARONI URBANI & M.L. DE ANDRADE

by weak synapomorphic characters convergent with other clades or

secondarily disappearing in other, closely derived branches. Such a

species-level analysis is too difficult for a so high number of species

although one might expect that it will weaken or strengthen cur-

rently accepted synapomorphies. In our genus-level approach, the

classificatory value of true or presumed generic apomorphies was

evaluated in terms of Consistency and Retention Indices within the

same genus-level analysis as already stated in the Methods chapter.

We made an effort to use as many as possible of the characters

employed in the papers by BOLTON (1998 & 1999) (see the discus-

sion about the individual characters’ inclusion or exclusion). This

approach poses nonetheless a dilemma about the phylogenetic value

of many of the characters that we used. A practical example should

be sufficient to exemplify our doubts.

BOLTON (1998) used for the first time characters like presence or

absence of a katepisternal groove and deeply (as opposed to super-

ficially) impressed labium and other similar traits to define genera

and tribes for members of his “dacetonine tribe group”. One may be

tempted to discard these characters as phylogenetically irrelevant.

But Bolton’s work should not be detracted so simply. BOLTON

(1.c.) tried to strengthen his work by identifying more than one often

apparently insignificant characters overlapping in distribution.

Discovery of such overlapping character sets might influence

our phylogenetic reconstructions and even apparently insignificant

characters, by reciprocally supporting each other, might result in

plausible supraspecific synapomorphies. Since sharing of the same

trait among closely related branches is better explained by common

descent rather than by homoplasy, these apparently insignificant

traits can be considered as synapomorphic and, hence, can be used to

construct phylogenies, only if A) they are uniform within all the taxa

to be classified and B) they affect all and only the taxa in question.

Unfortunately, one of the major results of our study will be

that neither condition A nor B is ever met for most characters sup-

posed to be relevant for the dacetine phylogeny. Stated otherwise,

all current attempts of phylogenetic reconstructions for the Dacetini

(BARONI URBANI & DE ANDRADE, 1994; BOLTON, 1998; present study)

are weakened at the root by being drawn on characters of doubtful

phylogenetic significance. There is no mention of better characters

ANT TRIBE DACETINI 13

in the ant literature and we were unable to discover superior ones.

All these studies, however, find their justification in proposing a

classificatory model as sound and as credible as possible.

For instance, the most impressive known synapomorphy for

the tribe Phalacromyrmecini so far is the presence of a katepister-

nal groove. Rank attribution to a character (and hence to the taxon

possessing it) is always a subjective evaluation. For the Phalacro-

myrmecini our study will show that the katepisternal groove is not

synapomorphic since it is present in only one species of one genus

among the three genera supposed to constitute the tribe (see later,

the discussion of our character # 36). Nonetheless, assuming that a

scientist will discover a species of a genus whatever with a katepister-

nal groove, or with a deeply (as opposed to superficially) impressed

labium, should this hypothetical scientist place this species in a dif-

ferent genus and tribe? And, assuming that our hypothetical myr-

mecologist would really do so, how many other myrmecologists will

follow him/her accepting the new genus and tribe? Our guess is

none. Or, at least, we so hope.

Our optimistic guess is likely to be contradicted anyway by

numerous students of behaviour, ecology, histology, etc. ‘These

students will even not try to understand the possible phylogenetic

meaning of a katepisternal groove. Of course: they are not taxono-

mists. They will nonetheless use the newly proposed nomenclature

in order to show that they are knowledgeable even with the latest

novelties of taxonomy.

We cannot change this situation but we regret that mentally lazy

taxonomists may also follow the same course of action. During the

tormented history of the Dacetini classification, this already hap-

pened when 18 Dacetini genera like Smithistruma, Epitritus, Kyidris,

Glamyromyrmex, Dorisidris, etc., considered as synonyms by BARONI

URBANI & DE ANDRADE (1994), were erroneously revived by BOLTON

(1995) as everybody admits today.

4, RESULTS

4.1. LIST OF CHARACTERS AND OF THEIR RECORDED STATES

Before discussing the morphological characters used to construct

the phylogeny and classification of the Dacetini, a physiological trait,

14 | _ C. BARONI URBANI & M.L. DE ANDRADE

the mode of action of the mandibles, should be briefly considered.

Although without using it directly in his analysis, BoLTON (1999)

makes an extensive description and treatment of it emphasizing its

phylogenetic relevance.

The two modes of mandibular action observed among the

Dacetini are seemingly mutually exclusive and were improperly

named as “static pressure” as opposed to “kinetic” mandibles. Natu-

rally, every kind of movement, including those necessary to exercise

a pressure whatever, is kinetic and never static by definition.

The “static pressure” and “kinetic” mandibles, however, have

been observed in a too small number of species to be seriously

considered as a taxonomic character, but, in our character analy-

sis, we shall discuss the distribution of their presumed most impor-

tant morphological correlate which is assumed to be taxonomically

and phylogenetically relevant according to BOLTON (1999). This is

the maximum angle between the open mandibles. This biometric

character resulted too variable to be considered among the phylo-

genetically significant ones listed below but it will be nonetheless

discussed in detail for its presumed discriminatory value between

the “genera” Pyramica and Strumigenys (see later the definition of

the genus Strumigenys in Chapter 4.5).

Nonetheless it must be added that, from a purely cladistic point

of view, the whole discussion has little interest since the mandibular

kinetic alone — as its morphological correlate(s) alone - are inad-

equate to the separation of the two “genera” since one of the two

will inevitably result paraphyletic to the other.

The following is an annotated list of the morphological charac-

ters tentatively retained to infer a phylogenetically drawn classifica-

tion of the Dacetini. In the great majority of cases, all characters

referred to the worker caste in the literature and in the following list

may be assumed to repeat the (unknown) gyne condition as well.

1. Worker. Maxillary palps six-jointed (0), five-jointed (1),

four-jointed (2), three-jointed (3), two-jointed (4), one-jointed (5),

absent (6). This is character # 8 of BARONI URBANI & DE ANDRADE

(1994) and character # 1 of BOLTON (1999), transformed respectively

from five and three to seven steps to better consider genera not

considered by BARONI URBANI & DE ANDRADE (1. c.) and by BOLTON

(1. c.). DieTz (2004, character # 7) also increases the number of

ANT'TPRIBE-DACETINI 15

steps of this character as we did, but from three to six steps only.

The reason for this is that no outgroups with six jointed maxillary

palps are considered in his work. Our coding for Octostruma differs

from the count of BOLTON (2003) as a result of the dissection of a

specimen of O. stenognatha (1-jointed instead of 2-jointed maxillary

palps) as already reported in our 1994 paper (p. 19). Ishakidris was

coded as 2-jointed according to BOLTON (1984) and DieTz (2004).

BOLTON’s (2003: 283) record of 3-jointed maxillary palps for this

genus is probably due to a typing or printing mistake. Dietz’s table

2 contains a number of erroneous codings for this character (e. g.

for Acanthognathus, Strumigenys, Basiceros, a. 0.).

2. Worker. Labial palps, four-jointed (0), three-jointed (1),

two-jointed (2), one-jointed (3). This is character # 9 of BARONI

URBANI & DE ANDRADE (1994), char. # 2 of BOLTON (1999) and

char. # 8 of DIETZ (2004) (where labial palps are called “palpos

mandibulares”). The range of this character is amplified here to

include outgroups with four-jointed palps. BOLTON (1999 & 2003)

and Direrz (2004) differ each other in coding this character. In a

few doubtful cases that we were unable to examine we recorded the

counts of both authors.

3. Worker. Labrum, not T-shaped (0), or T-shaped (1). This

is character 3 of BOLTON (1999) but coded polymorphic (instead

of never T-shaped) for Pyramica because of presence of ‘T-shaped

labium in some species like Pyramica subedentata (Mayr) (Fig. 1).

We coded nonetheless the “T-shaped” structure as regularly present

in Strumigenys as Bolton did, although the morphology within the

genus is far from being unequivocal and constant (Fig. 2).

4. Worker. Labrum capable of full reflexion over the buccal

cavity (0), or not (1). This is char. # 4 of BOLTON (1999).

5. Worker. Labral shield not hyperthrophied, not conceal-

ing the entire buccal cavity (0), or enormous, hypertrophied, when

reflexed concealing entirely the buccal cavity (1). This is char. # 5

of BOLTON (1999). We hesitated to add this character to the matrix

since it is logically negatively correlated with character # 4. As a

further proof of this, within Bolton’s “dacetonine tribe-group” it

is distributed among the genera considered in a perfectly comple-

mentary way with character # 4, with the sole exception of Epo-

postruma. By considering character # 5, there is hence the risk of

overweighting the former char. # 4. Our conclusion is confirmed by

16 | C. BARONI URBANI & M.L. DE ANDRADE

00049792 | oo >

porre ___.. = nea . n. MB,

Fig. 1 - “T-shaped labrum” of Pyramica subedentata (Mayr), (top) and Strumigenys

extlirhina Bolton (bottom). ‘The “T-shaped” construction should be syn-

apomorphic for Strumigenys and absent in Pyramica according to BOLTON

(1999, 2000)

ANT TRIBE DACETINI 17

ZMB

Uni Basel

00049784 50 um ua ee

Fig. 2 - Labia of Strumigenys rogeri Emery (top) and S. harpyia Bolton (bottom)

showing intrageneric variation beyond the presumed synapomorphic “T”

shape.

18 | €. BARONI URBANI & M.L. DE ANDRADE

examination of BOLTON’s (1. c.) character matrix where 0’s and 1’s

are antithetically distributed between characters 4 and 5. One might

expect a priori that a hypertrophied labrum (char. # 5) should be

unable of complete reflexion (char. # 4).

0. Worker, Eabium without. (0); or with a deeply: incised,

transverse groove detined by a sharp ridee (1). Presence of: this

ridge is given by BOLTON (1998: 70) as synapomorphic for the

tribe Basicerotini. The ridge, however, is faint in some Basicerotini

species, like Octostruma stenognatha Brown & Kempf (Fig. 3) and

present in a distantly related genus like Stegomyrmex, and in some

species of the non-basicerotine ingroups Pyramica and Strumigenys

(Fig. 4), and Colobostruma (Fig. 5, bottom).

ooo4eeia ees BO an MALE

Fig. 3 - Labrum of Octostruma stenognatha Brown & Kempf with superficial (not

deeply incised) transversal ridge perfectly comparable to the one of some

Strumigenys and Pyramica of Fig. 4.

7. Worker. Labrum without mid-dorsal impression (0), or

with a dorsal impression or pair of impressions (1). This character

ANT TRIBE DACETINI 19

ig.

00046782 Uni Basel

4 - Labrum with deeply incised transversal groove, a basicerotine synapomor-

phy, according to BOLTON (1998) present also among non-basicerotine spe-

cies, like Strumigenys elongata Roger (top) and Pyramica nannosobek Bolton

(bottom).

C. BARONI URBANI & M.L. DE ANDRADE

Uni Basel

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ANT TRIBE DACETINI 24

is given by BOLTON (1998: 72) as uniquely derived for the Dacetini

and secondarily lost in Acanthognathus. 'There are no traces of the

impression also in the dacetine Colobostruma species examined for

the present study, and in some Pyramica species (Fig. 5, top). We

are unable to assess the state of this character in the holotype unique

of Pilotrochus.

8. Worker. Trigger hairs absent (0), or present (1). The term

“trigger hairs”, introduced by Brown & WILSON (1959) appears

to be another “ant term” unknown in entomology textbooks. The

hairs described by Brown & WILSON (l. c.) are obviously hair-

shaped mechanoreceptors and we doubt that the peg-like structures

of probable chemoreceptor function visible e. g in most Basicerotini

and also drawn e. g. by Kempr (1960) for Phalacromyrmex could be

considered as homologous of the former. Presence of “trigger hairs”

on the mouthparts should be a synapomorphy of the “dacetonine

tribe-group” according to BOLTON (1998: 69). Since the myrmeco-

logical term “trigger hairs” implies simply response to a stimulus

exerted on the ant mandibles without specifying its nature, we try

to follow BoLTon (1998) and code uniformly presence of probable

chemoreceptor and mechanoreceptor structures on the mouthparts.

This same character is used also by Dietz (2004, character # 6)

where it is also coded as universally present among all and only the

dacetine genera (s. 1.). In our matrix Pyramica was coded polymor-

phic for this character because of the absence or strong reduction of

these hairs in some species (Fig. 6). This anomaly is partially admit-

ted also by BoLToN (2000: 178). Some Colobostruma species also

exhibit no “trigger hairs” (Fig. 5, bottom). Among our outgroups,

the clypeal hairs of some Myrmecia species (OGATA, 1991) are mor-

phologically indistinguishable from the “trigger hairs” of some

dacetines. We coded nonetheless Myrmecia as “O” for this charac-

ter in the (vague) hypothesis that differences in behaviour between

Myrmecia and dacetines may account also for differences in homol-

ogy. Moreover, presence of trigger hairs is obviously homologous in

function among all ants provided with such hairs but we doubt of

its morphological homology when the hairs are located on different

sclerites as it is the case for several Dacetini. We thought it neces-

sary formulating this precision but, to reduce confrontation with Mr

Bolton on purely academic ground, we coded nonetheless the simple

presence or absence of “trigger hairs” as a unique dacetine tribal

DID C. BARONI URBANI & M.L. DE ANDRADE

group character as Bolton and Dietz did. At least patent absence of

“trigger hairs” of any kind, of course, was recorded as absence.

00046781 ii lm Ud Baeel

Fig. 6 - Pyramica myllorhapha (Brown) without morphologically differentiated

“trigger hairs” on the mouthparts. Presence of “trigger hairs” is consid-

ered as a synapomorphy of his “dacetonine tribe group” by BOLTON (1998

& 1999),

9. Worker. Mandibles at rest crossing (0) or crossing in their

distal part of the masticatory border and opposing in the basal one

(1), or opposing on their whole border (2). State 2 is the synapo-

morphy of the Dacetini (s. 1.) according to BARONI URBANI & DE

ANDRADE (1994) (character # 6) and of the “dacetonine tribe-group”

of BOLTON (1998: 67) and Dierz (2004, character # 1). Acceptance

of the meaning of this character might propose re-inclusion of the

Sicilian Miocene Hypopomyrmex bombiccit EMERY, 1891, among the

Dacetini (see e. g. Emery’s Table 1, Fig. 11). This latter hypothesis

was not tested in the following due to the poor preservation condi-

tions of the sole known Hypopomyrmex specimen. BARONI URBANI &

DE ANDRADE (1994), misinterpreting a figure by KEMPF (1960), con-

ANT TRIBE DACETINI 23

sidered as entirely opposing also the mandibles of Phalacromyrmex.

This is erroneous as shown by a more careful scrutiny of Kempf’s

figure and by the examination of the holotype and paratype of Ph.

fugax. BOLTON (1998) and Dietz (2004) repeated nonetheless the

same error. The Pilotrochus holotype appears to exhibit a condi-

tion very similar to the one of Phalacromyrmex. For this reason,

we thought it better to code both genera as state “1”, intermedi-

ate between the crossing and opposing conditions. Moreover, in

spite of the presumable hierarchic importance of this character, it

is not clear which one should be the formicid plesiomorphic condi-

tion. The mandibles are able of both, crossing and opposing in the

unspecialized ant genus Prionomyrmex and among wasps, the sister

family of ants.

10. Worker. Mandibles engaging through most of their length

(0), or only apically (1). This is char. # 7 of BARONI URBANI &

DE ANDRADE (1994) and character # 7 of BOLTON (1999). Poly-

morphism in Strumigenys added here in consideration of Strumi-

genys guttulata Forel as described by BOLTON (2000: 976, explicit

description, and Fig. 530, illustration) and S. horvatht Emery

(present study). DieTz (2004) does not consider this structure for

his phylogenetic analysis. The reason for it — we believe — lies in

the fact that this character is polymorphic among some genera like

Octostruma (see e. g. DIETZ’s 1. c. Figs. 27 A and C). In our matrix,

Octostruma and other genera where we observed polymorphism are

consequently coded as polymorphic for this character. Rhopalothrix

should be coded as polymorphic for this character after transfer to

it of Eurhopalothrix bruchi (Santschi) as suggested by Dietz (2004:

200). Our examination of the 5 specimens representing the type

series of bruchi does not support Dietz’s conclusion. ‘The mandibu-

lar morphology represents well an average Eurhopalothrix and both

mandibles at rest engage through most of their length. The distri-

bution of this character is perfectly equivalent to Dietz’s character

# 9 (forma das mandibulas: triangulares ou especializadas) since the

shape of the mandibles is just another expression of their capacity

to engage each other. Our coding differs nonetheless from the one

of Dierz (1. c.) for Colobostruma and Mesostruma whose mandibles

engage through most of their length and where we see no traces of

specialization.

24 C. BARONI URBANI & M.L. DE ANDRADE

11. Worker. Mandibles normally toothed (0), or with alter-

nating small and large teeth (1). This, according to BOLTON (1998:

72) and Dietz (2004, character # 10) is a synapomorphic trait of

the genera of the tribe Phalacromyrmecini. Alternating small and

large teeth, however, are present also in Pyramica bunki (Brown)

(BoLTON, 2000, Fig. 113), Pyramica kichyo (Terayama, Lyn &

Wu), Octostruma betschi Perrault (PERRAULT, 1988: Fig. 2), and

Octostruma balzani (Emery) from Ecuador (Fig. 7) equally exhibit

teeth alternating in size. Since the tribe Phalacromyrmecini com-

prises only three monotypic genera, this presumed tribal character

is present in three phalacromyrmecines and at least in four, closely

related, but non-phalacromyrmecine species. BARONI URBANI & DE

ANDRADE (2006a) already called the attention on these species not

fitting Bolton’s classification but BOLTON (2006a) states that the

cases above are not comparable to the phalacromyrmecine mor-

phology because of minute differences and because the Pyramica

species have a basal lamella (see also our discussion of the lamella

under character # 13). One cannot consider the mandibular denti-

tion and the lamella as two independently derived characters (as

Bolton and ourselves did to construct our phylogenies) and use one

of the two characters as an attribute of the other as did BOLTON

(2006a). BOLTON (2006a) blames our use of the dentition without

considering presence or absence of the lamella but our way is the

sole correct way of considering both structures as independently

derived characters and hence of using both characters in phyloge-

netic studies. As far as the morphological differences in dentition

are concerned, we must admit that probably there are no two ant

teeth looking exactly the same, but to see a phylogenetic meaning

in details of the magnitude of those used by Bolton, to use Bolton’s

words, one needs to be cleverer than “other mere mortals”.

12. Worker. 2-3 apical mandibular teeth overlapping (0), or

interlocking (1). This is char. # 8 of BOLTON (1999). In addition

to the distribution of this character as given by Bolton (l. c.), the

apical teeth interlock also in a number of Strumigenys (among

other possible examples, S. percrypta Bolton (BOLTON, 2000, Fig.

375) and S. rogert Emery (BOLTON, 2000, Fig. 368)) and Pyramica,

like P. vartana Bolton (BOLTON, 2000, Fig. 135) species (see also

our Fig. 8 for additional examples encountered during the present

study).

ANT ‘TRIBE: DACETINI DS

mena iii i A

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Fig. 7 - Pyramica kichijo Terayama, Lin & Wu (Dacetini) (top) and Octostruma

balzani (Emery) from Yasuni, Ecuador (Basicerotini) (bottom). Mandibles

with alternating large and small teeth. Mandibular teeth alternating in size

is considered as a phalacromyrmecine synapomorphy by BOLTON (1998).

26 | C. BARONI URBANI & M.L. DE ANDRADE

000498110 | Ae

00049812 * 50 um vi NB.

Fig. 8 - Strumigenys doriae Emery (top) and S. rogeri Emery (bottom). Mandibles

with apical teeth interlocking (not interlocking according to the table of

BoLTON, 1999).

ANT TRIBE DACETINI 2%

13. Worker. Basimandibular process absent (0), or present but

not bifurcated (1), or present, long and apically bifurcated (2). This

character results from merging chars. # 9 of BOLTON (1999) (= char.

# 13 of Dietz, 2004, i.e. presence or absence of the basimandibu-

lar process) and char. # 10 of BOLTON (1999) (i.e. basimandibular

process round and thick vs. a spur, dentiform, or lamellate). We

decided to merge Bolton’s two characters in one because A) the

variation in shape that we observed is difficult to reduce to Bol-

ton’s two binary categories (Fig. 9), and, B) Bolton’s coding of his

character # 10 as autapomorphic for Daceton renders it cladistically

uninformative. Our coding might still need to be further modified

in the future since an apparent basimandibular process is present

also among most Basicerotini, Stegomyrmex, Tatuidris and Calyp-

tomyrmex. This reality is admitted only in part by BOLTON (1998:

72) who states “Basicerotine species with a modified basal tooth are

exceptional and certainly best regarded as independent acquisitions”.

We follow hence BOLTON (l. c.) and code the basimandibular pro-

cess as absent among the Basicerotini in our matrix. Additionally, at

least in some dacetine species currently classified in Pyramica, like

P. argiola (Emery) and P. nannosobek Bolton (Fig. 10) the process

is so reduced to be barely distinguishable or invisible. Note that P.

argiola is one of the species dissected by BOLTON (1998) to define

his Dacetonin but no mention to the absence of basimandinbular

tooth is made in this paper. Analogously BOLTON (2000:12) defines

the Dacetini as unequivocally characterized by the presence of the

lamella. Only in a different context, i.e. in the species-level discus-

sion he admits (page 286) that in the argiola-group the lamella is

“small, dentiform to low triangular and inconspicuous” (description

which we regard as exaggerated if compared with the morphology

visible in Fig. 10), in the murphyi-group is “reduced to a very narrow

stripe not visible in full face view”, and in the mnemosyne-group is

“minute to vestigial, at most a mere ridge on the margin”. BOLTON

(2006b) regards his former (insufficiently) reductive statements as

a clear argument in favour of the validity of the basal lamella as

a tribal character. We would consider it as a proof of the contrary

but followed nonetheless BoLTON (1999, 2003) in coding the pro-

cess as uniformly present in Pyramica and considering these previ-

ously neglected cases as secondary losses of the process as suggested

by the presence of its remnants, though we are not always sure to

28 C. BARONI URBANI & M.L. DE ANDRADE

be able to see such remnants. BOLTON (2006a) states that absence

of the lamella in the closely related “Phalacromyrmecini” is “pre-

sumably plesiomorphic”. We don’t see the difference with the very

similar condition of Pyramica argiola, nannosobek, etc. cited above,

unless one already has in mind his own favourite phylogeny instead

of trying to construct the most probable one. But even BOLTON (1.

c.) apparently notices the contradiction and tries to avoid it by call-

ing the previous exceptions to his general rule “a few documented

cases of obvious secondary reduction or modification of function”.

We never saw these documents. In favour of our point of view we

must notice also that BOLTON (1988:71), while reviving his Dacetini

s. str. (under the name Dacetonini), emphasizes the importance of

the basimandibular process without mention of ”cases of obvious

secondary reduction”. On the internal margin of the mandibles of

Phalacromyrmex fugax there is a small, denticulate swelling. We find

it difficult to consider this structure as a typical example of absence

of the lamella in Phalacromyrmex and the morphology e.g. of Pyra-

mica argiola (Fig. 10) as a documented demonstration of secondarily

‘lost presence’ of it. The bifurcated condition of the lamella is a

classic, undoubted autapomorphy for Acanthognathus.

14. Worker. Basimandibular seta absent (0), or present (1). This

is a potential synapomorphy for Basicerotini and Phalacromyrmecini

in BoLTON (1998: 73) and is coded as such by Dietz (2004, char-

acter # 4). Actually a basimandibular seta is present also in Stego-

myrmex and in some Strumigenys and Pyramica species (Figs. 11,

12 & 13), though in the latter two genera the position of the seta is

slightly more distal than in Stegomyrmex and Basicerotini.

15. Worker. Number of antennal joints: 11-12 (0), or less than

11 (1). This is char. # 4 of BaronI URBANI & DE ANDRADE (1994),

character # 11 of BOLTON (1999) and char. # 20 of Dietz (2004),

coded in binary form as already done by BoLTon (1999) and DIETz

(2004). There is increasing consensus on the poor phylogenetic value

of the dacetine antennal count variation. Already BOLTON (1983)

concluded “the reduction in antennomere count has little or no

value at genus level...”. Experimental attempts of cladistic analyses

in which the actual antennal counts were given for each genus and

considered as unordered, ordered, “Dollo”, or irreversible, regularly

yielded very implausible phylogenetic reconstructions. We agree

with BOLTON (l.c.) that polarization of two states at the extremes of

ANT TRIBE DACETINI 29

00049803 = y.a@ hea

00046812 00049808 MB

Uni Basel

Fig. 9 - Variability of the basimandibular process among some dacetine ants. Dace-

ton armigeron (Perty) (top left), Strumigenys micretes Brown (top right),

Pyramica zeteki (Brown) (bottom left), Strumigenys lyroessa (Roger)

(bottom right).

30 C. BARONI URBANI & M.L. DE ANDRADE

00049806 ZMB

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ZMB

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Fig. 10 - Pyramica argiola (Emery) (top) and P. nannosobek Bolton (bottom) with

basimandibular process barely distinguishable or absent. P. argiola has no

visible process but it shows two internal longitudinal ridges, which could

be both interpreted as a remnant of the basimandibular process. Presence

of the process or of its or remnants should be synapomorphic for the tribe

Dacetini according to BOLTON (1999, 2003).

ANT TRIBE DACETINI i 31

ZMB

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00049753

Fig. 11 - Strumigenys paranetes Brown (top) (Dacetini) and Octostruma nr. batesi

(Emery) (Basicerotini) (bottom) showing the basimandibular seta (pointed

by the arrow). Presence of the seta should be synapomorphic for the tribe

Basicerotini according to BOLTON (1998).

C. BARONI URBANI & M.L. DE ANDRADE

00050613

00049759 | - 50um DEE

Fig. 12 - Strumigenys gloriosa Bolton. (Dacetini) (top) and £Eurhopalothrix

platisquama Taylor (Basicerotini) (bottom) showing the basimandibular

seta (pointed by the arrow). Presence of the seta should be synapomor-

phic for the tribe Basicerotini according to BOLTON (1998).

ANT TRIBE DACETINI 33

00050877 ZMB

Uni Basel

00050619 n AO. UN gEMB

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13 - Pyramica denticulata (Mayr) (Dacetini) (top) and Stegomyrmex vizot-

toi Diniz (Stegomyrmecini) (bottom) showing the basimandibular seta

(pointed by the arrow). Presence of the seta should be synapomorphic for

the tribe Basicerotini according to BoLTON (1998).

34 C. BARONI URBANI & M.L. DE ANDRADE

the sole known gap (10 antennomeres) among the ants considered

in the present study appears the most reasonable way of considering

this character.

16. Worker. Two-segmented antennal club absent or indistinct

(0), or well developed (1). This is char. # 12 of BOLTON (1999) and

a simplification of BARONI URBANI & DE ANDRADE’s (1994) character

H# 5.

17. Worker. Scape straight at base (0), or gently downcurved

at base (1). This is char. # 14 of BOLTON (1999).

18. Worker. Base of scape straight or at least complanar with

basal condyle (0), or scape bent at right angle near the base (1).

The curved or angular scape is a synapomorphy of the Basicero-

tini according to BOLTON (1998: 71). Separating this character from

the previous one might need some dialectic exercise. We kept the

two separate because of the presumed important phylogenetic mean-

ing of the latter if the independent evolution of the two traits will

be confirmed. Examples of non-basicerotine species exhibiting the

basicerotine condition are Pyramica decipula (Bolton), Pyramica

nannosobek Bolton (Figs. 14 & 15) and Colobostruma froggatti (Forel)

(partially visible in Fig. 38). If we understand properly DIETZ (2004)

this character should be equivalent or very similar to his character

# 22 but we are unable to verify his recording for different taxa. In

spite of differences in definition and coding, we did not consider

DIETZ (° e.) as referring to another, different character since we

already fear that consideration of our characters 17, 18 and 19 could

overweight the shape of the scape within our data.

19. Worker. Scape not clavate (0), or clavate (1). Presence of a

clavate scape is given by BOLTON (1998: 72) and DIETz (2004, chars.

# 21 and # 23) as synapomorphic for the tribe Phalacromyrmecini.

We are unable to see differences between the descriptions of DIETZ’s

char. # 21 “Clava da antena: ausente (0); presente (1). A clava é um

alargamento do escapo...estrutura distinta e caracteristica...da tribo

Phalacromyrmecini” and char. # 23 “Escapo clavado: nao clavado

(0); clavado (1)”. Differences, however, are impressive within their

respective coding by DIETz (l. c.). According to our observations, a

clavate scape, in addition than in the Phalacromyrmecini, is pres-

ent also in Stegomyrmex, Tatuidris, in Pyramica warditera Bolton

(BOLTON, 2000, Fig. 165) and P. reticeps (Kempf) (BOLTON, 2000,

ANT TRIBE DACETINI 35

sedie ee RALE

00047465 _ EDU RALE

Fig. 14 - Inadequacy of the scape shape as a tribal character for the Basicerotini.

Example 1. Morphological similarity between Pyramica decipula (Mayr)

(Dacetini) (top) and Rhopalothrix ciliata Mayr (Basicerotini) (bottom).

C. BARONI URBANI & M.L. DE ANDRADE

(0)

(©) )

ZMB

Uni Basel

00047467

bal character for the Basicero-

between Pyramica nannosobek

a di

es}

so os

Gi of

O. ©

Ss a

Oo =

a @

o =

+ di

o 2

Soa

)

icerotin1

(Santschi) (Basi

bruchi

) (top) and Eurhopalothrix

(Dacet

Bolton

(bottom)

ANT TRIBE DACETINI 37

Fig. 162). We are unable to explain DIETZ's explicit reference to the

Phalacromyrmecini for his character # 21 (p. 32) and his record-

ing“of this characte umn dais stables, astrale tne clavate scape

is given as unknown in Phalacromyrmex and present in a number

of non-phalacromyrmecine genera like the Attini, Platythyrea,

Stegomyrmex, Blepharidatta, Microdaceton, Daceton, Orectognathus,

Colobostruma, Mesostruma, Epopostruma, Pyramica, Strumigenys,

Basiceros, Eurhopalothrix, Octostruma, Protalaridris, Rhopalothrix,

Talaridris. See also our worries about redundancy of the scape char-

acters expressed under our description of character 18.

20. Worker. Torulus simple or with a small lobe at most (0),

or with hypertrophied dorsal lobe and strongly curved downwards

(1). This is given by BOLTON (1998: 70) and by Dietz (2004, char. #

15) as a synapomorphy of the Basicerotini. A hypertrophied torulus,

however, is present also in Tatuidris, Mesostruma eccentrica Taylor,

M. bella Shattuck, Colobostruma alinodis (Forel), and in some Stru-

migenys and Pyramica species (Figs. 16 & 17).

21. Worker. Second funicular joint normal (0), or hypertro-

phic (1). This is a clear autapomorphy for Orectognathus.

22. Worker. Antennal fossa and scrobe (when present) conflu-

ent (0), or separated from each other by at least a cuticular rim or

crest (1). This is another basicerotine synapomorphy according to

BoLTON (1998: 71) and Dierz (2004, char. # 16). The crest is indis-

tinct in some Eurhopalothrix species like E. bruchi (Santschi) (Fig.

16, top) and FE. heliscata Wilson & Brown (Fig. 18, top). On the

contrary, a distinct crest is visible in some Pyramica and Strumi-

genys species (Fig. 18, bottom). A number of Dacetini species have

been already illustrated by Bolton with photographs showing the

presumed basicerotine structure in these non-basicerotine genera (see

e. g. BOLTON, 2000: Figs. 352, 483, 516 for Strumigenys; BOLTON,

1999: Fig. 60 and BoLTON, 2000: Figs. 236, 241 for Pyramica; and

BOLTON 2000: Fig. 46 for Colobostruma sisypha Shattuck).

23. Worker. Antennal scrobe absent (0), present and above the

eyes (1), present and below the eyes (2), or with the eyes at its

posterior border (3). This is character # 2 of BARONI URBANI & DE

ANDRADE (1994) where it was coded as two-states only, character #

16 of BOLTON (1999) and character # 18 of DIieTz (2004) with addi-

tion of state 3 (present study). Character state 3 was added here in

38 C. BARONI URBANI & M.L. DE ANDRADE

specs ttt. ama pre RATA

Fig. 16 - Inadequacy of the torulus shape as a tribal character for the Basicero-

tini. Example 1. Morphological similarity between Eurhopalothrix bruchi

(Santschi) (Basicerotini) (top) and Strumigenys godmani Forel (Dacetini)

(bottom).

ANT. TRIBE DACETINI 39

ns esi _ . = sro _ é: MB

Fig. 17 - Inadequacy of the torulus shape as a tribal character for the Basicero-

tini. Example 2. Morphological similarity between Octostruma betschi

Perrault (Basicerotini) (top) and Strumigenys micretes Brown (Dacetin1)

(bottom).

40) C. BARONI URBANI & M.L. DE ANDRADE

00047418

00049772 ee 100. ZMB

Uni Basel

Fig. 18 - Variability of the antennal fossa as a tribal character for the Basicero-

tini. The basicerotine Eurhopalothrix heliscata Wilson & Brown with fossa

weakly separated from the scrobe (top) (the fossa should be separated

by a crest in all basicerotines) and Strumigenys godmani Forel (Dacetini)

with crest separating the fossa from the scrobe (bottom) (fossa and scrobe

should be confluent, without separating crest among Dacetini).

ANT TRIBE DACETINI 41

order to properly code Tatuidris not considered by BOLTON (l. c.)

and DieTtz (2004). State “2” appears only in the character descrip-

tion and not in the relative table 2 of DieTz (2004). Polymorphism

in Colobostruma added to the table because of the (possibly second-

ary) reduction or absence of the scrobe in some Colobostruma species

with flat head, as noted by Shattuck in BOLTON (2000:31).

24. Worker. Eyes absent (0), dorsolateral (1), lateral (2), or

ventral (3). This is character # 3 of BARONI URBANI & DE ANDRADE

(1994), unchanged, and character # 17 of BOLTON (1999) with the

addition of state 0 in order to account for some blind Rhopalothrix

species (BROWN & KEMPF, 1960: 231) and for the first record of a

blind “Pyramica” (see later the discussion under Strumigenys inopi-

nata (de Andrade)). Considering Rhopalothrix inopinata de Andrade

in Pyramica — as suggested by DieTz (2004) and provisionally

accepted for the present study — or leaving it in Rhopalothrix as it

was originally described, does not affect either the number or the

topology of the possible shortest trees. Only the tree length is one-

step shorter when znopinata is left in Rhopalothrix.

25. Worker. Occipital foramen not in a depression (0), or in a

deep depression surrounded by a continuous cuticular margination

(1). This should be another basicerotine synapomorphy accord-

ing to BOLTON (1998: 71) and Dietz (2004, char. # 19). There is

no completely marginate depression at least in Octostruma balzani

Emery and Rhopalothrix ciliata Mayr. On the contrary, some Stru-

migenys and Pyramica species have the occipital foramen in a much

more marginate depression than the one of the previous basicerotine

examples (see Figs. 19 & 20). In Pyramica minkara (Bolton), not

photographed for the present study, the phenomenon is even more

evident.

26. Worker. Head of normal shape (0), or pyriform (1). This is

the dacetine “typical” character of the classic literature (e. g. EMERY,

1924), equivalent to char. # 1 of BARONI URBANI & DE ANDRADE

(1994). It is also equivalent to the “anterior head capsule is nar-

rowed from side to side” of BOLTON (1998: 68) where it is given

as synapomorphy of the “dacetonine tribe-group”. BOLTON (1998),

carefully reflected in DieTz (2004), defines much better this char-

acter and suggests a “Mandibular-Torular Index (MTI)” to quanti-

tatively express it. As we already explained in the Methods chapter

(q. v.), as it was originally formulated, this is a Ratio and not an

42 C. BARONI URBANI & M.L. DE ANDRADE

00046780 Res

SANTO llr DS

Uni Basel

19 - Inadequacy of the occipital foramen as a tribal character for the Basicero-

tini. Pyramica tenuissima (Brown) (top) and Rhopalothrix ciliata Mayr

(bottom). Note the margination of the foramen absent or weak in Rho-

palothrix and the deep margination in Pyramica.

ANT TRIBE DACETINI 43

00046778 Ud Bagel

00046779 Ud HE,

Fig. 20 - Inadequacy of the occipital foramen as a tribal character for the Basicero-

tini. Strumigenys ekasura Bolton (top) and Octostruma balzani (Emery)

(bottom). Note the margination of the foramen absent or weak in Octo-

struma and the deep margination in Strumigenys.

44 C. BARONI URBANI & M.L. DE ANDRADE

Index. Bolton’s quotient, hence, should be called “Mandibular-Tor-

ular Ratio”. In our case, the Dacetini should have a ratio 0.5-1.3,

and most remaining Myrmicinae a ratio between 1.5-3.0. Maintain-

ing the MTI as an index means its reformulation as MTI = (dis-

tance between the points where the outer margin of the fully closed

mandibles intersect the anterior clypeal margin) X 100 / (maximum

distance between the lower margins of the toruli). In our matrix,

hence, MTI > 150 (0), and MTT < 130 (1). In Tatuidris MTI = 90.

27. Worker. Occipital foramen posterior (0), or dorsal (1). The

dorsal position is a classical, excellent autapomorphy for Daceton.

28. Worker. Pronotal cervix without (0) thick, transverse rim,

or with a thick, transverse rim (1). This should be a synapomorphic

trait for Microdaceton according to BOLTON (1999).

29. Worker. Promesonotal suture mobile (0), or fused (1). The

dorsal position is character # 10 of BARONI URBANI et al. (1992).

30. Worker. Cuticular processes of promesonotum absent (0),

or present (1). This is character # 24 of BOLTON (1999) but coded

differently from BOLTON (Il. c.) in a number of genera for the fol-

lowing reasons: Epopostruma, polymorphic (instead of present only)

because of absence of process in E. curiosa Shattuck (BOLTON, 2000,

Fig. 88); Mesostruma, polymorphic (instead of present only) because

of absence of the process in M. brown Taylor (BOLTON, 2000, Fig.

57). Strumigenys, polymorphic (instead of absent only) because of

presence of the process in S. loriae Emery (BOLTON, 2000, Fig.

485).

31. Worker. Mid and hind tibial spurs present (0), or vestigial

to absent (1). Absence of tibial spurs is regarded as potentially syn-

apomorphic for the “dacetonine-group” by BOLTON (1998: 70). As

Bolton correctly writes, the commonness of the apomorphic condi-

tion among Myrmicinae renders it difficult attributing phylogenetic

value to this character. Tibial spurs are missing in Stegomyrmex and

in Calyptomyrmex, but present in Tatuidris.

32. Worker. Mesosternal hair beds invisible in profile (0), vis-

ible in profile (1), or hypertrophic in profile (2). Brown (1978) first

described this structure in Pilotrochus adding: “this organ appears to

be the external part of an exocrine gland or glands”. BARONI URBANI

& DE ANDRADE (1994: 61 & Fig. 36) recorded a similar structure in

Strumigenys assamensis and called it “mesopleural presumed glandu-

ANT TRIBE DACETINI 45

lar area”. BOLTON (1998) seized this belief under the name “meso-

pleural gland” and later (BOLTON, 1999) called it “hair-lined gland

of mesopleural anterior margin”. DIETZ (2004) also calls it “glandula

da mesopleura” referring to BoLTON (1998). As a matter of fact there

is no visible gland in this body region. There are broad hair beds

probably made out of sensilla trichoidea on the mesosternum. When

these hairs are particularly developed and/or abundant, they became

visible also on the profile (Fig. 21) and have been misinterpreted as

a glandular opening. This trait is given as a potential synapomor-

phy of Dacetonini + Phalacromyrmecini by BOLTON (1998: 73) and

is practically equivalent to character # 23 in BOLTON (1999) where

it appears as synapomorphic for Pyramica and Strumigenys only

(sic!). Dietz (2004, char. # 25) also uses this character and refers

to BOLTON (1998) but presents it as exclusive of the Phalacromyr-

mecini. The hairs, however, are well visible in profile also in some

Colobostruma, Mesostruma, Epopostruma, Orectognathus, Basiceros,

Eurhopalothrix, Rhopalothrix, and in Octostruma balzani (Emery).

33. Worker. Metapleural gland orifice absent, round or not

covered by a longitudinal integumental ridge never opening dorsally

to posterodorsally (0), or with a longitudinal slit or narrow crescent

opening dorsally to posterodorsally (1). This is given as myrmicine

synapomorphy by BOLTON (2003: 52) and should exclude Tatuidris

from the subfamily Myrmicinae.

34. Worker. Metapleural gland opening visible, 1. e. not covered

by the slit of character # 33 (Q), or invisible (1). This is character #

21 of BOLTON (1999).

35. Worker. Metapleural gland bulla [widely, according to

Bolton] separated from annulus of propodeal spiracle more than the

diameter of the spiracle [Dietz] (0), or [very close to the annulus

of the propodeal spiracle (Bolton)], at most equal to the diameter

of the spiracle and often touching the border of the bulla [Dietz]

(1). This is character # 22 of BOLTON (1999) and character # 26 of

Dietz (2004). Coding of this character, however, differs diametri-

cally between Bolton and Dietz. There are obvious difficulties in

standardizing some of Bolton’s words like “widely” or “very”. For

this reason, we followed Die Tz’s (2004) definition and we further

precise it by stating that we consider as spiracle only the true tra-

cheal opening without the surrounding bulging area. Having said

C. BARONI URBANI & M.L. DE ANDRADE

Fig «Qik =

ZMB

Uni Basel

pOpAgeIO 0 Am uit.

Octostruma balzani (Emery) (Basicerotini). Sensilla trichoidea grouped to

form mesosternal hair beds (top) in ventral view. The hair beds under

normal conditions and in ventral view are concealed by the procoxae; they

turn out to be visible here after dissection. When the sensilla are particu-

larly long or abundant they become visible also on the profile (bottom).

This structure, widespread among several genera included in this study,

was interpreted as an exocrine gland in the previous literature, and was

given as synapomorphic for the tribes Dacetini + Phalacromyrmecini

by BoLTon (1998) and as synapomorphic for the genera Strumigenys

and Pyramica alone by BOLTON (1999). According to DIeTz (2004), the

“gland” is autapomorphic for the Phalacromyrmecini.

ANT TRIBE DACETINI 47

that, our coding of this character is drawn from our own study of

the ants and differs from both, slightly from the one of Bolton in

the polymorphism of Colobostruma only, and radically from the one

of Dietz. We can easily and entirely defend our coding on the base

of the species that we studied.

36. Worker. Katepisternal oblique groove absent (0), or present

(1). Presence of the groove should be a synapomorphy of the tribe

Phalacromyrmecini according to BOLTON (1998: 72). Examination of

the holotype of Phalacromyrmex fugax Kempf shows that there is

no trace of groove in the type genus of the tribe, Phalacromyrmex

(Fig. 22). Our “discovery” is confirmed by the original description

by KEMPF (1960a) and by the redescription by BOLTON (1984). Both

authors, for Phalacromyrmex, speak only of oblique costulation, a

trait used also by BOLTON (1984: 378) to differentiate Phalacromyr-

mex (where the groove is absent) from Ishakidris (groove present).

A few pages later, BOLTON (1984: 381) adds that the “mesopleu-

ral organ... in Pilotrochus... is... apparently not subtended by the

open groove seen in Ishakidris”. ‘Vhe katepisternal groove, hence, is

coded as autapomorphic for I[shakidris in this paper. Its synapomor-

phic value for the Phalacromyrmecini pretended by BOLTON (1998)

was already discredited in an earlier paper by BOLTON (1984). But

BOLTON (2006a) defends again the tribal status of Phalacromyrmecini

on the generic presence of “some katepisternal system that appears

to channel the products of the mesopleural gland posteroventrally”.

Unfortunately the gland in question has been not yet discovered and

the hair-like structures suggesting its existence are sensilla trichoidea

widespread among the Dacetini (see our discussion under character

# 32). The holotype of Pilotrochus besmerus Brown equally shows no

traces of the katepisternal groove or, if such a “katepisternal system”

is present, there are three equivalent systems pointing respectively

frontward, upwards and backwards. This situation is perfectly visi-

ble also in BROWN’s (1978) Fig. 2 cited by BOLTON (2006a) to defend

his recording the presence of the groove. No katepisternal system of

any kind is visible in Phalacromyrmex (Fig. 22).

37. Worker. Lower mesopleura without (0), or with marked

longitudinal costulation (1). This seems to be the most impres-

sive autapomorphy for the genus Phalacromyrmex within the broad

sample of genera considered here.

48 C. BARONI URBANI & M.L. DE ANDRADE

Fig. 22 - Phalacromyrmex fugax Kempf, holotype worker, type species of the genus

Phalacromyrmex which is, in turn, the type genus of the “tribe Phalacro-

myrmecini”. Mesosoma in profile without traces of katepisternal oblique

groove; presence of the groove appears to be the most salient character

of the “tribe Phalacromyrmecini” according to BOLTON (1988). Distance

between two scale bars 0.1 mm.

38. Worker. Propodeal spiracle in profile at about midlength

of sclerite (0), or close to or at declivity (1). This is character # 25

of BOLTON (1999).

39. Worker. Petiole dorsoventrally unfused (0), or fused (1).

This is character # 16 of BARONI URBANI et al. (1992).

40. Worker. Petiole in posterior view with tergum and ster-

num differently shaped (0), or with tergum and sternum equally

convex, forming a circle (1). This is given as myrmicine syna-

pomorphy by BoLTon (2003: 52) and should exclude 7atuidris

from the subfamily Myrmicinae. We coded the character as pres-

ent in all Myrmecinae and absent in Tatuidris. Tergum and ster-

num are actually round also in Myrmecia and Pseudomyrmex.

These structures, however, differ from the one of Tatuidris and

from the Myrmicinae for being unfused instead of fused. Dis-

regarding the fused or unfused state, as BOLTON (2003) did,

this character results parsimony uninformative in our context.

ANT TRIBE DACETINI 49

41. Worker. Lateral outgrowths of pedicel absent (0), or

present (1). This is character # 28 of BOLTON (1999) coded dif-

ferently from BOLTON (l. c.) in some genera for the following rea-

sons: Mesostruma (polymorphic instead of present only) because of

absence in M. eccentrica Taylor (BOLTON, 2000, Fig. 56 and present

study) Epopostruma (polymorphic instead of present only) because

of absence of outgrowths in FE. quadrispinosa (Forel) (BOLTON, 2000,

Fig. 76 and present study). BOLTON’s (1. c.) matrix records the reg-

ular presence of outgrowths for Pyramica and Strumigenys. This

corresponds approximately to the truth only if one considers all out-

growths together, 1. e. integumental spines and spongiform append-

ages. We accepted this interpretation in our matrix but excluded

Bolton’s next character (character # 29, petiole and postpetiole

with or without spongiform tissue) as redundant. In addition, this

character should be coded differently from BOLTON (l. c.) in both,

Pyramica and Strumigenys. In Pyramica presence of the spongiform

appendages is polymorphic (instead of present only) because of

absence of spongiform appendages at least in Pyramica denticulata

(Mayr) (Fig. 23, top ) and in P. eggerst (Emery) (BOLTON, 2000:

184). Analogously, for Strumigenys, BOLTON (2000: 903) states that

the whole species group szalayi Emery has “spongiform append-

ages of waist very reduced or absent”. S. tigris Brown also has no

traces of spongiform appendages (Fig. 23, bottom). Bolton’s charac-

ter # 29 (spongiform appendages) alone, after the above corrections

concerning Pyramica and Strumigenys, results cladistically uninfor-

mative. Additionally, spongiform appendages are present also in

Colobostruma cerornata Brown (Brown, 1959: 2 [description] and

Fig. 2; BoLTON, 2000: 39 [description] and Fig. 45); nonetheless

Colobostruma is coded as without spongiform appendages in BOLTON

(1999: 1648) and in the present paper. BOLTON’s (1999) character #

29 is also equivalent to character # 27 of Dietz (2004). BoLTON’s

(1999) description of his character # 28 as “lateral or ventral out-

grows of any form” would practically change only the coding for a

number of basicerotine genera as polymorphic instead of absent only,

and further blunt the separation between Dacetini and Basicerotini,

a universally undesired feature, we suppose.

42. Worker. Postpetiolar tergum and sternum overlapping at

junction (0), or meeting end to end (1), This is given as myrmicine

synapomorphy by BOLTON (2003: 52) and should exclude Tatuizdris

from the subfamily Myrmicinae.

50 C. BARONI URBANI & M.L. DE ANDRADE

00050616 | Uni Basel

Fig. 23 - Absence of spongiform appendages in Pyramica denticulata (Mayr) (top)

and Strumigenys tigris Brown (bottom). Presence of spongiform append-

ages is given as synapomorphic for these two genera in BOLTON (1999).

ANT TRIBE DACETINI 51

43. Worker. Articulation between gaster and postpetiole

narrow (0), or broad (1). A narrow articulation is given as poten-

tially synapomorphic for the Dacetini + Phalacromyrmecini by

BoLTON (1998: 72). The narrow condition, however, is widespread

in a number of Myrmicinae including most outgroups considered in

the present paper. This character, for the ingroup species, is coded

polymorphic according to the variation observed while examining

the material available for the present study (e. g. Strumigenys hor-

wathi Emery and Pyramica crassicornis (Mayr) with broad articu-

lation and Basiceros disciger (Mayr) with narrow articulation (Fig.

24)). Inclusion or exclusion of this character from calculations or its

coding with or without polymorphism affect the length but not the

number and the topology of the shortest trees. This is also character

# 28 of Dinrz (2004).

44. Worker. Postpetiolar presclerites not set in a concavity

or depression (0), or arising from the base of a broad, deeply con-

cave depression (1). This is a synapomorphy for the Basicerotini

according to BOLTON (1998: 71) and Dietz (2004, char. # 24). The

postpetiolar presclerites of some Pyramica species like P. denticu-

lata (Mayr) (Fig. 25) or P. eggerst (Emery) (Fig. 26) arise from a

deep depression of size and shape perfectly comparable to the one

of some Basicerotini.

45. Worker. Pretergite of first gastral segment neck-like (0), or

subsessile to sessile (1). State 0 of this character is given as poten-

tially synapomorphic for the Dacetini and Phalacromyrmecini by

BoLTON (1998: 72). We added it to our data matrix notwithstand-

ing some perplexities about its meaning (e.g. the neck-like condition

seems to be plesiomorphic among myrmicines and, neck-like — a

morphological trait — is not perfectly antonymic of sessile) and pres-

ence in some taxa. Within the sample of genera considered in the

present paper, the sessile condition was coded as synapomorphic for

the Basicerotini in spite of numerous cases in which species belong-

ing to the two tribes appear to be indistinguishable (see e.g. Fig.

27, among other possible examples). An anonymous referee insisted

that the sessile condition applies also to Tatuidris, a condition that

we were unable to verify by dissection of the specimens available for

the present study. Inclusion or exclusion of this character for parsi-

mony analyses of our data does not affect the number and topology

of the shortest tree(s).

C. BARONI URBANI & M.L. DE ANDRADE

Fig. 24 -

00050582 00050635 00049797 ui BE,

Comparably broad articulations between postpetiole and gaster in the

dacetines Strumigenys horwathi Emery (A), Pyramica crassicornis (Mayr)

(B) and in the basicerotine Basiceros disciger (Mayr) (C). A broad articula-

tion is supposed to be synapomorphic for the basicerotines.

ANT TRIBE DACETINI 59

00047440 Uni Basel

00047442 Uni Basel

Fig. 25 - Similarity of structure of the postpetiolar presclerites among Dacetini and

Basicerotini: Pyramica denticulata (Mayr) (Dacetini, top) and Rhopalothrix

ciliata Mayr (Basicerotini, bottom).

54 C. BARONI URBANI & M.L. DE ANDRADE

ZMB

Uni Basel

00047443 Uni Basel

Fig. 26 - Similarity of structure of the postpetiolar presclerites among Dacetini and

Basicerotini: Pyramica eggersi (Emery) (Dacetini, top) and Octostruma bal-

zant (Emery) (Basicerotini, bottom).

ANT TRIBE DACETINI 55

46. Worker. “Limbus” (i. e. anterior transverse cuticular ridge

of the first gastral tergum) absent (0), or present (1). This is char-

acter # 32 of BoLTON (1999). In Bolton’s matrix the character is

synapomorphic for Pyramica and Strumigenys. On the other hand

BOLTON (1998: 71) states that the first gastral tergite and sternite of

the Basicerotini are synapomorphically marginate basally, immedi-

ately behind the postpetiole and adds that “the limbus... is an apo-

morphy... of the strumigenyite group of Dacetini...and... it 1s not a

homologue of the basal margination developed in Basicerotini”. Not

only we are sceptic about this a priori declaration of homoplasy, but

we are also unable to see differences between the tergal morphology

of some basicerotines and the one of some dacetines (Fig. 27). With

the best of our will we are unable to see BOLTON’s (1998) differentia-

tion between the presence of this structure “basally” (= Basicerotini

only), or “prebasally” (= Dacetini only and not homologous of the

former). Our Fig. 27 should be a good support for our interpreta-

tion among many other possible ones. In addition, a cuticular ridge

is present also on the first gastral tergum of Colobostruma cerornata

Brown and Phalacromyrmex fugax Kempf (present study). DIETZ

(2004, character # 30) apparently also disagrees with BOLTON (1998)

and considers this character as a synapomorphy of the basicerotines

+ phalacromyrmicines in his character description but, in his table 2,

he codes nonetheless the ridge as present in the basicerotine genera

only. In our matrix we coded simply the presence or absence of the

sole cuticular ridge that we were able to see.

47. Worker. Suture between first gastral tergite and sternite

anteriorly rounded (0), or angulated (1). This is character # 33 of

Bowron, (1999),

48. Worker. Base of the first gastral sternum in profile rounded

(0), or truncated (1). This is character # 34 of BOLTON (1989) where

the truncated condition results as synapomorphic for Strumigenys

and Pyramica. The truncated condition, however, is present also in

all Basicerotini, Colobostruma cerornata Brown, Ishakidris and Stego-

myrmex. There are, on the other hand, species of both Pyramica (e.

g. P. mutica (Brown)) and Strumigenys (e. g. S. nigra Brown, S.

tigris Brown (BOLTON, 2000: Fig. 514)) without truncation.

49. Worker. First gastral tergum and sternum smooth or at least

not sharply punctuated (0), or with dense and deep punctures (1). A

56 C. BARONI URBANI & M.L. DE ANDRADE

Uni Basel

00047458 ZMB

00046792 weer DO UM ud Bal

Fig. 27 - Similarity in morphology of the “limbus”, (an anterior cuticular ridge on

the first gastral tergum) in the dacetine Pyramica denticulata (Mayr) (top)

and the basicerotine Octostruma stenognatha Brown & Kempf (bottom).

This structure is interpreted as synapomorphic for the Basicerotini and,

independently derived from this, synapomorphic also for Strumigenys +

Pyramica in BoLTON (1998). The difference between the two synapomor-

phies is supposed to lie on the fact that no Dacetini have the margination

extending to the sternum. In this figure the dacetine Pyramica exhibits

a much stronger sternal margination than the basicerotine Octostruma.

Another putative difference between Dacetini and Basicerotini (char.#

45) is a neck-like anterior articulation of the gaster believed to be present

among Basicerotini only. Note the practically identical morphology of the

two species illustrated here.

ANT TRIBE DACETINI DS

punctuated first gastral segment is given as synapomorphic for the

Basicerotini by BOLTON (1999: 71). In the same paper BOLTON (Il.

c.) admits that in some species this sculpture may be “secondarily

reduced or effaced”. As stated at the beginning of this paper, we

coded what we observed without a priori decisions on its phyloge-

netic meaning. There are no traces of punctures in Basiceros (=Octo-

struma) onorei Baroni Urbani & de Andrade (Fig. 28, top), in some

Eurhopalothrix species already illustrated by ‘TAYLOR (1990) and in

Octostruma balzani (Emery) from Ecuador (Fig. 28, bottom). DIETZ

(2004) also retains this character as basicerotine synapomorphy (his

character # 29), but admittedly following BOLTON (I. c.) and without

checking the species available to him. In addition, there are a few

Pyramica species with gastral sculpture perfectly comparable to the

one of some basicerotines (Fig. 29).

50. Worker. Bizarre pilosity absent or poorly developed (0), or

present and well developed (1). This is character # 27 of BOLTON

(1999) but coded differently for Colobostruma (polymorphic instead

of absent only) because of the presence of bizarre hairs in C. ceror-

nata Brown (BOLTON, 2000, Fig. 45). We are not sure about what

“bizarre” should or should not include but we followed BOLTON

(1999) in coding this character as present in Strumigenys and Pyra-

mica. As a consequence of this we coded it present also for Pilotro-

chus and Tatuidris since the hairs of these genera are virtually

identical to those e.g. of Pyramica medusa Bolton (BOLTON, 2000,

Fig. 296) and Strumigenys caniophanes Bolton (BOLTON, 2000, Fig.

490). On the other hand, at least Pyramica mitis Brown is com-

pletely destitute of standing hairs of any kind (BOLTON, 2000: 442).

Pyramica, as a consequence of this, was coded as polymorphic. We

suppose that this character should be equivalent to DIETz’s (2004)

character # 31 “Pélos especiais no escapo, pigidio e esternito do 1°

segmento do gaster”.

51. Male. Mandibles normally developed (0), or reduced

(1). This is character # 34 of DieTz (2004) differently coded, i.e.

developed instead of unknown for Stegomyrmex (Diniz, 1990: 279),

and Orectognathus (EMERY, 1924: 318 and personal observations),

polymorphic instead of reduced only in Pyramica (mandibles well

developed at least in P. rostrata (Emery) and P. baudueri (Emery)),

polymorphic instead of reduced only in Eurhopalothrix because of a

C. BARONI URBANI & M.L. DE ANDRADE

ZMB

Uni Basel

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00)

ZMB

Uni Basel

00050644

1 tergum and sternum in Basiceros

Lack of sculpture on the first gastra

Fig

(Emery) from Ecuador

balzani

sp. (top) and in

UDO

Octostruma) onore

(bottom)

(

ANT TRIBE DACETINI 59

sentence of MANN (1922: 41) attributing “mandibles well developed”

to the male of Eurhopalothrix gravis (Mann).

Paternce — - nni II

Fig. 29 - Comparable sculpture of the first gastric tergite in the dacetine Pyra-

mica margaritae (Forel) (left) and in the basicerotine Eurhopalothrix bruchi

(Santschi) (right). A sculptured gaster is supposed to be synapomorphic

for the Basicerotini and absent among Dacetini.

52. Gyne and male. Anterior wing with (0) or without (1) cell

RS. This is character # 37 of DieTz (2004). Due to the exiguity of

the material available to us, except for a few outgroups not consid-

ered by DieTz (2004), this character is coded entirely fide DIETZ

(2004). This character is cladistically informative only because of

the differences between outgroup and ingroup taxa. Within the

dacetinomorph genera, this character remains invariant or unknown

and, as such, it is without interest for our purpose.

53. Gyne and male. First anal vein of the anterior wing present

(0), or absent (1). This is character # 38 of DieTz (2004). Due to the

exiguity of the material available to us, except for a few outgroups

C. BARONI URBANI & M.L. DE ANDRADE

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62 C. BARONI URBANI & M.L. DE ANDRADE

not considered by Dietz (2004), this character is coded essentially

fide DieTtz (2004: table 2); Dietz’s table, however, records the anal

vein as present in Eurhopalothrix, but Dietz’s generic diagnosis (p.

66) states that in Eurhopalothrix the anal veins can be “ausentes,

incompletas, ou na forma de veias espectrais”. For this reason we

coded the anal vein as polymorphic in Eurhopalothrix. In addition,

Dietz (l. c.) codes the anal vein as absent in Acanthognathus but

we considered it as polymorphic because of a drawing by BROWN &

KEeMPF (1969. Fig. 7) depicting a male of A. rudis with anal vein.

54. Gyne and male. Pterostigma present (0), or absent (1).

Absence of pterostigma is an autapomorphy of the genus Eurho-

palothnx according to DIETZ's (2004) character # 36.

The distribution of these character states among the taxa

included in the present study is given in Table 1.

4.2. CHARACTERS USED IN OTHER PHYLOGENETIC STUDIES AND

EXCLUDED FROM THE PRESENT ONE

As already stated in the paragraph devoted to the outgroup

selection we did not consider the 7 autapomorphies and the 4 ple-

siomorphies attributed by BOLTON (1998: 68) to Tatuidris. These are

parsimony non-informative by definition (see e. g. HENNIG, 1950,

WILEY, 1980, and Mappison & Mappison, 2002, among many

other possible sources). Autapomorphies and plesiomorphies may

be very useful in appreciating how distinctive a taxon can be but

contribute nothing to its placement within a phylogeny. Stated oth-

erwise, considering all the characters listed by BOLTON (1998) and

even adding many more similar new ones would not affect our phy-

logenetic reconstruction.

Other characters appearing in the literature and excluded from

the present study are listed in the following.

Presence of a preocular carina (synapomorphy of the “daceto-

nine tribe-group” in BOLTON (1998:69)). Equal to character # 14 of

DieTz (2004). We are unable to see traces of the carina in a number

of dacetines like, e.g. Epopostruma natalae Shattuck, Microdaceton

tibialis Weber, Daceton armigerum (Latreille), Acanthognathus ocel-

latus (Mayr). The carina, on the other hand, is present in a number

of non-dacetine myrmicines like several Attini.

ANT) TRIBE DACETINI 63

BoLTON’s (1999) character # 6. Number and position of trigger

hairs. In Bolton’s matrix this character is unique (1. e.) autapomor-

phic for Microdaceton and Acanthognathus with a different character

state for each of these genera. Unfortunately, coding all the other

genera considered by Bolton as uniformly with paired trigger hairs

is far from being satisfactory. In some instances (e. g. in Pyramica

myllorhapha (Brown)) the labral hairs are so small that we strongly

hesitate to differentiate them from normal hairs and to call them

trigger hairs. In addition, the remaining genera considered in the

present paper but excluded by BOLTON (1999) show a wide array of

different morphologies, which, wrestling with character definitions,

could lead only to a longer set of doubtful generic autapomorphies

at most. The simple presence of “trigger hairs” on the mouthparts

is given as synapomorphic for the “dacetonine-group” by BOLTON

(1998) and retained as such also by us (our char. # 8).

BoLToN’s (1999) character # 13 and Dietz (2004) character

# 17. Scape in “normal resting position” above or below the eye.

This character is either equivalent to Bolton’s char. # 16 states 1-2

(“scrobe above... below eye”) or impossible to assess on preserved

material in which the antennae have been mounted.

BOLTON’s (1999) character # 15 (scape apical section angled or

not angled). Either we misunderstand Bolton’s description of it or

we should code this character in a way rather different from BOLTON

(1. c.). It can be excluded without losses for Bolton’s phylogeny

since in Bolton’s original resulting tree (BOLTON, 1999, Fig. 3) this

character has CI = 0.33 and does not contribute at all to Bolton’s

phylogeny (RI = 0.0).

BoLToN’s (1999) character # 17. “Eyes... not ventrolateral” vs.

“eyes... ventrolateral“ is excluded in favour of BARONI URBANI & DE

ANDRADE’s (1994) information richer char. # 3, eyes absent, dorso-

lateral, lateral, or ventral.

BoLTON’s (1999) character # 18. Gap between mandibles and

head capsule in profile. This character, in Bolton’s matrix, is shared

by Daceton and Microdaceton only, although the two genera appear

as distantly related in Bolton’s phylogeny. We excluded it simply

because we are unable either to understand its description or we'd

code it in a different way. Bolton’s original coding, anyway, does

not support his phylogenetic reconstructioin. As a consequence of

64 C. BARONI URBANI & M.L. DE ANDRADE

this, later on in the same paper (BOLTON, 1999:1678), this character

state is given as autapomorphic for Microdaceton.

BOLTON’s (1999) character # 19. Presence or absence of the

basimandibular gland. In BOLTON (1998:73) this character is given

as synapomorphic for the tribes Dacetini and Phalacromyrmecini. In

BOLTON’s (1999) matrix, this character is recorded for Pyramica, Stru-

migenys and Microdaceton only. We excluded it simply because either

we are unable to understand its description or we’d code it in a way

different from both BOLTON (1998) and BOLTON (1999) (e. g. poly-

morphic for Strumigenys, Pyramica and Rhopalothrix). As a further

proof of our claim, we remember that this same character in DIETZ

(2004) (char. # 5, table 2) is coded differently from BOLTON (1999)

for the following genera: Microdaceton, Pyramica, and Strumigenys.

BOLTON’s (1999) character # 20. Presence of apicofemoral and

apicotibial glands (or, more generically, “series of paired exocrine

glands”). This trait is given as synapomorphic for Strumigenys and

Pyramica. But, a few pages later, BOLTON (l.c.: 1665) adds “the [api-

cofemoral|] gland is apparently absent or at least has no externally

visible bulla, in some whole groups and a number of individual spe-

cies”. And, for the apicotibial gland “variation is as for the femoral

dorsal gland”. The observed polymorphism of these glands in both,

Pyramica and Strumigenys, renders their use for the cladistic analy-

sis parsimony uninformative.

BOLTON’s (1999) character # 26 “Metapleuralal gland bulla low

and widely separated from propodeal spiracle...”, versus “abutting

propodeal spiracle”. This is a repetition of BOLTON’s (1999) char-

acter # 22 “Metapleural gland bulla widely separated from annulus

of propodeal spiracle...”. It differs from it for being coded in three

steps instead of two. The newly added third step (metapleural gland

bulla high and close to spiracle) appears only once as redundant

autapomorphy for Acanthognathus. Otherwise chars. 22 and 26 have

exactly the same score and appear synapomorphic for five dacetine

genera. A very similarly defined character (“Propodeal spiracle low

. abutting the... metapleural gland bulla”) was already used by

BOLTON (1998: 73) as synapomorphy for the tribes Basicerotini and

Phalacromyrmecini contradicting in this way BOLTON’s (1999) defi-

nition. We regard this multiple use and different coding of similarly

defined characters as an additional reason for exclusion from our

data matrix.

ANT TRIBE DACETINI 65

BOLTON’s (1999: 1649) character # 29. Presence of spongiform

appendages of petiole and postpetiole. "This was originally formu-

lated as a plausible synapomorphy for Strumigenys and Pyramica

(but see also discussion under our character 41). Presence or absence

of spongiform appendages on the pedicel was already included in

BOLTON’s character # 28 (1999: 1649) under the more generic defi-

nition of “lobes or outgrowths of any form” and, under this defini-

tion, is also considered in our analysis as character # 41.

BOLTON’s (1999) character # 30. “Tergite of petiole and post-

petiole without lateral cuticular processes”. Excluded since we are

unable to understand the difference with Bolton’s char. # 28 “Peti-

ole and postpetiole without lateral (tergal) or ventral (sternal) lobes

or outgrowths of any form...”.

BOLTON’s (1999) character # 31. Postpetiolar spiracle lateral or

ventral. We are sceptic about this character due to the difficulty in

observing it. The spiracle is practically in the same position among

most of the species that we examined. When the petiole is laterally

expanded or bears a lateral lamella, the spiracle appears to be more

ventral than when the expansions or the lamellae are absent. Coding

this character as Bolton did and considering it with the others in

our search for the shortest tree(s) does not affect the number and

topology of the shortest trees but increases the tree length.

BoLTON (1999: 1681). Labrum “mediodorsally with a very

broadly and deeply concave depression in its proximal half”. This

should be an Epopostruma synapomorphy. In contrast with this

assumption, the morphology of Mesostruma turneri (Forel) is perfectly

comparable e.g. to the one of Epopostruma alata Shattuck (Fig. 30).

BOLTON’s (1998: 70) synapomorphy for the Basicerotini “scape

neck articulation” is another formulation of and correlated with his

previous character “torulus” described at the same page and already

included in our data matrix.

DieTz’s (2004) character # 3, “Forma do labro”. The labrum

should be longer than broad in all dacetine genera and broader than

long in other ants. This rule of thumb applies well to a majority of

species but there are numerous exceptions like broad labia in Dace-

ton (present study), Rhopalothrix (BROWN & KEMPF, 1960, Figs. 57 &

58), Basiceros (nr. singulare, present study), Pyramica (e. g. BOLTON,

2000, Fig. 215 and P. nannosobek, our Fig. 4, bottom), and Strumi-

66 C. BARONI URBANI & M.L. DE ANDRADE

ALL di 4

zMB

Uni Basel

a ns | pur n. MB

Fig. 30 - Comparable morphology of the labral concavity (a presumed Epopostruma

synapomorphy) in Mesostruma turneri (Forel) (top) and Epopostruma alata

Shattuck (bottom).

ANT TRIBE DACETINI 67

genys (BOLTON, 1999, Fig. 93, and S. harpyia, present study). Since

there are little doubts about the monophyly of the dacetine genera,

a monophyly already supported by a number of synapomorphies, we

thought it better to exclude this redundant but questionable character.

DieTz’s (2004) character # 9 “Forma das mandibulas”. Elon-

gated or specialized (i.e. non triangular) mandibles are coded as pres-

ent in a number of dacetine genera not corresponding to any tribal

grouping proposed so far. In addition we'd code as triangular at

least the mandibles of Colobostruma and of some Mesostruma species

(see e. g. the figures in BOLTON, 2000). Accepting Dietz’s revived

combination of Eurhopalothrix bruchi in Rhopalothrix also the latter

genus should have been coded as polymorphic (instead of special-

ized only). The specialization of the mandibles, moreover, is per-

fectly correlated with their capacity of engaging apically only or not.

As such this character would overweight our character # 10 (q. v.).

DieTz’s (2004) character # 11, “Forma das mandibulas lamin-

ares”. Presence or absence of an apical fork is coded in a very sub-

jective way (for instance absence of apical fork in Daceton, unknown

presence in Colobostruma, Pyramica and Microdaceton) that we are

unable of understanding. Since Dietz states that the dacetine apical

forks “probably evolved separately” we consider this statement alone

as a reason good enough to exclude this character.

DieTz’s (2004) character # 12, “Numero de dentes das forquil-

las apicais”. The two-state coding of this character (three vs. four

teeth) by Dietz is difficult to maintain and is contradictory even

within DIETZ’s text and matrix. In DIETz’s table 2, for instance,

only Rhopalothrix and Protalaridris share the four-toothed state,

but, according to Brown (1980, with whom we concur), Protalari-

dris possess only one apical spine and according to the same DIETZ

(2004: 187), Protalaridris should have a five-toothed fork... Even

solving somehow the problem of a correct coding for Protalaridris,

there are other relevant and pertinent morphologies not accounted

for and difficult to classify in this character definition. Examples of

such morphologies are the two-toothed fork of Daceton (coded “?”

by Dietz, |. c.), the two-toothed and three-toothed fork of Strumig-

enys (coded as three-toothed only by DIETZ, l. c.), a. o.

Dirtz’s (2004) character # 32. “Espinhos nos tarsos anteriores”.

Presence of some undefined type of tarsal spines should be a syn-

68 C. BARONI URBANI & M.L. DE ANDRADE

apomorphy for the Phalacromyrmecini and Basicerotini, though the

author admits that similar but not homologous spines are present

also in Daceton (and other genera, our study). Fig. 31 shows simi-

larity of structures in Strumigenys and Octostruma. 'This and other

similar cases encountered during the present study prevent consid-

eration of this character as phylogenetically sound.

Fig. 31 - Tarsal spines in the dacetine Strumigenys grandidieri Forel (left) and in the

basicerotine Octostruma balzani (Emery), a synapomorphy for Basicerotini

+ Phalacromyrmecini according to DIieTz (2004).

Dietz’s (2004) character # 33. “Habito de forrageamento”. We

concur with the description of this character that we consider as

plausible in spite of obvious miscoding in Dietz’s matrix (e. g. Pyra-

ANT TRIBE DACETINI 69

mica and Strumigenys epigaeic or arboreal only) but we consider our

behavioural knowledge of these ants as by far too scanty to allow

phylogenetic inferences.

Dietz’s (2004) character # 35. “Asa — conexào de cu-a proximo

a 1M (0); distante de 1M basalmente (1); distante de 1M distal-

mente (2)”. State 0 is recorded only among outgroups and state 2

should be autapomorphic for Eurhopalothrix. We are unable to see

differences in this trait between the wings of the specimens of £.

procera (Emery) and Octostruma balzani (Emery) available to us.

DieTz’s (2004) character # 40. “Presenca de asas”. ‘This is an

autapomorphy of Dietz’s outgroup genus Blepharidatta (not consid-

ered in the present study) and, as such, totally irrelevant in this

context.

DietTz’s (2004) character # 41. “Segunda linha de dentes”.

This is an autapomorphy of the genus Eurhopalothrix and, as such,

redundant of our character 54 in this context. This trait, moreover,

00049805 00049804 00050634 uf pe,

Fig. 32 - Basimandibular depression in Protalaridris near armata Brown (A), Rho-

palothrix ciliata Mayr (B) and Octostruma balzani (Emery) (C).

70 C. BARONI URBANI & M.L. DE ANDRADE

is not synapomorphic for Eurhopalothrix since it is absent at least in

an undescribed “Eurhopalothrix” species available for this study and

to be described in chapter 6 (Species-level additions) as Basiceros

papuanum (Fig. 46).

Basimandibular depression absent (0), or present and distally

marginate (1). This character (Fig. 32), shared by Rhopalothrix,

Protalaridris and Talaridris is mentioned by DieTz (2004) in the

generic descriptions of the first two genera only but is not used in

his phylogenetic analysis. We observed it in Talaridris as well and a

similar structure is visible also in some Octostruma. Some Pyramica

species also have some kind of basimandibular depression.

Larval hairs bifid or not. This is character # 10 of BARONI

URBANI & DE ANDRADE (1994). This is a classic synapomorphy for

all “dacetiform” genera resulting from the literature but verified in

a too small number of taxa to be attributed unmistakable phyloge-

netic value.

4.3. RESULTS OF THE CLADISTIC ANALYSIS

Our parsimony analysis yields 414 equally shortest trees of length

208 (considering polymorphism as multiple speciation events), Con-

sistency Index (CI) 0.746, Retention Index (RI) 0.785, and Rescaled

Consistency Index (RC) 0.592. The strict consensus tree is given in

Fig, 397

Fig. 34 depicts the same consensus tree as the one of Fig. 33

but expressed as phylogram, i.e. with the branch lengths propor-

tional to the number of synapomorphies supporting each branch.

In it the synapomorphic characters for each branch are also added

to the different branches with the same character numbering as in

the previous text. The tree has the same number of branches as the

input genera in our data and horizontal square brackets on top of

the drawing indicate genera merged as a result of our analysis with

their oldest available synonym. Synonymies of genera are inferred

on the base of missing or too weak synapomorphies as shown on

the tree and discussed in detail under the genus group systematics

(chapter 4.5). Weakness and strength of the individual synapomor-

phies will be judged in terms of Consistency Index and Retention

Index on the Consensus Tree of Figs. 33 and 34.

ANT TRIBE DACETINI 71

Myrmecia

Pseudomyrmex

Myrmica

Stegomyrmex

Calyptomyrmex

Tatuidris

Acanthognathus

Colobostruma

Mesostruma

Epopostruma

Microdaceton

Daceton

Orectognathus

Pyramica

Strumigenys

Phalacromyrmex

Pilotrochus

Ishakidris

Basiceros

Eurhopalothrix

Octostruma

Protalaridris

Talaridris

Rhopalothrix

Fig. 33 - Strict consensus tree of the 414 equally most parsimonious trees drawn

from the characters and taxa discussed in the text. Length of the indi-

vidual trees 208.

we C. BARONI URBANI & M.L. DE ANDRADE

Myrmecia

Pseudomyrmex

Myrmica

Stegomyrmex

POU A

Calyptomyrmex

SA WWW Nn o

PONIO WO

Tatuidris

Ad awe

WRI ea GIO

Acanthognathus

TAL eevee seni

CSIGY

Sees"

Colobostruma

wet SF

ze 'Mesostruma 8

Lane ai

DI er a

fT?

SD W® IEpopostruma è

De Bs OS

NU jf

By. rte berry en Microdaceton

Lund. OPE OPES

PON

N o

Daceton

a | SANA

Orectognathus

i w

Pyramica =

SITI TRI e PAY . Q

wraw !Strumigenys g

De At 4 SO Pe

DANO Phalacromyrmex

WGI

NUDO

Pilotrochus

LAI

Rez}

ishakidris

Basiceros

Eurhopalothrix

Octostruma Do

UTALALDADAANPINA DM ee: ©

DOVUTA WD I Protalaridris 3

oo GY

Lea

Talaridris

Rhopalothrix

471

cae)

Fig. 34 - Same strict consensus tree as the one of Fig 33 expressed as phylogram,

i. e. with the branch lengths proportional to the number of derived char-

acters changing on them. Character numbering as in text. The number of

taxa is the same as in the inputted data. The square brackets embracing

groups of taxa show synonymies to be proposed in this paper because of

missing or too weak synapomorphies for one or more contained genera.

ANT TRIBE DACETINI BS

We refrain from attributing great phylogenetic value to this tree.

The main reasons for our evaluation are its construction, drawn on

a great number of phylogenetically insignificant characters and our

incapacity to find better ones. The scarcity and weakness of the

known characters on which our phylogeny is based can be better

appreciated on the tree of Fig. 35, a 50% majority consensus tree

drawn from 100 bootstrap replicates.

The phylogeny of Figs. 33-35, however, is an excellent discrim-

inant criterion to decide what can be actually said or not on dacetine

classification.

We insist that a good number of the characters from which

our phylogenetic reconstruction is drawn are of doubtful generic or

tribal value and are highly variable within the taxa considered. The

phylogenetic value of a character may be a rather subjective topic

but at least its variability will be described and taken into account

in the classificatory scheme proposed below.

Another phylogenetic issue worth discussing here is the plesi-

omorphic condition of the dacetine mandible morphology. In our

former Dacetini study (BARONI URBANI & DE ANDRADE 1994: 11) we

contrasted a widespread belief for which elongate mandibles should

be the plesiomorphic dacetine state and short mandibles the apo-

morphic one (Brown & WILSON, 1959). Our conclusion was based

on parsimony evidence drawn from analysis of morphological data

and on the effortless consideration that any conceivable Dacetini

sister-group should also have short mandibles.

BoLTON (1999: 1644) equated the two terms “long mandibulate”

and “short mandibulate” already established in the literature with his

“kinetic” and “static pressure mandibles”. Our lack of enthusiasm

for the use of these terms has been already justified at the begin-

ning of chapter 4.1. But BOLTON (l.c.) adds: “Philip Ward (Univer-

sity of California) has recently added support to the Brown-Wilson

hypothesis”. This statement is explained by the tracing by Ward

of the most parsimonious evolution of the mandibular length on

Bolton’s own cladogram (BoLTON, 1999: fig. 3). Irrespectively from

the quality of the cladogram in question, this result is a straightfor-

ward consequence of the nature of the studied sample and wonder-

ing about the size of the ancestral mandibles, in this context, would

be absurd. In fact, in a 9-taxa tree where all 8 most basal branches

74 C. BARONI URBANI & M.L. DE ANDRADE

100

72 79 | 51

Chi

100

Myrmecia

Pseudomyrmex

Myrmica

Stegomyrmex

Calyptomyrmex

Tatuidris

Acanthognathus

Colobostruma

Mesostruma

Epopostruma

Microdaceton

Daceton

Orectognathus

Pyramica

Strumigenys

Phalacromyrmex

Pilotrochus

Ishakidris

Basiceros

Eurhopalothrix

Octostruma

Protalaridris

Talaridris

Rhopalothrix

Fig. 35 - 50% Majority Rule consensus tree resulting from 100 bootstrap replicate

analyses of the data of Table 1. The figures on the branches give their

relative frequencies among the replicates in percent.

ANT TRIBE DACETINI PD

represent long mandibulate taxa (as is Bolton’s cladogram), there is

no arithmetical possibility for the hypothetical common ancestor to

be short mandibulate. This conclusion is trivial and the sole hypo-

thetical ancestor worth speculation is not this one but an earlier one

connecting the observed long mandibulate clade to the short man-

dibulate sister clade. Dr. Ward’s conclusions, in this case, are either

easily predictable and hence unimportant, or erroneous by trying to

extrapolate a deduction outside the observed variation range.

Performing again the same analysis on a broader sample of long

and short mandibulate taxa like the one considered in the present

study, confirms our former hypothesis and the common sense induc-

tion that long mandibulate Dacetini should have originated from a

short mandibulate ancestor (Fig. 36).

The information contained in the tree of Figs. 33-36 forces to

take nomenclatorial action on a number of problems, which, in hier-

archically decreasing taxonomic rank, are the following:

4.4. FAMILY-GROUP SYSTEMATICS

Tribe Agroecomyrmecini Carpenter

Agroecomyrmicini [sic] CARPENTER, 1930: 34. Type genus tAgroecomyrmex by

inference. Tribe of Myrmicinae.

Agroecomyrmecini Carpenter, BRowN & KEMPF, 1968: 184. First correct spelling.

Agroecomyrmecinae Carpenter, BOLTON, 2003: 51. Subfamily of Formicidae.

BoLTON (1998) lists a set of 7 autapomorphies and 4 plesiomor-

phies for Tatuidris aimed to prove the exclusion of this genus from

his “dacetonine tribe-group”. Autapomorphies, however, are cladis-

tically uninformative and plesiomorphies cannot be used to infer

phylogenetic relationships (see e. g. HENNIG, 1950; WILEy, 1980;

and Mappison & Mappison, 2002). Later on, BOLTON (2003: 51)

places Tatuidris, together with two poorly known fossil genera,

Agroecomyrmex and Eulithomyrmex, in a new separate subfamily,

the Agroecomyrmecinae. All the characters supposed to characterize

and separate the Agroecomyrmecinae from the Myrmicinae as given

by Bolton appear sporadically also among other Myrmicinae genera,

though in a convergent way according to BOLTON (I. c.).

76 C. BARONI URBANI & M.L. DE ANDRADE

| 3} mMyrmecia

3-———iPseudomyrmes

i w x

eM |] AT YA

=== DStegomyrmex

Re DCalyptomyrmex

g@ Acanthognathus

A E Colobostruma

E Mm Mesostruma

a mEpopostruma

Im Microdaceton

@Daceton

moOrectognathus

E E3Pyramica

)

Ci Saba

E DPilotrochus

goo shakidrs

E gm Strumigenys

(zi .

ZIO Basiceros

R4OYS

4 imEurhopalothrix

Mn = ea Octostruma

TUTTE TW E Iqupy[0]

E g Protalaridris

(fim Talaridris

Fig: 36 - Same tree as in Figs. 33 & 34 on which the most parsimonious recon-

struction of the evolution of the size of the mandibles is traced. Graphic

display by accelerated transformation. Contrarily to a common belief sup-

ported by BOLTON (1999), long mandibulate Dacetini appear to originate

from a short mandibulate ancestor.

ANT TRIBE DACETINI TV

The following character states shared by Tatuidris with the

other myrmicine genera considered for the present study result as

plausibly synapomorphic for the whole subfamily Myrmicinae:

1. Promesonotal suture fused (from an unfused ancestor). CI 1.00,

RI 1.00. All known myrmicines have a fused promesonotal

suture.

2. Petiole dorsoventrally fused (from an unfused ancestor). CI

1.00, RI 1.00. All known myrmicines have a dorsoventrally

fused petiole.

3. Loss or fusion of the second radial cell (from an ancestor with

recognizable second radial cell). CI 1.00, RI 1.00. Among the

ingroup taxa considered in the present study, only Stegomyrmex

is polymorphic for this trait.

4. Eyes position: lateral (vs. dorsolateral). CI 0.67, RI 0.80. The

ancestral position dorsolateral reappears homoplastically among

some dacetine taxa.

Since all potentially valid characters tending to exclude Tatui-

dris (and, by inference, Agroecomyrmex and Eulithomyrmex) from the

Myrmicinae were included in our data matrix and failed to fulfil

their supposed role, we feel compelled to re-propose the following

rank revival within the Myrmicinae:

Subfam. Myrmicinae: Tribus Agroecomyrmicini CARPENTER,

1930: 34. Type genus: Agroecomyrmex. Tribal status revived.

=Subfamily Agroecomyrmecinae Carpenter, BOLTON, 2003: 51

(downgraded to tribal rank in the present paper).

And the consequent subfamilial transfers

tAgroecomyrmex WHEELER, 1910: 265 et auctorum omnium

recentiorum. Genus ad Myrmicinae. Nec Agroecomyrmex, genus ad

Agroecomyrmicinae, BOLTON, 2003: 51. Subfamilial transfer.

tEulithomyrmex CARPENTER, 1935: 91 et auctorum omnium

recentiorum. Genus ad Myrmicinae. Nec Eulithomyrmex, genus ad

Agroecomyrmicinae, BOLTON, 2003: 52. Subfamilial transfer.

Tatuidris Brown & Kempr, 1968: 186 et auctorum omnium

recentiorum. Genus ad Myrmicinae. Nec Tatuidris, genus ad Agro-

ecomyrmicinae, BOLTON, 2003: 52. Subfamilial transfer.

78 C. BARONI URBANI & M.L. DE ANDRADE

Notice that our conservative appreciation of Tatuidris within the

Myrmicinae holds in spite of equivalent consideration of all the cla-

distically informative characters that, according to BoLTON (2003),

should prove its exclusion.

In our phylogenetic reconstruction, there are six synapomor-

phies bringing Tatuidris within the Myrmicinae in the same cluster

as all the Dacetini and excluding as less related other undoubted

Myrmicinae genera like Myrmica, Stegomyrmex, and Calyptomyr-

mex. These synapomorphies are the following:

1. Mandibles at rest opposing at least in part (instead of crossing).

CI 1.00, RI 1.00. This is one of the strongest dacetine synapo-

morphies.

2. MITI < 130 (as opposed to MTI > 150). CI 1.00, RI 1.00. This

is another strong dacetine synapomorphy.

3. Reduction of the maxillary palps from 2-jointed to 1-jointed. CI

0.85, RI 0.80. The one-jointed condition should be the original

dacetine morphology, though a number of dacetine genera and

individual species underwent secondarily inverted evolutionary

trends.

4. Reduction of the male mandibles. CI 0.75, RI 0.80. This appears

to be a general dacetine trait with a few known exceptions in

Pyramica and Eurhopalothrix.

5. Presence of a two-segmented antennal club. CI 0.33, RI 0.71.

This appears to be a universal dacetine trait with the sole excep-

tions of the small genera Microdaceton, Daceton, and Orectogna-

thus.

6. Reduction of the number of antennal joints from 11-12 to 9 or less.

CI 0.20, RI 0.50. Acanthognathus, Daceton, Phalacromyrmex and

Basiceros should represent evolutionary inversions according to

our phylogenetic reconstruction, but we already wrote about the

doubtful phylogenetic meaning of this character while describing it.

Our result contrasts, however, with the results of a “prelimi-

nary analysis” based on sequence data from seven nuclear genes by

WARD et al. (2005), from a phylogenetic tree inferred from Bayes-

ian posterior probabilities drawn from portions of five nuclear genes

and one mitochondrial gene by MOoREAU et al. (2006a), and from an

analogous tree drawn from seven nuclear gene fragments by BRADY

ANT TRIBE DACETINI 79

et al. (2006a). According to these results, Tatuzdris should belong to

a clade opposed to most other known ants.

We believe that two important aspects of these results should

be remembered.

The first is that assuming that molecular-drawn phylogenies

are always correct or at least better than morphology based ones

— as several myrmecologists seem to believe — is equivalent to the

belief that all computer-drawn results must be true. BRADY et al.

(2006b: Table 7) unfortunately omit fossil-based minimum ages for

the Dacetini from their data but (2006a:1) point out correctly to

“apparent conflicts between fossil, morphological, and molecular

data”. Actually, the main source of conflict among this triplet of

factors is the molecular data. We consider as alarming the fact that

genetic similarity measures contradict both, obvious morphological

similarity suppositions and morphology-based parsimony measures.

Second, molecular data are radically different from morphologi-

cal data by the difficulty of arguing in favour or against their results.

Morphological results may appear more or less plausible according

to the characters on which they are drawn, but the sole chance to

discuss the validity of results obtained from molecular data is to

compare them with results obtained from morphological data (the

so-called congruence criterion). It is obvious that in this case there

is total lack of congruence between the available molecular and mor-

phological information. Both aspects need to be further improved.

We assume, moreover, that the differences within the most

recent molecular ant phylogenetic reconstructions by BRADY (2003),

OHNISHI et al. (2003), AsTtRUC et al. (2004), SAux et al. (2004),

WARD et al. (2005), MOREAU et al. (2006a), and BRADY et al. (20062),

should be sufficient to impose some prudence before blindly prefer-

ring molecular to morphological results.

However interesting the Tatuidris molecular outcome may be,

we consider it as Dr. Ward, co-author of two of the previously cited

molecular analyses (WARD et al., 2005; BRADY et al., 2006a), consid-

ered it in a recent web document (Warp, 2005), where he writes

that Tatuidris “are rather specialized ants, and it seems plausible to

me [1. e. P. S. Ward] that their divergent evolution includes extreme

modification of the myrmicine groundplan”. In the analysis by

MOREAU et al. (2006a) and Brapy et al. (2006a), Tatuidris appears

80 C. BARONI URBANI & M.L. DE ANDRADE

as the sister genus of Paraponera, a phylogenetic position difficult

to digest when considering our present morphological understanding

of these two genera.

This presumed strict phylogenetic relationships between Tatui-

dris and Paraponera, however, is based on Bayesian posterior proba-

bilities, while maximum likelihood bootstrap (MOREAU et al., 2006a)

and maximum parsimony bootstrap analyses (MOREAU et al., 2006a;

Brapy et al., 2006b) failed to support the presumed sister pair (7at-

uidris, Paraponera) and any of the nodes intermediate between this

pair and the basal node of Formicidae (MOREAU et al., 2006b).

We wish that these results, at this stage, will be considered as a

stimulating molecular challenge to our organismic knowledge of ants.

Within our data set, assignment of Tatuidris to a unique clade

with the other dacetines is supported by 72% of the bootstrap rep-

licates and its classification among Myrmicinae by 100% bootstrap

replicates (Fig. 35), confirming in this way WARD's (2005) morpho-

logically based intuitive hypothesis.

Our data, nonetheless, might be insufficient to take a durable

decision about the monophyly or diphyly of the opposable mandi-

bles among the Myrmicinae. Our cladogram of Fig. 33 strongly sug-

gests a unique origin of the mandible opposability. But we are aware

that to obtain a firmer certainty (supported e. g. by high bootstrap

frequencies) one might need to consider all myrmicine genera and

many more additional characters. Once the sister-group relationship

between the dacetines and the agroecomyrmicines will be confirmed,

considering them as two separate tribes or only one tribe becomes

irrelevant and just a matter of taste ad a phylogenetically irrelevant

issue. The poor characterization of the tribe Dacetini (q. v.) after

exclusion of the Agraocomyrmecini may be considered as an argu-

ment in favour of merging the two tribes together. We maintain the

two tribes separate in this paper, at least provisionally, in order to

promote nomenclatorial stability.

Assuming that the characters observed in the extant Tatuidris

can be attributed by inference to the two fossil genera Agroecomyr-

mex and Eulithomyrmex, the tribe Agroecomyrmecini is character-

ized by the two following uniquely derived characters:

Petiolar tergum and sternum in posterior view differently shaped

(CI 1.00, RI 0.00).

ANT TRIBE DACETINI 81

Eyes at the posterior border of the antennal scrobe (CI 0.71,

RI 0.83).

BoLToN (1998, 2003) lists other potentially autapomorphic

characters not considered for the present analysis since they are

parsimony uninformative but which may prove useful in further

diagnosing these ants as a separate tribe.

Tribe Dacetini Forel

Dacetonini Foret, 1892: 344. Type genus Daceton Perty, by inference. Tribe of

Myrmicinae.

<Dacetonini Forel, BOLTON, 1998: 71.

Dacetonii FOREL, 1893a: 164.

Dacetii Forel, Emery, 1895c: 770.

Dacetini Forel, EMERY, 1914: 34 & 39.

Dacetiti Brown, 1952: 10. Type genus: Daceton Perty, by original designation.

BARONI URBANI & DE ANDRADE, 1994: 9. Junior synonym of Dacetini.

Dacetini Forel, BARONI URBANI & DE ANDRADE, 1994: 9.

<Dacetini Forel, BoLTON, 2000: 12.

Basicerotini Brown, 1949c. Type genus Basiceros Schulz by inference. Synonymy

reinstated.

Basicerotini BRown, BARONI URBANI & DE ANDRADE, 1994: 10. Junior synonym of

Dacetini.

Basicerotini Brown, BoLTon, 1998: 70. Tribe of Myrmicinae.

Arestognathiti BRown, 1952: 10. Misspelling for Orectognathiti.

Orectognathiti Brown, 1952: 10. Type genus: Orectognathus Smith, by original

designation. BARONI URBANI & DE ANDRADE, 1994: 9. Junior synonym of Dacetini.

Epopostrumiti Brown, 1952: 10. Type genus: Epopostruma Forel, by original

designation. BARONI URBANI & DE ANDRADE, 1994: 9. Junior synonym of Dacetini.

Strumigeniti Brown, 1952: 10. Type genus: Strumigenys Forel, by original

designation. BARONI URBANI & DE ANDRADE, 1994: 9. Junior synonym of Dacetini.

Phalacromyrmecini WHEELER & WHEELER, 1976: 60. Unavailable name.

Phalacromyrmecini DLussky & FEDOSEEVA, 1988: 80. First available name (reference to a

description). Type genus Phalacromyrmex Kempf by inference. Synonymy reinstated.

Phalacromyrmecini, BARONI URBANI & DE ANDRADE, 1994: 10. Junior synonym of

Dacetini.

Phalacromyrmecini, BOLTON, 1994: 106. Tribe of Myrmicinae.

Basicerotini is a junior synonym of Dacetini.

The possibility to separate from the remaining Dacetini a small

set of peculiar genera corresponding to the Basicerotini was already

considered and discarded by EMERY (1924: 313) who wrote:

82 C. BARONI URBANI & M.L. DE ANDRADE

“Un premier groupe comprend les genres à fosses antennaires placées au-dessus

des yeux: Acanthognathus, Microdaceton, Orectognathus, Strumigenys, Pentastruma,

Epitritus (Acanthognathus a la scrobe réduite a peu près de rien).

Un deuxiéme groupe comprend les genres Basiceros, Rhopalothrix et

Epopostruma, qui ont la scrobe placée au dessus de l'oeil. ...

Le genre Daceton, bien qu’il ait laréte frontale prolongée un peu au dessous de

l’oeil, me semble se rattacher au premier groupe”.

When Brown (1949c) separated along the same line as EMERY

(1924) his new tribe Basicerotini, he did not use the relative position

of the antennal scrobe to define it but referred only to some differ-

ences in hair morphology. BARONI URBANI & DE ANDRADE (1994)

proposed the synonymy of Basicerotini with Dacetini after showing

the inconsistency of this character.

BOLTON (1998) apparently agreed with this since he abandoned

all references to the hair morphology but revived the tribe Basicero-

tini on the base of 10 newly defined synapomorphies. One of them

is the position of the antennal scrobe already described and dis-

carded by Emery (1924). All BOLTON’s (1998) characters are either

considered for the present analysis or excluded from the analysis

after justification of their exclusion.

Here we propose again that the name Basicerotini should be

considered as a junior synonym of Dacetini for a cladistic reason, a

practical reason, and several general taxonomic reasons.

The cladistic reason is that maintaining the Basicerotini as a

tribe separate from the Dacetini would render the first paraphyletic

to the second or, at least to the Dacetini as they are understood by

Boltom-isee Wie. 32),

The practical reason is well exemplified by the case of Rho-

palothrix inopinata de Andrade, a species originally described as a

basicerotine and now transferred to the dacetine Strumigenys in spite

of the fact that its sole clearly visible synapomorphic character is a

basicerotine character (see later under the treatment of Strumige-

nyS).

The general taxonomic reasons are that the homogeneity of the

morphological boundaries of this hypothetical tribe result much less

precise than what is commonly accepted for valid ant tribes. The

following analysis of all eleven potentially synapomorphic traits of

the Basicerotini should further justify our appreciation.

ANT TRIBE DACETINI 83

Base of scape bent at right angle. CI 1.00, RI 1.00. Universal

among the Basicerotini, a similar scape is known also for a few

Dacetini species of the genera Colobostruma and Pyramica.

Antennal fossa separate from the antennal scrobe. CI 1.00, RI

1.00. The separation is indistinct in the basicerotines Eurhopalo-

thrix bruchi and E. heliscata. On the contrary the antennal fossa

and scrobe are clearly separate in some Strumigenys, Pyramica,

and Colobostruma.

Pretergite of the first gastral segment sessile. CI 1.00, RI 1.00.

Formally this character state, considered as plesiomorphic by

BoLToN (1998), is the sole unequivocal synapomorphy of the

Basicerotini. ‘This prominent position in our character analysis 1s

derived essentially from our coding of it. BOLTON (1998: 72) con-

sidered the complementary state of this same trait (1. e. pretergite

neck-like) as synapomorphic for his Dacetini + Phalacromyrmecini.

First gastral segment densely punctuated. CI 1.00, RI 1.00.

There are a few Eurhopalothrix and Octostruma species with the

first gastral segment smooth. On the other hand some Stegomyr-

mex and Pyramica species have a definitely punctuated first gas-

tral segment.

Hypertrophied torulus. CI 0.83, RI 0.83. A hypertrophied tor-

ulus can be found also in some scattered species of Pyramica,

Strumigenys, Mesostruma, Colobostruma and Tatuidris.

Labrum with transversal groove. CI 0.80, RI 0.83. This trait,

universal within the Basicerotini, is found also in some Pyramica,

Strumigenys and Colobostruma species among Dacetini s. str., and

in Stegomyrmex.

Male mandibles not reduced in size. CI 0.75, RI 0.80. Most

(not all) known dacetine and agroecomyrmecine males under-

went a reduction of the mandibles that should have secondarily

re-grown among most (not all) known basicerotine males. ‘The

paucity of known males of these ants parallels the plausibility of

this hypothesis.

Postpetiolar presclerites arising from a deep concavity. CI 0.67,

RI 0.86. As we already wrote in the characters’ description, this

same morphology is known also for some Pyramica species.

Base of first gastral sternum truncate. CI 0.67, RI 0.71. Origi-

nally defined by BOLTON (1999) as synapomorphic for Strumige-

84 C. BARONI URBANI & M.L. DE ANDRADE

nys and Pyramica, in addition than among all basicerotines, this

character state is present also in Ishakidris, Stegomyrmex, and in

some but not all Colobostruma, Pyramica, and Strumigenys spe-

cies.

- Antennal scrobe below the eyes. CI 0.60, RI 0.83. This trait,

constant among basicerotines, is shared with the dacetines Colo-

bostruma, Mesostruma, and Epopostruma.

- Articulation between gaster and postpetiole broad. CI 0.33, RI

0.66. The complementary state of this character (1. e. articulation

narrow) was originally proposed by BOLTON (1998:72) as poten-

tially synapomorphic for the Dacetini + Phalacromyrmecini.

On the contrary, within the Basicerotini the articulation can be

narrow in some Basiceros and outside them it is broad in Tatui-

dris and in some Pyramica and Strumigenys.

One should not forget that the goodness of fit of some of the

above traits on the resulting phylogeny and classification as shown

by high CI values is a trivial product of our coding these traits

as invariant among in- and/or out-group taxa as Bolton did. As a

matter of fact, exceptions to these a prior: homogeneity (= mono-

phyly) hypotheses are regularly described and photographically doc-

umented in our initial characters’ description (chapter 4.1).

Phalacromyrmecini is a junior synonym of Dacetini.

The first to recognize similarities between the three genera

included in this tribe, 1. e. Phalacromyrmex, Ishakidris, and Pilotro-

chus, was BOLTON (1984) who wrote that there is “no advantage to

adding yet another formal [i.e. tribal] name to the confusion”. In

this paper these genera were correctly compared with the dacetine

genus Glamyromyrmex (= Pyramica). 'The name Phalacromyrme-

cini was introduced without definition or description by WHEELER

& WHEELER (1976) and repeated (among others) by DLussky &

FEDOSEEVA (1988) who rendered the name available by referring

to BOLTON’s (1984) description (International Code of Zoological

Nomenclature 13 (a) (11)).

BARONI URBANI & DE ANDRADE (1994) considered the Phala-

cromyrmecini as a junior synonym of Dacetini. Unexpectedly, this

view was immediately contrasted by BOLTON (1995) without giving

ANT TRIBE DACETINI 85

reasons for it and later by BOLTON (1998) who defined the tribe by

means of three synapomorphies. In the present paper we already

showed that one of them (presence of a katepisternal groove) was

artificially introduced by BOLTON (1998) only to support separa-

tion of the Phalacromyrmecini. In a former paper, in fact, BOLTON

(1984) uses the presence of this same character in [shakidris to dif-

ferentiate it from the other two genera now attributed to the Phala-

cromyrmecini where the groove is absent.

Other potential “Phalacromyrmecini” synapomorphies evidenced

by our analysis are the following:

- Mandibles with alternating small and large teeth. CI 1.00, RI

1.00. If there is no variation of this character within the tribe

(one should remember perhaps that this tribe comprises only

three species classified in three different monotypic genera),

mandibles with teeth alternating in size are known also in an

equivalent number of Pyramica and Octostruma species (see our

character description). The high CI and RI values of this char-

acter, hence, are due to our following Bolton’s interpretation of

it totally ignoring ascertained outgroup variability (see p. 24 for

our discussion of char. # 11).

- Mesosternal hair beds visible. CI 0.91, RI 0.67. We reaffirm

that the consistency of this character state within this pre-

tended three-species tribe is due to the exiguity of species

included in the tribe. The same character state is widespread

among Dacetini and Basicerotini sensu BOLTON (1998) and was

used also as a synapomorphy for Strumigenys and Pyramica by

BOLTON (1999).

- Eyes ventral. CI 0.67, RI 0.80. Another trait shared with Stru-

migenys and Pyramica as well.

- Clavate scape. CI 0.50, RI 0.60. A clavate scape is known also

in Stegomyrmex, Calyptomyrmex, Tatuidris, and some Pyramica

species.

Each of these characters has a variability outside this three-spe-

cies tribe far beyond what is normally accepted for other ant tribes.

BoLTON’s (2006) additional argument in favour of retaining the

Phalacromyrmecini, i.e. absence of the basimandibular process is

untenable. In fact the presence of the process is supposed to be

86 C. BARONI URBANI & M.L. DE ANDRADE

apomorphic for the Dacetini and as BOLTON (I. c.) states, its absence

among Phalacromyrmecini is “presumably plesiomorphic”.

Limits and definition of the Dacetini

After merging the Basicerotini and the Phalacromyrmecini in it,

the tribe Dacetini may be characterized by the following five poten-

tial synapomorphies, only the first of which, in our opinion, can be

considered phylogenetically relevant.

- Presence of “trigger hairs”. CI 1.00, RI 1.00. We wrote earlier in

this paper that the dacetine “trigger hairs”, as used by BOLTON

(1998) and in the present study may be homologous in function

but they are obviously not homologous in morphology when they

arise from different sclerites. Nonetheless, in spite of being wide-

spread among Dacetini, at least a few Colobostruma and Pyramica

species are deprived of such hairs. Given their rarity, these cases

could be interpreted as secondary losses.

- Labrum not capable of full reflexion. CI 0.50, RI 0.87. This char-

acter matches perfectly its scope as dacetine synapomorphy, except

for its absence in Colobostruma, Mesostruma and Epopostruma.

- Scape subcylindrical. CI 0.50, RI 0.60. We already discussed the

phylogenetic meaning of this character for its alternative state

(scape clavate) as potential synapomorphy for the Phalacromyr-

mecini.

- Presence of limbus. CI 0.50, RI 0.80. Evidentiation of this char-

acter seems to be an artefact of character optimization. The

limbus, in fact, is absent among several Dacetini genera although

it is present only in Stegomyrmex outside this clade.

- Propodeal spiracle close to propodeal declivity. CI 0.33, RI 0.60.

‘The spiracle is at about midlength of the sclerite in Daceton and,

among the outgroups, it is again close to the declivity in Calyp-

tomyrmex.

‘The above synapomorphic weaknesses of the Dacetini repropose

the opportunity of separating them from the Agroecomyrmecini. If,

on one hand, there is no doubt that only one Dacetini tribe includ-

ing the Agroecomyrmecini would automatically result much better

defined and stable, we see no urgent practical reasons to take this

decision immediately.

ANT TRIBE DACETINI 87

4.5. GENUS-GROUP SYSTEMATICS

In the following taxa discussion we shall report only the most

recent or the most significant references and synonyms. A more

exhaustive list of the older ones can be found in BARONI URBANI &

DE ANDRADE (1994), BOLTON (1995) and BOLTON (2000).

All our classificatory inferences are drawn from our Fig. 33,

which, we repeat, we consider essentially as an operational picture

of our factual Dacetini knowledge.

The following ingroup genera, listed in alphabetical order,

appear as actually or potentially taxonomically sound by possessing

one or more plausible synapomorphy each:

Acanthognathus Mayr

Acanthognathus Mayr, 1887: 567. Type species Acanthognathus ocellatus Mayr by

monotypy.

Synapomorphies resulting from our analysis:

Worker (and gyne) metapleural gland very close to propodeal

spiracle. CI 0.62, RI 0.40. This trait appears also in Epopostruma,

Microdaceton, Protalaridris, a. o. as coded also by BOLTON (1999,

Table 1) for his Dacetonini.

Worker (and gyne) scape straight at base. CI 0.50, RI 0.33. The

straight condition of Acanthognathus is shared with Daceton.

Worker (and gyne) antennae 11-jointed. CI 0.20, RI 0.50 for

the 11-12 jointed state.

This list, however, does not evidentiate the main synapomor-

phy for the species of the genus, i.e. our char. #13, state 2, the

basimandibular process long and apically bifurcated. 'The reason for

this omission is a purely logical pitfall: since state #2 appears only

in Acanthognathus and state #1 is present in all the other genera of

the clade, it is impossible to ascertain whether the ancestor of the

whole clade presented state #1 or #2 and hence if #2 is exclusive of

Acanthognathus or common to Acanthognathus and the hypothetical

ancestor of the whole clade.

The extant Acanthognathus species were revised and keyed by

Brown & KEMPF (1969). Afterwards BARONI URBANI & DE ANDRADE

(1994) described one fossil species from Dominican amber.

88 C. BARONI URBANI & M.L. DE ANDRADE

Basiceros Schulz

Ceratobasis SMITH, 1860: 78. Type species Meranoplus singularis Smith by monotypy.

Nec Ceratobasis Lacordaire, 1848 (Coleoptera).

Basiceros SCHULZ, 1906: 156 (replacement name for Ceratobasis).

Aspididris WEBER, 1950: 3. Type species Aspididris militaris Weber by monotypy.

Synonymy by Brown, 1974: 132.

Creightonidris Brown, 1949c: 89. Type species Creightonidris scambognatha Brown

by original designation. Synonymy by DIETZ, 2004: 48.

Rhopalothrix Mayr, 1870: 415. Type species Rhopalothrix ciliata Mayr designated

by WHEELER, 1911b: 172. New synonymy.

Heptastruma WEBER, 1934: 54. Type species Heptastruma wheelert Weber by original

designation. Synonymy with Rhopalothrix by Brown & KEMPF, 1960: 230. New

synonymy.

Acanthidris WEBER, 1941: 188. Type species Acanthidris isthmicus Weber by original

designation. Synonymy with Rhopalothrix by Brown & KEMPF, 1960: 230. New

synonymy.

Rhopalothrix subg. Octostruma FoREL, 1912b: 196. Type species Rhopalothrix simoni

designated by WHEELER, 1913: 82. New synonymy.

Octostruma Forel, Brown, 1948: 102.

Talaridris WEBER, 1941: 184. Type species Talaridris mandibularis Weber by original

designation. New synonymy.

Eurhopalothrix Brown & KEMPF, 1960: 202. Unavailable name without designation

of type species.

Eurhopalothrix Brown & Kempr, 1961: 44. Type species Rhopalothrix bolaut Mayr

by original designation. New synonymy.

Protalaridris BROWN, 1980: 36. Type species Protalaridris armata Brown by original

designation. New synonymy.

The genus Basiceros may be identified and characterized by the

set of synapomorphies that we already listed and discussed for the

tribe Basicerotini. If tribal rank for them appears exaggerated to say

the least, recognizing to these synapomorphies generic rank seems to

be more plausible and satisfactory.

No explicit synapomorphy characterizes Basiceros. ‘The good

score of the pretergite of the first gastral segment sessile (as opposed

to pedunculate) is due essentially to our coding of it. See our Fig.

27 as an example in which this character can not be detected.

Other useful but also not universally distributed characters are:

- Antennal fossa separate from the antennal scrobe. Exceptions

Basiceros (= Eurhopalothrix = Rhopalothrix) bruchi and helisca-

tum. This trait is present also in other Dacetini outside the clade

(species of Strumigenys and Epopostruma).

ANT TRIBE DACETINI 89

- First gastral segment densely punctuated. Exceptions Basiceros

(= Octostruma) balzani and onoret. ‘Vhis trait is present also in

some Strumigenys.

We are aware that even after downgrading the tribe Basicerotini

to genus level, the ensuing genus Basiceros results more ill-defined

than most other ant genera, but we regard the present proposal as

the one coupling the minimum nomenclatorial change with the max-

imum practical utility.

All the new genus-level synonymies above flow directly either

from the total lack of synapomorphies or from the implausibility of the

known ones as they result in our Fig. 34. These “genera”, however,

are present in a number of practical keys where they are separate on

the basis of the number of antennal joints. By being a meristic char-

acter, transitional forms in antennomere counts cannot be expected

and this makes such counts excellent practical characters. Their

poor phylogenetic value was already stressed explicitly or implic-

itly by BOLTON (1999), DieTz (2004) and ourselves (present paper)

by coding their 8-steps variability in a reduced binary form only.

Further evidence for the generic synonymies proposed above

results from the following critical summary of all known but insuf-

ficient synapomorphies on which these generic names were based:

Rhopalothrix: loss of the anal vein. CI 0.75, RI 0.50. Besides

the reduced number of known wing venations in the whole tribe,

the anal vein is missing also in some Eurhopalothrix, Strumigenys,

and Acanthognathus.

Octostruma: no synapomorphies for this genus result from our

data.

Talaridris: reduction of the number of labial palps from two to

one CI 0.70, RI 0.67. One-jointed labial palps are known, among

others, also in Strumigenys, Acanthoghathus, Basiceros, Eurhopalothrix

and Octostruma. 'The apomorphic condition attributed to Talaridris

is a consequence of the fact that one-jointed palps are a secondary

reduction from 2 jointed, the state assigned to the ancestor of the

whole clade.

Eurhopalothrix: loss of the pterostigma. CI 1.00, RI 0.00. This

is a potentially excellent character discovered by DIETZ (2004). Note

that DIETZ (I. c.) was able to study the males of 7 Eurhopalothrix

90 C. BARONI URBANI & M.L. DE ANDRADE

out of a total of 39 known species and that he examined gynes of 29

species, although it is not specified if these were all winged or not.

DiETz (2004: 34) adds also presence of a second row of mandibu-

lar teeth [among gynes] as apomorphic for Eurhopalothrix but the

gyne of Basiceros papuanus n. sp. (a species with 7 antennal joints

- as Eurhopalothrix should be - to be described later in the pres-

ent paper) has no trace of the second row of teeth. Assuming that

Eurhopalothrix could be separate from the other genera discussed

here on the basis of the lack of pterostigma, Basiceros would result

paraphyletic to it.

Protalaridris: metapleural gland bulla close to the annulus. CI

0.62, RI 0.40. This same character state re-appears in some Eurho-

palothrix, Octostruma and in a number of other, less related Dacetini.

The generic synonymies above imply the following new com-

binations:

Note: The name Basiceros is composed of the two Greek words “Bào1g”

(=basis, foot), feminine, and ”képag” (=horn), neuter. It is hence a neuter noun and

established species names like singularis or militaris must be changed to singulare

and militare.

Basiceros acutipilis (Kempf). New combination for Rhopalothrix acutipilis

Kemper, 1962: 28.

Basiceros allopeciosum (Brown & Kempf). New combination for Eurhopalothrix

allopeciosa BROWN & KEMPF, 1960: 206.

Basiceros apharogonium (Snelling). New combination for £Eurhopalothrix

apharogonia SNELLING, 1968: 1.

Basiceros armatum (Brown). New combination for Protalaridris armata

Brown, 1980: 37.

Basiceros australe (Brown & Kempf). New combination for Eurhopalothrix

australis BROWN & KEMPF, 1960: 218.

Basiceros balzani (Emery). New combination for Rhopalothrix balzani EMERY,

1894: 217 and Octostruma balzani (Emery), BROWN, 1949c: 92.

Basiceros batesi (Emery). New combination for Rhopalothrix batest EMERY,

1894: 218 and Octostruma batest (Emery), BRowN, 1949c: 92.

Basiceros betschi (Perrrault).. New combination for Octostruma betschi

PERRAULT, 1988: 303.

Basiceros biroi (Szabo). New combination for Rhopalothrix biroi SZABÒ, 1910:

365 and Eurhopalothrix birot (Szabo), BROWN & KEMPF, 1960: 222.

Basiceros bolaut (Mayr). New combination for Rhopalothnx bolaui MAYR,

1870: 415 and Eurhopalothrix bolaui (Mayr), Brown & KEMPF, 1960: 210.

Basiceros brevicorne (Emery). New combination for Rhopalothrix brevicornis

EMmERY, 1897: 572 and Eurhopalothrix brevicornis (Emery), BROWN & KEMPF, 1960: 215.

Basiceros browni (Taylor). New combination for Eurhopalothrix browni

TAYLOR, 1990: 404.

ANT TRIBE DACETINI 91

Basiceros bruchi (Santschi). New combination for Rhopalothrix bruchi SANTSCHI,

1922: 256 and Eurhopalothrix bruchi (Santschi), Brown & KEMPF, 1960: 214.

Basiceros caledonicum (Brown & Kempf). New combination for Eurhopalothrix

caledonica BROWN & KEMPF, 1960: 220.

Basiceros chapmani (Taylor). New combination for Eurhopalothrix chapmani

‘TayLor, 1990: 406.

Basiceros ciliatum (Mayr). New combination for Rhopalothrix ciliata MAYR,

1870: 415.

Basiceros cinnameum (Taylor). New combination for Eurhopalothrix cinnamea

TTANILOR, 1970: 350;

Basiceros clypeatum (Brown & Kempf). New combination for Eurhopalothrix

clypeata BROWN & KEMPF, 1960: 205.

Basiceros coronatum (‘Vaylor). New combination for Eurhopalothrix coronata

TaAyLOR, 1990: 407.

Basiceros depressum (Ketterl et al.). New combination for Eurhopalothrix

depressa KETTERL et al., 2004: 45.

Basiceros diadema (Brown & Kempf). New combination for Rhopalothrix

diadema BROWN & KEMPF, 1960: 239.

Basiceros dubium (Taylor). New combination for Eurhopalothrix dubia

‘TayLor, 1990: 409.

Basiceros emeryi (Forel). New combination for Rhopalothrix emeryi FOREL,

1912a: 58 and Eurhopalothrix emeryi (Forel), BRowN & Kempr, 1960: 230.

Basiceros floridanum (Brown & Kempf). New combination for Eurhopalothrix

floridana Brown & KEMPF, 1960: 207.

Basiceros grave (Mann). New combination for Rhopalothrix gravis MANN,

1922: 40 and Eurhopalothrix gravis (Mann), BROWN & KEMPF, 1960: 211.

Basiceros greensladet (Taylor). New combination for Eurhopalothrix greensladei

TAYLOR, 1968a: 342.

Basiceros heliscatum (Wilson & Brown). New combination for Eurhopalothrix

heliscata WILSON & Brown, 1985: 410.

Basiceros hoplites (Taylor). New combination for Eurkhopalothrix hoplites

‘TAYLOR, 1980b: 231.

Basiceros theringi (Emery). New combination for Rhopalothrix Iheringi EMERY,

1888: 361 and Octostruma theringi (Emery), BROWN, 1949c: 92.

Basiceros impressum (Palacio). New combination for Octostruma impressa

PaLacio, 1997: 411.

Basiceros inca (Brown & Kempf). New combination for Octostruma inca

BROWN & KEMPF, 1960: 185.

Basiceros insidiator (Taylor). New combination for Eurhopalothrix insidiatrix

TAYLOR, 1980b: 238.

Basiceros isabellae (Mann). New combination for Rhopalothrix isabellae MANN,

1919: 357 and Eurhopalothrix isabellae (Mann), Brown & Kempr, 1960: 225.

Basiceros isthmicum (Weber). New combination for Acanthidris isthmicus

WEBER, 1941: 188.

Basiceros jennya (Taylor). New combination for £Eurhopalothrix jennya

TAYLOR, 1990: 413.

92 C. BARONI URBANI & M.L. DE ANDRADE

Basiceros kusnezovi (Brown & Kempf). New combination for Rhopalothrix

kusnezovi BROWN & KEMPF, 1960: 238.

Basiceros lenkoi (Kempf). New combination for Eurhopalothrix lenkoi KEMPF,

1967; 358:

Basiceros omnivagum (Taylor). New combination for Eurhopalothrix omnivaga

TAYLOR, 1990: 413.

Basiceros orbis (Taylor). New combination for Rhopalothrix orbis ‘TAYLOR,

1968a: 336.

Basiceros mandibulare (Weber). New combination for Talaridris mandibularis

WEBER, 1941: 185.

Basiceros petiolatum (Mayr). New combination for Rhopalothrix petiolata

Mayr, 1887: 580 and Octostruma petiolata (Mayr), BRown, 1949c: 92.

Basiceros philippinum (Brown & Kempf). New combination for Eurhopalothrix

philippina BROWN & Kemper, 1960: 224.

Basiceros piluliferum (Brown & Kempf). New combination for Eurhopalothrix

pilulifera BROWN & KEMPF, 1960: 208.

Basiceros platisgquama (Taylor), New combination for Eurhopalothrix

platisquama TAYLOR, 1990: 417.

Basiceros plaumanni (Brown & Kempf). New combination for Rhopalothrix

plaumanni BROWN & KEMPF, 1960: 235.

Basiceros procerum (Emery). New combination for Rhopalothrix procera

Emery, 1897: 572 and Eurhopalothrix procera (Emery), BROWN & KEMPF, 1960: 225.

Basiceros punctatum (Szabo). New combination for Rhopalothrix punctata

SzaBo, 1910: 366 and Eurhopalothrix punctata (Szabo), Brown & KEMPF, 1960: 221.

Basiceros rothschildi (Taylor). New combination for Eurhopalothrix rothschildi

TayLoR, 1990: 418.

Basiceros rugiferum (Mayr). New combination for Rhopalothrix rugifer MAYR,

1887: 579 and Octostruma rugifera (Mayr), Brown, 1949c: 92.

Basiceros rugiferoide (Brown & Kempf). New combination for Octostruma

rugiferoides BROWN & KEMPF, 1960: 200.

Basiceros seguense (Taylor). New combination for Eurhopalothrix seguensis

TAYLOR, 1990: 421.

Basiceros simoni (Emery). New combination for Rhopalothrix simoni EMERY,

1890: 67 and Octostruma simoni (Emery), Brown, 1949c: 92. Junior synonym of

Octostruma iheringi Emery, BROWN & KEMPF, 1960: 187. Octostruma simoni (Emery),

Dietz, 2004: 127.

Basiceros speciosum (Brown & Kempf). New combination for Eurhopalothrix

speciosa BROWN & KEMPF, 1960: 203.

Basiceros spectabile (Kempf). New combination for Eurhopalothrix spectabilis

KEMPF, 1962: 27.

Basiceros stannardi (Brown & Kempf). New combination for Rhopalothrix

stannardi BROWN & KEMPF, 1960: 236.

Basiceros stenognathum (Brown & Kempf). New combination for Octostruma

stenognatha BROWN & KEMPF, 1960: 196.

Basiceros stenoscapum (Palacio). New combination for Octostruma stenoscapa

PaLacio, 1997: 414.

Basiceros szentivanyi (Taylor). New combination for Eurhopalothrix szentivanyi

TAYLOR, 1968a: 346.

ANT TRIBE DACETINI 93

Basiceros weberi (Brown & Kempf). New combination for Rhopalothrix weber

Brown & KEMPF, 1960: 234.

Basiceros wheeleri (Mann). New combination for Rhopalothrix wheeleri MANN,

1922: 43. Octostruma wheeleri (Mann), Brown, 1949c: 92.

The known species of Basiceros have been recently keyed and

redescribed by Dietz (2004) under the name of their new generic

synonyms proposed above.

Daceton Perty

Daceton PERTY, 1833: 136. Type species Formica armigera Latreille, by monotypy.

Autapomorphies for this monotypic genus included in our data

matrix are the following:

Worker (and gyne) foramen occipitale dorsal, CI 1.00, RI 0.00.

Worker (and gyne) scape straight at base. CI 0.50, RI 0.33. The

straight condition of Daceton is shared with Acanthognathus.

Worker (and gyne) orifice of the metapleural gland visible. CI

0.33, RI 0.50. The visible condition of Daceton appears to be unique

among the Dacetini. The low CI and RI values are due to variation

among the outgroups.

Worker (and gyne) propodeal spiracle median (instead of poste-

rior). CI 0.33, RI 0.60. The median location of Daceton appears to

be unique among the Dacetini. The low CI and RI values are due

to variation among the outgroups.

Worker (and gyne) antennae 11-jointed. CI 0.20, RI 0.50 for

the 11-12 jointed state.

The above apomorphy list is largely sufficient to regard Daceton

as a separate genus.

The genus contains only one known species, D. armigerum

(Latreille).

Epopostruma Forel

Strumigenys subg. Epopostruma ForeL, 1895: 422. Type species Strumigenys

(Epopostruma) quadrispinosa Forel, designated by WHEELER, 1911b: 163.

Epopostruma Forel, EMmERY, 1897: 573. Raised to genus.

94 C. BARONI URBANI & M.L. DE ANDRADE

Hexadaceton Brown, 1948: 120. Type species Hexadaceton frosti Brown, by original

designation. Synonymy by Brown, 1973b.

Colobostruma WHEELER, 1927: 32 (subgenus ad Epopostruma). Type species

Epopostruma leae Wheeler by monotypy. New synonymy.

Colobostruma Brown, 1948: 118. Raised to genus.

Alistruma Brown, 1948: 117. Type species Epopostruma foliacea Emery, by original

designation. Synonymy with Colobostruma by Brown, 1959. New synonymy.

Clarkistruma Brown, 1948: 124. Type species Epopostruma alinodis Forel, by original

designation. Synonymy with Colobostruma by Brown, 1959. New synonymy.

Mesostruma Brown, 1948: 18. Type species Strumigenys turneri Forel, by original

designation. Synonymy with Colobostruma by BARONI URBANI & DE ANDRADE,

1994: 15; revived from synonymy BOLTON, 1999: 1680. New synonymy.

Our search was unable to point out at synapomorphies for this

genus in its narrow sense, as it was understood in the most recent

papers. Neither synapomorphies described in the literature nor

emerged in this study as a result of character optimization result

from our search. BOLTON (1999: 1681) gives the following two

~ 0005060 | —— 100 ym ui Beer

Fig. 37 - Epopostruma quadrispinosa (Forel), worker without preocular groove. Pres-

ence of the groove is given as synapomorphic for Epopostruma in BOLTON

(1999).

ANT TRIBE DACETINI 95

generic synapomorphies: presence of the labral concavity and of a

vertical preocular groove. We previously pointed out that the labral

concavity was given by BoLTON (1999:1681) as synapomorphic for

Epopostruma and by BOLTON (1998:72) for all the Dacetini (see the

introduction). Moreover, the labral concavity of some Epopostruma

is very similar to the one of Mesostruma (Fig. 30). On the other

hand, the Epopostruma’s vertical preocular groove is well visible

only in some Epopostruma species illustrated by Shattuck’s photo-

graphs (in BOLTON, 2000) but much less or not at all in others (see

also our Fig. 37).

Without clear synapomorphies distinguishing it, Epopostruma 1s

destined to remain in the literature as the oldest available generic

name of a small clade with Colobostruma and Mesostruma as junior

synonyms as already suggested by BARONI URBANI & DE ANDRADE

(1994).

According to BOLTON (1999: 1680) Mesostruma (with 8 Austra-

lian species) should differ from Colobostruma (with 16 Australian

species) for only one synapomorphy: “Mandibles elongate triangu-

lar, with a larger apical and smaller preapical tooth; proximal of

this the margin is edentate and lamellate”. Elongate triangular man-

dibles are present also in Colobostruma cerornata Brown (BOLTON,

2000, page 39 [description] and Fig. 38 [figure]) and in C. froggatti

(Forel) (Fig. 38). The dentition of the mandibles is identical at least

in Colobostruma sisypha Shattuck (SHATTUCK in BOLTON (2000, Fig.

39)) and Mesostruma eccentrica Taylor (SHATTUCK in BOLTON (2000,

Fig. 52)). (BOLTON (1999: 1679) lists five synapomorphies for Colo-

bostruma as a genus different from Mesostruma. ‘These, of course,

loose their meaning after demonstrating the paraphyly of the second

versus the first. Both generic names together (i.e. Colobostruma

and Mesostruma) are characterized by an excellent synapomorphy

(hypertrophied labium) but are in their turn paraphyletic to the

oldest available name for the whole clade: Epopostruma for which

there are no known convincing synapomorphies.

The following four synapomorphies resulting from our data for

the broader Epopostruma (i.e. including also Colobostruma and Meso-

struma) are all the product of character optimization; three of them

are character-states found also in an ancestor and hence representing

an evolutionary reversal:

96 C. BARONI URBANI & M.L. DE ANDRADE

00050612 | — 100 um I z MB

Uni Basel

Fig. 38 - Colobostruma froggatti (Forel) showing elongate mandibles, a supposed

apomorphy for the genus Mesostruma according to BOLTON (1999).

Worker (and gyne) antennal scrobe below the eyes. CI 0.71,

RI 0.83. This trait, according to our phylogenetic reconstruction,

should have been secondarily lost in some Epopostruma species actu-

ally included in Colobostruma and re-appears homoplastically among

all basicerotine genera. We consider this pretended secondary loss as

plausibly due to head flattening in some species and as a result the

most significant synapomorphy for the genus.

Worker (and gyne) labrum capable of full reflexion. CI 0.50, RI

0.87. AIl remainder dacetine genera have a labium incapable of full

reflexion. The low CI and RI values are due to variability among

the outgroups. The validity of this character is further weakened

by the fact that the capability of full reflexion is widespread among

ants and its presumed apomorphic value, in this case, would require

a tertiary re-gain of this property.

Worker (and gyne) labrum not T-shaped. CI 0.50, RI 0.33.

ANT TRIBE DACETINI 97

This is a widespread dacetine character absent among three imme-

diate Epopostruma outgroup genera.

Worker (and gyne) 2-segmented antennal club. CI 0.33, RI

0.71. AIl remainder dacetine genera have a 2-segmented antennal

club. The low CI and RI values are due to variability among the

outgroups.

The species previously included in Epopostruma, Colobostruma,

and Mesostruma are keyed and described under these three generic

names by SHATTUCK (in BOLTON, 2000).

Our synonymies imply the following new or reaffirmed generic

transfers:

Epopostruma alinodis ForEL, 1913. Clarkistruma alinodis (Forel), BRown, 1948: 124.

Colobostruma alinodis (Forel) Brown & WILSON, 1959: 281. Combination in

Epopostruma reinstated.

Epopostruma australis (Brown). New combination for Colobostruma australis

Brown, 1959: 4.

Epopostruma bella (Shattuck). New combination for Mesostruma bella SHATTUCK

in BOLTON, 2000: 48.

Epopostruma biconcava (Shattuck). New combination for Colobostruma biconcava

SHATTUCK in BoLTON, 2000: 35.

Epopostruma biconvexa (Shattuck). New combination for Colobostruma biconvexa

SHATTUCK int BOLTON, 2000; 35,

Epopostruma bicornis (Shattuck). New combination for Colobostruma bicorna (sic)

SHATTUCK in BOLTON, 2000: 45. Note: the Latin word cornu is a neuter noun

and cannot be declined in the feminine form. The feminine adjective for two-

horned is bicornis (Horatius, Vergilius).

Epopostruma browni (Taylor). New combination for Mesostruma browni ‘TAYLOR,

1962: 1.

Epopostruma cerornata (Brown). New combination for Colobostruma cerornata

Brown, 1959: 1.

Epopostruma eccentrica (Taylor). New combination for Mesostruma eccentrica

TaAvcor, 1973 31.

Epopostruma elliotti (Clark). New combination for Epitritus elliotti CLARK, 1928:

42, and Clarkistruma elliotti (Clark), BROWN, 1948: 124, and Colobostruma elliotti

(Clark), TayLor & Brown, 1985: 60.

Epopostruma foliacea Emery, 1897. Alistruma foliacea (Emery), BRown, 1948:

117. Colobostruma foliacea (Emery) BoLTon, 1995: 146. Combination in

Epopostruma reinstated.

Epopostruma froggatti Forel, 1913. Alistruma froggatti (Forel), BRown, 1948: 117.

Colobostruma froggatti (Forel) TayLor & Brown, 1985: 60. Combination in

Epopostruma reinstated.

Epopostruma inornata (Shattuck). New combination for Mesostruma inornata

SHATTUCK in BOLTON, 2000: 51.

98 C. BARONI URBANI & M.L. DE ANDRADE

Epopostruma lacuna (Shattuck). New combination for Colobostruma lacuna

SHATTUCK in BoLTON, 2000: 36.

Epopostruma laevigata (Brown). New combination for Mesostruma laevigata

Brown, 1952: 12

Epopostruma leae Wheeler. Colobostruma leae (Wheeler), Brown, 1948: 118.

Combination in Epopostruma reinstated.

Epopostruma loweryi (Taylor). New combination for Mesostruma loweryi "TAYLOR,

IC TRRTOSE

Epopostruma mellea (Shattuck). New combination for Colobostruma mellea

SHATTUCK in BoLTON, 2000: 37.

Epopostruma nancyae (Brown). New combination for Colobostruma nancyae BROWN,

1963b: 22;

Epopostruma papulata (Brown). New combination for Colobostruma papulata

Brown, 1965a: 21.

Epopostruma sisypha (Shattuck). New combination for Colobostruma sisypha

SHATTUCK in BOLTON, 2000: 37.

Epopostruma turneri (Forel). Strumigenys (Epopostruma) turnert FoREL, 1895: 424.

Epopostruma turneri (Forel), EMmERY, 1924: 330. Mesostruma turneri (Forel),

Brown, 1948: 119. Combination in Epopostruma reinstated.

Epopostruma unicornis (Shattuck). New combination for Colobostruma unicorna

(sic) SHATTUCK in BOLTON, 2000: 46. Note: the Latin word cornu is a neuter

noun and cannot be declined in the feminine form. The feminine adjective for

single horned is unicornis (Plinius, Tertullianus).

Ishakidris Bolton

Ishakidris BOLTON, 1984: 374. Type species Ishakidris ascitaspis Bolton, by original

designation.

The two genus-level apomorphies resulting for this genus are:

Worker (and gyne?) with visible katepisternal groove (CI 1.00,

RI 0.00).

Worker (and gyne?) with first gastral sternum truncated at base

(CI 0.67, RI 0.71). This latter character, already considered as syn-

apomorphic for Basiceros, is widespread in Strumigenys and irregu-

larly present also among other outgroups.

‘The pretended generic apomorphies above are likely to be a by-

product of the clustering together of the three “Phalacromyrmecini”

genera, a clustering that we already showed as being based on weak

characters (see the discussion under the synonymy of the tribe Phal-

acromyrmecini). We suspect that all the former Phalacromyrmecini

genera might be better understood as atypical Strumigenys species.

ANT TRIBE DACETINI 99

Microdaceton Santschi

Microdaceton SANTSCHI, 1913b: 478. Type species Microdaceton exornatum Santschi,

by monotypy.

The following characters result apomorphic for this genus:

Worker (and gyne) pronotal cervix with thick transverse rim. CI

1.00, RI 0.00.

Worker (and gyne) maxillary palps three-jointed. CI 0,85, RI

0.80. Among the Dacetini the three-jointed condition is known only

in Phalacromyrmex and Pilotrochus.

Worker (and gyne) labial palps two-jointed. CI 0.70, RI 0.67.

Among the Dacetini the two-jointed condition is known in Phala-

cromyrmex, Pilotrochus, Ishakidris and some Basiceros species.

Other plausible generic synapomorphies not included in our

data matrix are listed by BoLToN (1999: 1675).

The four Muicrodaceton species known so far are keyed and

described by BoLTON (2000)

Orectognathus Smith

Orectognathus SMITH, 1853: 227. Type species Orectognathus antennatus Smith, by

monotypy.

Only one, convincing, synapomorphy is known for this genus:

the hypertrophy of the second funicular joint of workers and gynes

(CI 1.00, RI 0.00).

The known species of this genus have been keyed and described

by TAYLOR (1980a).

Phalacromyrmex Kempf

Phalacromyrmex KEMPF, 1960a: 89. Type species Phalacromyrmex fugax Kempf, by

original designation.

Only one unequivocal synapomorphy for this genus results from

our data, the presence of mesopleural costulation. CI 1.00, RI 0.00.

100 C. BARONI URBANI & M.L. DE ANDRADE

We doubt, however the generic rank of this character. Other apo-

morphic characters resulting from our analysis are:

Worker (and gyne?) presence of a cuticular process of the

mesonotum. CI 0.86, RI 0.75. Cuticular projections of the mesono-

tum are known also in Epopostruma and Strumigenys.

Worker (and gyne?) metapleaural gland bulla close to the annu-

lus. CI 0.62, RI 0.40. The same character state is encountered in a

number of non-related Dacetini genera like Protalaridris, Microdace-

ton, Epopostruma, a. o.

Worker (and gyne?) antennae 11-jointed. CI 0.20, RI 0.50. A

character state shared with several outgroups and with Daceton and

Acanthognathus.

The pretended generic apomorphies above are likely to be a

by-product of the clustering together of the three “Phalacromyr-

mecini” genera, a clustering that we already showed as being based

on weak characters (see the discussion under the synonymy of the

tribe Phalacromyrmecini). We suspect that all the former Phala-

cromyrmecini genera might be better understood as atypical Stru-

migenys species.

Pilotrochus Brown

Pilotrochus Brown, 1978: 218. Type species Pilotrochus besmerus Brown, by original

designation.

The sole known apomorphy for this genus is the hypertrophic

development of the mesosternal hair beds, visible on the profile,

CI 1.00, RI 0.00. Mesosternal hair beds are widespread in ants.

Whether the hypertrophic development visible in Pilotrochus should

be considered as a genus-level apomorphy or not may be debated.

This doubt, together with the clustering of the three “Phalacromyr-

mecini” genera that we already showed as being based on doubt-

ful characters (see the discussion under the synonymy of the tribe

Phalacromyrmecini) weakens considerably the credibility of Pilotro-

chus as a valid monotypic genus. We suspect that all the former

Phalacromyrmecini genera might be better understood as atypical

Strumigenys species.

ANT TRIBE DACETINI 101

Strumigenys Smith

Strumigenys SMITH, 1860: 72. Type species Strumigenys mandibularis Smith, by

monotypy.

Labidogenys ROGER, 1862: 249. Type species Labidogenys lyroessa Roger by monotypy.

Labidogenys Roger as a synonym of Strumigenys, ROGER, 1863b: 40.

Pyramica RoGER, 1862: 251. Type species Pyramica gundlachi Roger by monotypy.

Pyramica Roger as a synonym of Strumigenys, ROGER, 1863b: 40. Pyramica Roger,

revived from synonymy, BOLTON, 1999: 1667. Synonymy with Strumigenys

reinstated.

Cephaloxys SMITH, 1865: 76. Type species Cephaloxys capitata Smith by monotypy.

Junior homonym of Cephaloxys Signoret, 1847 (Hemiptera). Replacement name

Smithistruma Brown, 1948: 104.

Epitritus EMERY 1869a: 136. Type species Epitritus argiolus Emery by monotypy.

Epitritus Emery as a synonym of Strumigenys, BARONI URBANI & DE ANDRADE,

1994: 32. Epitritus Emery revived from synonymy, BOLTON, 1995: 188. Epitritus

Emery as a synonym of Pyramica, BOLTON, 1999: 1667. Synonymy with

Strumigenys reinstated.

Trichoscapa EMERY, 1869b: 24 (subgenus ad Strumigenys). Type species Strumigenys

membranifera Emery by monotypy. Trichoscapa Emery as a synonym of

Strumigenys, DALLA ToRRE (1893: 145). Trichoscapa Emery as a synonym

of Cephaloxys, WHEELER (1922: 668). Trichoscapa Emery as a good genus,

Brown, 1948: 112. Trichoscapa as a synonym of Strumigenys, BaRONI URBANI

& DE ANDRADE, 1994: 32. Trichoscapa Emery revived from synonymy, BOLTON,

1995: 421. Trichoscapa Emery as a synonym of Pyramica, BOLTON, 1999: 1667.

Synonymy with Strumigenys reinstated.

Pentastruma FoREL, 1912a: 50. Type species Pentastruma sauteri Forel by monotypy.

Pentastruma Forel as a synonym of Strumigenys, BARONI URBANI & DE ANDRADE,

1994: 32. Pentastruma Forel revived from synonymy, BOLTON, 1995: 316.

Pentastruma Forel as a synonym of Pyramica, BOLTON, 1999: 1667. Synonymy

with Strumigenys reinstated.

Glamyromyrmex WHEELER, 1915b: 487. Type species Glamyromyrmex beebei Wheeler

by monotypy. Glamyromyrmex Wheeler as a synonym of Strumigenys, BARONI

URBANI & DE ANDRADE, 1994: 32. Glamyromyrmex Wheeler revived from

synonymy, BOLTON, 1995: 207. Glamyromyrmex Wheeler as a synonym of

Pyramica, BOLTON, 1999: 1667. Synonymy with Strumigenys reinstated.

Codiomyrmex WHEELER, 1916: 326. Type species Codiomyrmex thaxteri Wheeler by

monotypy. Codiomyrmex Wheeler as a synonym of Strumigenys, BARONI URBANI

& DE ANDRADE, 1994: 32. Codiomyrmex Wheeler revived from synonymy,

BoLTON, 1995: 146. Codiomyrmex Wheeler as a synonym of Pyramica, BOLTON,

1999: 1668. Synonymy with Strumigenys reinstated.

Tingimyrmex MANN, 1926: 104 (subgenus ad Strumigenys). Type species Strumigenys

mirabilis Mann by monotypy. Tingimyrmex Mann as a good genus, BROWN,

1948: 111. Tingimyrmex Mann as a synonym of Strumigenys, BARONI URBANI &

DE ANDRADE, 1994: 32. Tingimyrmex Mann revived from synonymy, BOLTON,

1995: 420. Tingimyrmex Mann as a synonym of Pyramica, BOLTON, 1999: 1668.

Synonymy with Strumigenys reinstated.

Codioxenus SANTSCHI, 1931: 278 (subgenus ad Epitritus). 'Type species Epztritus

simulans Santschi by monotypy. Codioxenus Santschi as a good genus, BROWN,

1948: 123. Codioxenus Santschi as a synonym of Strumigenys, BARONI URBANI &

102 C. BARONI URBANI & M.L. DE ANDRADE

DE ANDRADE, 1994: 32. Codioxenus Santschi revived from synonymy, BOLTON,

1995: 146. Codioxenus Santschi as a synonym of Pyramica, BOLTON, 1999: 1668.

Synonymy with Strumigenys reinstated.

Proscopomyrmex PATRIZI, 1946: 294. Type species Proscopomyrmex londianensis

Patrizi by monotypy. Proscopomyrmex Patrizi as a synonym of Strumigenys,

Brown, 1949b: 15.

Eneria DONISTHORPE, 1948: 598. Type species Eneria excisa Donisthorpe by original

designation. Eneria Donisthorpe as a synonym of Strumigenys, BROWN, 1949b:

{58

Smithistruma Brown, 1948: 104. Type species Cephaloxys capitata Smith. Nomen

novum pro Cephaloxys Smith 1865 nec Cephaloxys Signoret 1847. Smithistruma

Brown as a synonym of Strumigenys, BARONI URBANI & DE ANDRADE, 1994: 32.

Smithistruma Brown revived from synonymy, BOLTON, 1995: 383. Smuithistruma

Brown as a synonym of Pyramica, BoLTon, 1999: 1668. Synonymy with

Strumigenys reinstated.

Weberistruma Brown, 1948: 106 (subgenus ad Smithistruma). Type species

Strumigenys leptothrix Wheeler by original designation. Raised to genus by

Brown, 1949b. Weberistruma Brown as a synonym of Smithistruma, BROWN,

1973a: 35. Weberistruma as a synonym of Pyramica, BOLTON, 1999: 1668.

Undoubted synonym of a synonym of Strumigenys supported in this paper. New

synonymy.

Wessonistruma Brown, 1948: 106 (subgenus ad Smithistruma). Type species

Strumigenys pergandei Emery by original designation. Wessonistruma Brown as

a synonym of Smithistruma, BROWN, 1973a: 35. Wessonistruma as a synonym

of Pyramica, BOLTON, 1999: 1668. Undoubted synonym of a synonym of

Strumigenys supported in this paper. New synonymy

Serrastruma Brown, 1948: 107 (subgenus ad Smithistruma). Type species Strumigenys

simoni Emery by original designation. Serrastruma Brown as a good genus,

Brown, 1949b: 6. Serrastruma Brown as a synonym of Strumigenys, BARONI

URBANI & DE ANDRADE, 1994: 32. Serrastruma Brown revived from synonymy,

BoLTOoN, 1995: 382. Serrastruma Brown as a synonym of Pyramica, BOLTON,

1999: 1668. Synonymy with Strumigenys reinstated.

Neostruma Brown, 1948: 111. Type species Strumigenys crassicornis Mayr by original

designation. Neostruma Brown as a synonym of Strumigenys, BARONI URBANI

& DE ANDRADE, 1994: 32. Neostruma Brown revived from synonymy, BOLTON,

1995: 292. Neostruma Brown as a synonym of Pyramica, BOLTON, 1999: 1668.

Synonymy with Strumigenys reinstated.

Dorisidris BRowN, 1948: 116. Type species Strumigenys nitens Santschi by original

designation. Dorisidris Brown as a synonym of Strumigenys, BARONI URBANI

& DE ANDRADE, 1994: 32. Dorisidris Brown revived from synonymy, BOLTON,

1995: 177. Dorisidris Brown as a synonym of Pyramica, BOLTON, 1999: 1668.

Synonymy with Strumigenys reinstated.

Miccostruma Brown, 1948: 116. Type species Epitritus mandibularis Szabo by

original designation. Miccostruma Brown as a synonym of Smithistruma, BOLTON,

1983: 274. Miccostruma Brown as a synonym of Pyramica, BOLTON, 1999: 1668.

Undoubted synonym of a synonym of Strumigenys supported in this paper. New

synonymy.

Quadristruma BROWN, 1949a: 47. Type species Epitritus emmae Emery by original

designation. Quadriustruma Brown as a synonym of Strumigenys, BARONI URBANI

ANT TRIBE DACETINI 103

& DE ANDRADE, 1994: 32. Quadristruma Brown revived from synonymy, BOLTON,

1995: 377. Quadristruma Brown synonymy with Strumigenys reinstated, BOLTON,

1999: 1672.

Kyidris Brown, 1949b: 3. Type species Kyidris mutica Brown by original designation.

Kyidris Brown as a synonym of Strumigenys, BARONI URBANI & DE ANDRADE,

1994: 32. Kyidris Brown revived from synonymy, BOLTON, 1995: 219. Kyzdris

Brown as a synonym of Pyramica, BOLTON, 1999: 1668. Synonymy with

Strumigenys reinstated.

Polyhomoa Azuma, 1950: 36. Type species Polyhomoa itoi Azuma by monotypy.

Polyhomoa Azuma as a synonym of Kyidris, CREIGHTON, 1950: 93. Polyhomoa

Azuma as a synonym of Pyramica, BOLTON, 1999: 1668. Undoubted synonym of

a synonym of Strumigenys supported in this paper. New synonymy.

Chelystruma Brown, 1950a: 33 (subgenus ad Glamyromyrmex). Type species

Glamyromyrmex lilloana Brown by monotypy. Chelystruma Brown as a good

genus, Kempr, 1959: 338. Chelystruma as a synonym of Strumigenys, BARONI

URBANI & DE ANDRADE, 1994: 32. Chelystruma Brown revived from synonymy,

Botton, 1995: 145. Chelystruma Brown as a synonym of Pyramica, BOLTON,

1999: 1668. Synonymy with Strumigenys reinstated.

Borgmeierita BROWN, 1953: 23. Type species Codiomyrmex excisus Weber by original

designation. Borgmeierita Brown as a synonym of Glamyromyrmex, BROWN,

1973a: 35. Borgmeierita Brown as a synonym of Pyramica, BOLTON, 1999: 1668.

Undoubted synonym of a synonym of Strumigenys supported in this paper. New

synonymy.

Platystruma Brown, 1953: 112 (subgenus ad Smithistruma). Type species Strumigenys

depressiceps Weber by original designation. Platystruma Brown as a synonym of

Smithistruma, BRowNn, 1973a: 35. Platystruma Brown as a synonym of Pyramica,

BoLton, 1999: 1668. Undoubted synonym of a synonym of Strumigenys

supported in this paper. New synonymy.

Gymnomyrmex BORGMEIER, 1954: 279. Type species Gymnomyrmex splendens

Borgmeier by original designation. Gymnomyrmex Borgmeier as a synonym of

Strumigenys, BARONI URBANI & DE ANDRADE, 1994: 32. Gymnomyrmex Borgmeier

revived from synonymy, BOLTON, 1995: 211. Gymnomyrmex Borgmeier as a

synonym of Pyramica, BOLTON, 1999: 1668. Synonymy with Strumigenys

reinstated.

Dysedrognathus TAyLor, 1968b: 132. Type species Dysedrognathus extemenus Taylor

by original designation. Dysedrognathus Taylor as a synonym of Strumigenys,

BARONI URBANI & DE ANDRADE, 1994: 32. Dysedrognathus Taylor revived from

synonymy, BoLTon, 1995: 183. Dysedrognathus Taylor as a synonym of Pyramica,

BoLTON, 1999: 1668. Synonymy with Strumigenys reinstated.

Asketogenys Brown, 1972: 23. Type species Asketogenys acubecca Brown, by original

designation. Asketogenys Brown as a synonym of Strumigenys, BARONI URBANI

& DE ANDRADE, 1994: 32. Asketogenys Brown revived from synonymy, BOLTON,

1995: 75. Asketogenys Brown as a synonym of Pyramica, BOLTON, 1999: 1668.

Synonymy with Strumigenys reinstated.

Cladarogenys Brown, 1976: 33. Type species Cladarogenys lasia Brown, by original

designation. Cladarogenys Brown as a synonym of Strumigenys, BARONI URBANI

& DE ANDRADE, 1994: 32. Cladarogenys Brown revived from synonymy, BOLTON,

1995: 145. Cladarogenys Brown as a synonym of Pyramica, BoLTON, 1999: 1668.

Synonymy with Strumigenys reinstated.

104 C. BARONI URBANI & M.L. DE ANDRADE

In our study this genus results characterized by the following

synapomorphies:

Worker (and gyne) reduction of the labial palps from 2 to 1.

CI 0.70, RI 0.67. This trait is shared also with Acanthognathus and

with some Basiceros species previously included in Talaridris.

Worker (and gyne) presence of a basimandibular process. CI

0.67, RI 0.86. The synapomorphic state of this character (sometimes

of doubtful detection, see Fig. 10) is an artefact of character opti-

mization due to the fact that the basimandibular process is absent

in Phalacromyrmex, Pilotrochus, Ishakidris, Basiceros, and the hypo-

thetical ancestor of Acanthognathus (process present in highly trans-

formed form) and the remaining Dacetini genera (process present).

Because of its widespread presence in a number of Dacetini genera,

presence of the basimandibular process is a poor discriminant trait

to identify Strumigenys.

Worker (and gyne) mesosternal hair beds visible in profile. CI

0.91, RI 0.67. In spite of its perfect match within Strumigenys, this

character is irregularly distributed among several dacetine genera

including the three monotypic genera Phalacromyrmex, Pilotrochus

and Ishakidris. Moreover, we regard the character itself as insecure:

mesosternal hair beds are widespread and coding their visibility in

profile as phylogenetically significant needs some imagination and

abstraction capacity.

Worker (and gyne) eyes ventral. CI 0.67, RI 0.80. This trait

is shared with all and only the former Phalacromyrmecini, but was

coded as polymorphic in “Pyramica” as a result of our transfer to

Strumigenys of the blind species inopinata (see below, the list of new

combinations in Strumigenys) originally described in “Rhopalothrix”.

The position of the eyes, otherwise, is likely to represent the main

synypomorphy of the genus.

Gyne (and male?) secondary loss of the anal vein. CI 0.75, RI

0.50. This. trait appears to be constant among Strumigenys but it

reappears in some Acanthognathus and Basiceros species.

Strumigenys (including Pyramica) results weakly defined from

our analysis, unable to focus on one single clear-cut synapomor-

phy. On the other hand, by appearing as branch in a tetratomy, its

sister-group with which it could eventually be merged is also not

clearly identified. For these reasons and for nomenclatorial conser-

ANT TRIBE DACETINI 105

vationism we prefer to provisionally maintain Strumigenys as a valid

genus defined by a combination of characters, a procedure seldom

accepted in phylogenetic studies.

Finally, no apomorphies separating Pyramica from Strumige-

nys appeared as a result of our character optimization and we were

unable to include a single one in our data.

The reason for this exclusion is that no one of the characters

listed by BOLTON (1999, 2000) to separate the two genera appear to

hold even after a superficial scrutiny.

The following four characters should separate the two genera

according to BOLTON (1999, 2000):

1) “Mandibles in ventral view broad at extreme base, their artic-

ulations located at about the midlength of the labio-maxillary

complex”. In BOLTON (1999) this is given as apomorphic for

Pyramica as opposed to “Mandibles in ventral view narrow at

extreme base, apparently arising from the apex of the labio-

maxillary complex” in Strumigenys. This character is not used

to separate the two genera in the key by BOLTON (2000: 15). We

are greatly embarrassed in tracing a boundary between the two

categories defined by BOLTON (l. c.) and find it difficult under-

standing the character definition even on the basis of some of

Bolton’s: examples (seee. e. BOLTON 19997 Figs. 359 & 31). Our

Figs. 39 - 41 exemplify cases of comparable mandibular mor-

phology for both Pyramica and Strumigenys.

2) “Mandibles at full gap open to only 60-90°” in Pyramica, vs.

“Mandibles at full gape open to 170° or more” in Strumigenys.

Documenting this character may be difficult for the risks to tear

the adductor muscles, an action that would result in too great

opening angles and for the tendency to contract the muscle bun-

dles yielding smaller opening angles in drying specimens. Fig.

42 records two examples drawn from a wider array of species

contradicting BOLTON’s (1999 & 2000) use of this character and

observed on relaxed material. Another case was already docu-

mented in a web publication by de ANDRADE & BARONI URBANI

(2005)

3) “Labral lobes hypertrophied” (synapomorphic) versus “arising

from across entire width of labium” (plesiomorphic) in Pyra-

mica; or “labrum with distal lobes reduced... or vestigial” (ple-

106 C. BARONI URBANI & M.L. DE ANDRADE

ZMB

Uni Basel

00047453 en Neen

Fig. 39 - Comparable width and position of the base of the mandibles in Pyramica

semicompta (Brown) (top) and Strumigenys lyroessa (Roger) (bottom). The

arrows show the position of the mandibular articulations and of the labial

palpi as indicator of the distal border of the labio-maxillary complex.

ANT TRIBE DACETINI LO

00047445 uf betel

00047456 Uni Basel

Fig. 40 - Comparable width and position of the base of the mandibles in Pyramica

denticulata (Mayr) (top) and Strumigenys lanuginosa Wheeler (bottom).

The arrows show the position of the mandibular articulations and of the

labial palpi as indicator of the distal border of the labio-maxillary complex.

108 C. BARONI URBANI & M.L. DE ANDRADE

00047449 yz

ZMB

00047454 100 um Uni Basel

‘ig. 41 - Comparable position of the base of the mandibles in Pyramica eggersi

(Emery) (top) and Strumigenys godmani Forel (bottom). ‘The arrows show

the position of the mandibular articulations and of the labial palpi as indi-

cator of the distal border of the labio-maxillary complex.

ANT TRIBE DACETINI 109

00047429 Ud HE,

00046784 "i 100 pm > ui ba

Fig. 42 - Pyramica zeteki (Brown) (top) and Pyramica subedentata (Mayr) (bottom)

opening the left mandible at 90° (= 180° for both mandibles). Pyramica

species were supposed to never open their mandibles wider than 90°.

110 C. BARONI URBANI & M.L. DE ANDRADE

siomorphic) “not arising from across entire width of labium”

(synapomorphic) in Strumigenys. Figures 43 - 44 should be suf-

ficient to exemplify the indefensibility of this partition. Another

case contradicting Bolton’s classification was already published

in a web document by DE ANDRADE & BARONI URBANI (2005).

4) [?Opening of the] “buccal cavity relatively short and wide, lateral

margins of cavity... not converging anteriorly” (plesiomorphic)

in Pyramica, as opposed to “relatively long and narrow, lateral

margins of cavity... converging anteriorly”, synapomorphic for

Strumigenys. Figures 45 - 46 show four cases of contradictory

distribution of this character among two Strumigenys and two

“Pyramica” species.

In addition and as a general consideration, any attempt to regard

Strumigenys and Pyramica as valid genera separated by the mandibu-

lar kinetic or by any of its morphological correlates would inevitably

render one of the two fictitious genera invalid for being paraphyletic

to the other as it was already shown under chapter 4.1.

The erroneous separation of Pyramica from Strumigenys yields

to the following, unfortunately necessary list of reaffirmed or new

combinations and replacement names for new homonyms in Strumi-

genys. Only the most important references are reported in our list.

Additional ones can be found in BOLTON (2000). Needless to say,

the tormented nomenclatorial fate of the species listed below (some

of them were attributed to three different genera in less than 20

years) can be regarded as an additional proof of the opportunity of

merging all Strumigenys satellite genera in one. The following is a

list of species affected by nomenclatorial changes as compared with

the nomenclature adopted by BOLTON (2000) and not a Strumigenys

species list. The species list can be easily extracted from the mono-

graph of BOLTON (2000).

Strumigenys abdita WESSON & WESSON, 1939: 106. Smithistruma abdita (Wesson &

Wesson), SMITH, 1951: 827. Pyramica abdita, BOLTON, 1999. 1673. Combination

in Strumigenys reinstated.

Strumigenys abditivata (Bolton). New combination for Pyramica abditivata

BOLTON, 2000: 231.

Strumigenys acheron (Bolton). New combination for Pyramica acheron BOLTON,

2000: 416.

Strumigenys acubecca (Brown). New combination for Asketogenys acubecca BROWN,

1972: 23 and Pyramica acubecca, BOLTON, 1999: 1672.

ANT TRIBE DACETINI al

I "8 _"£z_ rl. eS as MB

00049791 | ( «(yp Z MB.

Fig. 43 - Uniformity of shape of the labium among putative Pyramica and Strumi-

genys species. Pyramica xenognatha (Kempf) (top) and Strumigenys exili-

rhina Bolton (bottom).

112 C. BARONI URBANI & M.L. DE ANDRADE

00049786 Ud Ra

00050876. 50 um ada

Fig. 44 - Uniformity of shape of the labium among putative Pyramica and Strumi-

genys species. Pyramica subedentata (Mayr) (top) and Strumigenys micretes

Brown (bottom).

ANT TRIBE DACETINI 19/5.

sep WAI o PIT

Uni Basel

Fig. 45 - Pyramica decipula Bolton (top) with narrow opening and subparallel mar-

gins of the buccal cavity and Strumigenys emmae (Emery) (bottom) with

broad opening and convergent margins of the buccal cavity. ‘The opposite

state of this character should characterize the two genera.

114 C. BARONI URBANI & M.L. DE ANDRADE

00049750. gee

Uni Basel

Gpose7s2. ov ‘100 um ZMB

Uni Basel

Fig. 46 - Pyramica subedentata (Mayr) (top) with narrow opening and subparallel

margins of the buccal cavity and Strumigenys chapmani Brown (bottom)

with broad opening and convergent margins of the buccal cavity. The

opposite state of this character should characterize the two genera.

ANT TRIBE DACETINI 115

Strumigenys aello (Bolton). New combination for Pyramica aello BOLTON, 2000: 416.

Strumigenys aethegenys (Bolton). New combination for Pyramica aethegenys

BOLTON, 2000: 179.

Strumigenys africana (Bolton). New combination for Glamyromyrmex africanus

BOLTON, 1983: 322 and Pyramica africana, BOLTON, 1999: 1672.

Strumigenys agnosta (Bolton). New combination for Pyramica agnosta BOLTON,

2000: 293.

Strumigenys agosti (Bolton). New combination for Pyramica agostit BOLTON, 2000:

412.

Strumigenys ailaoshana (Xu & Zhou). New combination for Pyramica ailaoshana

Xu & ZHou, 2004: 440.

Strumigenys alberti FOREL, 1893b. 380. Smithistruma alberti (Forel) BROWN 1953: 93.

Pyramica alberti, BOLTON, 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys alecto (Bolton). New combination for Pyramica alecto BOLTON, 2000:

429.

Strumigenys ambatrix (Bolton). New combination for Pyramica ambatrix BOLTON,

2000? B55"

Strumigenys anarta (Bolton). New combination for Smithistruma anarta BOLTON,

1983: 314 and Pyramica anarta, BOLTON, 1999: 1673.

Strumigenys anderseni (Bolton). New combination for Pyramica anderseni BOLTON,

2000: 474.

Strumigenys angulata SMITH, 1931: 697. Smithistruma angulata (Smith) Brown, 1953:

54. Pyramica angulata, BoLTON, 1999. 1673. Combination in Strumigenys

reinstated.

Strumigenys anorbicula (Bolton). New combination for Pyramica anorbicula

BoLTON, 2000: 287.

Strumigenys appalachicolensis (Deyrup & Lubertazzi). New combination for

Pyramica appalachicolensis DEyRuUP & LUBERTAZZI, 2001: 15.

Strumigenys appretiata (Borgmeier). New combination for Glamyromyrmex

appretiatus BORGMEIER, 1954: 282 and Pyramica appretiata, BOLTON, 1999: 1672.

Strumigenys arahana (Bolton). New combination for Smithistruma arahana

BoLTON, 1983: 300 and Pyramica arahana, BOLTON, 1999: 1673.

Strumigenys archboldi (Deyrup & Cover). New combination for Smithistruma

archboldi DEYRUP & COVER, 1998: 217 and Pyramica archboldi, BOLTON, 1999: 1673.

Strumigenys arges (Bolton). New combination for Pyramica arges BOLTON, 2000: 462.

Strumigenys argiola (Emery). Epitritus argiolus EmeRY, 1869a: 136. Strumigenys

argiola (Emery), BARONI URBANI, 1998: 163. Pyramica argiola, BOLTON, 1999:

1672. Combination in Strumigenys reinstated.

Strumigenys arizonica (Ward). New combination for Smithistruma arizonica WARD,

1988: 121 and Pyramica arizonica, BOLTON, 1999: 1673.

Strumigenys asaphes (Bolton). New combination for Pyramica asaphes BOLTON,

2000: 171.

Strumigenys assamensis de Andrade in BARONI URBANI & DE ANDRADE, 1994: 61.

Smithistruma assamensis (de Andrade), BOLTON, 1995: 384. Pyramica assamensts,

BoLTON, 1999. 1673. Combination in Strumigenys reinstated.

Strumigenys atopogenys (Bolton). New combination for Pyramica atopogenys

BoLTON, 2000: 417.

116 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys atropos (Bolton). New combination for Pyramica atropos BOLTON,

2000: 457.

Strumigenys auctidens (Bolton). New combination for Pyramica auctidens BOLTON,

2000: 179,

Strumigenys augustandrewi (Longino). New combination for Pyramica augustandrewi

Longino, 2006: 140.

Strumigenys azteca (Kempf). New combination for Glamyromyrmex aztecus KEMPF,

1960b: 444 and Pyramica azteca, BoLTON, 1999: 1672.

Strumigenys baudueri (Emery). Epitritus baudueri EMERY, 1875: 474. Strumigenys

baudueri (Emery), Mayr, 1887: 571 et auctorum omnium recentiorum usque

ad 1948. Smithistruma baudueri (Emery), BRown, 1948: 105. Pyramica baudueri,

BoLTON, 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys beebei (Wheeler). New combination for Glamyromyrmex beebet

WHEELER, 1915b: 488 and Pyramica beebet, BOLTON, 1999: 1672.

Strumigenys behasyla (Bolton). New combination for Smithistruma behasyla

BoLTON, 1983: 286 and Pyramica behasyla, BOLTON, 1999: 1673.

Strumigenys belial (Bolton). New combination for Pyramica belial BOLTON, 2000:

oak.

Strumigenys bellatrix (Bolton). New combination for Pyramica bellatrix BOLTON,

2000: 336.

Strumigenys benten (Terayama, Lin & Wu). New combination for Smithistruma

benten 'TERAYAMA, LIN & Wu, 1996: 329.

Strumigenys bequaerti SANTSCHI, 1923: 286. Serrastruma bequaerti (Santschi)

ConsanI, 1951: 171. Pyramica bequaerti, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys bimarginata WESSON & WEsSON, 1939: 95. Smithistruma bimarginata

(Wesson & Wesson) SMITH, 1951: 827. Pyramica bimarginata, BOLTON, 1999:

1673. Combination in Strumigenys reinstated.

Strumigenys boltoni (Deyrup). New combination for Pyramica boltoni DEYRUP,

2006: È.

Strumigenys brevicornis MANN, 1922: 38. Neostruma brevicornis (Mann), BRown, 1948:

111. Pyramica brevicornis, BOLTON, 1999: 1672. Combination in Strumigenys

reinstated.

Strumigenys brontes (Bolton). New combination for Pyramica brontes BOLTON,

2000: 463.

Strumigenys browni (Bolton). New combination for Pyramica brown BOLTON,

2000: 168.

Strumigenys bubisnoda (Bolton). New combination for Pyramica bubisnoda BOLTON,

2000: 393.

Strumigenys bunki (Brown). New combination for Smithistruma bunki BROWN,

1950b: 41 and Pyramica bunki, BoLTON, 1999: 1673.

Strumigenys californica (Brown). New combination for Smithistruma californica

Brown, 1950b: 40 and Pyramica californica, BoLTON, 1999: 1673.

Strumigenys canina (Brown & Boisvert). New combination for Pentastruma canina

Brown & Boisvert, 1979: 203 and Pyramica canina, BOLTON, 1999: 1673.

Strumigenys capitata (Smith). Cephaloxys capitata SMITH, 1865: 77. Strumigenys

capitata (Smith), Mayr, 1866: 517. Smithistruma capitata (Smith), BRown, 1948:

ANT TRIBE DACETINI igi

105. Pyramica capitata, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys carinognatha (Bolton). New combination for Pyramica carinognatha

BOLTON, 2000: 418.

Strumigenys carnassa (Bolton). New combination for Pyramica carnassa BOLTON,

2000: 475.

Strumigenys carolinensis (Brown). New combination for Smithistruma carolinensis

Brown, 1964: 185 and Pyramica carolinensis, BOLTON, 1999: 1673.

Strumigenys cascanteae (Longino). New combination for Pyramica cascanteae

LONGINO, 2006: 138.

Strumigenys cassicuspis (Bolton). New combination for Pyramica cassicuspis

BoLTON, 2000: 216.

Strumigenys castanea (Brown). New combination for Smithistruma castanea BROWN,

1953: 107 and Pyramica castanea, BOLTON, 1999: 1673.

Strumigenys cavinasis (Brown). New combination for Smithistruma cavinasis

Brown, 1950b: 42 and Pyramica cavinasis, BOLTON, 1999: 1673.

Strumigenys charybdys (Bolton). New combination for Pyramica charybdys BOLTON,

2000: 396.

Strumigenys chiricahua (Ward). New combination for Smithistruma chiricahua

WARD, 1988: 119 and Pyramica chiricahua, BOLTON, 1999: 1673.

Strumigenys chyatha (Bolton). New combination for Smithistruma chyatha BOLTON,

1983: 288 and Pyramica chyatha BOLTON, 1999: 1673.

Strumigenys cincinnata (Kempf). New combination for Smithistruma cincinnata

Kemper, 1975: 419 and Pyramica cincinnata, BOLTON, 1999: 1673.

Strumigenys circothrix (Ogata & Onoyama). New combination for Smithistruma

circothrix OGATA & ONOYAMA, 1998: 280 and Pyramica circothrix. BOLTON, 1999: 1673.

Strumigenys clotho (Bolton). New combination for Pyramica clotho BOLTON, 2000: 413.

Strumigenys cloydi (Pfitzer). New combination for Smithistruma cloydi PFITZER,

1951: 198 and Pyramica cloydi, BOLTON, 1999: 1673.

Strumigenys clypeata ROGER, 1863a: 213 [clipeata!]. Smithistruma clypeata (Roger)

SmitH, 1951: 827. Pyramica clypeata, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys comis (Kempf). New combination for Gymnomyrmex comis KEMPF,

1959: 342 and Pyramica comis, BOLTON, 1999: 1672.

Strumigenys concolor SANTSCHI, 1914b: 375. Serrastruma concolor (Santschi), WEBER,

1925: 6. Pyramica concolor, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys connectens KeMPF, 1958: 59. Pyramica connectens, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys conspersa EMERY, 1906: 169. Smithistruma conspersa (Emery), BROWN,

1953: 100. Pyramica conspersa, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys convexiceps SANTSCHI, 1931: 277. Pyramica convexiceps, BOLTON, 1999:

1672. Combination in Strumigenys reinstated.

Strumigenys crassicornis MAYR, 1887: 569. Neostruma crassicornis (Mayr) Brown, 1948:

111. Pyramica crassicornis, BOLTON, 1999: 1672. Combination in Strumigenys

reinstated.

118 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys creightoni SMITH, 1931: 705. Smithistruma creightoni (Smith), SMITH,

1951: 827. Pyramica creightoni, BoLToN, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys crementa (Bolton). New combination for Pyramica crementa BOLTON,

2000: 182.

Strumigenys cryptura (Bolton). New combination for Glamyromyrmex crypturus

BoLTON, 1983: 327 and Pyramica cryptura, BOLTON, 1999: 1672.

Strumigenys dagon (Bolton). New combination for Glamyromyrmex dagon BOLTON,

1983: 325 and Pyramica dagon Bolton, 1999: 1672.

Strumigenys dapsilis (Bolton). New combination for Pyramica dapsilis BOLTON,

2000-2147)

Strumigenys daspleta (Bolton). New combination for Pyramica daspleta BOLTON,

2000: 445.

Strumigenys datissa (Bolton). New combination for Smithistruma datissa BOLTON,

1983: 289 and Pyramica datissa, BOLTON, 1999: 1673.

Strumigenys dayui (Xu). New combination for Epitritus dayui Xu, 2000: 297.

Strumigenys decipula (Bolton). New combination for Pyramica decipula BOLTON,

2000: 183.

Strumigenys deinognatha (Bolton). New combination for Pyramica deinognatha

BOLTON, 2000: 419.

Strumigenys deinomastax (Bolton). New combination for Pyramica deinomastax

BoLTON, 2000: 160.

Strumigenys denticulata Mayr, 1887: 570. Pyramica denticulata, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys dentinasis (Kempf). New combination for Gymnomyrmex dentinasis

Kempr, 1960b: 450 and Pyramica dentinasis, BOLTON 1999: 1672.

Strumigenys dentiscapa (Bolton). New combination for Pyramica dentiscapa

BoLTON, 2000: 396.

Strumigenys depilosa (Bolton). New combination for Pyramica depilosa BOLTON,

2000: 312.

Strumigenys depressiceps WEBER, 1934: 47. Smithistruma depressiceps (Weber),

Brown, 1953: 112. Pyramica depressiceps, BoLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys deyrupi (Bolton). New combination for Pyramica deyrupi BOLTON,

2000: 119.

Strumigenys dictynna (Bolton). New combination for Pyramica dictynna BOLTON,

2000: 294.

Strumigenys dietrichi SMITH, 1931: 696. Smithistruma dietrichi (Smith), SMITH, 1951:

827. Pyramica dietrichi, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys disjuncta (Bolton). New combination for Pyramica disjuncta BOLTON,

2000: 414.

Strumigenys dispalata (Bolton). New combination for Pyramica dispalata BOLTON,

2000: 213.

Strumigenys dohertyi EMERY, 1897: 576. Smithistruma dohertyi (Emery), BROWN, 1948:

105. Pyramica dohertyi, BoLTON, 1999: 1673. Combination in Strumigenys

reinstated.

ANT TRIBE DACETINI 119

Strumigenys dontopagis (Bolton). New combination for Pyramica dontopagis

BoLTON, 2000: 172.

Strumigenys doryceps (Bolton). New combination for Pyramica doryceps BOLTON,

20007 2447

Strumigenys dotaja (Bolton). New combination for Serrastruma dotaja BOLTON,

1983: 339 and Pyramica dotaja, BOLTON, 1999: 1673.

Strumigenys dyschima (Bolton). New combination for Pyramica dyschima BOLTON,

2000: 451.

Strumigenys eggersi EMERY, 1890: 69. Pyramica eggersi, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys electrina de Andrade in BARONI URBANI & DE ANDRADE, 1994: 38.

Smithistruma electrina (de Andrade), BOLTON, 1995: 384. Pyramica electrina,

BoLTON, 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys elegantula (Terayama & Kubota). New combination for Smithistruma

elegantula "TERAYAMA & KuBOTA, 1989: 788 and Pyramica elegantula, BOLTON,

1999: 1673.

Strumigenys emarginata Mayr, 1901: 26. Smithistruma emarginata (Mayr), BROWN,

1948: 105. Pyramica emarginata, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys emeswangi (Bolton). New combination for Pyramica emeswangi

BoLTON, 2000: 458.

Strumigenys emiliae ForEL, 1907: 11. Smithistruma emiliae (Forel), Brown, 1953:

104. Pyramica emiliae, BoLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys enkara (Bolton). New combination for Smithistruma enkara BOLTON,

1983: 301 and Pyramica enkara, BOLTON, 1999: 1673.

Strumigenys enopla (Bolton). New combination for Pyramica enopla BOLTON, 2000:

185.

Strumigenys epinotalis WEBER, 1934: 46 [Strumigenys studiosi epinotalis|. Smithistruma

epinotalis (Weber), Brown, 1953: 101. Pyramica epinotalis, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys epipola (Bolton). New combination for Pyramica epipola BOLTON,

Zion: ‘3997

Strumigenys erikae (Longino). New combination for Pyramica erikae LONGINO,

2006: 141.

Strumigenys erynnes (Bolton). New combination for Pyramica erynnes BOLTON,

2000: 351.

Strumigenys euryale (Bolton). New combination for Pyramica euryale BOLTON,

2000: 432.

Strumigenys excisa (Weber). New combination for Codiomyrmex excisus WEBER,

1934: 51 and Pyramica excisa, BoLTON, 1999: 1672.

Strumigenys exiguaevitae Baroni Urbani. New name for Pyramica hoplites BOLTON,

2000: 347, transferred to Strumigenys in the present paper. Nec Strumigenys

hoplites BROWN, 1973c: 266. Derivatio nominis: from the Latin exzgua = short +

vita = life, referred to the short life of the specific name hoplites in combination

with the generic name Pyramica.

Strumigenys extemena (Taylor). New combination for Dysedrognathus extemenus

TAYLOR, 1968b: 133 and Pyramica extemena, BOLTON, 1999: 1672.

120 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys exunca (Bolton). New combination for Pyramica exunca BOLTON,

2000: 302.

Strumigenys fautrix (Bolton). New combination for Pyramica fautrix BOLTON,

20007355:

Strumigenys fenkara (Bolton). New combination for Smithistruma fenkara BOLTON,

1983: 302 and Pyramica fenkara, BoLTON, 1999: 1673.

Strumigenys filirrhina (Brown). New combination for Smithistruma filirrhina

Brown, 1950b: 37 and Pyramica filirrhina, BOLTON, 1999: 1673.

Strumigenys filitalpa (Brown). New combination for Smithistruma filitalpa BROWN,

1950b: 39 and Pyramica filitalpa, BOLTON, 1999: 1673.

Strumigenys fisheri (Bolton). Pyramica fisheri Bouron, 2000: 330. New

combination.

Strumigenys flagellata (Taylor). New combination for Codiomyrmex flagellatus

TayLor, 1962: 7 and Pyramica flagellata, BOLTON, 1999: 1672.

Strumigenys formosa (Terayama, Lin & Wu). New combination for Epitritus

formosus TERAYAMA, LIN & Wu, 1995: 85 and Pyramica formosa, BOLTON, 1999:

1672.

Strumigenys formosimonticola (Terayama, Lin & Wu). New combination for

Smithistruma formosimonticola TERAYAMA, LIN & Wu, 1996: 331 and Pyramica

formosimonticola, BOLTON, 1999: 1673.

Strumigenys fridericimuelleri FOREL, 1886: 213. Smithistruma fridericimuelleri (Forel),

Brown, 1953: 98. Pyramica fridericimuelleri, BOLTON, 1999: 1673. Combination

in Strumigenys reinstated.

Strumigenys fulda (Bolton). New combination for Smithistruma fulda BOLTON,

1983: 282 and Pyramica fulda, BOLTON, 1999: 1673.

Strumigenys furtiva (Bolton). New combination for Pyramica furtiva BOLTON,

2000: 156.

Strumigenys gatuda (Bolton). New combination for Smithistruma gatuda BOLTON,

1983: 292 and Pyramica gatuda, BOLTON, 1999: 1673.

Strumigenys gemella Kempr, 1975: 416. Pyramica gemella, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys geoterra (Bolton). New combination for Serrastruma geoterra BOLTON,

1983: 341 and Pyramica geoterra, BOLTON 1999: 1673.

Strumigenys glenognatha (Bolton). New combination for Pyramica glenognatha

BOLTON, 2000: 160.

Strumigenys grytava (Bolton). New combination for Pyramica grytava BOLTON,

2000; 220.

Strumigenys gundlachi (Roger). Pyramica Gundlachi ROGER, 1862: 253. Strumigenys

Gundlachi (Roger), ROGER 1863b: 40 et auctorum omnium recentiorum usque ad

1999. Pyramica gundlachi, BoLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys gyges (Bolton). New combination for Pyramica gyges BOLTON, 2000:

424.

Strumigenys hadrodens (Bolton). New combination for Pyramica hadrodens BOLTON,

2000: 161.

Strumigenys halosis (Bolton). New combination for Pyramica halosis BOLTON,

2000: 162.

ANT TRIBE DACETINI 121

Strumigenys hathor (Bolton). New combination for Pyramica hathor BOLTON,

2000: 346.

Strumigenys hemisobek (Bolton). New combination for Pyramica hemisobek BOLTON,

2000: 452.

Strumigenys hensekta (Bolton). New combination for Smithistruma hensekta

BoLTON, 1983: 293 and Pyramica hensekta, BOLTON, 1999: 1673.

Strumigenys hexamera (Brown). Epitritus hexamerus BROWN, 1958: 70. Strumigenys

hexamera (Brown) BARONI URBANI & DE ANDRADE, 1994: 13. Pyramica hexamera,

BoLTON 1999: 1672. Combination in Strumigenys reinstated.

Strumigenys hirashimai (Ogata). New combination for Epitritus hirashimai OGATA,

1990: 197 and Pyramica hirashimai, BOLTON, 1999: 1672.

Strumigenys hiroshimensis (Ogata & Onoyama)). New combination for Smithistruma

hiroshimensis OGATA & ONOYAMA, 1998: 281 and Pyramica hiroshimensis, BOLTON,

1999: 1673.

Strumigenys hyalina (Bolton). New combination for Pyramica hyalina BOLTON,

20002 128,

Strumigenys hyphata (Brown). New combination for Smithistruma hyphata

Brown,1953: 110 and Pyramica hyphata, BOLTON, 1999: 1673.

Strumigenys impidora (Bolton). New combination for Smithistruma impidora

BOLTON, 1983: 294 and Pyramica impidora, BOLTON, 1999: 1673.

Strumigenys incerta (Brown). New combination for Smithistruma incerta BROWN,

1949b: 10 and Pyramica incerta, BOLTON, 1999: 1673.

Strumigenys inopina (Deyrup & Cover). New combination for Smithistruma inopina

DeyRUP & Cover, 1998: 215 and Pyramica inopina, BOLTON, 1999: 1673.

Strumigenys inopinata (de Andrade). New combination for Rhopalothrix

inopinata de Andrade in BARONI URBANI & DE ANDRADE, 1994: 54. DIETZ

(2004) repeatedly suggests the transfer of this species to Pyramica without

proposing the new combination. His reasons are a set of characteristic (not

synapomorphic) Strumigenys characters visible in our 1994 figures. ‘The unique,

damaged inopinata holotype exhibits one presumed, important “basicerotine”

synapomorphy according to BOLTON (1998), a deep transverse labial ridge, and

some circumstantial ones, like bizarre pilosity on the scape, scape shape, extreme

reduction of the eyes, etc. We already showed in the present paper (see our

discussion of character # 6) that the transverse labial ridge may be absent in

some “basicerotine” species and present in other Strumigenys species. After some

hesitation we propose the transfer of inopinata to Strumigenys for possession of

one Strumigenys + “Phalacromyrmecini” weak synapomorphy: the presence of

the mesepisternal hair beds. These beds, otherwise, are visible also in a few

“Octostruma” species. S. inopinata, moreover, possesses spongiform appendages

on the peduncle and gaster, another trait that we don’t consider as synapomorphic

but only typical of Strumigenys. Nonetheless it must be noted that retention

of inopinata in Rhopalothrix (= Basiceros), as it was originally described, would

strengthen Strumigenys generic boundaries (see our generic definition above).

Strumigenys inquilina (Bolton). New combination for Serrastruma inquilina BOLTON,

1983: 342 and Pyramica inquilina, BOLTON, 1999: 1673.

Strumigenys insula (Bolton). New combination for Pyramica insula BOLTON, 2000: 400.

Strumigenys inusitata (Lattke). New combination for Trichoscapa inusitata LATTKE,

1992: 142 and Smithistruma inusitata (Lattke), BOLTON, 1995: 384 and Pyramica

inusitata, BOLTON, 1999: 1673.

12 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys jacobsoni MENOZZI, 1939: 180. Smithistruma jacobsoni (Menozzi),

Brown, 1948: 105. Pyramica jacobsoni, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys jamaicensis Brown, 1959: 6. Pyramica jamaicensis, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys japonica ITo, 1914: 40. Smithistruma japonica (Ito), BRown, 1948:

105. Pyramica japonica, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys karawajewi Brown, 1948: 44. Smithistruma karawajewi (Brown),

Brown, 1948: 105. Pyramica karawajewi, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys kempfi (Taylor & Brown). New combination for Smithistruma Rempfi

TayLor & Brown, 1978: 35 and Pyramica kempfi, BOLTON, 1999: 1673.

Strumigenys kerasma (Bolton). New combination for Smithistruma kerasma BOLTON,

1983: 303 and Pyramica kerasma, BOLTON, 1999: 1673.

Strumigenys khakaura (Bolton). New combination for Pyramica khakaura BoLTON,

2000: 348.

Strumigenys kichijo (Terayama, Lin & Wu). New combination for Smithistruma

kichijo TERAYAMA, LIN & Wu, 1996: 335 and Pyramica kichijo, BOLTON, 1999: 1673.

Strumigenys kompisomala (Bolton). New combination for Pyramica kompisomala

BoLTON, 2000: 234.

Strumigenys kyidriformis (Brown). New combination for Smithistruma kyidriformis

Brown, 1964: 188 and Pyramica kyidriformis, BOLTON, 1999: 1673.

Strumigenys lachesis (Bolton). New combination for Pyramica lachesis BOLTON,

2000: 389.

Strumigenys laevinasis SMITH, 1931: 701. Smithistruma laevinasis (Smith), SMITH,

1951: 827. Pyramica laevinasis, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys laevipleura KEMPF, 1958: 64. Pyramica laevipleura, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys lalassa (Bolton). New combination for Pyramica lalassa BOLTON,

2000: 189.

Strumigenys lasia (Brown). New combination for Cladarogenys lasia BROWN, 1976:

34 and Pyramica lasia, BOLTON 1999: 1672.

Strumigenys laticeps (Brown). New combination for Epitritus laticeps BROWN, 1962:

77 and Pyramica laticeps, BOLTON, 1999: 1672.

Strumigenys leptothrix WHEELER, 1929: 55. Smithistruma leptothrix (Wheeler),

Brown, 1948: 107. Weberistruma leptothrix (Wheeler), Brown, 1949: 8. Pyramica

leptothrix, BOLTON, 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys lilloana (Brown). New combination for Glamyromyrmex lilloana

Brown, 1950a: 34 and Pyramica lilloana, BOLTON, 1999: 1672.

Strumigenys longinoi (Bolton). New combination for Pyramica longinoi BOLTON,

2000; 173,

Strumigenys loveridgei (Brown). New combination for Codiomyrmex loveridget

Brown, 1953: 21 and Pyramica loveridgei, BOLTON, 1999: 1672.

Strumigenys lucifuga (Bolton). New combination for Pyramica lucifuga BOLTON,

2000: 314.

ANT TRIBE DACETINI 493

Strumigenys ludovici FoREL, 1904: 369. Serrastruma ludovici (Forel), WEBER 1952:

6. Pyramica ludovici, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys lujae Forel in WASMANN, 1902: 294. Serrastruma lujae (Forel), WEBER,

1952: 6. Pyramica lujae, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys lygatrix (Bolton). New combination for Pyramica lygatrix BOLTON,

2000: 204.

Strumigenys malaplax (Bolton). New combination for Smithistruma malaplax

BoLTON, 1983: 304 and Pyramica malaplax, BoLTON, 1999: 1673.

Strumigenys marchosias (Bolton). New combination for Pyramica marchosias

BoLTON, 2000: 290.

Strumigenys margaritae ForeL, 1893b: 378. Smithistruma margaritae (Forel),

SMITH, 1951: 827. Pyramica margaritae, BoLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys marginata (Santschi). New combination for Epitritus marginatus

SANTSCHI, 1914a: 114 and Miccostruma marginata (Santschi), BRown, 1948:

123 and Smithistruma marginata (Santschi), BOLTON, 1983: 312 and Pyramica

marginata, BOLTON, 1999: 1673.

Strumigenys masukoi (Ogata & Onoyama). New combination for Smithistruma

masukot OGATA & ONOYAMA, 1998: 283 and Pyramica masukoi, BOLTON, 1999: 1673.

Strumigenys maxtillaris Baroni Urbani. New name for Epitritus mandibularis SZABO,

1909: 27, transferred to Strumigenys in the present paper. Nec Strumigenys

mandibularis SMITH, 1860: 72. Derivatio nominis: from the Latin mazillaris,

practically a synonym of the preoccupied name mandibularis. Muiccostruma

mandibularis (Szabo), BROWN, 1948b: 123. Smithistruma mandibularis (Szabo),

BoLTon, 1983: 283. Pyramica mandibularis, BOLTON, 1999: 1673.

Strumigenys maynet FoREL, 1916: 427. Serrastruma maynei (Forel), Brown, 1952:

77. Pyramica mavynei, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys mazu (Terayama, Lin & Wu). New combination for Smithistruma

mazu 'TERAYAMA, LIN & Wu, 1996: 337 and Pyramica mazu, BOLTON, 1999:

1673.

Strumigenys media (Wilson & Brown). New combination for Kyidris media WILSON

& Brown, 1956: 445 and Pyramica media, BOLTON, 1999: 1672.

Strumigenys medusa (Bolton). New combination for Pyramica medusa BOLTON,

2000: 436.

Strumigenys megaera (Bolton). New combination for Pyramica megaera BOLTON,

2000: 437.

Strumigenys mekaha (Bolton). New combination for Smithistruma mekaha BOLTON,

1983: 305 and Pyramica mekaha, BOLTON, 1999: 1673.

Strumigenys membranifera EMERY, 1869b: 24. Trichoscapa membranifera (Emery),

Brown, 1948: 113. Pyramica membranifera, BOLTON, 1999: 1673. Combination

in Strumigenys reinstated.

Strumigenys memorialis (Deyrup). New combination for Smithistruma memorialis

Deyrup, 1998: 81 and Pyramica memorialis, BOLTON, 1999: 1673.

Strumigenys metazytes (Bolton). New combination for Pyramica metazytes BOLTON,

2000» 121,

124 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys metopia (Brown). New combination for Neostruma metopia BROWN,

1959: 11 and Pyramica metopia, BoLTON, 1999: 1672.

Strumigenys metrix (Bolton). New combination for Pyramica metrix BoLTON, 2000:

222.

Strumigenys miccata (Bolton). New combination for Serrastruma miccata BOLTON,

1983: 348 and Pyramica miccata, BOLTON, 1999: 1673.

Strumigenys microthrix (Kempf). New combination for Smithistruma microthrix

‘KeMPF, 1975: 422 and Pyramica microthrix, BOLTON, 1999: 1673.

Strumigenys minima (Bolton). New combination for Epitritus minimus BOLTON,

1972: 205 and Pyramica minima, BOLTON, 1999: 1672.

Strumigenys minkara (Bolton). New combination for Smithistruma minkara

BOLTON, 1983: 306 and Pyramica minkara, BOLTON, 1999: 1673.

Strumigenys minuscula (Kempf). New combination for Gymnomyrmex minusculus

Kemper, 1962: 24 and Pyramica minuscula, BOLTON, 1999: 1672.

Strumigenys mira (Bolton). New combination for Pyramica mira BOLTON, 2000: 324.

Strumigenys mirabilis MANN, 1926: 105. Tingimyrmex mirabilis (Mann), BROWN, 1948:

111. Pyramica mirabilis, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys missouriensis SMITH, 1931: 701. Smithistruma missouriensis (Smith),

SMITH 1951: 828. Pyramica missouriensis, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys mitis (Brown). New combination for Pyramica mitis Brown in BOLTON,

2000: 442.

Strumigenys mnemosyne (Bolton). New combination for Pyramica mnemosyne

-BOLTON, 2000: 446.

Strumigenys moloch (Bolton). New combination for Pyramica moloch BOLTON,

2000: 235.

Strumigenys morisitat (Ogata & Onoyama). New combination for Smithistruma

morisitat OGATA & ONOYAMA, 1998: 284 and Pyramica morisitai, BOLTON, 1999: 1673.

Strumigenys mormo (Bolton). New combination for Pyramica mormo BOLTON,

2000: 290.

Strumigenys mumfordi WHEELER, 1932: 160. Smithistruma mumfordi (Wheeler),

Brown, 1953: 124. Pyramica mumfordi, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys murphy (Taylor). New combination for Epitritus murphy: TAYLOR,

1968b: 130 and Pyramica murphyi, BOLTON, 1999: 1672.

Strumigenys mutica (Brown). New combination for Kyidris mutica Brown, 1949: 3

and Pyramica mutica, BOLTON, 1999: 1672.

Strumigenys myllorhapha (Brown). New combination for Neostruma myllorhapha

Brown, 1959: 12 and Pyramica myllorhapha, BOLTON, 1999: 1672.

Strumigenys nannosobek (Bolton). New combination for Pyramica nannosobek

BoLTON, 2000: 453.

Strumigenys necopina (Bolton). New combination for Pyramica necopina BOLTON,

20001223

Strumigenys nepalensis de Andrade in BARONI URBANI & DE ANDRADE, 1994: 57.

Smithistruma nepalensis (de Andrade) BOLTON, 1995: 385. Pyramica nepalensts,

BoLTON, 1999: 1673. Combination in Strumigenys reinstated.

ANT TRIBE DACETINI 125

Strumigenys nigrescens WHEELER, 1911a: 28 [Strumigenys alberti var. nigresens].

Smithistruma nigrescens (Wheeler), BROWN, 1953: 96. Pyramica nigrescens,

BoLTON, 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys ninda (Bolton). New combination for Smithistruma ninda BOLTON,

1983: 284 and Pyramica ninda, BOLTON, 1999: 1673.

Strumigenys nimravida (Bolton). New combination for Pyramica nimravida

BoLTON, 2000: 332.

Strumigenys nitens SANTSCHI, 1932: 413. Codiomyrmex nitens (Santschi), WEBER, 1934:

52. Dorisidris nitens (Santschi), Brown, 1948: 116. Pyramica nitens, BOLTON,

1999: 1672. Combination in Strumigenys reinstated.

Strumigenys noara (Bolton). New combination for Pyramica noara BOLTON, 2000:

304.

Strumigenys nongba (Xu & Zhou). New combination for Pyramica nongba Xu &

ZHOU, 2004: 440.

Strumigenys nubila LATTKE & GoiTia, 1997: 387. Pyramica nubila, BOLTON: 1999.

1673. Combination in Strumigenys reinstated.

Strumigenys nykara (Bolton). New combination for Smithistruma nykara BOLTON,

1983: 307 and Pyramica nykara, BOLTON, 1999: 1673.

Strumigenys oconitrilloae (Longino). New combination for Pyramica oconitrilloae

LONGINO, 2006: 139.

Strumigenys ocypete (Bolton). New combination for Pyramica ocypete BOLTON,

2000: 420.

Strumigenys ogyga (Bolton). New combination for Pyramica ogyga BOLTON, 2000:

324.

Strumigenys ohioensis KENNEDY & SCHRAMM, 1933: 98. Smithistruma ohioensis

(Kennedy & Schramm) SMITH, 1951: 828. Pyramica ohioensis, BOLTON, 1999:

1673. Combination in Strumigenys reinstated.

Strumigenys olsoni (Bolton). New combination for Pyramica olsoni BOLTON, 2000: 353.

Strumigenys orchibia (Brown). New combination for Smithistruma orchibia BROWN,

1953: 105 and Pyramica orchibia, BOLTON, 1999: 1673.

Strumigenys ornata Mayr, 1887: 571. Smithistruma ornata (Mayr), SMITH, 1951:

828. Pyramica ornata, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys oxysma (Bolton). New combination for Smithistruma oxysma BOLTON,

1983: 315 and Pyramica oxysma, BOLTON, 1999: 1673.

Strumigenys paniaguaae (Longino). New combination for Pyramica paniaguae

LONGINO, 2006: 137.

Strumigenys paradoxa (Bolton). New combination for Pyramica paradoxa BOLTON,

2000: 210.

Strumigenys parsauga (Bolton). New combination for Pyramica parsauga BOLTON,

2000: 157.

Strumigenys pasisops (Bolton). New combination for Pyramica pasisops BOLTON,

2000: 192.

Strumigenys pedunculata (Brown). New combination for Smithistruma pedunculata

Brown, 1953: 118 and Pyramica pedunculata, BOLTON, 1999: 1673.

Strumigenys peetersi (Bolton). New combination for Pyramica peetersi BOLTON,

2000: 474.

126 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys pergander EMERY, 1895a: 326. Smithistruma pergandei (Emery),

Brown, 1948: 106. Pyramica pergandei, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys perissognatha (Bolton). New combination for Pyramica perissognatha

BoLTON, 2000: 238.

Strumigenys phasma (Bolton). New combination for Pyramica phasma BOLTON,

2000: 403. |

Strumigenys pholidota (Bolton). New combination for Pyramica pholidota BOLTON,

2000: 225.

Strumigenys pilinasis FOREL, 1901: 339. Smithistruma pilinasis (Forel), SMITH, 1951:

828. Pyramica pilinasis, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys piliversa (Bolton). New combination for Pyramica piliversa BOLTON,

2000: 339.

Strumigenys placora (Bolton). New combination for Smithistruma placora BOLTON,

1983: 308 and Pyramica placora, BOLTON, 1999: 1673.

Strumigenys podarge (Bolton). New combination for Pyramica podarge BOLTON,

2000: 421.

Strumigenys prex (Bolton). New combination for Pyramica prex BoLTON, 2000:

174.

Strumigenys probatrix (Brown). New combination for Smithistruma probatrix

Brown, 1964: 186 and Pyramica probatrix, BOLTON, 1999: 1673.

Strumigenys pulchella Emery, 1895a: 327. Smithistruma pulchella (Emery),

SMITH, 1951: 828. Pyramica pulchella, BoLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys pydrax (Bolton). New combination for Pyramica pydrax BOLTON,

2000: 466.

Strumigenys raptans (Bolton). New combination for Pyramica raptans BOLTON,

2000: 162.

Strumigenys ravidura (Bolton). New combination for Glamyromyrmex ravidurus

BoLTON, 1983: 331 and Pyramica ravidura, BOLTON, 1999: 1672.

Strumigenys reflexa WESSON & WESSON, 1939: 102. Smithistruma reflexa (Wesson &

Wesson), SMITH, 1951: 828. Pyramica reflexa, BOLTON, 1999: 1673. Combination

in Strumigenys reinstated.

Strumigenys reliquia (Ward). New combination for Smithistruma reliquia WARD,

1988: 117 and Pyramica reliquia, BOLTON, 1999: 1673.

Strumigenys reticeps (Kempf). New combination for Codiomyrmex reticeps KEMPF,

1969: 286 and Pyramica reticeps, BOLTON, 1999: 1672.

Strumigenys rhea (Bolton). New combination for Pyramica rhea BoLTON, 2000:

403.

Strumigenys robertsoni (Bolton). New combination for Pyramica robertsoni BOLTON,

2000: 305.

Strumigenys rogata (Bolton). New combination for Pyramica rogata BOLTON, 2000:

7S:

Strumigenys rohwert SMITH, 1935: 214. Smithistruma rohweri (Smith), SMITH, 1951:

838. Pyramica rohweri, BoLTON, 1999: 1673. Combination in Strumigenys

reinstated.

ANT TRIBE DACETINI 107

Strumigenys roomi (Bolton). New combination for Epitritus roomi BOLTON, 1972:

206 and Pyramica roomi, BOLTON, 1999: 1672.

Strumigenys rostrata EMERY, 1895a: 329. Smithistruma rostrata (Emery), SMITH, 1951:

828. Pyramica rostrata, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys rostrataeformis (Brown). New combination for Smithistruma

rostrataeformis BROWN, 1949: 12 and Pyramica rostrataeformis, BoLTON, 1999: 1673.

Strumigenys rudinodis STARCKE, 1941: ii [Strumigenys inezae var. rudinodis].

Weberistruma rudinodis (Starcke), Brown, 1953: 26. Smithistruma rudinodis

(Starcke), BoLTON, 1995: 385. Pyramica rudinodis, BOLTON: 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys rugithorax (Kempf). New combination for Gymnomyrmex rugithorax

Kempr, 1959: 339 and Pyramica rugithorax, BOLTON, 1999: 1672.

Strumigenys runa (Bolton). New combination for Pyramica runa BOLTON, 2000:

447,

Strumigenys rusta (Bolton). New combination for Smithistruma rusta BOLTON,

1983: 3138 and Pyramica rusta, BOLTON, 1999: 1673.

Strumigenys sahura (Bolton). New combination for Glamyromyrmex sahurus

BoLTon, 1983: 326 and Pyramica sahura, BOLTON, 1999: 1672.

Strumigenys sardonica (Bolton). New combination for Pyramica sardonica BOLTON,

2000: 326.

Strumigenys sautert (Forel). New combination for Pentastruma sauteri FOREL,

1912a: 51 and Pyramica sauteri, BOLTON, 1999: 1673.

Strumigenys schleeorum Baroni Urbani in BARONI URBANI & DE ANDRADE, 1994:

35. Smithistruma schleeorum (Baroni Urbani), BOLTON, 1995: 385. Pyramica

schleeorum, BOLTON, 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys schulzi EMERY, 1894: 213. Smithistruma schulzi (Emery), BRown, 1953:

108. Pyramica schulzi, BOLTON, 1999: 1673. Combination in Strumigenys

reinstated.

Strumigenys scolapax (Bolton). New combination for Pyramica scolapax BOLTON,

2000: 439.

Strumigenys scylla (Bolton). New combination for Pyramica scylla BOLTON, 2000:

439.

Strumigenys semicompta (Brown). New combination for Codiomyrmex semicomptus

Brown, 1959: 9 and Pyramica semicompta, BOLTON, 1999: 1672.

Strumigenys serket (Bolton). New combination for Pyramica serket BOLTON, 2000:

349.

Strumigenys serradens (Bolton). New combination for Pyramica serradens BOLTON,

2000: 404.

Strumigenys serraformis (Bolton). New combination for Pyramica serraformis

BoLTon, 2000: 405.

Strumigenys serrula SANTSCHI, 1910: 390 [Strumigenys lujae var. serrula]. Serrastruma

serrula (Santschi), WEBER, 1952: 5. Pyramica serrula, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys seti (Bolton). New combination for Pyramica seti BOLTON, 2000: 349.

Strumigenys sharra (Bolton). New combination for Smithistruma sharra BOLTON,

1983: 295 and Pyramica sharra, BOLTON, 1999: 1673.

128 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys shattucki (Bolton). New combination for Pyramica shattucki BOLTON,

2000: 475.

Strumigenys stagodens (Bolton). New combination for Pyramica siagodens BOLTON,

2000: 163.

Strumigenys simoni EMERY, 1895a: 42. Smithistruma simoni (Emery), BROWN, 1948:

107. Serrastruma simoni (Emery), BROWN: 1952: 82. Pyramica simoni (Emery),

BoLTON: 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys simulans (Santschi). New combination for Epitritus simulans SANTSCHI,

1931: 278 and Codioxenus simulans (Santschi), BROWN, 1948: 123 and Pyramica

simulans, BOLTON, 1999: 1672.

Strumigenys sinensis (Wang). New combination for Pyramica sinensis Wang in

BoLTON, 2000: 390.

Strumigenys sistrura (Bolton). New combination for Glamyromyrmex sistrurus

BoLTON, 1983: 329 and Pyramica sistrura, BOLTON 1999: 1672.

Strumigenys splendens (Borgmeier). New combination for Gymnomyrmex splendens

BORGMEIER, 1954: 279 and Pyramica splendens, BOLTON, 1999: 1672.

Strumigenys stauroma (Bolton). New combination for Pyramica stauroma BOLTON,

2000: 226.

Strumigenys stenotes (Bolton). New combination for Pyramica stenotes BOLTON,

2000: 192.

Strumigenys subedentata Mayr, 1887: 570. Pyramica subedentata, BOLTON: 1999:

1673. Combination in Strumigenys reinstated.

Strumigenys sublucida (Brown). New combination for Smi:ithistruma sublucida

Brown, 1953: 99 and Pyramica sublucida, BOLTON, 1999: 1673.

Strumigenys subsessa (Bolton). New combination for Pyramica subsessa BOLTON,

2000: 306.

Strumigenys substricta (Kempf). New combination for Glamyromyrmex substrictus

Kempr, 1964: 68 and Pyramica substricta, BOLTON, 1999: 1672.

Strumigenys sulumana (Bolton). New combination for Serrastruma sulumana

BOLTON, 1983: 352 and Pyramica sulumana, BOLTON, 1999: 1673.

Strumigenys stheno (Bolton). New combination for Pyramica stheno BOLTON, 2000:

440.

Strumigenys symmetrix (Bolton). New combination for Pyramica symmetrix

BoLTon, 2000: 356.

Strumigenys synkara (Bolton). New combination for Smithistruma synkara BOLTON,

1983: 309 and Pyramica synkara, BOLTON, 1999: 1673.

Strumigenys tacta (Bolton). New combination for Smithistruma tacta BOLTON,

1983: 317 and Pyramica tacta, BOLTON, 1999: 1673.

Strumigenys takasago (Terayama, Lin & Wu). New combination for Kyidris

takasago TERAYAMA, LIN & Wu, 1995: 87 and Pyramica takasago, BOLTON, 1999:

1672.

Strumigenys talpa WEBER, 1934: 63. Smithistruma talpa (Weber), SMITH 1951: 828.

Pyramica talpa, BOLTON, 1999: 1673. Combination in Strumigenys reinstated.

Strumigenys tanymastax (Brown). New combination for Smithistruma tanymastax

Brown, 1964: 183 and Pyramica tanymastax, BOLTON, 1999: 1673.

Strumigenys taphra (Bolton). New combination for Pyramica taphra BOLTON,

2000: 448.

ANT TRIBE DACETINI 129

Strumigenys tarbosyne (Bolton). New combination for Pyramica tarbosyne Bolton,

2000: 422.

Strumigenys tathula (Bolton). New combination for Pyramica tathula BOLTON,

2000.3 52,

Strumigenys tenuipilis EMERY, 1915: 264 [Strumigenys baudueri var. tenuipilis].

Smithistruma tenuipilis (Emery), BROWN, 1953: 132. Pyramica tenuipilis, BOLTON,

1999: 1673. Combination in Strumigenys reinstated.

Strumigenys tenuissima (Brown). New combination for Smithistruma tenuissima

Brown, 1953: 133 and Pyramica tenuissima, BOLTON, 1999: 1673.

Strumigenys teratrix (Bolton). New combination for Pyramica teratrix BOLTON,

2000: 163.

Strumigenys terayamai (Bolton). New combination for Pyramica terayamai BOLTON,

2000: 468.

Strumigenys terroni (Bolton). New combination for Smithistruma terroni BOLTON,

1983: 299 and Pyramica terroni, BOLTON, 1999: 1673.

Strumigenys tethepa (Bolton). New combination for Pyramica tethepa BOLTON,

2000: 328.

Strumigenys tethys (Bolton). New combination for Pyramica tethys BOLTON, 2000:

406.

Strumigenys tetragnatha (‘Taylor). New combination for Codiomyrmex tetragnathus

TayLOR, 1966: 225 and Pyramica tetragnatha, BOLTON, 1999: 1672.

Strumigenys thaxteri (Wheeler). New combination for Codiomyrmex thaxteri

WHEELER, 1916: 327 and Pyramica thaxteri, BoLTON, 1999: 1672.

Strumigenys theia (Bolton). New combination for Pyramica theta BOLTON, 2000:

407.

Strumigenys themis (Bolton). New combination for Pyramica themis BoLTON, 2000:

407.

Strumigenys thuvida (Bolton). New combination for Glamyromyrmex thuvidus

BoLTON, 1983: 332 and Pyramica thuvida, BOLTON, 1999: 1672.

Strumigenys tiglath (Bolton). New combination for Epitritus tiglath BOLTON, 1983:

357 and Pyramica tiglath, BOLTON, 1999: 1672.

Strumigenys tigrilla (Brown). New combination for Miccostruma tigrilla BROWN,

1973a: 32 and Smithistruma tigrilla (Brown), BOLTON, 1983: 284 and Pyramica

tigrilla, BoLTON, 1999: 1673.

Strumigenys tisiphone (Bolton). New combination for Pyramica tisiphone BOLTON,

2000: 390.

Strumigenys tlaloc (Bolton). New combination for Pyramica tlaloc BOLTON, 2000:

244.

Strumigenys tolomyla (Bolton). New combination for Smithistruma tolomyla

BoLTOoN, 1983: 310 and Pyramica tolomyla, BOLTON, 1999: 1673.

Strumigenys transversa SANTSCHI, 1913: 258. Smithistruma transversa (Santschi),

Brown, 1948: 105. Pyramica transversa, BOLTON, 1999: 1673. Combination in

Strumigenys reinstated.

Strumigenys trauma (Bolton). New combination for Pyramica trauma BOLTON,

2000: 408.

Strumigenys trieces BRowN, 1960: 50. Pyramica trieces, BouTon, 1999: 1673.

Combination in Strumigenys reinstated.

130 C. BARONI URBANI & M.L. DE ANDRADE

Strumigenys truncatidens (Brown). New combination for Smithistruma truncatidens

Brown, 1950b: 43 and Pyramica truncatidens, BoLTON, 1999: 1673.

Strumigenys trymala (Bolton). New combination for Glamyromyrmex trymalus

BoLTON, 1983: 333 and Pyramica trymala, BOLTON, 1999: 1672.

Strumigenys tukulta (Bolton). New combination for Glamyromyrmex tukultus

BoLTon, 1983: 334 and Pyramica tukulta, BOLTON, 1999: 1672.

Strumigenys turpis (Bolton). New combination for Pyramica turpis BOLTON, 2000:

176.

Strumigenys umboceps (Bolton). New combination for Pyramica umboceps BOLTON,

2000: 227.

Strumigenys urrhobia (Bolton). New combination for Pyramica urrhobia BOLTON,

2000: 228.

Strumigenys vartana (Bolton). New combination for Pyramica vartana BOLTON,

ZOO0 2 95)

Strumigenys vescops (Bolton). New combination for Pyramica vescops BOLTON,

2000: 409.

Strumigenys victrix (Bolton). New combination for Pyramica victrix BOLTON,

2000: 356.

Strumigenys villiersi (Perrault). New combination for Gymnomyrmex villiersi

PERRAULT, 1986: 1 and Pyramica villiersi, BOLTON, 1999: 1672.

Strumigenys vodensa (Bolton). New combination for Smithistruma vodensa BOLTON,

1983: 317 and Pyramica vodensa, BOLTON: 1999: 1673.

Strumigenys warditeras (Bolton). New combination for Pyramica warditeras

BoLTON, 2000: 246.

Strumigenys weberi (Brown). New combination for Smithistruma weberi BROWN,

1959: 7 and Pyramica weberi, BoLTON, 1999: 1673.

Strumigenys wheeleriana Baroni Urbani. New name for Glamyromyrmex wheeleri

SMITH, 1944: 266, transferred to Strumigenys in the present paper. Pyramica

wheeleri, BOLTON, 1999: 1672. Nec Epitritus wheeleri DONISTHORPE, 1916: 121,

junior synonym of Strumigenys emmae (Emery). Nec Strumigenys wheeleri MANN,

1921: 466, secondary homonym of Epitritus wheeleri DONISTHORPE, 1916: 121.

Strumigenys wilsoniana Baroni Urbani. New name for Pyramica wilsoni Wang in

BoLTON, 2000: 469. Nec Strumigenys wilsont Brown, 1969: 28.

Strumigenys wrayt (Brown). New combination for Smithistruma wrayi BROWN,

1950b: 38 and Pyramica wrayi, BOLTON, 1999: 1673.

Strumigenys xenochelyna (Bolton). New combination for Pyramica xenochelyna

BoLTON, 2000: 165.

Strumigenys xenognatha KEMPF, 1958: 64. Pyramica xenognatha, BOLTON, 1999: 1673.

Combination in Strumigenys reinstated.

Strumigenys xenomastax (Bolton). New combination for Pyramica xenomastax

BoLTON, 2000: 471.

Strumigenys valeogyna (Wilson & Brown). New combination for Kyidris yaleogyna

WILSON & Brown, 1956: 443 and Pyramica yaleogyna, BOLTON, 1999: 1672.

Strumigenys yangi (Xu & Zhou). New combination for Pyramica yangi Xu &

ZHOU, 2004: 445.

Strumigenys zeteki (Brown). New combination for Neostruma zeteki BRown, 1959:

10 and Pyramica zeteki, BOLTON, 1999: 1672.

ANT TRIBE DACETINI 131

5. KEY TO THE EXTANT GENERA OF AGROECOMYRMECINI AND

DACETINI

The key to the Dacetini genera by BOLTON (2000), in spite of the

facilitation due to the exclusion of Basiceros and some small genera,

contains a number of invalid statements if one broadens the number

of taxa examined. For instance, presence or absence of the limbus as

used in the first couplet to separate his “strumigenyte genera” from

the other dacetines does not hold if Phalacromyrmex and Basice-

ros are also considered. Several elements of BOLTON’s (2000) key

cannot be used in our context. On the other hand, uncertainties are

implicitly recognized also by Bolton while qualifying characters as

“extremely rare” or “extremely rarely absent”. Several other char-

acters were already shown during the present study to hold for a

majority of species only.

The following key should be of broader validity due to the more

rigorous and operational generic definitions adopted for the present

study. Although imprecise, the key still contains also some probabi-

listic statements similar to those employed by Bolton since they may

facilitate identification in a number of cases.

1. Eyes present and posterior to the antennal scrobe. Petio-

lar tergum and sternum differently shaped. No specialized

large mechanoreceptors on the mouthparts. Neotropical

TR Wah aC MN) MAERSK) ROE OS af amie! 6| N ARONA A DON Tatuidris

- Eyes generally present and never posterior to the anten-

nal scrobe, or, very rarely (Strumigenys inopinata) absent.

Petiolar tergum and sternum equally convex. Mouthparts

nearly always with some large, specialized mechanorecep-

tors (exceptions a few Strumigenys and Epopostruma spe-

2. Pretergite of first gastral segment subsessile to sessile. Base

of scape bent at right angle near the base. First gastral seg-

ment nearly always sculptured. Neotropical, Indomalayan,

PATASIP ALLA: « -« 4-4 sip REI See IO ee eee eee Basiceros

- Pretergite of first gastral segment neck like. Base of scape

straight or at least complanar with the basal condyle, rarely

bent at right angle. First gastral segment almost never

sculptured (exceptions, a few Strumigenys species) ......... 3

C. BARONI URBANI & M.L. DE ANDRADE

Mandibles with alternating small and large teeth. Scape

Clavare. Bas timandibouiar process absent, e... ire 2A. 4

Mandibular dentition different, or, if alternating large and

small teeth present, the scape is never clavate. Basimandi-

bularproccsstncart always presenti (A'S 15 rt i PIA 6

Katepisternal oblique groove deeply impressed. Vertexal

angles pointed backwards, mesosoma dorsally marginate.

ES Aha ee ne as oe ee ae a ree Ishakidris

None .of.the. characters .above. . . 5

Mesosternal hair beds hypertrophied. Long and flexuous

pilosity on the whole body. Head sculpture deeply reticu-

late. Ampennac ereht pointed..\dadagascar su Pilotrochus

Mesopleurae obliquely costulate. Pilosity rare and short.

Frontal lobes strongly developed. Antennae eleven jointed.

GAZA AI Rete Lek itihds Hs AL: Phalacromyrmex

Eyes ventral or, very rarely, absent. Labial palps one

jointed. Basal process of the mandibles never hypertrophied.

World 4fopics and: temperate aredsen 0. a, Sep, Strumigenys

Eyes always present, dorsal or lateral. Labial palps gener-

ally two or three jointed, if one jointed, the basal process

GE Me Mane Dies ASIA PMCTITOD ICE. ur, int I %

Basimandibular process hypertrophic and bifurcated at the

apex, situated below the labrum with closed mandibles.

Maxallaty palps.absenta Neotropicale. 1. nio Acanthognathus

Basimandibular process normally developed, situated above

the labrum with closed mandibles. Maxillary palps present

nea MIDA DA LA ra Ai 10 8

8. Second funicular joint hypertrophic, longer than the last seg-

meno Indomalayan and Australian... S.2 Orectognathus

- Second funicular joint normally developed, shorter than

RES ORE Lice... ee. Moraes, Gal ewes Aad AY 9

Antennae eleven jointed. Eyes dorsal. Occipital foramen

oneal Nmeoriyamieal. tun, sated hoe ne eee teow brina e Daceton

ANT TRIBE DACETINI SS

- Antennae four to eight jointed. Eyes dorsolateral. Occipital

foramen: POSterror yf es eee ee a ee Rena 0 10

10. Large pre-genal cavity visible in profile behind the base of

the mandibles. Palp formula 3,2. Labrum not capable of full

reflexion over the buccal cavity. Antennal scrobes absent.

Ethiopian.g® sie Boe ees ee ee e Microdaceton

- Gap between mandibles and head capsule reduced in pro-

file. Palp formula 5,3. Labrum capable of full reflexion

over the buccal cavity. Antennal scrobes below the eyes.

Arstrallan$ 8:13 eee ese sete ct Ape ae Epopostruma

6. SPECIES-LEVEL ADDITIONS

Basiceros floridanum (Brown and Kempf)

Cuba: Sancti Spiritus Province: way between ‘Topes de Col-

lantes and Trinidad, 19.XII.2000, 1 worker, M. L. de Andrade &

C. Baroni Urbani, Winkler extraction from litter (MSNG). Ciego

de Avila Province: way between Ciego de Avila and Majagua,

17.XII.2000, 1 worker, C. Baroni Urbani & M. L. de Andrade,

Winkler extraction from litter (MSNG).

This species was previously known only from Florida, from

where there is an extensive list of locality records, and from Mexico

(DEvRUP et al., 1997). Brown & KEMPF (1960) suggest that the

species may have been introduced to Florida from Latin America.

Deryrup et al. (1997) substantially agree with this supposition but

note that the lack of West Indian records should be considered as a

counter-argument for the hypothesis. The two Cuban records above

fill this gap.

Basiceros onorei Baroni Urbani & de Andrade n. sp.

Type material: holotype worker (unique) from Ecuador labelled:

Banos de Agua Santa, Prov. Tungurahua, 01°24’S 78°25°W, 1860 m, sendero Bella

Vista, leaf-litter, 26.VIII.2004, C. Baroni Urbani & M. L. de Andrade (PUCE).

Derivatio nominis. This species is named after Prof Dr Giovanni

Onore who facilitated in multiple ways our Dacetini field work in Ecuador.

134 C. BARONI URBANI & M.L. DE ANDRADE

Diagnosis. A Bastceros species belonging to the petiola-

tum-group as defined by BROWN & KEMPF (1960) (in Octostruma) and

differing from all species of this group, inca, jheringi, stenoscapum,

petiolatum and wheeleri (BROWN & KEMPF, 1960; PaLACIO, 1997), by

the following combination of characters: occipital margin with a row

of four clavate hairs, each upper scrobe margin with one clavate

hair, pronotum and mesonotum with a pair of clavate hairs each,

gaster with 4 rows of erect, clavate hairs (2,2,2,4), sides of the basal

face of the propodeum strongly marginate, and propodeum, pleurae

and gaster largely smooth, sub-opaque to shining.

Worker description (Fig. 47). Head dorsum weakly convex in

full dorsal view. Postero-lateral sides of the head with the two ante-

rior thirds diverging posteriorly into a round angle and the poste-

rior third gently converging into a weakly concave vertexal margin.

Frontal lobes weakly expanded and convex. Antennal fossae ven-

trally with a marked carina visible in full-face view, straight, cover-

ing the lower margin of the scrobes and ending below the eye. Eyes

small, with 4 ommatidia in the longest row, placed on the upper

margin of the antennal scrobes. With head in profile the scrobe very

distinct, with the upper margin behind the eye broad, lamellaceous,

semitransparent and the lower one strongly carinate. Anterior cly-

peal border medially with a broad concavity. Scapes slightly com-

pressed dorsoventrally, with strong subbasal bend. Antennae with

eight joints. Apical funicular joint slightly longer than the rest of

the funiculus. Mandibles triangular, with a series of 5 teeth and two

irregular denticles before the subround apex.

Mesosoma anteriorly convex and posteriorly sloping in pro-

file. Dorsum of the mesosoma medially with a broad, superficial,

longitudinal sulcus spacing from the pronotum to the mesonotum.

Propodeal suture superficially impressed. Posterior half of the basal

face of the propodeum gently concave. Sides of the basal face of the

propodeum strongly marginate. Propodeal teeth large, lamellaceous,

transparent, apically pointed and with broad base. Upper bases of

propodeal teeth strongly marginate, connected each other and form-

ing a clear carina dividing the basal and declivous faces. Lower

base of propodeal teeth ending on the middle of the declivous face.

Posterior half of the declivous propodeal face weakly marginate.

Propodeal spiracle large and below the lower base of the propodeal

tooth.

ANT TRIBE DACETINI 135

00050585

00047414 | RAR

Fig. 47 - Basiceros onorei n. sp. from Ecuador. Worker, head in dorsal view (top)

and entire profile (bottom).

136 C. BARONI URBANI & M.L. DE ANDRADE

Petiole with a long neck and with the node high anteriorly and

sloping posteriorly. Ventral surface of the petiolar neck anteriorly

with a small lamellaceous tooth pointed forwards. Postpetiole almost

flat in side view; in dorsal view the anterior and posterior borders

well marked by a thicker and anteriorly semitransparent carina.

Gaster oval. Base of the first gastral tergite clearly marginate.

Base of the first gastral sternite superficially marginate.

Sculpture. Head reticulate-punctuate and with thin, irregular

longitudinal rugosities. Pronotum and mesonotum irregularly reticu-

late and very superficially punctuate, the reticulation less marked on

the posterior half of the mesonotum. Propodeum and pleurae smooth

and shining, in addition the pleurae with a few minute punctures.

Gaster smooth and shining, in addition the posterior border of the

first gastral tergite, the anterior and posterior borders of the first

gastral sternite and all remaining tergites and sternites with well

impressed, large punctures.

Pilosity. Body with appressed, short, thin, decumbent hairs,

very rare on the dorsum of the propodeum and pleurae. Posterior

margin of the head dorsum with a row of four clavate erect hairs.

Upper antennal scrobes with one clavate erect hair each. Pronotum,

mesonotum and posterior half of the petiole and postpetiole with a

pair of clavate, erect hairs each. First gastral tergite with four rows

of clavate erect hairs, the first up to the third rows with two clavate

hairs and the fourth row with four hairs close to the posterior border.

Remaining gastral tergites with four clavate hairs, thinner than on

the first tergite. First gastral sternite medially with erect, truncate

or weakly clavate hairs; posterior half of the first gastral sternite and

remaining gastral sternites with clavate hairs much thinner than on

the tergites. Apex of the tibiae with a few clavate hairs.

Colour. Dark brown with slightly lighter antennae and legs.

Measurements (in mm) and indices: TL 2.84; HL 0.67; HW 0.74; SL 0.45;

Mie O27 Ee 006: WE 0.7621 LO

Discussion. BSB. onoret differs from the 5 known species of

the petiolatum-group of “Octostruma” as defined by BROWN & KEMPF

(1960) by the combination of characters listed in the diagnosis. B.

onoret shares with theringi the head dorsum with irregular rugosities

but differs from theringi mainly by the presence of standing hairs

on the mesosoma and gaster. B. onoret shares with petiolatum the

ANT TRIBE DACETINI 137

mesosoma and gaster superficially smooth but differs from it mainly

by having 6 standing hairs instead of 16 on the head dorsum and by

the head weakly instead of broadly convex.

Basiceros papuanum de Andrade n. sp.

Type material: holotype worker (left antenna and left hind tibia and

tarsus missing) from Papua New Guinea labelled: Papua NG: Morobe, Wau, 1150

m, 17.V.1992, G. Cuccodoro, # 2C (MHNG). Paratype: 1 dealate gyne (right

funiculus missing), same data and collection as the holotype.

Derivatio nominis. “Papuanus” is a neologism indicating the

provenance of this species from Papua New Guinea.

Diagnosis. A Bastceros species belonging to the “brevi-

cornis-group” of “Eurhopalothrix”® as defined by BROWN & KEMPF

(1960) and TayLoR (1968a), resembling 5. brevicorne but differing

from it by the following combination of characters: erect specialized

hairs on the head dorsum 10 (8+2) instead of 16 (8+4+4), first gas-

tral tergite without standing hairs instead of with 3 pairs, and larger

size (TL 2.1 mm instead of 1.7-1.8 mm).

Worker description (Fig. 48). Head dorsum gently convex in

full dorsal view. Postero-lateral sides of the head with the two ante-

rior thirds diverging posteriorly to a subround angle and the poste-

rior third gently converging into a weakly concave vertexal margin.

Frontal lobes weakly expanded and convex. Antennal fossae ventrally

with a marked carina visible in full-face view, straight, covering the

lower margin of the scrobes and ending below the eye. Eyes small,

with 2 ommatidia in the longest row, placed on the upper margin

of the antennal scrobes. With head in profile the scrobes are very

distinct, with the upper margin behind the eye with a thin margin,

and the lower one thicker and lamellaceous. Anterior clypeal border

medially with a broad concavity. Scapes slightly compressed dorso-

ventrally, with subbasal strong bend. Antennae with seven segments.

Apical funicular joint slightly longer than the rest of the funiculus.

Mandibles triangular, with a series of 12 teeth, teeth 3-5-7-9 much

longer than the remaining ones.

Mesosoma in profile with pronotum and mesonotum forming a

convex arch and basal face of the propodeum strongly sloping poste-

riorly. Propodeal suture very superficially impressed. Area between

C. BARONI URBANI & M.L. DE ANDRADE

yey

00050606 Uni Basel

00050605 —- 109 wm Ud BE,

48 - Basiceros papuanum n. sp. from Papua New Guinea. Head of the gyne in

dorsal view (top) and entire profile of the worker (bottom).

ANT TRIBE DACETINI 139

basal and declivous faces of the propodeum with a small triangular

sub-lamellaceous tooth prolonging to the declivous face as a thin

lamella.

Petiole with a long neck and with high, convex node. Petiolar

node slightly more than 1/2 broader than long. Ventral surface of

the petiolar neck with a small lamellaceous tooth pointed forwards

anteriorly. Postpetiole convex in side view and about 1/2 broader

than long and broadly connected to the gaster.

Gaster oval. Base of the first gastral tergite clearly marginate.

Base of the first gastral sternite superficially marginate.

Sculpture. Head, mesosoma, petiole, postpetiole and gaster

densely foveolate, the foveae slightly sparser on the first gastral ster-

nite. Legs strongly punctuate.

Pilosity. Body with appressed, short, decumbent hairs, very

rare on the anterior face of pronotal dorsum, on the lower pro- and

mesopleurae, and on the propodeum, thicker on the remaining part

of the pronotum and mesonotum, thinner and slightly longer on

the posterior part of the first gastral tergite, longer and decumbent

on the posterior half of the first gastral sternite. Frons with two

rows of specialized hairs, the anterior row composed by $ clavate

hairs forming an arch connecting the eyes and the posterior row

composed by 2 hairs on the middle of the vertex. Remaining gastral

tergites and sternites with few, subdecumbent, spatulate hairs. In

addition the second, third and fourth gastral sternites with subde-

cumbent, long and slightly spatulate hairs. Distal outer face of the

tibiae and upper outer face of tarsi with one or two thick spatulate

hairs each.

Colour. Ferruginous-brown.

Measurements (in mm) and indices: Worker (holotype): TL 2.10; HL 0.48;

HW 0.53; SL 0.29; ML 0.13; EL 0.04; WL 0.49; CI 110.4; SI 54.7; MI 27.1.

Gyne description. Similar to the worker but differing in the fol-

lowing details. Eyes large. Ocelli present. Scape and mandibles as

in Fig. 48. Mesosoma robust and flat in profile. Parapsidal furrows

weakly impressed. Sides of the scutellum converging posteriorly to

form a rounded posterior border. Basal face of the propodeum very

short and in the same plane as the declivous one. Propodeal tooth

and lamellae less developed.

140 C. BARONI URBANI & M.L. DE ANDRADE

Pilosity. Similar to the worker but the anterior half of the

mesonotum with two pairs of short, thin, erect hairs on the centre

and each side of the mesonotum with a similar hair as the dorsal

ones. Pre-scutellum and scutellum with a clavate hair on each side,

thicker on the scutellum.

Gyne (paratype) PL 2540 HE 2095: BW 0762; SL 0:33; ML 0.20; EL. 0.13;

WES: CE iy ot 532 MM 364

Discussion. The brevicorne group is known from the lt-

erature to be composed by six species, australe, biroi, brevicorne,

caledonicum, philippinum and punctatum (BROWN & KEMPF, 1960;

TAYLOR, 1968a). Among these six species, the new species described

here as papuanum, resembles brevicorne in general shape. B. pa-

puanum is the sole species of the group having the specialized hairs

on the head dorsum distributed in two rows of 8 and 2 hairs each

instead of 8:4:4 or 2 or without hairs. "TayLor (1967) mentions a

dealate gyne from N. E. New Guinea (Kunai Creek, Bulolo River

Valley) resembling punctatum and brevicorne or an undescribed spe-

cies. Later, TAYLOR (1970) suggests that the gyne from Kunai Creek

seems unlikely to be punctatum. This gyne from Kunai Creek dif-

fers from brevicorne mainly by the head with 12 hairs divided in

two rows of 8:4 instead of 8:4:4, by the first gastral tergite without

specialized hairs and by the larger size. It differs from gyne of pa-

puanum by the larger values of HL and WL and by the specialized

hairs on the head dorsum 8:4 instead of 8:2. Considering the cephalic

chaetotaxy the Kunai Creek specimen could be attributed either to

a new species very close to papuanum, or to true papuanum, if one

considers that the type material of papuanum might have lost both

lateral hairs of the second row of hairs on the middle of the vertex.

Basiceros sp. nr. ciliatum (Mayr)

Ecuador, Pichincha, S. José de Guaramal, 1,900 m, August 3,

2004, 1 dealate gyne, G. Osella (PUCE).

This specimen should be referred to B. ciliatum Mayr, the gyne

of which is still unknown. Since this unique gyne is not accompa-

nied by workers, our attribution remains tentative.

ANT TRIBE DACETINI 141

Strumigenys veddha de Andrade n. sp.

Type material: holotype worker (unique) from Sri Lanka labelled:

Ceylan North central, Alut Oya, 3.II.[19]70, Mussard, Besuchet & L6bl (MHNG).

Derivatio nominis: from the Veddhas, the name of one of the Sri

Lanka’s indigenous inhabitants. It is used as a noun in apposition.

Diagnosis. A Strumigenys belonging to the lyroessa-group

and to the prosopis-complex as defined by BOLTON (2000) and dif-

fering from the six species of this complex by the CI = 73.3 instead

of > 77, by the cephalic dorsum without standing hairs and by the

much more superficial sculpture on the head and mesosoma, and,

in particular, from the species /oricata Bolton, panopla Bolton, pro-

sopis Bolton and strenosa Bolton by the petiole with a pair of stand-

ing hairs, and from anchiplex Boltonn and propinqua Bolton by the

smaller values of SI < 57 instead of > 60, by the lack of pronotal

humeral hairs and by the first gastral tergite with basal and apical

pairs of erect hairs only.

Worker description (Fig. 49). Head strongly converging anteri-

orly, with round vertexal corners and moderately flattened dorsoven-

trally. Frontal lobes weakly expanded and convex. Antennal fossae

ventrally with a developed carina visible in full-face view, straight,

covering the lower margin of the scrobes and ending in front of the

eyes. Eyes small, with 5 ommatidia in the longest row, placed over

the ventral margin of the antennal scrobes, and not visible in dorsal

view. Scrobe distinct in profile, with marked upper and lower mar-

gins only. Lateral clypeal margin gently converging anteriorly to a

straight margin. Scapes less than half of the head length, slightly

thicker on the posterior half and surpassing the eyes posteriorly.

Antennae with six segments. Apical funicular joint longer than

the rest of the funiculus. Mandibles short, broad proximally and

tapered distally; their outer margin convex basally. Apical fork of

the mandible with two spiniform teeth, the apicodorsal one larger;

space between the two apical spiniform teeth bearing one intercalary

denticle. Preapical dentition consisting of 1 spiniform tooth.

Mesosoma in profile slightly flat and gently sloping posteriorly.

Propodeal teeth subtended by a broad lamella.

Petiole with a long neck and round node. Ventral surface of the

petiole with a broad spongiform lamina. Petiolar node with posterior

sides and posterior margin surrounded by spongiform processes.

Postpetiole gently convex in profile. Anterior, lateral and posterior

142 C. BARONI URBANI & M.L. DE ANDRADE

00050860

00050580 100 um Uni Basel

Fig. 49 - Strumigenys veddha n. sp. from Sri Lanka. Worker, head in dorsal view

(top) and entire profile (bottom).

ANT TRIBE DACETINI 143

faces of the postpetiole surrounded by spongiform processes broader

posteriorly. Ventral surface of the postpetiole with large and dense

spongiform process.

Gaster oval and with a few, short costulae. Base of the first gas-

tral tergite and sternite with spongiform pad larger on the tergite.

Sculpture. Head and anterior half of propleurae finely reticulate-

punctuate and superficially shining. Mesosomal dorsum and petiolar

node largely smooth and shining, with very sparse, faint, minute

reticulation-punctuation. Posterior half of the propleurae and remain-

ing pleurae smooth. Postpetiole and gaster smooth and shining.

Pilosity. Head dorsum with appressed, thin, short hairs only.

Upper scrobes margin and leading edge of scape with similar but

thicker and slightly longer hairs than on the head dorsum. Apico-

scrobal hair missing. Mesosoma, petiole, postpetiole and gaster with

hairs similar to those on the head dorsum but much sparser. Meso-

soma and postpetiole without erect hairs. Petiole with a pair of stiff

standing hairs. First gastral tergite with two pairs of erect stiff hairs,

one pair near the base, the other near the apex.

Colour. Light yellowish.

Measurements (in mm) and Indices: TL 2.12; HL 0.58; HW 0.425; SL 0.24;

NIESOtE-0.06; WL 0.56; C173.3; SI 56.5: MT 29.3,

Discussion. Among the 6 species of the prosopis-complex

S. veddha resembles especially loricata. Veddha and loricata share the

cephalic dorsum finely reticulate-punctuate and the promesonotum

with sculpture similar to the one on the head but less impressed. But

veddha differs from loricata mainly by the longer head (CI = 73.3

instead of > 82) and by the first gastral tergite with two pairs of stiff

hairs, one near to the base and another near to the apex (/oricata has

no standing hairs on the first tergite). Among the six species of the

prosopis-complex only veddha and panopla have the first gastral tergite

with one pair of stiff hairs near the base and another pair near the apex.

Strumigenys onorei Baroni Urbani & de Andrade n. sp.

Type material: holotype worker from Ecuador labelled: Bafios de Agua

Santa, Prov. Tungurahua, 01°24’S 78°25’W, 1860 m, sendero Bella Vista, leaf-litter,

26.VIII.2004, Juan Manuel Vieira Correa (PUCE). Paratype: 1 worker, same data

and collection as the holotype.

144 C. BARONI URBANI & M.L. DE ANDRADE

Derivatio nominis. This species is named after Prof Dr Giovanni

Onore as a sign of gratitude for his multiple helps during our fieldwork in

Ecuador.

Diagnosis. A Strumigenys belonging to the gundlachi-

group and gundlachi-complex as defined by BOLTON (2000), resem-

bling enopla (Bolton), but differing from it by the smaller SI values

(< 79.5 instead of > 84), by the shorter propodeal spines, by the

postpetiole with the ventral spongiform process larger and by the

standing hairs on head and gaster shorter.

Worker description (Fig. 50). Head strongly converging anteri-

orly and with round vertexal corners. Frontal lobes slightly expanded

and convex. Antennal fossae ventrally with a marked carina visible

in full-face view, straight, covering the lower margin of the scrobes

and ending close to the upper border of the eye. Eyes with 3-4

ommatidia in the longest row, placed over the ventral margin of the

antennal scrobes, and partially visible in dorsal view. With the head

in profile the scrobe distinct, with the upper margin more marked

than the lower one. Lateral clypeal margins gently converging ante-

riorly into a straight margin. Scapes slightly compressed dorsoven-

trally, with weak sub-basal bend, about 2/3 of the head length and

surpassing the eyes posteriorly. Antennae with six segments. Apical

funicular joint slightly longer than the rest of the funiculus. Man-

dibles elongate. Apical fork of the mandibles with two teeth and

with two intercalary denticles. Preapical dentition consisting of a

row with 4-6 minute denticles.

Mesosoma in profile gently sloping posteriorly. Propodeal

teeth small and triangular; declivous propodeal face with a narrow

lamella.

Petiole with a long pedicel and with the node high and convex.

Ventral surface of the petiole without spongiform lamina. Petiolar

node with marked posterior margin and without spongiform pro-

cess. Postpetiole gently convex in profile. Postpetiole with marked

anterior face; lateral and posterior faces surrounded by narrow

spongiform processes slightly broader on the posterior face. Ventral

surface of the postpetiole with spongiform process shorter than the

height of the node in profile.

Gaster oval and with few, short costulae. Base of the first gas-

tral tergite with narrow, spongiform pad.

ANT TRIBE DACETINI 145

preme __..— .. _’ _ GAME,

00050624 uABS,

Fig. 50 - Strumigenys onorei n. sp. from Ecuador. Worker, head in dorsal view

(top) and entire profile (bottom).

146 C. BARONI URBANI & M.L. DE ANDRADE

Sculpture. Head, mesosoma, petiole and postpetiole reticulate-

punctuate. Lower mesopleurae and gaster smooth.

Pilosity. Head and mesosoma with subdecumbent or decumbent,

weakly remiform hairs, rarer on the pronotum and mesonotum,

missing on the propodeum. Apicoscrobal hair long and flagellate.

Cephalic dorsum with two pairs of standing hairs, one close to

the vertexal margin and the other close to the highest point of the

vertex. Pronotal humeral hair long and flagellate. Mesonotal dorsum

with 1 pair of erect, flagellate hairs. The holotype shows on the left

side of the mesonotum one erect stiff hair before the flagellate one.

This hair is missing in the unique paratype worker. Petiole, post-

petiole and first gastral tergite with erect, sparse, weakly remiform

hairs slightly longer on the gaster. Posterior half of the first gastral

sternites and remaining sternites with appressed and erect pointed

hairs.

Colore Darks broe wimp lacs:

Measurements (in mm) and indices: TL 2.10-230; HL 0.49-0.54; HW 0.39-

0: 4650S 0) ot-0) 34a Vile, Or37 013.9) EleO.06;- WIE 0752-0:58: Cl 79.6; Sle 791-795;

Mle 7222375558

Discussion. The 15 species of the gundlachi-complex can

be divided in 4 clusters of closely related species. S. onore: can be

placed in the gundlachi s. str. cluster comprising the following 5

species: denticulata Mayr, eggersi Emery, enopla (Bolton), gundlachi

(Roger) and jamaicensis Brown. Among these five species, onore?

resembles enopla, and both species share the SI > 79-100. Of the

other species of the gundlachi complex, only some specimens of

jamaicensis and denticulata may have SI 80-81, but jamaicensis differs

from onorei and enopla by the strongly developed preapical dentition

(inconspicuous in onorei and enopla) and by the larger spongiform

process of postpetiole, while denticulata differs from both, onorez

and enopla by the pair of erect hairs on the mesonotum shorter and

stiff instead of long and flagellate and by the spongiform process

of postpetiole much more reduced or absent. S. onore: differs from

all the 5 species mentioned above by having the smallest propodeal

teeth and probably also by the mesonotum with 2 pairs of erect

hairs instead of one (see the description). By using the identification

key by BoLTON (2000) for the Neotropical species of Pyramica, S.

onoret will fall in the couplet 26 where it can be differentiated from

ANT TRIBE DACETINI 147

jamaicensis by using all characters of the first couplet and exclud-

ing only “ head slightly shorter and broader, CI 77-85”, and from

enopla by the following characters: SI < 80, standing hairs on the

head about as long as the eye length, spongiform process of the

postpetiole at least 1/3 as height to the exposed cuticle of side of

postpetiole disc. hey

Strumigenys longimala de Andrade n. sp.

Type material: holotype worker from Ecuador labelled: Yasuni

Scientific Station, Prov. Orellana 002 40% 46002 5.476423 6032 4 Wo 2301 leat-

litter, 28.VIII.2004, M. L. de Andrade & C. Baroni Urbani (PUCE). Paratypes:

2 worker and 1 dealate gyne same data and collection as the holotype (PUCE, 1

paratype worker MHNG).

Derivatio nominis: from the Latin longus (= long) and mala (=

mandible), referred to the length of the mandibles, among the longest of the genus.

Diagnosis. A Strumigenys belonging to the trudifera-group

as defined by BOLTON (2000), but differing from the sole species of

the group (i. e. trudifera Kempf & Brown), in the worker and gyne

by the smaller values of SI < 104.5 instead of 108.0, by the propo-

deal spines shorter, and by the flagellate hairs on the gaster sparser;

in the worker only by the smaller values of MI (<103) instead of

>110; and in the gyne only by MI 91.6 instead of 102.8.

Worker description (Fig. 51). Head strongly converging anteri-

orly and with round vertexal angles. Frontal lobes slightly expanded

and convex. Anterior half of the dorsal margin of the antennal

scrobes broad and ending in a subround denticle over the eyes.

Antennal fossae ventrally with a broad carina visible in full-face

view, straight and medially denticulate, covering the lower margin

of the scrobes and ending close to the upper border of the eye. Eyes

with 3 ommatidia in the longest row, placed over the ventral margin

of the antennal scrobes, and partially visible in dorsal view. With

the head in profile the scrobes are distinct, almost reaching the ver-

texal angles, with ventral margin and posterior half of the dorsal

margin superficially marked. Lateral clypeal margins gently convex

and anteriorly concave in the middle. Postbuccal groove impressed.

Scapes subcylindric, slightly more than 2/3 of the head length and

surpassing the eyes posteriorly. Antennae with six segments. Apical

funicular joint as long as the rest of the funiculus. Mandibles very

148 C. BARONI URBANI & M.L. DE ANDRADE

long, about as long as the head length. Apical fork of the mandibles

with two teeth and without intercalary denticles. Preapical dentition

consisting of one long, spiniform tooth only.

Mesosoma in profile convex anteriorly and slightly raised poste-

riorly before the propodeal spines. Propodeal spines long but shorter

than the the basal face of the propodeum; declivous propodeal face

with a thin lamella.

Petiole with a long pedicel and with convex node. Ventral sur-

face of the petiole with a thin lamina. Petiolar node with marked

posterior margin and with a small spongiform process on the poste-

rior lateral third. Postpetiole convex in profile, with marked anterior

and posterior faces; lateral faces surrounded by broad spongiform

process. Ventral surface of the postpetiole with very broad spongi-

form process.

Gaster oval and with costulae on its anterior third. Base of the

first gastral tergite and sternite with spongiform pad broader on the

sternite.

Sculpture. Head, mesosoma, petiole and postpetiole reticu-

late-punctuate. Gaster and area between meso- and metapleurae

smooth.

Pilosity. Head and mesosoma with subdecumbent or decum-

bent, narrow spatulate hairs, rarer on the mesosoma. Apicoscrobal

hair long and flagellate. Cephalic dorsum with two pairs of standing

hairs. Leading edge of the scape with 2 long hairs that curve toward

the base of the scape. Pronotal humeral hair long and flagellate.

Mesonotum with 1 pair of erect, flagellate hairs. Petiole, postpeti-

ole, first and second gastral tergites with flagellate hairs. Posterior

half of the first gastral sternite and remaining sternites with sparse,

erect pointed hairs.

Colori; 2 Dark brown.

Measurements (in mm) and indices: TL 2.78-2.94; HL 0.64-0.66; HW 0.44-

0.46; SL 0.46-0.48; ML 0.65-0.68; EL 0.05-0.06; WL 0.62-0.64; CI 68.7-69.7; SI

104.3-104.5; MI 101.5-103.0.

Gyne description. Similar to the worker but differing in the

following details. Eyes large. Ocelli present. Mesosoma robust and

weakly convex in profile. Parapsidal furrows impressed. Sides of

the scutellum with a thin lamella converging posteriorly to form

an obtuse angle. Basal face of the propodeum declivous posteriorly.

ANT TRIBE DACETINI 149

00050874

00050873 | 300 um Ud Bagel

Fig. 51 - Strumigenys longimala n. sp. from the Yasuni Scientific Station (Ecua-

dor). Worker, head in dorsal view (top) and entire profile (bottom).

150 C. BARONI URBANI & M.L. DE ANDRADE

Propodeal spines large and continuing to the declivous face as a

lamella.

Ventral surface of the petiole with a broader lamina.

Sculpture. Head and mesosoma with irregular rugosities, more

regular and denser on the mesosoma.

Pilosity. Similar to the worker but the mesonotum with 3 pairs

of flagellate hairs. Pre-scutellum and scutellum with a flagellate hair

on each side.

Measurements (in jmim) and idicess Wl 3226; Eli 0772: AW 0552: Si 0750;

MISS 06 BECO Ors dJER0 7617229613929,

Discussion. S. longimala is obviously the sister species of

trudifera (Brazil, Colombia and Venezuela) with which it shares all

most remarkable traits. Both species constitute the trudifera-group

characterized by the very long mandibles and by the scape with 2

hairs curved towards the base. S. longimala and trudifera can be

easily separated from all other species of the genus by the values of

SI, MI (see diagnosis) and by the number of flagellate hairs on the

first gastral tergite. The worker and gyne of longimala have 16 and

22 flagellate hairs respectively on the first gastral tergite while those

of trudifera have 20 and 26.

Strumigenys aduncomala de Andrade n. sp.

Type material: holotype worker (unique) from India labelled: Upper

Shillong, 1900 m, 13.V.1976, Meghalaya, W. Wittmer & C. Baroni Urbani (NHMB).

Derivatio nominis: from the Latin aduncus (= curved), and mala (=

mandible), referred to the very curved shape of the mandibles.

Diagnosis. A Strumigenys belonging to the caniophanes-

group and to the caniophanes-complex as defined by BOLTON (2000),

resembling lacunosa Lin & Wu, but differing from it by the dorso-

lateral margin of the head with 2 instead of 3 hairs projecting later-

ally, by its smaller size (TL = 2.52 mm instead of 3.1 mm) and by

the lower MI (43.7 instead of 50).

Worker description (Fig. 52). Head converging anteriorly, with

round vertexal corners. Frontal lobes weakly expanded and convex.

Antennal fossae ventrally with a narrow carina visible in full-face

view, straight, not covering the lower margin of the scrobes and

ANT TRIBE DACETINI sd

ending much before the eyes. Eyes small, with 3-4 ommatidia in the

longest row, placed over the ventral margin of the antennal scrobes,

and visible in dorsal view. With head in profile the scrobe distinct,

with weakly marked upper and lower margins. Lateral clypeal mar-

gins gently converging anteriorly into a straight margin. Scapes sub-

cylindrical, about 1/2 of the head length and surpassing the eyes

posteriorly. Antennae with six joints. Apical funicular joint moder-

ately constricted basally and longer than the rest of the funiculus.

Mandibles curved. Apical fork of the mandible with two spiniform

teeth, the apicodorsal larger; space between the apical teeth bearing

one intercalary denticle. Preapical dentition consisting of one tooth.

Mesosoma in profile weakly convex anteriorly and sloping pos-

teriorly. Propodeal teeth pointed; declivous propodeal face marginate

only. Mesepisternal hair beds large but smaller than the maximum

width of the fore coxa.

Petiole with a long neck and round node. Ventral surface of the

petiole with a broad spongiform lamina. Petiolar node with posterior

sides and posterior margin surrounded by spongiform process. Post-

petiole gently convex in side view. Anterior, lateral and posterior

faces of the postpetiole surrounded by spongiform processes broader

posteriorly. Ventral surface of the postpetiole with large and dense

spongiform process.

Gaster oval and with thin, short costulae. Base of the first gas-

tral tergite and sternite with spongiform pad larger on the tergite.

Sculpture. Head, dorsum of the mesosoma, propleurae and peti-

ole densely reticulate-punctuate. Postpetiole with very sparse, super-

ficial punctures. Meso- and metapleurae and gaster smooth.

Pilosity. Head, mesosoma, petiole and postpetiole with appressed,

thin, hairs. Dorsolateral margin of the head in full-face view with 2

freely laterally projecting flagellate hairs, one in apicoscrobal posi-

tion and one just posterior to the eye. Cephalic dorsum with a trans-

verse row of 4 erect fine hairs along the occipital margin, and with

2 pairs of similar hairs anterior to this, the anterior pair shorter and

on the frons and the posterior pair on the middle of the occipital

lobes. Pronotal humeral hairs flagellate and long. Dorsum of the

pronotum with two pairs of erect fine hairs. Dorsum of the mesono-

tum with 4 pairs of erect flagellate hairs, the anterior and posterior

pairs shorter. Dorsum of the propodeum with rare subdecumbent

193 C. BARONI URBANI & M.L. DE ANDRADE

FAV

_ Uni Basel

00050584 | ni Bai

Fig. 52 - Strumigenys aduncomala n. sp. from Meghalaya (India). Worker, head in

dorsal view (top) and entire profile (bottom).

ANT TRIBE DACETINI 153

fine hairs. Petiolar node with three pairs of fine hairs, the posterior

pair longer and flagellate. Postpetiole and gaster with long flagellate

hairs. Femora and tibiae with a few erect or suberect hairs. Mid-

and hind basitarsi with 2 long fine erect hairs.

Colo: wien ero was hy

Measurements (in mm) and indices: TL 2.52; HL 0.64; HW 0.46; SL 0.34;

Mi, ‘0:23; EL. 0.04; WL 0/69; CI 71.9: ST 73.9; MI 43.7.

Discussion. Among the 12 species of the caniophanes-

complex, aduncomala resembles more lacunosa. S. aduncomala and

lacunosa share the apical funicular segment moderately constricted

basally, the dorsum of the pronotum with at least 2 pairs of erect

hairs and the dorsum of the mesonotum with about 4 pairs of erect

hairs. At first glance S. aduncomala superficially resembles exilirhina

Bolton in general body shape, for having the dorsolateral margin

of the head in full-face view with 2 projecting hairs, and for the

basal constriction of the apical funicular antennomere. But exilirhina

belongs to the mayri-group whose component species are charac-

terized mainly by the anterior clypeal margin with a U-shaped or

V-shaped median notch and by the apical antennomere strongly

constricted basally. S. aduncomala has the anterior clypeal margin

straight and the apical antennomere moderately constricted basally.

Strumigenys caniophanoides de Andrade n. sp.

Type material: holotype worker (unique) from Bhutan labelled:

Phuntsholing, 2/400 m, 16.1V.1972, Nat. Hist. Museum Basel-Bhutan Expedition

(NHMB).

Derivatio nominis: from the existing Strumigenys species name

caniophanes and the Greek etdo0cg (= look), referred to the similarity between

cantophanes and the new species.

Diagnosis. A Strumigenys belonging to the caniophanes-

group and to the caniophanes-complex as defined by BOLTON (2000),

resembling caniophanes Bolton and paraposta Bolton, but differing

from caniophanes by the mandibles with a preapical denticle (with-

out denticle on caniophanes), by the rugae on the side of mesosoma

irregular and sparser, and from paraposta by its larger size (TL 3.38-

3.42 mm instead of 2.9-3.0 mm) and by the dorsolateral margin of

the head with more than 5 hairs projecting laterally, and from both,

cantophanes and paraposta, by the SI 80.0-81.7 instead of 68-78.

154 C. BARONI URBANI & M.L. DE ANDRADE

Worker description (Fig. 53). Head with sides converging anteri-

orly and with round occipital corners. Frontal lobes weakly expanded

and convex. Antennal fossae ventrally with a marked carina visible

in full-face view, straight, covering the lower margin of the scrobes

and ending in front of the eyes. Eyes small, with 5 ommatidia in the

longest row, placed over the ventral margin of the antennal scrobes,

and visible in dorsal view. With head in profile the scrobe distinct,

with weakly marked upper and lower margins. Clypeal dorsum con-

cave medially. Lateral clypeal margin gently converging anteriorly

into a straight margin. Scapes subcylindrical, about 3/5 of the head

length and surpassing the eyes posteriorly. Antennae with six joints.

Apical funicular joint slightly shorter than the rest of the funiculus.

Mandibles elongate, basally and apically weakly convex and medi-

ally straight. Apical fork of the mandibles with two spiniform teeth

and one intercalary denticle. Preapical dentition consisting of one

tooth near the apex.

Mesosoma in profile weakly convex anteriorly and sloping pos-

teriorly. Propodeal teeth pointed; declivous propodeal face margin-

ate only.

Petiole with a long neck and with low and long node. Ven-

tral surface of the petiole with a broad spongiform lamina. Petiolar

node with posterior margin surrounded by spongiform process that

in profile covers the posterior third of the node’s sides. Postpetiole

gently convex in side view. Anterior, lateral and posterior faces of

the postpetiole surrounded by spongiform processes broader poste-

riorly. Ventral surface of the postpetiole with large and dense spon-

giform process.

Gaster oval and with thin, short costulae. Base of the first gas-

tral tergite and sternite with spongiform pad larger on the tergite.

Sculpture. Head coarsely and densely reticulate-punctuate.

Mesosoma reticulate-punctuate and with irregular longitudinal

rugosities; the rugosities slightly more regular on the pleurae and

missing on the lower mesopleurae. Petiole and postpetiole densely

reticulate-punctuate. Gaster smooth. Outer face of fore coxae with

transversal rugosities.

Pilosity. Head, mesosoma, petiole and postpetiole with appressed,

thin hairs. Dorsolateral margin of the head in full-face view with

more than 5 freely laterally projecting hairs. Cephalic dorsum at

ANT TRIBE DACETINI 195

TAREE e i GAME,

900 HM ud Bagel

Fig. 53 - Strumigenys caniophanoides n. sp. from Bhutan. Worker, head in dorsal

view (top) and profile (bottom).

156 C. BARONI URBANI & M.L. DE ANDRADE

level of the eyes with fine standing hairs of different lengths. Prono-

tal humeral hairs long and flagellate. Dorsum of the mesosoma, pet-

iole, postpetiole and gaster with erect, fine hairs of different lengths,

longer on the gaster. Femora, tibiae and tarsi with numerous erect,

fine hairs of different lengths, two of which on the tibiae and two

of which on the tarsi much longer than the others.

Colour. reddish brown.

Measurements (in mm) and indices: TL 3.42; HL 0.82; HW 0.60; SL 0.49;

ML 0.40; EL 0.08; WL 0.90; CI 73.2; SI 81.7; MI 48.8.

Additional material: Nepal: Prov. Kosi, Distr. Sankhuwasawa,

Vallée d’ Arun, vic. Num, 1100 m, 21.1V.1984, 1 worker, I. Lobl &

A. Smetana (MHNG).

Measurements (in mm) and indices of the Nepalese worker: TL 3.38; HL 0.82;

HW 0.60; SL 0.48; ML 0.40; EL 0.08; WL 0.88; CI 73.2; SI 80.0; MI 48.8.

Discussion. Among the 12 species of the caniophanes-

complex S. caniophanoides is particularly similar to caniophanes and

paraposta. S. caniophanoides shares with caniophanes the large size

and the dorsolateral sides of the head in full dorsal view with more

than 5 projecting hairs. S. caniophanoides shares with paraposta the

pleurae with irregular rugae, the lower mesopleurae largely smooth

and the presence of preapical dentition on the mandibles.

Strumigenys hindu. de Andrade n. sp.

Type material: holotype worker (unique) from Nepal labelled: Pokhara,

820 m, 15-18.VI.1976, Nepal, W. Wittmer, C. Baroni Urbani (NHMB).

Derivatio nominis: the Hindu religion, one of the two major

religions of Nepal is used here as a noun in apposition.

Diagnosis. A Strumigenys belonging to the godeffroyi-

group and to the godeffroyi-complex as defined by BOLTON (2000),

resembling uberyx Bolton and buddhista de Andrade (q. v.), but

differing from these two species by its larger size (TL = 2.5 mm

instead of 1.8-1.9 mm) and by the SI = 102.6 (instead of < 91).

Worker description (Fig. 54). Head sides converging anteri-

orly, with round vertexal corners. Frontal lobes weakly expanded

and convex. Antennal fossae ventrally with a marked carina vis-

ANT TRIBE DACETINI 157

ible in full-face view, straight, covering the lower margin of the

scrobes and ending close to the eyes. Eyes small, with 2 ommatidia

in the longest row, placed over the ventral margin of the antennal

scrobes, and slightly visible in dorsal view. With the head in pro-

file the scrobe distinct, with superficially marked upper and lower

margins. Clypeal dorsum gently concave medially. Lateral clypeal

margin gently converging anteriorly into a straight margin. Scapes

subcylindrical, about 2/3 of the head length and surpassing the eyes

posteriorly. Antennae with six joints. Apical funicular joint much

longer than the rest of the funiculus. Mandibles curved. Apical fork

of the mandibles with two spiniform teeth and with two intercalary

denticles. Preapical dentition consisting of one spiniform tooth near

the apex.

Mesosoma in profile weakly convex anteriorly and sloping pos-

teriorly into the gently convex basal face of the propodeum. Pro-

podeal teeth pointed and ventrally connected to a medially convex

lamella.

Petiole with a long neck and with the node high, dorsally almost

flat and anteriorly tumuliform. Ventral surface of the petiole with a

broad spongiform lamina. Petiolar node with posterior margin and

whole sides surrounded by spongiform process. Postpetiole gently

convex in side view. Anterior, lateral and posterior margins of the

postpetiole surrounded by spongiform processes. Ventral surface of

the postpetiole with large and dense spongiform process.

Gaster oval and with thin, short costulae. Base of the first gas-

tral tergite and sternite with spongiform pad larger on the tergite.

Sculpture. Head reticulate and minutely punctuate. Pronotum,

pleurae and declivous face of the propodeum smooth. Mesonotum,

basal face of the propodeum and petiole punctuate, the punctures

fainter and sparser on the petiole. Postpetiole and gaster smooth.

Pilosity. Head, mesosoma, petiole and postpetiole with appressed,

thin hairs. Dorsolateral margin of the head in full-face view with the

apicoscrobal flagellate hair only. Cephalic dorsum with three rows of

erect hairs, the first row with 4 hairs close to the occipital margin,

the second row with 4 standing hairs and the third row with 2 hairs

on the frons. Upper scrobal margin with narrow spoon-shaped hairs

curved anteriorly. Clypeal border with hairs similar to those of the

upper scrobe. Scapes with hairs also similar to those on the upper

C. BARONI URBANI & M.L. DE ANDRADE

158

ZMB

Uni Basel

LO)

©)

LO)

Uni Basel

300 um

00050591

dorsal view (top)

head in

)

Worker

from Nepal

le (bottom)

indu n

ire pro

1genys

Strum

and ent

54 -

ANT TRIBE DACETINI 159

scrobes and curved posteriorly. Pronotal humeral hairs long and

flagellate. Dorsum of the pronotum with a pair of erect long hairs.

Mesonotum with a pair of erect flagellate hairs. Petiole, postpetiole

and gaster with few erect flagellate hairs. Dorsal face of hind femora

with an erect fine hair on the basal third. Outer face of mid and

hind tibiae with an erect, long flagellate hair. Hind basitarsi with

1-2 pairs of erect, long flagellate hairs.

Colour. light brown.

Measurements Gn mm) and indices: TE 2,50% HE 0/62; HW 0.38: SL. 0.39;

WE 0.26; ET .0.03: Wily 066; CP 61233451 102-6;.MIl 41-9.

Discussion. S. hindu, uberyx and buddhista are very simi-

lar each other but the characters listed in their respective species

diagnosis separate them clearly.

Strumigenys buddhista de Andrade n. sp.

Type material: holotype worker (unique) from Nepal labelled: Pokhara,

820 m, 15-18.VI.1976, Nepal, W. Wittmer, C. Baroni Urbani (NHMB).

Derivatio nominis: name derived from the Buddhism, the second

major religion in Nepal.

Diagnosis. A Strumigenys belonging to the godeffroyi-

group and to the godeffroyi-complex as defined by BOLTON (2000),

resembling uberyx and hindu, but differing from uberyx by the man-

dibles longer and with less convex sides and by the anterior face of

the petiolar node more protruding anteriorly, and from hindu by its

smaller size (TL 1.96 mm instead of 2.50 mm), by the smaller SI

(90.9 instead of SI = 102.6), and from both, uberyx and hindu, by

the standing hairs on the head dorsum, less numerous.

Worker description (Fig. 55). Head sides converging anteriorly,

with round vertexal corners. Frontal lobes weakly expanded and

convex. Antennal fossae ventrally with a marked carina visible in

full-face view, straight, covering the lower margin of the scrobes

and ending close to the eyes. Eyes small, with 2 ommatidia in the

longest row, placed over the ventral margin of the antennal scrobes,

and slightly visible in dorsal view. With the head in profile the

scrobe distinct, with superficially marked upper and lower margins.

Clypeal dorsum weakly concave medially. Lateral clypeal margin

160 C. BARONI URBANI & M.L. DE ANDRADE

gently converging anteriorly into a straight margin. Scapes subcylin-

drical, about 3/5 of the head length and surpassing the eyes posteri-

orly. Antennae with six segments. Apical funicular joint longer than

the rest of the funiculus. Mandibles diverging anteriorly and curved

on the two anterior thirds. Apical fork of the mandibles with two

spiniform teeth and with one intercalary denticle. Preapical denti-

tion consisting of one spiniform tooth near the apex.

Mesosoma in profile with weakly convex mesonotum and with a

longitudinal carina starting from the mesonotum and ending on the

anterior half of the basal face of the propodeum; the carina thicker

and shaped as a small triangular protuberance on the propodeum.

Posterior half of the propodeum declivous. Propodeal teeth pointed

and ventrally connected to a lamella.

Petiole with a long neck and with the node high, weakly convex

dorsally and broadly tumuliform anteriorly. Ventral surface of the

petiole with a broad spongiform lamina. Petiolar node with posterior

margin and posterior sides surrounded by spongiform process. Post-

petiole gently convex in profile. Anterior, lateral and posterior margin

of the postpetiole surrounded by spongiform processes. Ventral sur-

face of the postpetiole with large and dense spongiform process.

Gaster oval and with few, thin, very short costulae. Base of the

first gastral tergite and sternite with a spongiform pad larger on the

tergite.

Sculpture. Head minutely reticulate-punctuate. Pronotum, pleu-

rae and declivous face of the propodeum smooth; in addition the

pronotum with a few sparse punctures. Mesonotum, basal face of the

propodeum and petiole punctuate. Postpetiole and gaster smooth.

Pilosity. Head, mesosoma, petiole and postpetiole with

appressed hairs, denser on the petiole. Dorsolateral margin of the

head in full-face view with only the apicoscrobal flagellate hairs.

Cephalic dorsum with three rows of erect hairs, the first row with

4 hairs close to the occipital margin, the second row with 2 stand-

ing hairs and the third row with 2 standing hairs on the frons.

Upper scrobal margin with narrowly spoon-shaped hairs curved

anteriorly. Clypeal border with similar hairs as the upper scrobe.

Scapes with similar hairs as the upper scrobes and curved poste-

riorly. Pronotal humeral hairs long and flagellate. Dorsum of the

pronotum with a pair of erect, long flagellate hairs. Mesonotum

ANT TRIBE DACETINI 161

ig.

00050853 ant eee

00050592 en: GOO is a ee

55 - Strumigenys buddhista n. sp. from Nepal. Worker, head in dorsal view

(top) and entire profile (bottom).

162 C. BARONI URBANI & M.L. DE ANDRADE

with a pair of erect, flagellate hairs. Petiole, postpetiole and gaster

with few erect flagellate hairs. Outer face of mid and hind tibiae

with an erect, long, flagellate hair. Hind basitarsi with 1 pair of

erect, long, flagellate hairs.

Colour. light brown.

Measurements. (ine) aac indices: PE, 196; FE 050; HW 0:33) Si. 30;

Mik ‘0:29 -NET 0:02 10:54 ©h-66,0; (SÌ 90/9: MI 46.0;

Discussion. Strumigenys buddhista is very similar to

uberyx and hindu. "These three species have similar body sculpture

but the number of standing hairs on the head dorsum helps to sepa-

rate them. The three rows of standing hairs on the head dorsum

are placed as follows: 1 row close to the vertexal margin, 1 row at

the same line as the dorsolateral flagellate hairs and 1 row at the

level of the eyes. In buddhista the rows have 4,2,2 hairs, in uberyx

the rows have 4,4,4 hairs and in hindu the rows have 4,4,2 hairs.

Besides the differences in the number of standing hairs on the head

dorsum, buddhista differs from uberyx and hindu by the higher value

of MI (46 instead of MI 41-42 as in uberyx and hindu). In addi-

tion, buddhista and uberyx share the small size and buddhista and

hindu share the anterior face of the petiole protruding anteriorly and

tumuliform.

Strumigenys nageli Baroni Urbani & de Andrade n. sp.

Type material: holotype worker from Ecuador labelled: Esmeralda,

Rioverde, Via San Lorenzo, Km 67, leaf-litter, 22.VIII.2004, C. Baroni Urbani

& M. L. de Andrade (PUCE). Paratypes: 4 workers, same data as the holotype

(PUCE, MSNG).

Pres tvatio mominis: this species is named after Prof Dr Peter Nagel.

Diagnosis. A Strumigenys belonging to the elongata-group as

defined by BOLTON (2000), resembling spathula Lattke & Goitia

but differing from it by the apicoscrobal hairs short and spatulate

instead of long and flagellate, by the humeral hairs long and thicker

in their distal half, instead of long and filiform and by the spatulate

standing hairs on the gaster, longer and thinner.

Worker description (Fig. 56). Head sides converging anteriorly,

with round vertexal corners. Frontal lobes weakly expanded and

convex. Antennal fossae ventrally with a narrow carina visible in

ANT TRIBE DACETINI 163

full-face view, straight, not covering the lower margin of the scrobes

and ending close to the eyes. Eyes small, with 4 ommatidia in the

longest row, placed over the ventral margin of the antennal scrobes,

and visible in dorsal view. With head in profile the scrobe distinct,

with marked upper margin only. Clypeus triangular. Lateral clypeal

margin gently converging anteriorly to a straight or gently concave

margin. Scapes subcylindrical, about 2/3 of the head length and

largely surpassing the eyes posteriorly. Antennae with six joints.

Apical funicular joint much longer than the rest of the funiculus.

Mandibles elongate. Apical fork of the mandible with two spiniform

teeth and without intercalary denticles.

Mesosoma in profile weakly convex anteriorly and sloping

posteriorly to the convex basal face of the propodeum. Propodeal

teeth pointed. Declivous face of the propodeum with a very faint

margin.

Petiole with a long neck, the node high and convex dorsally.

Ventral surface of the petiole with a narrow lamina on the anterior

third. Petiolar node with posterior margin surrounded by spongi-

form process. Postpetiole convex in profile. Anterior and posterior

margins of the postpetiole surrounded by spongiform processes, the

process narrow on the anterior margin. Ventral surface of the post-

petiole with dense spongiform process.

Gaster oval and with thin, short costulae. Base of the first gastral

tergite and sternite with spongiform pad irregular on the sternite.

Sculpture. Head, mesosoma, petiole and postpetiole reticulate-

punctuate, this sculpture less marked on the postpetiole. Centre of

the lower mesopleurae smooth. Gaster smooth.

Pilosity. Head, mesosoma, petiole and postpetiole with subde-

cumbent or decumbent, spatulate hairs, very sparse on the meso-

soma and rare on the pedicel. Apicoscrobal hair spatulate. Cephalic

dorsum with one pair of standing spatulate hairs close to the occipi-

tal margin. Humeral hairs long, increasing in width from mid-length

to near the apex and apically gently pointed or truncate. Mesonotal

dorsum with 1 pair of erect hairs similar to the humeral ones but

shorter. Petiole, postpetiole and gaster with erect, long, spatulate

hairs very sparse on the pedicel.

164 C. BARONI URBANI & M.L. DE ANDRADE

000497980 zM B

Uni Basel

Fig. 56 - Strumigenys nageli n. sp. from Ecuador. Worker, head in dorsal view

(top) and entire profile (bottom).

ANT TRIBE DACETINI 165

Colour. brown with slightly darker gaster.

Measurements (in mm) and indices: TL 2.44-2.58; HL 0.63-0.66; HW 0.49-

0.50; SL 0.45; ML 0.38-0.40; EL 0.05-0.06; WL 0.60-0.65; CI 75.7-77.7; SI 90.0-

91.8; MI 60.3-60.6.

Discussion. S. nageli is very similar to spathula from

Venezuela, Trinidad and Central America but the characters listed

in the diagnosis permit easy separation of the two species.

Strumigenys aequinoctialis de Andrade n. sp.

Type material: holotype and paratype workers from Ecuador

(Pichincha) labelled: km 38 road Calacali - La Independencia, 21.VIII.2004, 2000

m, leaf litter, C. Baroni Urbani & M. L. de Andrade (MSNG).

Derivatio nominis: from the Latin aequinoctialis (=equinoctial), the

closest concept to the Equator in Roman times.

Diagnosis. A Strumigenys belonging to the schulzi-group as

defined by BOLTON (2000), resembling umboceps (Bolton), but dif-

fering from it by its standing hairs much longer, by its propodeal

spines longer, by its vertex with 4 longer, erect hairs, by its larger

size and darker colour.

Worker description (Fig. 57). Head strongly converging ante-

riorly and with round vertexal corners. Head in profile with poste-

rior half strongly tumuliform. Frontal lobes slightly expanded and

convex. Antennal fossae ventrally with a carina visible in full-face

view, straight, covering the lower margin of the scrobes and ending

before the upper border of the eye. Eyes with 4-5 ommatidia in the

longest row, placed over the ventral margin of the antennal scrobes.

With the head in profile the scrobe is superficial, with the upper

and lower margins weakly marked. Anterior clypeal margin broadly

convex. Scapes slightly compressed dorsoventrally, with sub-basal

bend, about half of the head length and surpassing the eyes posteri-

orly. Antennae with six segments. Apical funicular joint longer than

the rest of the funiculus. Mandibles short, triangular. Basal tooth

broad, subtriangular and followed by 11 teeth or denticles. Tooth 1

(basal), 2 and 3 the longest, subequal in size, tooth 4 smaller than

1-3, tooth 5 slightly shorter than teeth 1-3, teeth 6-10 diminishing

in size apically, apical tooh small but pointed.

166 C. BARONI URBANI & M.L. DE ANDRADE

Mesosoma in profile convex anteriorly and gently sloping poste-

riorly. Propodeal teeth pointed and subtended by a carina.

Petiole with node high and convex and with long peduncle. No

spongiform processes on the petiole, except a very thin whitish carina

on its posterior border. Postpetiole broadly convex, with developed

spongiform processes ventrally. Anterior and posterior postpetiolar

borders with a narrow spongiform crest, the crest broader posteriorly.

Gaster oval and with few, short costulae. Base of the first gas-

tral tergite with narrow limbus. Base of the first gastral sternite with

narrow spongiform pad.

Sculpture. Head, mesosoma and petiole reticulate-punctuate.

Mesopleurae largely smooth. Postpetiole and gaster smooth and

shining, except a few reticulations on the posterior half of the post-

petiolar dorsum.

Pilosity. Head, mesosoma and petiole with suberect or subdecum-

bent, spatulate hairs, slightly shorter on the clypeus, sparser on the

mesosoma and rare on the petiole. Apicoscrobal hair absent. Cephalic

dorsum with 4 long, erect hairs, slightly thicker on the apical half.

Leading edge of the scapes with spatulate hairs curved basally and api-

cally. Lateral clypeal margins with spatulate hairs curved anteriorly.

Pronotal humeral hair present, long and weakly flagellate. Sides of the

mesonotum with 1 erect, long, truncated, hair. Petiole, postpetiole,

and gaster with erect-suberect, truncated hairs, rare on the petiole.

Colour. Holotype dark brown; paratype reddish brown

with darker gaster.

Measurements (in mm) and indices: TL 2.30-2.60; HL 0.62-0.72; HW 0.45-

0.52; SL 0.31-0.36; ML 0.11-0.15; EL 0.06-0.09; WL 0.63-0.74; CI 72.2-72.6; SI

68.8-69.2; MI 17.7-20.8.

Discussion. $. aequinoctialis and its closest relative, umbo-

ceps (Bolton) occupy an isolate position within the genus for their

high-domed cephalic capsule. S. aequinoctialis is easily distinguished

from umboceps for the four long hairs behind the highest point of

the vertex as already described in the diagnosis. "The two aequinoc-

tialis specimens differ from the holotype unique of umboceps in the

MCZC also for the presence of standing hairs on the mesosoma.

Since there are differences in the number of hairs also between the

aequinoctialis holotype and paratype, one cannot exclude that the

umboceps unique had lost all its standing hairs during manipulation.

ANT TRIBE DACETINI 167

00050861 100 um vi BE,

00047415

Fig. 57 - Strumigenys aequinoctialis n. sp. from Ecuador. Worker, head in dorsal

view (top) and entire profile (bottom).

168 C. BARONI URBANI & M.L. DE ANDRADE

+Strumigenys pilosula de Andrade n. sp.

Type material: holotype, winged gyne (unique) in Dominican amber

(GOPC) No. H 10-190.

Derivatio nominis: pilosulus is the diminutive of the Latin adjective

pilosus (= hairy) and is referred to the smaller number of macrochaetae that

differentiate this species from its closest relative, S. schleeorum Baroni Urbani.

Diagnosis. A Strumigenys species resembling the fossil

schleeorum but differing from it by having 4 standing hairs close to

the vertexal margin instead of 2, by a pair of standing hairs close to

the ocelli (no such hairs in schleeorum), and by the leading edge of

scape without freely projecting hairs instead of with freely project-

ing hairs.

Gyne description (Figs. 58, 59, 60). Head strongly converging

anteriorly and with round vertexal corners. Frontal lobes slightly

expanded and convex. Antennal fossae ventrally with a marked

carina visible in full-face view, straight, covering the lower margin

of the scrobes and ending close to the upper border of the com-

pound eyes. Compound eyes large, very protruding, occupying

large part of the posterior half of the antennal scrobe, and largely

visible in dorsal view. Ocelli developed. With the head in profile

the scrobes are superficial, with the upper margin indistinct and

the lower one marked anteriorly only. Lateral clypeal margin con-

verging anteriorly into an anterior convex margin weakly tumu-

liform medially. Scapes weakly dorsoventrally compressed, with

sub-round bend, slightly less than 1/2 of the head length and

slightly surpassing the eyes posteriorly. Antennae with six joints.

Apical funicular joint slightly longer than the rest of the funiculus.

Mandibles about 1/2 of the head length, touching each other only

at the apex when opposed, with slightly convex external borders.

Internal border of the mandibles with a broad basal lamella fol-

lowed by a row of minute, poorly differentiated denticles and by

a single spiniform tooth. Labral lobes large and partially visible

between the mandibles.

Mesosoma robust and gently convex in profile. Parapsidal fur-

rows weakly impressed. Scutellum with the sides converging pos-

teriorly and with the posterior border rounded. Basal face of the

propodeum long and declivous posteriorly; its sides marginate and

its dorsum gently concave. Area between basal and declivous faces

ANT TRIBE |DACETINI

169

58 - Strumigenys pilosula n. sp. from Dominican amber. Gyne,

tance between two scale bars 0.1 mm.

profile. Dis-

170 C. BARONI URBANI & M.L. DE ANDRADE

Fig. 59 - Strumigenys pilosula n. sp. from Dominican amber. Gyne, head in dorsal

view. Distance between two scale bars 0.1 mm.

ANT TRIBE DACETIN BAI

Fig. 60 - Strumigenys pilosula n. sp. from Dominican amber. Gyne, head and

mesosoma in profile showing the pilosity. Distance between two scale

bars 0.1 mm.

wor C. BARONI URBANI & M.L. DE ANDRADE

of the propodeum with a lamellaceous, pointed, transparent tooth

that prolongs to the declivous face as a broad lamella.

Petiole with a long neck and with the node high and convex.

Ventral surface of the petiole without spongiform lamina. Petiolar

node with marked posterior margin and without spongiform process.

Postpetiole convex in profile and without spongiform processes.

Gaster oval. Base of the first gastral tergite with developed ante-

rior transverse cuticular ridge.

Sculpture. Head, mesosoma, petiole and postpetiole minutely

reticulate, the reticulation more superficial on the postpetiolar

dorsum. Mesopleurae, lower metapleurae and gaster smooth.

Pilosity. Dorsum of the head and scapes with sparse, weakly

remiform hairs, subdecumbent on the head and appressed on the

scapes. Leading edge of the scapes without free hairs. Border of

the clypeus with few, free, weakly remiform hairs pointed medially.

Apicoscrobal hair slightly pointed. Cephalic dorsum with 4 standing

hairs close to the vertexal margin and a pair of hairs close to the

ocelli. Pronotal humeral hairs long. Sides of the mesonotum with

2 pairs of hairs, the anterior pair much longer and thicker than the

posterior one. Dorsum of the mesonotum with 2 pairs of erect hairs,

one pair close to the anterior border and the second close to the

parapsidal furrows. Petiole with a pair of pointed hairs dorso-lat-

erally. Postpetiole with 2 pairs of pointed hairs, the anterior pair

dorso-lateral and longer, the posterior pair dorsal and close to the

posterior border. Ventral surface of the postpetiole with a thick hair.

Gaster with few erect pointed hairs.

Cov oar dight brown.

Measurements (in. nam) and indices: 'TL22.30; HE 0.58; HW 0.30; SL 027;

ML 0.14; EL 0.14; WL 0.67; CI 51.7; SI 90.0; MI 24.1.

Widierial examaned, ‘The unique pilosula example as

embedded in a small, cut and polished, yellowish amber sample 0.5

x 0.5 x 1.5 cm containing only this specimen. ‘The amber sample

bears the No. H 10-190 in the GOPC. The preservation conditions

are good, although the right side of the ant is slightly flattened.

Discussion. S. pilosula resembles another previously

known Dominican fossil, S. schleeorum (compare the figures of the

present study with Figs. 22 & 23 by BARONI URBANI & DE ANDRADE,

1994). Both species share a peculiar mandibular shape, with rela-

ANT TRIBE DACETINI 173

tively short, curved and pointed mandibles without differentiated

apical teeth touching each other only at the apex when closed. This

structure is unknown among extant Strumigenys and appears to

characterize a now extinct Dominican amber clade.

TStrumigenys poinari Baroni Urbani & de Andrade n. sp.

Type material: holotype worker in Dominican amber (GOPC No. H

10-220). Paratypes: 4 workers in the same amber piece and collection as the holotype.

Derivatio nominis: this species is named after Dr. George O. Poinar,

Jr. who permitted us to study this and many more interesting amber samples.

Diagnosis. A Strumigenys belonging to the rostrata-group

as defined by BOLTON (2000), resembling carolinensis (Brown), but

differing from it by the presence of 4 suberect hairs on the vertexal

margin, by the pronotal humeral hair gently spatulate instead of

long and flagellate, and by the hind basitarsi without flagellate hairs.

Worker description (Figs. 61 & 62). Head strongly converging

anteriorly and with round vertexal corners. Frontal lobes slightly

expanded and convex. Antennal fossae ventrally with a carina visible

in full-face view, straight, covering the lower margin of the scrobes

and ending before the upper border of the eye. Eyes with about

5 ommatidia in the longest row, placed over the ventral margin

of the antennal scrobes. With the head in profile the scrobes are

distinct, with the upper and lower margins superficially marked.

Lateral clypeal margins gently converging anteriorly into a convex

margin. Scapes slightly compressed dorsoventrally, with sub-basal

bend gently convex, slightly less than 1/2 of the head length and

surpassing the eyes posteriorly. Antennae with six segments. Apical

funicular joint longer than the rest of the funiculus. Mandibles

short. Internal border of the mandibles with broad and triangular

basal lamella followed by 12 teeth or denticles. Tooth 1 (basal) and

2 small, subequal in size, tooth 3 the longest, tooth 4 half size than

3, tooth 5 slightly shorter than 3, teeth 6 and 7 similar to 1 and 2

and followed by 4 denticles and by a small apical tooth.

Mesosoma in profile gently sloping posteriorly. A short lon-

gitudinal ruga runs on the dorsum of the pronotum and mesono-

tum. Propodeal teeth large, triangular and subtended by a broad

lamella.

174 C. BARONI URBANI & M.L. DE ANDRADE

Fig. 61 - Strumigenys poinari n. sp., Dominican amber fragment including the

holotype (A) and paratypes B-D (top) and paratype E at another location

in the same amber fragment (bottom). Distance between two scale bars

0.1 mm.

ANT TRIBE DACETINI 175

Fig. 62 - Strumigenys poinari n. sp. from Dominican amber. Holotype worker in

profile. Distance between two scale bars 0.1 mm.

Petiole with node high and convex and with developed spongiform

processes. Ventral surface of the petiole with spongiform lamina. Post-

petiole convex in profile and with developed spongiform processes.

Gaster oval and with few, short costulae. Base of the first gas-

tral tergite with broad limbus. Base of the first gastral sternite with

spongiform pad.

Sculpture. Head, mesosoma and petiole reticulate-punctuate, the

reticulation-punctuation larger on the head. In addition the mesono-

tum with sparse, very thin, longitudinal rugosities, much sparser

on the mesonotum and propodeum. Mesopleurae and metapleurae

largely smooth and shining. Dorsum of the postpetiole minutely

punctuate and superficially shining. Gaster smooth and shining.

Pilosity. Head and mesosoma with subdecumbent or decumbent,

spatulate hairs, rarer on the mesosoma. Apicoscrobal hair absent.

176 C. BARONI URBANI & M.L. DE ANDRADE

Cephalic dorsum with 4 suberect, spatulate hairs close to the ver-

texal margin. Leading edge of the scapes with spatulate hairs curved

basally and apically. Lateral clypeal margins with spatulate hairs

curved anteriorly. Pronotal humeral hair short and slightly spatulate.

Sides of the mesonotum with 1 erect, curved, thick, flagellate hair.

Petiole, postpetiole and first gastral tergite with sparse, curved, thick

flagellate hairs. First gastral sternites with suberect spatulate hairs.

Cot Gr 5? dark brown:

Measurements (in mm) and indices: TL 2.15-2.20; HL 0.58-0.59; HW 0.39-

0.40; SL 0.27-0.28; ML 0.13; EL 0.05; WL 0.54-0.56; CI 66.1-69.0; SI 67.5-71.8;

MI 22.0-22.4.

Material examined. Holotype and 4 paratype work-

ers, all embedded in the same yellow amber sample 1.6 x 2.1 x 0.7

cm containing 5 workers of Strumigenys, a Diplorhoptrum worker

(gaster and large part of postpetiole and right legs missing), an

unidentified small insect and many debris. GOPC H 10-220. The

preservation conditions of the Strumigenys specimens are good.

Discussion. S. poinart resembles S. carolinensis in gen-

eral habitus and mandibular dentition but the two species can be

easily separate on the basis of the pilosity (see the diagnosis). S.

carolinensis is known only from North and South Carolina and

from Florida. Clearly Nearctic relationships are an uncommon trait

among Dominican amber ants. There are no obvious relationships

between JS. poinari and two other Strumigenys previously described

from Dominican amber (S. pilosula de Andrade, present paper, and

S. schleeorum Baroni Urbani & de Andrade, (BARONI URBANI &

ANDRADE, 1994)). On the contrary S. poinari and the third known

Dominican fossil Strumigenys, S. electrina de Andrade might belong

to a unique small clade. The two species, however, differ from each

other in a number of details like presence of apicoscrobal hairs (elec-

trina only), larger size of poinart, etc.

7. ACKNOWLEDGEMENTS

We are particularly grateful to several persons who facilitated

our work in several different ways. Prof Giovanni Onore, Juan Vieira

and David Donoso of the Pontificia Universidad Catélica of Quito

generously offered material and multiple helps during our collecting

ANT TRIBE DACETINI 177

in Ecuador. This fresh material represents the primary source for

the dissections used in this paper.

Daniel Mathys and Marcel Diggelin of the Microscopy Labo-

ratory of the Basle University made with their usual know how and

patience all the SEM micrographs.

Particularly warm thanks are due to Dr Roberto Poggi for his

patience and competence in editing our manuscript.

The help of the following collection curators was essential to the

completion of our study.

ANIC, Dr Stephen O. Shattuck.

FMNH, Mr Philip P. Parrillo.

GOPC, Dr: George O; Poinar, Jr.

MHNG, Dr Bernhard Merz.

MIZA, Dr John Lattke.

NHMB, Dr Michel Brancucci and Dr Daniel Burckhardt.

PUCE, Prof Dr Giovanni Onore. Deposited in PUCE are also

the ant collections of David Donoso and Juan Manuel Vieira Correa.

We are particularly grateful to these two young and enthusiastic

myrmecologists for their offer of material for study and for several

invaluable helps during our fieldwork in Ecuador.

Moreover, we feel obliged to especially recognize the help

received from certain persons in studying material kept in some

institutions where we encountered previously unknown difficulties:

BMNH, Prof Dr Quentin Wheeler and Mr Stuart Hime. We

addressed our first requests to examine part of the material studied

by Mr Bolton to Ms Ryder and to Mr Hime but in vain. It was

only after writing to Professor Wheeler that we obtained a reply and

the ants from Mr Hime.

MCZC, Mr Stefan P. Cover, who, in order to send us speci-

mens from the Harvard collection, wrestled for over one year with

new restrictions from the Museum (e. g. loan of specimens for the

sole comparison purpose is now prohibited) and from U. S. federal

directives (e.g. technical hitches in sending parcels after the World

Trade Center September 11, 2001 attacks; the Museum violating

new regulations from the US Department of Fish and Wildlife), and

others. Also a mail to Prof E. O. Wilson remained unanswered on

this subject. Other U. S. institutions contacted by us at the same

time appeared to be not affected by these restrictions.

178 C. BARONI URBANI & M.L. DE ANDRADE

MZSP, Prof Dr C. R. F. Brandào, who, in spite of his com-

mitments as Director of the Museu de Zoologia in San Paulo, was

able to organize the sending of the specimens under his care after

six months confrontation with new Brazilian Federal laws imposing

unparalleled bureaucratic formalities.

If, on one hand, we can only repeat our gratitude to all those

who spent their time in order to permit us to study specimens under

their care, on the other we consider the bureaucratic and practical

difficulties encountered somewhere as perilous for the free access to

scientific information and we wish that the authorities responsible

of them may undertake as soon as possible the necessary steps to

improve the situation faced by us.

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9: ABSTRACT

The authors perform a critical analysis of potentially related genera currently

included or excluded from the tribe Dacetini and of the characters used to justify

inclusions or exclusions in these and other, related family—group taxa.

A major methodological difference between the present study and the analysis

of BoLTON (1999) will explain the diversity of results. For the present study a trait is

ANT TRIBE DACETINI 189

recorded as present, absent, or polymorphic in a given taxon according to its observed

presence or absence. BOLTON (1999) analysis does not consider known presences or

absences when he thinks that these are due to homoplasy. This favours subjective

judgment and, in this way, it might allow construction and defence of countless

hypothetical clades including species without the synapomorphies characteristic of the

clade since these relevant characters may be supposed to have been secondarily lost.

Among the results of the morphological analysis, the so called “mesopleural

gland” supposed to be characteristic of the Dacetini is described as constituted by

differently developed brushes of hair beds (sensilla trichoidea) widespread among

several ant genera and unrelated to visible glandular structures.

A cladistically constructed phylogeny of the genera examined is supported by

few undoubted synapomorphies and is considered mainly as a decisional criterion

to build a classification based on unequivocal characters only, a condition entirely

missing in the current classification.

The following changes result as a consequence of this analysis and of the

principle that at least one unequivocal apomorphy must characterize a taxon:

The subfamily Agroecomyrmecinae is a junior synonym of Myrmicinae. Its

downgraded status as a separate tribe, Agroecomyrmecini, sister tribe of Dacetini

within the Myrmicinae, is maintained provisionally to preserve nomenclatorial

continuity. As a matter of fact the Dacetini alone appear much more ill defined

than a broader tribe including Agroecomyrmecini+Dacetini.

The tribal names Basicerotini and Phalacromyrmecini are considered as

synonyms of Dacetini.

A number of genera already considered as synonyms of Strumigenys by BARONI

URBANI & DE ANDRADE (1994), emended by BoLTON (1995), and considered again

as synonyms of a single emended name by BOLTON (1999) are re-established as

Strumigenys’ synonyms. These are: Pyramica, Epitritus, Trichoscapa, Cephaloxys,

Pentastruma, Glamyromyrmex, Codiomyrmex, Tingimyrmex, Codioxenus, Smithistruma,

Weberistruma, Wessonistruma, Serrastruma, Neostruma, Dorisidris, Miccostruma,

Kyidris, Polyhomoa, Chelystruma, Borgmeierita, Platystruma, Gymnomyrmex,

Dysedrognathus, Asketogenys, Cladarogenys. 'Vhe following genera are new synonyms

of Epopostruma: Colobostruma, Alistruma, Clarkistruma, Mesostruma. The following

genera are new synonyms of Basiceros: Rhopalothrix, Heptastruma, Acanthidris,

Octostruma, Talaridris, Eurhopalothrix, Protalaridris.

The following species are described as new: Basiceros onorei from Ecuador,

B. papuanum from Papua New Guinea, Strumigenys veddha from Sri Lanka, S.

aduncomala from India, S. caniophanoides from Bhutan, S. hindu from Nepal, S.

buddhista from Nepal, S. nageli from Ecuador, S. onorei from Ecuador, S. longimala

from Ecuador, S. aequinoctialis from Ecuador, S. pilosula from Dominican amber

(Miocene), S. poinari from Dominican amber (Miocene).

10. RIASSUNTO

La tribù dei Dacetini: limiti e generi che la compongono, con descrizioni di

specie nuove (Hymenoptera, Formicidae).

Si analizzano criticamente alcuni generi attualmente inclusi od esclusi dalla

tribù dei Dacetini ed i caratteri impiegati per giustificare le rispettive inclusioni od

esclusioni in questo ed altri prossimi taxa di livello sopragenerico.

190 C. BARONI URBANI & M.L. DE ANDRADE

Una cospicua differenza metodologica tra questo studio e quello di BoLTON

(1999) spiega largamente la diversità dei risultati. In questo studio ciascun carattere

è stato considerato presente, assente o polimorfico in un taxon a seconda delle

presenze ed assenze realmente osservate. BoLTON (1999), al contrario, non tiene

cura delle presenze ed assenze da lui ritenute a priori risultato di convergenza o di

perdita secondaria. Questo modo di vedere favorisce la valutazione soggettiva e, di

conseguenza, permette la creazione e la difesa d’innumerevoli taxa comprendenti

specie in cui le apomorfie del taxon in questione sono assenti perché quest'ultime

potrebbero essere state perdute secondariamente.

Tra i risultati delle osservazioni morfologiche è da notare che la cosiddetta

“ghiandola mesopleurale”, correntemente considerata caratteristica di molti Dacetini,

viene descritta come costituita da gruppi di sensilli tricoidei presenti in molte

formiche disparate e dissociati da strutture ghiandolari visibili.

Un’analisi di parsimonia dei dati morfologici permette la costruzione di un

cladogramma dei generi studiati, cladogramma giustificato da poche sinapomorfie

indiscutibili e proposto essenzialmente come criterio decisionale per costruire una

classificazione del gruppo non equivoca, una caratteristica del tutto assente nella

classificazione corrente.

Le seguenti modifiche classificatorie e nomenclatoriali conseguono direttamente

dall’analisi del cladogramma e dall’applicazione del principio per cui un taxon deve

essere caratterizzato da almeno un’apomorfia inequivocabile.

La sottofamiglia Agroecomyrmecinae è un sinonimo juniore di Myrmicinae. Il

suo stato subordinato di tribù gemella dei Dacetini all’interno dei Myrmicinae viene

mantenuto provvisoriamente per conservare una certa continuità nomenclatoriale. I

Dacetini da soli, infatti, appaiono molto più superficialmente caratterizzati di una

tribù più ampia comprendente gli attuali Dacetini + Agroecomyrmecini.

Le tribù Basicerotini e Phalacromyrmecini vengono considerate sinonimi

dei Dacetini. Numerosi generi già considerati sinonimi di Strumigenys da BARONI

URBANI & DE ANDRADE (1994) e resuscitati da BOLTON (1995) e poi considerati

sinonimi di un altro genere resuscitato da BOLTON (1999) sono riproposti come

sinonimi di Strumigenys. Questi sono: Pyramica, Epitritus, Trichoscapa, Cephaloxys,

Pentastruma, Glamyromyrmex, Codiomyrmex, Tingimyrmex, Codioxenus, Smithistruma,

Weberistruma, Wessonistruma, Serrastruma, Neostruma, Dorisidris, Miccostruma,

Kyidris, Polyhomoa, Chelystruma, Borgmeierita, Platystruma, Gymnomyrmex,

Dysedrognathus, Asketogenys, Cladarogenys. 1 generi seguenti sono nuovi sinonimi

di Epopostruma: Colobostruma, Alistruma, Clarkistruma, Mesostruma. I generi

seguenti sono nuovi sinonimi di Basiceros: Rhopalothrix, Heptastruma, Acanthidris,

Octostruma, Talaridris, Eurhopalothrix, Protalaridris.

Le specie seguenti, studiate per la presente revisione, vengono descritte

come nuove: Basiceros onorei dell’ Ecuador, B. papuanum di Papua Nuova Guinea,

Strumigenys veddha dello Sri Lanka, S. aduncomala dell’ India, S. caniophanoides del

Bhutan, S. hindu del Nepal, S. buddhista del Nepal, S. nageli dell Ecuador, S. onoret

dell’ Ecuador, S. longimala dell'Ecuador, S. aequinoctialis dell’ Ecuador, S. pilosula

dell’ambra dominicana (Miocene), S. poinari dell ambra dominicana (Miocene).

ANT TRIBE DACETINI

CONTENTS

Introduction

Material and methods

. Outgroup comparison

Character choice and taxonomic rank

Results

. List of characters and of their recorded states

. Characters used in other phylogenetic studies and excluded

from the present one

. Results of the cladistic analysis

. Family-group systematics

. Genus-group systematics

Key to the extant genera of Agroecomyrmecini and Dacetini

Species-level additions

Acknowledgements

References

Abstract

Riassunto

191

133

176

178

188

189

£98

TARUN K. PAL*

ON A COLLECTION OF HETEROMERA AND CUCUJOIDEA

OF THE MUSEO CIVICO DI STORIA NATURALE

“G. DORIA”, GENOVA

(COLEOPTERA; POLYPHAGA)

INTRODUCTION

I received a collection of undetermined beetles from the Museo

Civico di Storia Naturale “G. Doria”, Genova which had been col-

lected in continental and insular Africa, South and Southeast Asia,

Pacific Islands etc. more than one century ago.

Several species of Heteromera and Cucujoidea were amongst

this material, including two new species of Inopeplus Smith and one

new species for each of the genera Indoleptipsius Pal, Aphanocephalus

Wollaston, Parafallia Arrow, Paramaschema Heller, Cephalophanus

John and Euxestus Wollaston.

The inventory of the material and descriptions of the eight new

species are presented in this work. The material examined is now

housed in the following institutions:

MSNG - Museo Civico di Storia Naturale “G. Doria”, Genova;

ZSI - Zoological Survey of India, Kolkata.

SYSTEMATIC ACCOUNT

Order COLEOPTERA

Suborder POLYPHAGA

Superfamily HETEROMERA

* Zoological Survey of India, M-Block, New Alipore, Kolkata- 700 053, India.

E-mail: tkpal51@rediffmail.com

194 T.K. PAL

Family OTHNIIDAE

Genus Ocholissa Pascoe

Ocholissa laticeps Grouvelle

Ocholissa laticeps Grouvelle, 1908, Annls. Soc. ent. Fr., 77: 410.

Material: Birmania, Bhamò, X.1886, L. Fea, 1 ex. (MSNG);

Tenasserim, Thagatà, 1V.1887, L. Fea, 1 ex. (MSNG); Giava, Tci-

bodas, X.1874, O. Beccari, 1 ex. (ZSI).

Size: length 3.05-3.08 mm.

Distribution: India (Karnatak); Myanmar (new record); Indo-

nesia (new record).

Family INOPEPLIDAE

Genus Inopeplus Smith

Inopeplus biocellatus (Motschulsky)

Euryplatys biocellatus Motschulsky, 1859, Etud. ent., 8: 98.

Inopeplus biocellatus: Grouvelle, 1908, Annls. Soc. ent. Fr.,77: 462; Sengupta, Pal &

Mukhopadhyay, 1977, Orient. Ins., 11: 399.

Material: Mentawei, Sipora, Sereinu, V-VI.1894, E. Modigliani,

1 ex. (MSNG); Mentawei, Si Oban, IV-VIII.1894, E. Modigliani,

1 ex. (ZSI).

Size: length 3.80-4.50 mm.

Distribution: India (Arunachal Pradesh, Meghalaya, West

Bengal); Indonesia: Java (new record).

Inopeplus pacificus n. sp.

General appearance (Fig. 1) elongated, flattened, shiny, black-

ish, elytra with pale orbicular spot, last four abdominal segments

(occasionally three) exposed, mouthparts partially exposed.

Head broader than long, apical margin truncate, fronto-clypeal

eo Ces

* Casale

Agra

A COLLECTION OF HETEROMERA AND CUCUJOIDEA

1.0 mm.).

Fig. 1 - Inopeplus pacificus n. sp., dorsal view (scale

196 TK PAD

suture distinct and nearly straight, apical margin of frons with

transverse depression; no longitudinal impressed line on vertex, a

semicircular depression on vertex surrounding inner margin of eye

less distinct, a short oblique depression arising from orbital margin;

puncturation on vertex fine, rounded, moderately dense, separated

by 2-4 diameter; eyes moderately large and finely facetted. Antenna

moderately long and slender, scape moderately large and curved,

pedicel shorter and narrower than scape, segment 3 more or less

similar in proportion to pedicel, segments 4-9 subequal, about as

broad as long and slightly wider than 3, segment 10 slightly elon-

gate, segment 11 elongate and acuminate at apex; scape and pedicel

reddish-brown and segment 3-11 dark brown.

Prothorax transverse, flattened, widest beyond middle and

markedly narrowed towards base; lateral margin smooth, distinctly

sinuate in posterior third, finely bordered from base to posterior

two-thirds; small paired depressions on posterior third of pronotal

disc; puncturation on pronotum roundish, finer and sparser than on

vertex, separated by about 3-6 diameter.

Scutellum transverse, rounded apically and impunctate.

Elytra broader than long, widest near apex, puncturation fine,

slightly sparser than on pronotum; a rounded pale spot on posterior

half of each elytron and the border of which less distinct, last four

(occasionally three) abdominal segments exposed.

Ventral surface blackish, shiny.

Aedeagus (Fig. 3 a,b) somewhat trilobed-type; median lobe

broad, elongate with tubular tip, basal end moderately dilated;

tegmen forming a ring with large dorsal cap, basal angles produced

into paired struts; parameres elongate, feebly bilobed, outer lobe

little narrower and apically more extended, a few setae on either lobe.

Measurements of holotype: total length 2.76 mm, width of head

across eyes 0.67 mm, length of antenna 1.03 mm, length and width

of prothorax 0.42 mm and 0.67 mm, length and width of elytra 0.96

mm and 1.05 mm.

Holotype: Is. Goodenough, 1.1890, L. Loria (MSNG).

Paratypes:. same data as holotype, 10 ex. (8: MSNG, 2: ZSI);

N. Guinea mer., Waicunina, VI.1890, L. Loria, 3 ex. (2: MSNG,

1: ZSI); Mentawei, Si Oban, IV-VIII.1894, E. Modigliani, 2 ex.

(Zain

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 197

Etymology: the species-name refers to its occurrence in

the Islands in the Pacific Ocean.

Remarks: the species comes close to an insular species,

I. andamanicus Pal & Datta, 1982 but can be distinguished by the

presence of paired admedian depressions on basal third of prono-

tum, a suborbicular pale spot near middle of either elytron, eyes

comparatively longer and tempora of head not longer than eyes. The

aedeagus of J. andamanicus is not studied and hence could not be

compared with J. pacificus.

Inopeplus sthulalinga n. sp.

General appearance (Fig. 2) elongated, flattened, shiny, choco-

late-brown, unicolourous, last three abdominal segments (occasion-

ally four) exposed, mouthparts partially exposed.

Head broader than long, apical margin truncate, fronto-clypeal

suture distinct and nearly straight, apical margin of frons with trans-

verse depression, no longitudinal impressed line on vertex; punctur-

ation on vertex coarse, rounded to little elongate, moderately dense,

separated by 1-3 diameter; eyes moderately large and moderately

coarsely facetted, orbital margin of vertex narrowly depressed, no

oblique depression arises from orbital margin, tempora longer than

eyes. Antenna moderately long and slender, scape moderately large

and curved, pedicel shorter and narrower than scape, segment 3

more or less similar in proportion to pedicel, segments 4-10 about

as broad as long and slightly transverse, broader than preceding

segment and feebly wider towards apex, segment 11 elongate and

broadly pointed apicad.

Prothorax slightly elongate, flattened, widest little behind front

across anterior denticles and narrowed towards base; lateral margin

with one anterior and two posterior denticles, finely bordered from

base to anterior denticles; puncturation on pronotum roundish, finer

and sparser than on vertex, slightly denser laterally, separated by

about 1.5-4 diameter.

Scutellum transverse, rounded apically and impunctate.

Elytra about as broad as long, broadened posteriorly, glabrous,

an outwardly directed metathoracic spine present near middle of

198 T.K. PAL

Fig. 2 - Inopeplus sthulalinga n. sp., dorsal view (scale = 1.0 mm.).

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 199

elytra, last three (occasionally four) abdominal segments exposed.

Ventral surface brown, shiny.

Aedeagus (Fig. 3 c, d) somewhat trilobed-type; median lobe

broad, elongate, somewhat broader apically, basal end dilated;

tegmen forming a ring with large dorsal cap, base on sides produced

into paired struts; parameres short, elongate, apex broadly rounded

with a few setae. |

Measurements of holotype: total length 2.72 mm, width of head

across eyes 0.63 mm, length of antenna 0.94 mm, length and width

of prothorax 0.48 mm and 0.44 mm, length and width of elytra 1.00

mm and 0.93 mm.

Holotype: Is. S. Thomé, Agua-Izé, 400-700 m, XII.1900, L.

Fea (MSNG).

Paratypes: same data as holotype, 3 ex. (MSNG); Is. Principe,

Roca Inf. D. Henrique, I-IV.1901, 200-300 m, L. Fea, 11 ex. (7:

MSNG, 4: ZSI); Is. Principe, Roca Inf. D. Henrique, V.1901, L.

Fea, 2 ex. (MSNG).

Etymology: the specific-name is the Bengali adjective

‘sthulalinga’, that refers to the broad penis (median lobe) of male

of this species.

Remarks: This species shows apparent similarity with J.

patkoicus Pal, 1992 (northeast India) in prothoracic denticulation but

can be differentiated by slightly elongate prothorax (vs. transverse

in patkoicus), impunctate and immaculate elytra (vs. finely punctate

elytra with a pair of ovoid spots in basal half in patkoicus); apex of

median lobe of aedeagus broad and parameres distinctly different.

Superfamily CUCUJOIDEA

Family MONOTOMIDAE

Genus Mimemodes Reitter

Mimemodes sp.

Material: New Guinea, Dilo, VI-VII.1890, L. Loria, 1 ex.

(MSNG).

200 T.K. PAL

Fig. 3 - Aedeagi. a, Inopeplus pacificus n. sp., dorsal view; b, [nopeplus pacificus n.

sp., enlarged view of right paramere; c, Inopeplus sthulalinga n. sp., dorsal

view; d, Inopeplus sthulalinga n. sp., enlarged view of right paramere.

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 201

Size: length 2.70 mm.

Remarks: this species shows certain resemblances with

one Indian species, M. kimbhutus Sengupta, 1976, but differs by its

longer tempora (more than half as long as eye) and larger eyes. The

long and distinct tempora of this species is not often seen in other

species of Mimemodes.

Genus Indoleptipsius Pal

Indoleptipsius poggii n. sp.

General appearance (Fig. 4) elongated, moderately depressed,

mandibles partly exposed, eyes non-projecting, antennal club 1-seg-

mented, yellowish-brown, dorsal puncturation moderately coarse,

little shiny and almost glabrous, elytra exposes one tergite of abdomen.

Head transverse, widest across eyes; eyes moderately large and

about one-third as long as head, moderately coarsely facetted; no

distinct sculpture on dorsal side except oblique, linear supra-anten-

nal depressions; puncturation on vertex moderately coarse and less

dense, separated by about 2-3 diameter, frontal area rather finely

punctate. Antenna slightly longer than head, scape moderately large

and broadly elongate, pedicel short and narrower than scape, seg-

ments 3-8 narrower, subequal and more or less transverse, segment

9 little wider than preceding segment; segment 10 (club) slightly

elongate and about as long as preceding four segments.

Prothorax about as broad as long, slightly narrower posteriorly,

front angles obtuse, hind angles rounded and not well marked, sides

slightly uneven with minute notches at intervals; disc with a median

elongate impunctate area which broadened posterad, puncturation on

disc moderately coarse and not dense, separated by about 1.5-3 diameter.

Scutellum elongate, glabrous and apical margin rounded.

Elytra about 1.7x as long as broad, sides feebly arched, apex of

each elytron separately rounded, moderately large punctures in eight

regular rows along fine impressed strial lines, interstices not raised

or ribbed; pygidium with moderately coarse and sparse punctures,

sparsely pubescent.

On ventral side prosternum and metasternum medially impunc-

T.K. PAL

202

TETTE rene ont.

aS Agen Aerts»: hE in he hen reg neyo

; ogee’ © ORM o ote no

ARR Gp pent ecg >a WE

6 POO OOOOH OEE OT ci 3: 4 te.

0-00 Oo Oe oe Orrin g Jo “eg

<P 4-90 terre © ced

wie

RAR, ee LS eS

1.0 mm.).

Fig. 4 - Indoleptipsius poggit n. sp., dorsal view (scale

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 203

tate, sparse punctures only on sides; abdominal ventrites I-IV

impunctate medially and sparsely punctate on sides, last ventrite

sparsely punctate-pubescent.

Aedeagus (Fig. 5) uninverted cucujoid-type; median lobe short

and broad, median orifice at tip, basal foramen large; from the ven-

tral edge of foramen arises a ring and paired long slender struts

come out of this ring; tegmen forming a ring-piece and placed on

ventral side of the struts.

Fig. 5 - Indoleptipsius poggii n. sp., aedeagus, dorsal view.

204 TRS PAW

Measurements of holotype: total length 1.60 mm, width of head

across eyes 0.38 mm, length of antenna 0.39 mm, length and width

of prothorax 0.40 mm and 0.40 mm, length and width of elytra 0.80

mm and 0.46 mm.

Holotype d: New Guinea, Dilo, VI-VII.1890, L. Loria

(MSNG).

Paratypes: same data as holotype, 11 ex. (9: MSNG, 2: ZSI);

New Guinea, Ighibirei, VII-VIII.1890, L. Loria, 6 ex. (MSNG);

New Guinea mer., Bujakori, VIII.1890, L. Loria, 2 ex. (ZSI); New

Guinea S.E., M.ti Astrolabe, II. 1893, L. Loria, 7 ex. (6: MSNG,

1: ZSI); New Guinea, Ramoi, II.1875, O. Beccari, 2 ex. (MSNG);

New Guinea, Hatam, VII.1875, O. Beccari, 1 ex. (MSNG).

Etymology: a patronym for Dr. Roberto Poggi, Director,

Museo Civico di Storia Naturale, Genova, in honour of his contri-

bution to the systematics of beetles, and for his generous support to

the present study.

Remarks: this species differs from the other lone species

of the genus, J. ushae Pal, 1996a by the sides of prothorax almost

straight (vs. slightly arcuate in ushae), indistinctly serrated and serra-

tion notches distantly placed (vs. distinctly serrated with closely set

serrulations in ushae), puncturation on vertex and pronotum finer

and pronotum much less punctate with a medially impunctate area.

Genus Thione Sharp

Thione sp.

Material: Borneo, Sarawak, 1865-66, coll. G. Doria, 1 ex.; Isole

Key, 1873, O. Beccari, 1 ex. (MSNG).

Size: length 4.00 mm - 4.48 mm.

Genus Shoguna Lewis

Shoguna feae Grouvelle

Shoguna feae Grouvelle, 1896, Annali Mus. civ. St. nat. Genova, (2) 16 (=36): 41.

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 205

Material: Borneo, Sarawak, 1865-66, coll. G. Doria, 1 ex.

(MSNG).

Size: length 4.60.

Distribution: Myanmar; Indonesia (Sumatra, Borneo). I have

examined specimens also from Andaman Is., India.

Family DISCOLOMIDAE

Genus Aphanocephalus Wollaston

Aphanocephalus naius n. sp.

General appearance (Fig. 6 a) oblong, ovoid, about 1.28x as long

as broad, convex dorsally and flattened ventrally, shiny, pronotum and

elytra glabrous, puncturation obscure; dark brown except narrow paler

stripes on sides of pronotum, antennae and legs paler than dorsum.

Head small, exposed part distinctly transverse, almost concealed

from above by pronotum, fronto-clypeal suture situated just in front of

antennal insertions, dorsum pubescent; eyes moderately large, almost

covered dorsally by pronotum; antenna (see Fig. 6 b) short, about

as long as pronotum, scape moderately large, pedicel narrower and

little elongate, segment 3 elongate, segments 4-7 short and subequal,

segment 8 slightly wider than 7, club large, slightly elongate, about

one-fourth as long as antenna and with single pre-apical annulation.

Prothorax strongly transverse (1.00:2.40), narrowed in front,

front margin unemarginate, front angles bluntly rounded, lateral

margin feebly rounded and finely bordered, two lateral pits situated

close to anterior angle and posterior third, posterior angles acute,

basal margin distinctly sinuate on either side of middle; a narrow

elongate, yellowish sublateral stripe on either side of pronotum;

puncturation obscure and surface glabrous.

Elytra slightly elongate (1.05:1.0), more than thrice (3.1:1.0) as

long as prothorax, basal margin emarginated and fitting closely with

prothorax, humeral angles blunt, widest near middle; sides evenly

rounded, finely bordered and a little explanate, four lateral pits on

border in basal two-thirds; surface glabrous and puncturation obscure.

Ventral side paler; prosternum impunctate or obscurely punctate;

metasternum with fine and sparse punctures, impunctate mesally;

206 T.K. PAL

se Pat

ate re fe

L'AMICO LIA

CEI CORE

Fot

nang E PRE EPPO EE Hee ag. gt

Miei

Vee,

ls,

“o

Fig. 6 - Aphanocephalus naius n. sp.: a, dorsal view (scale = 1.0 mm.); b, exposed

part of head and antenna (scale = 1.0 mm.); c, aedeagus, lateral view.

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 20.7

puncturation on abdominal ventrite fine and sparse, scanty pubescence.

Aedeagus (Fig. 6 c) on one side, median lobe forming a broad

tube with acuminate and bifid tip, tegmen forming a cap-piece

enveloping median lobe, its proximal end broad cap-like with sharp

projections on sides, its distal end bifid and blunt.

Measurements of holotype: total length 1.92 mm, length of

antenna 0.50 mm, length and width of prothorax 0.50 mm and 1.20

mm, length and width of elytra 1.58 mm and 1.50 mm.

Holotype: Is. Fernando Poo, Basilé, 400-600 m, VIII-IX.1901,

Perea (MSNG)

Paratypes: same data as holotype, 40 ex. (34: MSNG, 6: ZSI).

Etymology: the species-name is derived from the Greek

word ‘nai’ (vat), meaning yes.

Remarks: this species shows resemblances with an Afri-

can species, A. decellei John, 1964, by its largely globular facies and

apparent similarity of aedeagal structure. But it can be differentiated

by its deep brown pronotum with a narrow-elongate yellowish sub-

lateral stripe on either side, bifid apex of median lobe with broadly

acuminate tips (vs. rather spinous tips in decellez).

Aphanocephalus sp.

Material: M.te Camerun, Buea, 800-1200 m.s.m., VI-VII.1902,

L. Fea, 7 9° (head of 1 ex. broken) (MSNG).

Size: length 1.72-1.90 mm.

Remarks: this species shows apparent similarity in facies

and colouration with the Indian species, A. stkkima Pal, 1996b. But

the specific identity could not be ascertained for lack of a male spec-

imen and examination of its aedeagus.

Genus Parafallia Arrow

Parafallia habra n. sp.

General appearance (Fig. 7) oblong, ovoid, about 1.3x as long

as broad, convex dorsally and flattened ventrally, shiny, moderately

208 Ke PAL

punctate, pronotum and elytra almost glabrous and head with scanty

pubescence; yellowish- to reddish-brown, legs paler.

Head small, exposed part distinctly transverse, partly concealed

by pronotum, fronto-clypeal suture situated in front of antennal

Fig. 7 - Parafallia habra n. sp., dorsal view (scale = 1.0 mm.).

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 209

insertions, dorsum with fine and sparse pubescence; eyes moderately

large, somewhat projecting and coarsely facetted; antenna moder-

ately long, longer than pronotum, scape moderately large, pedicel

short, narrower and about as broad as long, segment 3 clearly elon-

gate, segments 4-8 short, subequal and feebly wider towards apex,

club large, elongate, nearly one-third as long as antenna, with two

feeble preapical annulations.

Prothorax strongly transverse (0.56:1.00), narrowed in front,

front margin deeply emarginate and bisinuate, front angles broadly

pointed, lateral margin feebly arched and finely bordered, two lat-

eral pits situated close to front angle and above base, hind angles

prominent and almost right angle, basal margin rounded; punctura-

tion on pronotum fine and sparse, setae hardly seen, coarser punc-

tures arranged more closely above basal margin.

Scutellum transverse, triangular and impunctate.

Elytra broader than long (0.96:1.00), about 2.3x as long as pro-

thorax, basal margin emarginate and fitting with prothoracic base,

humeral angles not prominent and blunt, widest near middle; sides

evenly rounded, finely bordered and a little explanate; puncturation

coarser than on pronotum, moderately sparsely arranged, separated

by about 3-6 diameter, punctures bordering margin comparatively

coarser.

Ventral side paler, sterna with moderately coarse and sparse

punctures laterally and impunctate mesally, puncturation on abdom-

inal ventrites moderately coarse and sparse, almost glabrous.

Aedeagus (Fig. 8 a,b) on one side, median lobe forming a tubu-

lar-shape; tegmen forming a cap-piece partially enveloping median

lobe, distal end spatulate and studded with long setae.

Measurements of holotype: total length 1.75 mm, width of head

across eyes 0.47 mm, length of antenna 0.60 mm, length and width

of prothorax 0.52 mm and 0.92 mm, length and width of elytra 1.21

mm and 1.25 mm.

Holotype d: Sumatra, Lago Toba, II & XI.1891, E. Modigliani

(MSNG).

Paratype: Sumatra, D. Tolong, XI.1890, E. Modigliani, 1 ex. (ZSI).

Etymology: the species-name is derived from the Greek

word ‘habros’ (aBpdc) abros, meaning pretty or nice.

Remarks: this species shows resemblances with another

KY PAL

210

Sumatran species, P. minima John, 1968 in facies but can be differ-

entiated by its front margin of prothorax emarginated and bisinuate

(vs. emarginated and simply concave in minimus), a row of small

punctures bordering posterior margin of pronotum (vs. no regular

Fig. 8 - Parafallia habra n. sp., aedeagus: a, lateral view; b, tip of tegminal cap.

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 200

row of subbasal punctures in minimus), elytral punctures, especially

in anterior half finer and sparser, antennal club with two transverse

fine annulations (vs. no distinct annulation in minimus). Aedeagus of

P. minimus has not been studied and hence could not be compared

with that of P. habra.

Genus Paramaschema Heller

Paramaschema farak n. sp.

General appearance (Fig. 9) broadly oblong, convex dorsally

and flattened ventrally, sides explanate and margins wavy, shiny,

coarsely punctate-pubescent; reddish-brown with head, median part

of pronotum, and sutural as well as sides of elytra darker.

Head small, exposed part transverse, partly concealed by prono-

tum, fronto-clypeal suture situated in front of antennal insertions,

vertex moderately coarsely punctate, clypeus impunctate, dorsum

moderately setose; eyes moderately large, somewhat projecting and

coarsely facetted, considerably covered by pronotum; antenna mod-

erately long, longer than pronotum, scape moderately large, pedicel

shorter and narrower than scape, segments 3-5 longer than pedicel

and subequal, segments 6-7 shorter than preceding segment, club

large, elongate, balloon-shaped, slightly shorter than one-third as

long as antenna, without preapical annulation.

Prothorax strongly transverse (0.56:1.00) and widest near middle,

narrowed towards front, front margin deeply emarginate and concave,

front angles blunt, sides outwardly curved, basal margin trisinuate;

pronotal disc convex and broadly explanate on sides; puncturation

on disc coarse and moderately sparse, separated by about 2-4 diam-

eter, punctures slightly denser on sides, pronotum sparsely setose;

median part of disc dark brown and sides yellowish.

Scutellum transverse, triangular and impunctate.

Elytra broader than long (0.84:1.00), about 2.4x as long as protho-

rax; basal margin emarginate, bisinuate and fitting with prothoracic

base, humeral angles blunt; widest in anterior half, sides explanate,

margin undulated; puncturation on elytral disc coarse, dense, sepa-

rated by about 0.75-1.5 diameter, punctures obscure on explanate

Tae PAY

212

i data

1.0 mm.).

Fig. 9 - Paramaschema farak n. sp., dorsal view (scale

2453

A COLLECTION OF HETEROMERA AND CUCUJOIDEA

sides, setae moderately sparse but distinct; sides and sutural part of

elytra darker leaving an elongate yellowish space on each elytron.

Ventral side reddish-brown, paler than dark area of dorsum; pro-

sternum with moderately dense punctures mesally and impunctate on

sides, metasternum almost impunctate; abdominal ventrites impunc-

tate and finely setose; elytral epipleura broad, its surface slightly wavy.

Aedeagus (Fig. 10) on one side, median lobe forming a tube, its

distal end bilobed; tegmen forming a cap-piece partially enveloping

median lobe, its distal end spatulate and beset with setae.

Pr i e eee

—,

Fig. 10 - Paramaschema farak n. sp.: aedeagus, lateral view.

214 T.K. PAL

Measurements of holotype: total length 2.10 mm, width of head

across eyes 0.53 mm, length of antenna 0.78 mm, length and width

of prothorax 0.60 mm and 1.20 mm, length and width of elytra 1.47

nm, and, 1,,75,mm,

Holotype d: Mentawei, Si Oban, IV-VIII. 1894, E. Modigliani

(MSNG).

Paratype: Mentawei, Sipora, Sereinu, V-VI.1894, E. Modiglia-

mi diccxe(ZSl):

Etymology: the specific name is the Hindi adjective ‘farak’

(meaning different) refers to very different appearance of the body

and male genitalia of the species.

Remarks: this species shows some similarity with the

Japanese species, P. tumidium John, 1954 in slightly bicoloured

dorsum but can be differentiated by the sides of pronotum lighter

(vs. antero-median part of pronotum lighter in tumidium), elongated

lighter patch from base to more than posterior third of each elytron

(vs. lighter antero-median half of elytra in tumidium), no protuber-

ance on humeral portion of elytra, punctures on pronotum and elytra

considerably sparser. Further, unlike other species of Paramaschema

the tip of median lobe of this species is broad and bilobed.

Genus Cephalophanus John

Cephalophanus chhakka n. sp.

General appearance (Fig. 11) rather orbicular, convex dorsally

and flattened ventrally, vestiture constituting minute setae, moder-

ately long antenna with a distinct club; six moderately large, globu-

lar yellowish spots on blackish elytra, sides of pronotum paler than

middle.

Head small, exposed part transverse, largely concealed by pro-

notum and hardly seen from above in dorsal view, fronto-clypeal

suture situated in front of antennal insertions, dorsum punctate-

pubescent; eyes moderately large, coarsely facetted, covered almost

completely from above by pronotum; antenna moderately long,

longer than pronotum, scape moderately large, pedicel short, nar-

rower and about as broad as long, segment 3 elongate, segments

215

pastttt tag,

A COLLECTION OF HETEROMERA AND CUCUJOIDEA

Pare ga

aye

wae

ste

bai

wees

a ee Hw yn ©

ase tive

5 “<<

; exposed part of head and

antenna above dorsal view of the body (scale = 1.0 mm.).

Fig. 11 - Cephalophanus chhakka n. sp., dorsal view

216 RS PATS

4-7 short, subequal, a little elongate or about as broad as long, seg-

ments 8-9 feebly wider than preceding segment and slightly trans-

verse, club large, elongate and balloon-shaped, about one-fourth as

long as antenna, without preapical annulation.

Prothorax strongly transverse (0.54:1.00), narrowed in front,

front margin considerably emarginate and concave, front angles

blunt, lateral margin feebly arched and narrowly bordered, two lat-

eral pits situated close to front angle and slightly beneath middle,

hind angles prominent and slightly acute, basal margin clearly sinu-

ate on either side of middle; pronotal disc convex and moderately

explanate at sides; puncturation on disc fine and sparse, punctures

separated by about 3-6 diameter, pronotum sparsely setose; median

part of pronotum blackish and sides yellowish.

Scutellum small, transverse, obscurely punctate and setose..

Elytra broader than long (0.92:1.00), about 2.7x as long as pro-

thorax, basal margin slightly sinuate on either side of middle and

fitting with prothoracic base, humeral angles blunt, widest near

middle; sides somewhat explanate and margins almost rounded;

punctures on elytral disc moderately coarse, separated by about 1-2

diameter, setae moderately dense and distinct, little denser on sides;

elytra blackish, three large orbicular yellowish spots on each ely-

tron- two in anterior half and one in posterior half, six smaller spots

arranged close to border.

Ventral side reddish-brown medially and paler on sides; pro- and

metasternum with fine and dense punctures; abdominal ventrites clearly

punctate but slightly finer than on metasternum; ventrum not setose.

Measurements of holotype: total length 3.74 mm, width of head

across eyes 0.80 mm, length of antenna 1.36 mm, length and width

of prothorax 1.12 mm and 2.04 mm, length and width of elytra 3.08

mini. and. 3.32. nana:

Holotype 9: Sumatra, Si-Rambé, XII.1890-III.1891, E. Modi-

gliani (MSNG).

Paratypes: same data as holotype, 5 ex. (3: MSNG, 2: ZSI);

Sumatra, M.te Singalang, VII.1878, O. Beccari, 1 ex. (MSNG).

Etymology: the specific name is the Bengali adjective

‘chhakka’, meaning six, that refers to the six large orbicular spots

on elytra.

Remarks: this species shows some similarity with the

A COLLECTION OF HETEROMERA AND CUCUJOIDEA DET,

North Bornean species, C. Reninganus John, 1967 and C. octopuncta-

tus John, 1967 in partially lighter pronotum and presence of orbicu-

lar lighter patches on dark brown elytra. But this species can be

distinguished from the above and other known species of Cephalo-

phanus in having interruption of lateral lighter band of pronotum in

antero-median part, presence of three large yellowish orbicular spots

on each elytron in addition to six small spherical marginal spots,

elytral punctures are slightly coarser and denser than in C. Reninga-

nus and C. octopunctatus. In the paratype of C. chhakka from Mount

Singalang the lateral lighter bands of pronotum are confined near

the hind angles leaving remaining part of pronotum dark brown.

Family CERYLONIDAE

Genus Euxestus Wollaston

Euxestus grandis n. sp.

General appearance (Fig. 12) hemispherical, about 1.3x as long

as broad, convex dorsally and flattened ventrally, shiny, dorsum

with fine and moderately sparse punctures, almost glabrous; red-

dish-brown, legs paler.

Head small, exposed part distinctly transverse, partly concealed

by pronotum, fronto-clypeal suture situated in front of antennal

insertions, clypeus broad and front margin rounded, labrum partly

visible, frons and vertex evenly convex; puncturation on vertex mod-

erately fine, irregular, separated by about 1-3 diameter; eyes mod-

erately large, somewhat projecting and coarsely facetted. Antenna

moderately long, longer than pronotum, resting in antennal cavities

in repose, scape moderately large and curved, pedicel short, narrower

and about as broad as long, segment 3 elongate, segments 4-8 short,

subequal and transverse, segment 9 slightly wider than preceding

segment; club large, about as broad as long and somewhat oblique,

with subapical transverse impression and apical setose lobe.

Prothorax strongly transverse (0.42:1.00), narrowed in front,

front margin slightly emarginate, front angles obtusely rounded; lat-

eral margin feebly arched, smooth and finely bordered; hind angles

slightly acute from above, basal margin slightly bisinuate on either

218 T.K. PAL

side of scutellum; puncturation on pronotum finer than on vertex,

moderately sparsely arranged, separated by about 2-4 diameter, setae

scanty and obscure.

Fig. 12 - Euxestus grandis n. sp., dorsal view (scale = 1.0 mm.).

Scutellum transverse, triangular and impunctate.

Elytra broader than long (0.94:1.00), about 2.7x as long as pro-

thorax, basal margin emarginate and fitting with prothoracic base,

humeral angles not prominent and blunt, widest near anterior third;

sides evenly rounded; some minute punctures arranged in widely

separated linear rows, setae obscure.

Ventral side not paler than dorsum; sides of prosternum with

well developed cavities for reception of antennae, median part of

prosternal process longitudinally ridged; metasternum with sparse

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 219

punctures laterally and almost impunctate mesally; abdominal ven-

trites with fine and moderately sparse punctures, glabrous.

Measurements of holotype: total length 2.60 mm, width of head

across eyes 0.74 mm, length of antenna 0.76 mm, length and width

of prothorax 0.65 mm and 1.52 mm, length and width of elytra 1.80

mm and 1.90 mm.

Holotype: Mentawei, Sipora, Sereinu, V-VI.1894, E. Modiglia-

ni (MSNG). |

Paratypes: same data as holotype, 1 ex. (MSNG); Sumatra,

Siboga, X-1890-III.1891, E. Modigliani, 1 ex. (ZSI).

Etymology: The species name is derived from the Latin

word ‘grandis’, meaning large, that refers to its comparatively larger

body size.

Remarks: This species differs from two Sumatran species,

E. translucidus Motschulsky, 1859 and E. erithacus (Chevrolat, 1863)

by more convex and globose shape, prothorax more diverging pos-

teriorly and more transverse (1.0:2.5 vs. 1.0:1.8 in translucidus and

erithacus), linear arrangement of elytral punctures (vs. confused in

translucidus and erithacus), antennal club segment obliquely trans-

verse (inner margin shorter than outer margin vs. bilaterally sym-

metrical in translucidus and erithacus), and larger body size (2.6 mm.

long vs. 1.9-2.2 mm. long in translucidus and erithacus).

Family SPHAEROSOMATIDAE

Genus Sphaerosoma Leach

Sphaerosoma sp.

Material: Sumatra, Padang, 1890, E. Modigliani, 1 ex. (MSNG).

Size: length 2.05 mm.

ACKNOWLEDGEMENTS

I express my sincere gratitude to Dr. Roberto Poggi, Director,

Museo Civico di Storia Naturale “G. Doria”, Genova for kindly

providing an opportunity to examine this interesting collection of

220 TK) PAL

beetles and for generous support. I am indebted to Dr. J. R. B.

Alfred, Director, Zoological Survey of India, Kolkata for providing

necessary facilities for the work.

REFERENCES

CHEVROLAT A., 1863 - Coléoptères de Pile de Cuba. Notes, synonymies et descrip-

tions d’espèces nouvelles - Annls. Soc. ent. Fr., Paris, (4), 3: 589-620.

JOHN H., 1954 - Notizen ùber Paramaschema (Col., Notiophygidae). Eine Ergan-

zung und eine Neubeschreibung - Idea, Djakarta, 10 (1-2): 13-15.

JoHN H., 1964 - Neue Spezies von Aphanocephalus Woll. (Coleoptera: Discolomi-

dae) - Rev. Zool. Bot. afr., Tervuren, 69 (1-2): 137-164.

JOHN H., 1967 - Neue Spezies der Gattungen Cephalophanus John und Aphanoce-

phalus Woll. (Coleoptera: Discolomidae) - Pacif. Ins., Honolulu, 9 (2): 257-264.

JoHN H., 1968 - Discolomidae des Amsterdam Museum - Ent. Bericht., Amsterdam,

28: 190-195.

MorscHuLSsKy V. de, 1859 - Insectes des Indes Orientales et de contrées analogues

- Etud. ent., Helsingfors, 8: 25-146.

Pat T. K., 1992 - A new Inopeplid species (Coleoptera: Cucujoidea) from Patkoi

Hill Range of Arunachal Pradesh - 7. Bombay nat. Hist. Soc., 89 (2): 238-

241.

PaL T. K., 1996a - Indoleptipsius ushae, a new genus and species of Rhizophagidae

(Coleoptera: Cucujoidea) from Arunachal Pradesh, India - Hexapoda, Madras,

8 (1): 1-7.

PaL T. K., 1996b - Three new species of Discolomidae (Coleoptera: Clavicornia)

from Sikkim, India - Hexapoda, Madras, 8 (1): 9-17.

PaL T.K. & Datta A. K., 1982 - Inopeplidae (Coleoptera) from Andaman Islands,

India - Rec. zool. Surv. India, Calcutta, 79: 469-473.

SeNGUPTA T., 1976 - On the Genus Mimemodes (Rhizophagidae: Coleoptera) with

descriptions of new species from India - 7. zool. Soc. India, Calcutta, 28 (1-2):

67-73.

ABSTRACT

Determination of a collection of heteromeroid and cucujoid beetles of the

Museo Civico di Storia Naturale “G. Doria”, Genoa revealed 15 species belonging

to 6 families, viz., Othniidae, Inopeplidae, Monotomidae, Discolomidae, Ceryloni-

dae and Sphaerosomatidae. These include 2 new species of Inopeplidae: Inopeplus

pacificus (Pacific Islands, Mentawai), Inopeplus sthulalinga (West African Islands);

1 new species of Monotomidae: Indoleptipsius poggi (New Guinea); 4 new species

of Discolomidae: Aphanocephalus naius (West African Islands), Parafallia habra

(Sumatra), Paramaschema farak (Mentawai), Cephalophanus chhakka (Sumatra); and

1 new species of Cerylonidae: Euxestus grandis (Mentawai, Sumatra). The inventory

of the material including descriptions of the new species is presented.

A COLLECTION OF HETEROMERA AND CUCUJOIDEA 221

RIASSUNTO

Su una collezione di Heteromera e Cucujoidea del Museo Civico di Storia

Naturale “G. Doria” di Genova (Coleoptera, Polyphaga).

A seguito dell'esame di una serie di esemplari conservati da oltre un secolo

nelle collezioni del Museo di Genova, l'Autore elenca una quindicina di specie,

descrivendone ed illustrandone otto nuove:

Inopeplidae = Inopeplus pacificus n. sp. (Is. Goodenough, Nuova Guinea e

Mentawei), I. sthulalinga n. sp. (Is. S. Thomé e Is. Principe)

Monotomidae = Indoleptipsius poggit n. sp. (Nuova Guinea)

Discolomidae = Aphanocephalus naius n. sp. (Is. Fernando Poo), Parafal-

lia habra n. sp. (Sumatra), Paramaschema farak n. sp. (Mentawei), Cephalophanus

chhakka n. sp. (Sumatra)

Cerylonidae = Euxestus grandis n. sp. (Mentawei e Sumatra).

223

RoBERTO BATTISTON* & PaoLO FONTANA**

RICERCHE ZOOLOGICHE DELLA NAVE OCEANOGRA-

FICA “MINERVA?” (C.N.R.) SULLE ISOLE CIRCUMSARDE.

XXX. EMBIIDINA

(INSECTA)

INTRODUZIONE

Attualmente nel mondo sono note almeno 250 specie per l’Or-

dine Embiidina distribuite in gran parte nelle regioni tropicali. Una

piccola parte è presente nel bacino del Mediterraneo (Ross 1966) e

soltanto 7 entità sono note per l’Italia. Gran parte di queste ultime è

endemica (FAILLA et al. 1995; FONTANA 2002; FONTANA et al. 2002).

La presenza di questi insetti nei Musei e nelle collezioni europee è

scarsa e la loro conoscenza molto limitata, sebbene essi sembrino

localmente abbastanza comuni.

MATERIALI E METODI

Gli Embiotteri raccolti nelle piccole isole circumsarde nel corso

delle esplorazioni svolte con la nave oceanografica del C.N.R.

“Minerva” sono tutti conservati in alcool nella collezione del Museo

Civico di Storia Naturale di Genova (MSNG); si tratta di un buon

numero di esemplari (oltre 200), ma purtroppo nella stragrande

maggioranza dei casi sono state reperite solo forme immature o fem-

mine, che non permettono una identificazione sicura. In una sola

occasione è stato raccolto un maschio adulto, che è stato montato

su vetrino in Balsamo del Canada, secondo la tecnica descritta in

FONTANA et al. 2002.

* Museo Canal di Brenta, Via Garibaldi 27, 36020 Valstagna (VI);

e-mail: roberto.battiston@yahoo.it

** Dipartimento Agronomia Ambientale e Produzioni Vegetali, (Entomologia),

AGRIPOLIS, Viale dell’ Università 16, 35020 Legnaro (PD);

e-mail: paolo.fontana@unipd.it

224 R. BATTISTON & P FONTANA

RISE

Embia tyrrhenica Stefani, 1953

Materiale esaminato: Sardegna, Is. Molara, presso sorgente (SS),

8.VI.1989, leg. R. Poggi, 1 é (MSNG).

Distribuzione: specie mediterranea, nota fino ad ora per poche

località in Italia e in Croazia (Ross 1966; FONTANA et al. 2002).

sp. indet.

Materiale esaminato: 12 99 e 204 immaturi (di seguito indicati con “i.”).

Allo stato attuale delle nostre conoscenze, come detto, per questi

esemplari (tutti in MSNG) non si può giungere ad una sicura iden-

tificazione, ma si ritiene comunque utile fornire l’elenco completo

dei reperti, sperando che su questa base future ricerche in loco pos-

sano portare al reperimento dei maschi indispensabili per il ricono-

scimento.

La successione delle isole segue l’ordinamento adottato in BAC-

CETTI et al., 1989.

- Is. Razzoli, 26.IX.1985 e 13.XI.1986, 3 1., R. Poggi;

- Is, -La Presa, 16:X.19989,; 1-1., R. Poggi;

= is Sparc, Cala Grane, to. X.1989, 2 1, Ri Pope

= IssSpargiotto; 254 X%1985,..1.4.,.R. Poggi;

- Is. Maddalena, 12.XI.1986, 1 i1., M. Bologna; id., Dint. La

Maddalena, 23.IX.1985, 6 i., R. Poggi; id., Bacino artificiale,

13-XEk-1986, 11, LaTonob;;1d:)Spalmatore;:18;VI.1989, 2 14) Re

Poggi; id., I Pozzoni, 14.X.1989, 5 i., R. Poggi; id., Case Fan-

gotto, 14.X.1989, 1 i., R. Poggi; Cala Bassa Trinità, 14.X.1989,

hi Re Posso

= Is mSrsStefanop1 XXI 4986, 1,1 Ra Poss;

- Is. Caprera, Pendici M.te Teialone, 15.X.1989, 4 1., R. Poggi;

id., Forte Settentrionale, 15.X.1989, 2 1., R. Poggi;

- Is. delle Bisce, 11.XI.1986, 7 1., R. Poggi;

- Is. delle Rocche, 11.XI.1986, 6 1., R. Poggi;

ia. Power E bPV1986,0li; BR. Poggi id, 1LPV 1986, 1,9, RI

Argano;

EMBIIDINA DELLE ISOLE CIRCUMSARDE 225

Is. Poveri II, 11.1V.1986, 1 2, R. Poggi;

Is. Camere W, 11.IV.1986, 1 9, R. Argano & V. Cottarelli; id.,

1 OM. Bologna ids ParereRe*Popertiadk SV 98 2 TRI

Poggi;

Is. Camere E, 11.1V.1986, 4 1., R. Poggi;

Is. Figarolo, 28.VII.1986 e 11.IX.1987, 3 1., R. Poggi;

IS. Verde, Z.Vil 1987, 5 i2 Re Pees

Is. Tavolara, 27.IX.1985, 9.XI.1986, 13.IX.1987 e 8.VI.1989, 1

2,71. R. Poss 14 9 XIL19s6, 4 1., Mi Bboloona:

Is. Molara, 28.VII.1986, 28.X1.1986 e 8.VI.1989, 4 1., R. Poggi;

ih 2.IV1980: dnc olk. Araneae, OI rl eno pk

Is. Molarotto, 27.IX.1985, 10.IV.1986 e 12.IX.1987, 97 1., R.

Poggi; id., 10.IV.1986, 5 1., M. Bologna;

Is. Varaglione Sud, 30.VII.1986, 1 2, R. Poggi;

Is. Serpentara, 25.V1I.1987 e 9.VI.1989, 4 1., R. Poggi;

Is. der Cavoli, 9.VI.1989, 1 1.,.R. Poggi;

ISP Ta Vacca! 262V 121987 PI VI 1989 E TON WO sao reat. OR.

Poggi; id., 26.VI.1987, 1 2, V. Cottarelli;

lis il L'oro, 40.V.1988pdu 8, Regdeoget:

Is. S. Antioco, Stagno de Cirdu, 12.VI.1989, 1 1., R. Poggi;

Is. S. Pietro, Pendici M.te Guardia dei Mori, 8.VII.1990, 1 1.,

R. Poggi;

Is. Mal di Ventre, 28.VI.1987, 1 9, R. Poggi;

Is. Piana di Alghero, 29.VI.1987 e 15.VI.1989, 6. 1., R. Poggi;

is-der Potri!29.VIHI'987, 7 ER” Posen

Is. Asinara, 12.X.1989, 1 2, R. Poggi; id., Cala Arena, 1.VII.1987,

2 1., R. Poggi; id., Cala Realeyassed988ete® arti Gattarelli:

id., Cala Reale, 9.VII.1990, 1 i., R. Poggi;

Is) Scombro, 13.%.1989, Ps "RY Pores:

CONCLUSIONI

Le isole circumsarde, secondo la Lista delle Isole Italiane ITA,

ammontano a 182, gran parte delle quali sono per lo piu di piccole

dimensioni o scogli. Il Grande Atlante d’Italia dell’ Istituto Geogra-

fico De Agostini a scala 1:300.000 ne individua soltanto 57 compresi

DIDO R. BATTISTON & P FONTANA

3 scogli maggiori. La presenza di Embiidina è stata finora rilevata

da Aprile fino agli ultimi giorni di Novembre in 30 isole, più della

metà di quelle di più ampie dimensioni.

La maggior parte di queste era collegata alla Sardegna tramite

terre emerse fino ad almeno 15.000 anni fa, quando il livello del

mare ha iniziato a crescere dopo la glaciazione wurmiana; questo

spiega l'ampia diffusione del gruppo in tali ambienti. L'isola più

piccola su cui sono stati rinvenuti Embiidina è il piccolo arcipelago

granitico dei Varaglioni il cui scoglio più grande misura circa 0.027

km quadrati e dista dalla costa 3 km.

Interessanti rimangono quindi l’antichità e la vastità della colo-

nizzazione, per insetti dalle abitudini sedentarie e molto resistenti al

cambiamenti ambientali annuali e stagionali.

BIBLIOGRAFIA

BacceTT1 B., COBOLLI SBORDONI M. & Pocci R., 1989 - Ricerche zoologiche della

nave oceanografica “Minerva” (C.N.R.) sulle isole circumsarde. I. Introduzione

- Annali Mus. civ. St. nat. “G. Doria”, Genova, 87: 127-136.

FarLLa M. C., La Greca M., LomBarpo F., Messina A., SCALI V., STEFANI R. &

VIGNA TAGLIANTI A., 1995 - Blattaria, Mantodea, Isoptera, Orthoptera, Pha-

smatodea, Dermaptera, Embioptera. In: Minelli A., Ruffo S. & La Posta S.

(eds.), Checklist delle specie della fauna italiana, 36 (1994) - Ed. Calderini, M.

Bologna, 23 pp.

FONTANA P., 2002 - Contribution to the knowledge of mediterranean Embiidina

with description of a new species of the genus Embia Latreille, 1825 from Sar-

dinia (Italy) (Insecta Embiidina) - Atti Acc. Rov. Agiati, Rovereto, a. 252, ser.

VIII, 2 (B): 39-50.

FONTANA P., BuzzeETTI F. M., Coco A. & ODE B., 2002 - Guida al riconoscimento

e allo studio di Cavallette, Grilli, Mantidi e insetti affini del Veneto (Blattaria,

Mantodea, Isoptera, Orthoptera, Phasmatodea, Dermaptera, Embiidina) - Ed.

Museo Natur. Archeol. Vicenza, 592 pp.

Krauss H. A., 1911 - Monographie der Embien - Zoologica, Stuttgart, 60: 1-78.

Ross E. S., 1966 - The Embioptera of Europe and the Mediterranean Region -

Bull. Brit. Mus. (Nat. Hist.), Ent., London, 17: 275-326.

Ross E. S. , 2000 - Embia. Contributions to the Biosystematics of the Insect Order

Embiidina - Occ. Papers Calif. Acad. Science, San Francisco, 149: (part 1)1-53;

(part 2)1-36.

STEFANI R., 1953 - Nuovi Embiotteri della Sardegna - Boll. Soc. ent. ital., Genova,

83: 84-98.

EMBIIDINA DELLE ISOLE CIRCUMSARDE D27

RIASSUNTO

Sono presentati nuovi dati sugli Embiidina delle isole della Sardegna. Un’ana-

lisi del materiale conservato nel Museo Civico di Storia Naturale di Genova ha rile-

vato che la gran parte degli individui raccolti e conservati è composta da immaturi,

quindi non identificabili, fatta qui eccezione per un solo maschio di Embia tyrrhe-

nica Stefani 1953. Vengono proposte alcune considerazioni geografiche ed ecologiche

sulla distribuzione e la presenza stagionale degli Embiidina nelle isole circumsarde.

ABSTRACT

Zoological researches of the oceanographic ship “Minerva” (C.N.R.) on the

islands around Sardinia. XXX. Embiidina. (Insecta).

New data on Embiidina on the islands around Sardinia are presented. An

analysis of the material preserved in the Museo Civico di Storia Naturale di Genova

(Italy) shows that the largest part of the specimens preserved in the collection is

represented by immature specimens, therefore unidentifiable, with the exception of

a male of Embia tyrrhenica Stefani 1953. Some geographical and ecological consi-

derations are presented concerning the distribution and the seasonal occurrence of

Embiidina in the circumsardinian islands.

oes

4 sont maggiori. La pitesetiza di Embiidina è stata rose dei

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| i spiega” l'ampia diluizione del gruppo in tali ambienti. L'isola: più

piccola su cui sono Stati rinvenuti Embiidina è il piccolo arcip pelag ii

granitico dei Varaglioni il cui scoglio più grande misura Téa ean

ban ABLAAE REF dintaitlel gino sid qir}ore>20 ori to vartnnaze» ‘In: rigolooN

ewe - Interessanti ang nbs anne Dante GRe erm ane Real pace

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«0 SAR re IAAD IRINA 5100.146158 AGI. SAN CIARA

Avo iain sasalice date toored: jeogrel edt sedi aworle (qlat1) A

to: froitqnexs edi diiw sldsitisnsbinw stolst9iì ,ereminoge siwismmi vd botnse91q3t |

«tanoa Issigolose bas: ispifigstaoog smog £221 instore pamesttuo ordinal lo ‘eten a4

To spristiy990. lenoesse orli bas noinidirieib od? grinison00 bataossig 918 enoise1ab |

BIBLIOGRAFIA ee -abnalet agiaibisemi; ni srt ni sribidaet

Baccerri B. CORDELI Spokbon’ Mi è Podi: I R., 1989 - Ricerche zoologiché della”

nave oceanogreticn: “Minerva” (€. NR.) sulle bid *ircumsarde. I. Introd uxIONe 2

SC Annali Mus. civ. Si. wat. “G. Doria”, Genova, 87: 127-126 La a

i Far. tA M. C. La Gaeca M., “oupasoo F.. Messina A., ScaLi V., STEFANI R «ij

“Viens TAGLIANTI A.; 1995 » Biatraria, Mantodea; Isoptera, ©: hoprerai Phas]

smatodea, Dermaptera, Embioptera. int Minelli A., Ruffo S. & La Posta S. 3 È

(eds), Checklist delle pr della fauna italiana. 36 (1994) - Ed. Calderini, Mb

Bologna, 23 pp. 0 | i

Fovrana Pi: 2002 + Cai to per knowledge: of mediterranean Embidinag

yeh description af. a new species of the genus Embio Latreille e R25 from Sarei

“dunia (italy) (insects Embiidiria) ~ Atti Ace. Row. Agiati, Rovereto, a. 252, seri

"FRA AP VEL, 2 (By 39-50. ae

ek aes FostanaP.,' Buzzetti F.. M., Coco A & OnE 8,200 - Guide al ri ‘onoscimintii

‘e allo studio di Cavallette, ‘Grilli, Mantidi è insetti affini del Veneto (Blatter

43 Mantodea, Tsopterà, Otthoptera, Phasmatodea, Dermaptera, Embiidina) + Ed sd

- Musco Natur. Archeo, Vicenza, 592 pp, i È

ae Vi Hai H. AL 191f x Monographie der Embicn - Zoo log: a, Stuttgart, 60: ae i

Mu | aia Ross E. 8. 1966 ‘The Embioptera of pene dnd. the: Mediterranean. Region

fast Ball, Bint, Mus, (Nat. Hist.), Ent., London, 17: 275-326 ;

Ea a if ev fr. Mess B.S. 2000 - Embia. AULE, to the Biesyaternatics of the Insect Or

ce ki CENTRINO Rasbidina < Oce. Papers Calif. "ani Science, San Francisco, 149: (part 1}1-8

ee A {part 271-536. en

I DE ea te Liri L ade Nuovi Embiatteri delle Sarda Moll, Sac. ent, itals, Geno

ee 6-08. 7

ee eee kin ea Bie viti ARI eee i ve

bans te ia eet Pes pai

ne di Rim i

229

CHI-FENG LEE* & MANFRED JACH**

NEW SPECIES AND NEW RECORDS OF PSEPHENIDAE

FROM ASIA

(COLEOPTERA)

INTRODUCTION

Most genera of Psephenidae have been reviewed taxonomically.

Recently we received a number of unidentified specimens for identifi-

cation. Among this material, the Iranian psephenids are especially note-

worthy, because this family has never been recorded from that country.

In this paper we describe four new species from Iran, India,

Malaysia and Indonesia, and we provide new distribution records

nor 19 taxa;

MATERIAL AND METHODS

The terminology used herein follows LEE, YANG & SATO (1997,

1999) for Macroeubria Pic, LEE, YANG & SATÒ (1998) for Ectopria

LeConte and LEE, JACH & YANG (1998) for Schinostethus Waterhouse.

Specimens examined are deposited in the following museums or

institutions:

BMNH The Natural History Museum, London, United Kingdom

MSNG Museo Civico di Storia Naturale “Giacomo Doria”,

Genova, Italy

* Applied Zoology Division, Taiwan Agricultural Research Institute, Taichurg,

‘Taiwan.

** Naturhistorisches Museum, Vienna, Austria

Correspondence: Chi-Feng Lee, Applied Zoology Division, Taiwan Agricultural

Research Institute, Wudeng, 413 Taichung, Taiwan. E-mail: chifenglee@gmail.com

230 C.-F LEE & M. JACH

NMEG Naturkundesmuseum, Erfurt, Germany

NMNS National Museum of Natural Science, ‘Taichung,

Taiwan

NMPC National Museum (Natural History), Prague, Czech

Republic

NMW Naturhistorisches Museum, Vienna, Austria

In the species descriptions the following abbreviations are used:

LE = maximum length of elytron

LP = length of middle of pronotum

WE = maximum width of elytra

WP = maximum width of pronotum

Dicranopselaphus brevicornis Lee & Yang

Dicranopselaphus brevicornis Lee & Yang, 1996: 173

1 Sd: “Indonesia: Sulawesi Utara, Dumoga-Bone N. P. April-

May 1985, ‘Clarke’ Camp Lower montane forest, 1140 m, R. Ent.

Soc. Lond., Project Wallace B.M. 1985-10, Malaise trap” (BMNH);

1 9, same as previous, but with different date (6 December 1985)

and altitude (Lowland forest ca. 200m) (BMNH).

Distribution: Indonesia (Sumatra, Sulawesi: new record).

Dicranopselaphus emmanueli (Pic)

Grammeubria emmanueli Pic, 1918: 13

Dicranopselaphus emmanueli (Pic): LEE & YANG 1996; LEE, YANG & SATÒ 2000

1g, 2 22: “Sarawak, Kuching dist. Mt. Penrissen, 1000 m 24-

26.3.1994 Sv. Bily leg.” (NMPC); 1 dé: “Malaysia-W, Pahang 20 km

E of Ioph, 1500 m Cameron Highlands, Tanah Rata, 16-18.1V.2000,

PR. Cechovsky leg.” (NMPC); 1 d: “Malaysia W., Johor 40 km N of

Kota Tinggi Kampong Uhi Sedili env. 23-28.III.2004, Cechovsky

Poter les” (NMPC)

Distribution: Philippines (Luzon, Mindanao, Palawan), East

PSEPHENIDAE FROM ASIA 231

Malaysia (Sabah, Sarawak), West Malaysia (Pahang, Johor: new

records), Indonesia (Sumatra).

Dicranopselaphus flavus Lee & Yang

Dicranopselaphus flavus Lee & Yang, 1996: 174

16: “Urung Tama Sibolangit, E. Sumatra, 30.X.1999, H. Taki-

zawa leg.” (NMNS).

Distribution: East Malaysia (Sabah), Indonesia (Sumatra: new

record).

Dicranopselaphus imparis Lee & Yang

Dicranopselaphus imparis Lee & Yang, 1996: 565

16,1 £: “Ban Saleui, 1350 m, Xamneua, Laos 21-24.VI.2003,

M. Sato leg.” (NMNS); 1 9: “Phou Pan, 1750 m, Xamneua, Laos

16-21.VI.2003, M. Sato leg.” (NMNS).

Distribution: Vietnam, Laos (new record).

Dicranopselaphus malickyi Lee & Yang

Dicranopselaphus malickyi Lee & Yang, 1996: 192

16,39%" “Vietnam; Ni N of Hanei, Sa, Pay nw, 1250

NN Hoang Lien Son Nat. Park, 05.VI.-09.VI.1998, leg. A. Napo-

lov” (NMEG).

Distribution: Thailand, Myanmar, Vietnam (new record).

Ectopria iranensis n. sp. (Figs 1-3)

Holotype gd: “S Iran, Kushk N. Masiri, 1800 m 12.6.1973 /

Loc. no. 237 Exp. Nat. Mus. Praha” (NMPC).

Male. Length 3.3 mm, width 2.1 mm. Coloration brown; head

darker blackish-brown, prosternum and proepisternum yellowish-

232 C.-F LEE & M. JACH

brown. Antennae lost. Maxillary palpus (Fig. 3) slender, terminal

segment bifurcate, relative lengths of segments 2-4 about 2.2 : 1:

1.7. Labial palpus (Fig. 2) small, about 0.5 times as long as max-

illary palpus, segment 2 subequal to segment 3, terminal segment

bifurcate. WP/LP = 2.64; LE/WE = 1.21; WP/WE = 0.87.

Aedeagus (Fig. 1) 4.4 times as long as wide. Penis long, 0.78

times total length of aedeagus, apex narrowly rounded, gradu-

ally narrowed toward apex. Parameres widest at middle, abruptly

narrowed near base, lateral margin concave in apical third; mesal

margin connected at middle. Fibula 0.3 times total length of aedea-

gus; Y-shaped. |

Etymology. This species is named for the type locality.

Remarks. This species is characterized by its elongate

aedeagus and by its bifurcate maxillary and labial palpi.

Granuleubria pakistanica Jàch & Lee

Granuleubria pakistanica Jich & Lee, 1994: 228; LEE & YANG 1999

3 6g: “SE Iran, 1-2.4.1973 env. Sarbaz, valley of river Sarbaz /

Loc. no. 145 Exped. Nat. Mus. Praha” (NMPC).

Distribution: Nepal, Pakistan, Iran (new record).

Faechanax insignis (Fairmaire)

Eubrianax insignis Fairmaire, 1904: 87

Faechanax insignis (Fairmaire): LEE, SATO & YANG 2000b

2 dd: “Bang Phra Chong Buri Prov., E. Thailand, 18.V.2002,

S. Tsuyuki leg.” (NMNS).

Distribution: Vietnam, Laos, Myanmar, Thailand (new record).

Faechanax major major (Pic)

Eubrianax major Pic, 1913: 172

Faechanax major (Pic): LEE, SATO & YANG 2000b

2383

PSEPHENIDAE FROM ASIA

WU LO

maxil-

Figs 1-3: Ectopria iranensis n. sp., male. 1 - aedeagus; 2 - labial palpus;

lary palpus.

234 C.-F LEE & M. JACH

1 d: “Nepfal], Narayan/Chitwan 13 km W Sauraha, Kasara,

Chitwan-NP / 27°33’°07” N, 84°21’59”E, 20.VI.2005, 180 m, leg. A.

Weigel, dec. Forest” (NMEG).

Variation: this specimen has a yellowish-brown body, but the

apical halves of the elytra are dark brown.

Distribution: Nepal (new record), Thailand, Malaysia, Indone-

sia (Sumatra, Java), Philippines (Mindanao).

Macroeubria bella Lee, Yang & Sato

Macroeubria bella Lee, Yang & Satò, 1999: 190

1 2: “Vietnam, N. 100 km SSW Hanoi, 40 km SW Thano

Hoa, Ben En National Park, 50 m 23-28.V1II.1997, lg. A. Napolov”

(NMEG).

Distribution: Thailand, Vietnam (new record).

Macroeubria bicolor Lee, Yang & Satò

Macroeubria bicolor Lee, Yang & Satò, 1997: 139

1 d: “Laos: Lakxao, 13-14.III.2005, leg. C.- F. Lee” (NMNS).

Distribution: China (Fujian), Laos (new record).

Macroeubria contracta Lee, Yang & Sato

Macroeubria contracta Lee, Yang & Satò, 1999: 191

1 2: “Vietnam, N. 100 km SSW Hanoi, 40 km SW ThanoHoa, Ben

En National Park, 50 m, 23-28.VII.1997. leg. A. Napolov” (NMEG).

Distribution: Thailand, Vietnam (new record), Laos, Indonesia

(Sumatra).

Macroeubria monstrosa Lee, Yang & Satò

Macroeubria monstrosa Lee, Yang & Satò, 1999: 197

PSEPHENIDAE FROM ASIA 235

4 3d: “Giava [Java], Tcibodas, Ott. 1874, O. Beccari” (MSNG);

1 d: “Urung Tama Sibolangit, E. Sumatra, 17-18.IX.1998, H.

Takizawa leg.” (NMNS); 1 d, same locality and collector, 26.X.1999

(NMNS).

Variation: all individuals from Java are brown and their penis is

longer, reaching apex of parameres.

Distribution: East Malaysia (Sabah, Sarawak), Indonesia (Java,

Sumatra: new records).

Macroeubria shimadai n. sp. (Figs 4-7)

Holotype d: “[West Malaysia] Pahang St. Cameron Highlands,

Robinson Waterfall (window trap) 12-15.III.2002, Takashi Shimada

leg.” (NMW).

Male. Length 2.8 mm, width 1.9 mm. Coloration blackish

brown. Antennal segment 3 triangular (Fig. 5), segment 4 pectinate,

others lost. Maxillary palpus (Fig. 7) slender, terminal segment with

a tube-like process on apex of outer margin, relative lengths of seg-

ments 2-4 about 2.1 : 1 : 2.3. Labial palpus (Fig. 6) small, about

0.35 times as long as maxillary palpus; terminal segment medially

dilated, apex bifurcate; relative lengths of segments 2-3 about 1 :

1.5. Striae pattern similar to M. contracta; WP/LP = 2.55; LE/WE

= 1.14; WP/WE = 0.74.

Aedeagus (Fig. 4) 3.5 times as long as wide. Penis subequal to

lengths of parameres and basal piece; narrowed at apical 2/5, gradu-

ally widened from apical 2/5 to 3/5, apex rounded. Fibula 0.3 times as

long as penis, apical margin rounded with a small rounded process at

middle; baso-lateral apophyses very long. Parameres wide, dorsal baso-

lateral apophyses very long, reaching apical 1/3 of basal piece; ventral

baso-lateral apophyses short. Basal piece gradually widened from apex

to basal 1/3, moderately narrowed at basal 1/3, basal margin truncate.

Etymology. This species is named after Takashi Shimada,

who collected the holotype.

Remarks: this species is similar to M. bella with regard to

the maxillary and labial palpi, and aedeagus, but it differs from the

latter by the blackish-brown color, the longer aedeagus, the wider

parameres, and the rounded apex of the fibula.

236 C.-E LEE & M. JACH

Wie

\ |

)

‘

0.2mm

0.1 mm

SALUS A 7

0.2 mm

Figs 4-8: Macroeubria shimadai n. sp., male: 4 - aedeagus; 5 - antenna; 6 - labial

palpus; 7 - maxillary palpus; M. truncata, male: 8 - antenna.

PSEPHENIDAE FROM ASIA 237

Macroeubria truncata Lee, Yang & Sato

Macroeubria truncata Lee, Yang & Satò, 1999: 199

1 3: “Sibolangit, E. Sumatra, 26.X.1999, H. Takizawa leg.” (NMNS).

Variation: in contrast to the serrate antennae of the holotype

this specimen has pectinate antennae (Fig. 8).

Distribution: West Malaysia, Indonesia (Sumatra: new record).

Odontanax laosensis (Pic)

Eubrianax laosensis Pic, 1923: 9

Odontanax laosensis (Pic): LEE, SATO & YANG 2000a

1 36,1 9: “Nepal, Annapurna 1996, Sikles Mts. Garlang 1350

m, 18.4., Schmidt” (NMEG); 1 gd: “Vietnam, N; N of Hanoi,

SaPa vii. Env., 1250 m, NN Hoang Lien Son Nat. Park, 05.VI.-

09.VI.1998, leg. A. Napolov” (NMEG); 1 6: “Vietnam, N, N of

Hanoi, SaPa vii. Env., 1250 m NN Hoang Lien Son Nat. Park,

05.VI.-09.VI.1998, leg. A. Napolov” (NMEG).

Variation: one specimen from Vietnam has a brown body.

Distribution: Laos, Vietnam, India, Nepal (new record).

Schinostethus (Sundodrupeus) priscus Lee, Jach & Yang

Schinostethus (Sundodrupeus) priscus Lee, Jich & Yang, 1998: 317

3. Sd: \ “Malaysia: Sabah Kerkot Laut, 18.VI.1999,f leg. C.-F.

Lee” (NMNS); 1 d: “Sarawak, Kuching dist., Mt. Serapi 27.-

29.3.1994 J. Horak” (NMW).

Variation: medio-lateral parameral process varies in size and

position (Figs 9-10).

Distribution: East Malaysia (Sabah, Sarawak: new record).

Schinostethus(Sundodrupeus) sumatrensis n.sp.(Figs 11-14)

Holotype 6: “Urung Tama, Sibolangit, E. Sumatra, 17-

18.IX.1998, H. Takizawa leg.” (NMW).

C.-F LEE & M. JACH

238

Figs 9-10: Variation of aedeagi of Schinostethus (Sundodrupeus) priscus.

PSEPHENIDAE FROM ASIA 239

Male. Length 3.1 mm; width 2.4 mm. Coloration dark brown,

but eyes and antennae black. Relative lengths of rami of antennal

segments 3-7 about 0.11 : 0.26 : 0.67 : 0.90 : 1 (Fig. 14). Maxillary

palpus (Fig. 11) slender, terminal segment truncate, relative lengths

of segments 2-4 about 3.0 : 1 : 1.9. Labial palpus (Fig. 12) small,

about 0.5 times as long as maxillary palpus, segment 2 subequal to

segment 3; apical margin irregular. WP/LP = 2.61; LE/WE = 1.13;

WP/WE = 0.74.

Aedeagus (Fig. 13) 2.6 times as long as wide. Penis long, 0.85

times total length of aedeagus, apex rounded, gradually narrowed

toward middle, abruptly widened behind middle. Parameres widest

at connection with phallobase, slightly narrowed at apical 1/3; latero-

apical processes broadly rounded, medio-apical processes widely con-

nected with latero-apical process basally and laterally, apex curved

and acute.

Etymology. This species is named after the type locality.

Remarks: this species is close to S. sakai due to similar

color and aedeagus, but differs by its long penis, by the unique

articulation of the parameral processes, and by the short third anten-

nomere.

Schinostethus(Sundodrupeus) transversus _ n.sp.(Figs 15-19)

Holotype d: “India: Uttaranchal 30 km N Bageshwar W

Loharket [village] / 1800-1900 m, 24.VI.2003, leg. Z. Kejval & M.

Tryzna” (NMW). Paratypes: 7 males and 2 females, same label data

as holotype (NMW, NMNS).

Male: length 4.7-5.2 mm, width 3.1-3.4 mm. Coloration black-

ish-brown, except prothorax (including pronotum, prosternum, and

proepisternum) yellowish brown, pronotum medially darkened. Rel-

ative lengths of rami of antennal segments 3-7 about 0.20 : 0.33 :

0.62 : 0.92 : 1 (Fig. 18). Maxillary palpus (Fig. 15) slender, ter-

minal segment truncate, relative lengths of segments 2-4 about 3.2

=1 72:1) Labial'palpus (Fig. 16) small, about OS—times as Mong

as maxillary palpus, segment 2 subequal to segment 3; terminal

margin truncate. WP/LP = 2.61-2.74; LE/WE = 1.13-1.16; WP/

WE = 0.83-0.85.

240 C.-K LEE & M. JACH

11 0.1 mm

E ;

E N

& ©

HE du ed

Figs 11-14: Schinostethus (Sundodrupeus) sumatrensis n. sp., male. 11 - maxillary

palpus; 12 - labial palpus; 13 - aedeagus; 14 - antenna.

PSEPHENIDAE FROM ASIA 241

15 0.5 mm

Candice li

ia ee. 18 19

Figs 15-19: Schinostethus (Sundodrupeus) transversus n. sp. 15 - maxillary palpus;

16 - labial palpus; 17 - aedeagus; 18 - male antenna; 19 - female an-

tenna.

242 C.-F LEE & M. JACH

Aedeagus (Fig. 17) 2.6 times as long as wide. Penis long, 0.76

times total length of aedeagus, apex rounded, gradually narrowed

toward apex. Parameres widest at connection with phallobase, mod-

erately narrowed at apical 1/3; latero-apical processes narrowly

rounded, medio-apical process connected with latero-apical process

basally, apex curved and acute.

Female. Length 5.8 mm, width 4.1 mm. Coloration similar to

male, but black band on pronotum reduced to a small spot. Anten-

nal segment 3 elongate, segment 4 feebly serrate, segments 5-10

moderately serrate (Fig. 19). WP/L P= 2.69; LE/WE = 1.11; WP/

WE= 0.78.

Etymology: this species is named in reference to its trans-

verse pronotum.

Remarks: this new species is similar to Schinostethus fla-

bellatus in general appearance, but differs by its slender and curved

latero-apical parameral processes and by the more transverse pro-

notum.

ACKNOWLEDGEMENTS

We are indebted to M. Barclay (BMNH), J. Hajek (NMPC)

and R. Poggi (MSNG) for sending valuable specimens in study.

REFERENCES

FaIRMAIRE L., 1904 - Descriptions de Lamellicornes indo-chinois nouveaux ou peu

connus - Mission Pavie Indochine (1879-1895), Paris, 3: 86-90.

JicH M. & LEE C.-F., 1994 - Description of Granuleubria, a new genus of Eubriinae

from West and South Asia (Coleoptera: Psephenidae) - Kol. Rundsch., Wien,

64: 223-232.

LEE C.-F. & YANG P.-S., 1996 - Taxonomic revision of the Oriental species of

Dicranopselaphus Guérin-Méneville (Coleoptera: Psephenidae: Eubriinae) - Ent.

scand., Lund, 27: 169-196.

LEE C.-F. & Yanc P.-S., 1999 - Notes on Granuleubria Jach and Lee, with descrip-

tions of four new species and the immature stages of G. fontana, new species

(Coleoptera: Psephenidae: Eubriinae) - Col. Bull., Sacramento, 53: 259-274.

Lee C.-F., JicH M. & Yana P.-S., 1998 - Psephenidae: II. Synopsis of Schinoste-

thus Waterhouse, with descriptions of 14 new species (Coleoptera) (pp. 303-

326), in: Jach M. & Ji L. (eds.) - Water Beetles of China. Vol. II - Zool.-bot.

Gesell. in Osterreich and Wiener Coleopterolo-genverein, Wien, 371 pp.

PSEPHENIDAE FROM ASIA 243

Lee C.-F., Sato M. & YANG P.-S., 2000a - A revision of the Eubrianacinae (Co-

leoptera, Psephenidae) IV. Odontanax gen. nov. - Fap. Journ. syst. Ent., Mat-

suyama, 6: 151-170.

LeE C.-F., Sarto M. & YANG P.-S., 2000b - A revision of the Eubrianacinae (Co-

leoptera, Psephenidae) V. Faechanax gen. nov. - Elytra, Tokyo, 28: 119-129.

LEE C.-F., YANG P.-S. & Sato M., 1997 - The eastern Asian species of the genus

Macroeubria Pic (Coleoptera, Psephenidae, Eubriinae) - fap. Journ. syst. Ent.,

Matsuyama, 3: 129-160.

LEE C.-F., YANG P.-S. & SATÒ M., 1998 - Psephenidae: I. Notes on the East Asian

species of Ectopria LeConte (pp. 297-301), in: Jich M. & Ji L. (eds.) - Water

Beetles of China. Vol. II - Zool.-bot. Gesell. in Osterreich and Wiener Co-

leopterologenverein, Wien, 371 pp.

Lee C.-F., YanG P.-S. & SATO M., 1999 - The southeast Asian species of the genus

Macroeubria Pic (Coleoptera, Psephenidae, Eubriinae) - fap. Fourn. syst. Ent.,

Matsuyama, 5: 181-201.

Lee C.-F., YANG P.-S. & Sato M., 2000 - A synopsis of Dicranopselaphus (Co-

leoptera: Psephenidae, Eubriinae), with descriptions of nine new species - Ent.

Science, Tokyo, 3: 557-568.

Pic M., 1913 - Diagnoses de Dascillides et Cyphonids nouveaux - Echange, Moulis,

eo 171-173.

Pic M., 1918 - Nouveautés diverses - Meél. exot.-ent., Moulis, 29: 7-24.

Pic M., 1923 - Nouveautés diverses - Meél. exot.-ent., Moulis, 40: 3-32.

ABSTRACT

New distribution records are provided for the following species: Dicranop-

selaphus brevicornis Lee & Yang, D. emmanueli (Pic), D. flavus Lee & Yang, D.

imparis Lee & Yang, D. malickyi Lee & Yang, Granuleubria pakistanica Jich & Lee,

Jaechanax insignis (Fairmaire), 7. major major (Pic), Macroeubria bella Lee, Yang &

Sato, M. bicolor Lee, Yang & Satò, M. contracta Lee, Yang & Satò, M. monstrosa

Lee, Yang & Satò, M. truncata Lee, Yang & Satò, Odontanax laosensis (Pic) and

Schinostethus (Sundodrupeus) priscus Lee, Jich & Yang.

Four new species are described and illustrated: Ectopria iranensis n. sp., from

Iran, Macroeubria shimadai n. sp., from Malaysia, S. (Sundodrupeus) sumatrensis n.

sp., from Sumatra and Schinostethus (Sundodrupeus) transversus n. sp., from India.

244 C.-F LEE & M. JACH

RIASSUNTO

Nuove specie e nuovi reperti di Psephenidae dell’Asia (Coleoptera).

Si forniscono nuovi dati sulla distribuzione delle seguenti specie: Dicranop-

selaphus brevicornis Lee & Yang, D. emmanueli (Pic), D. flavus Lee & Yang, D.

imparis Lee & Yang, D. malickyi Lee & Yang, Granuleubria pakistanica Jach & Lee,

Faechanax insignis (Fairmaire), 7. major major (Pic), Macroeubria bella Lee, Yang &

Sato, M. bicolor Lee, Yang & Satò, M. contracta Lee, Yang & Satò, M. monstrosa

Lee, Yang & Satò, M. truncata Lee, Yang & Satò, Odontanax laosensis (Pic) e Schi-

nostethus (Sundodrupeus) priscus Lee, Jich & Yang.

Sono inoltre descritte ed illustrate quattro nuove specie: Ectopria iranensis

n. sp., dell Iran, Macroeubria shimadai n. sp., della Malaysia, S. (Sundodrupeus)

sumatrensis n. sp., di Sumatra and Schinostethus (Sundodrupeus) transversus n. sp.,

dell’ India.

RoBERT D. GORDON* & CLAUDIO CANEPARI**

245

SOUTH AMERICAN COCCINELLIDAE (COLEOPTERA),

PART XI: A SYSTEMATIC REVISION

OF HYPERASPIDINI (HYPERASPIDINAE)

TABLE OF CONTENTS

Introduction

Biology

Diagnostic characters

Methods

Discussion of type materials

List of South American Hyperaspidini

Systematics

Tribe Hyperaspidini

Key to genera of Hyperaspidini

Genus Diazonema Weise

Genus Menoscelis Mulsant

Genus Thalassa Mulsant

Genus Clypeaspis, n. gen.

Genus Prognataspis, n. gen.

Genus Peruaspis, n. gen.

Genus Hyperaspis Chevrolat

Section I

The donzeli group

The chapini group

pag.

246

247

248

250

251

253

256

258

259

271

275

278

280

282

286

297

303

* Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA

c/o U. S. National Museum Natural History, Washington, DC 20560 Mailing address:

Northern Plains Entomology, P. O. Box 65, Willow City, ND 58384 (U.S.A.)

Tel.: (701) 366-4504 e-mail: rdgordon@utma.com

** Museo Civico di Storia Naturale di Milano

Mailing address: Via Venezia 1, 20097 San Donato Milanese (MI) (Italy)

e-mail: ccanepari@libero.it

246 R.D. GORDON & C. CANEPARI

The onerata group 308

The joannae group 599

The howdeni group 347

Species not assigned to a Group 32

Species known only from females 358

Section II 371

The vredenburgi group 371

The conclusa group 381

Genus Tenuisvalvae Duverger 398

Acknowledgements 413

References 7 414

Abstract 417

Riassunto 418

Index to scientific names 419

‘Tables 422

INTRODUCTION

This study continues the revision of South American Coccinel-

lidae. Parts I-X dealt with a variety of taxa with particular emphasis

on Coccidulinae and Scymninae. Here we begin to deal with the

extensive list of taxa in Hyperaspidinae, beginning with the tribe

Hyperaspidini.

Hyperaspidinae are small, usually colorful, highly patterned lady

beetles primarily Western Hemisphere in distribution. As such, they

attracted the attention of early taxonomic researchers who created a

great many specific names as well as some confusion in categories

above the species level. MULSANT (1850) described several genera

and subgenera of “Hyperaspiens” as well as many species. He di-

two of which,

contain the taxa herein dealt

vided his “Hyperaspiens” into several “Branches,’

“Hyperaspiares” and “Thalassiaires,

with. Mulsant’s genus Cleothera, with two subgenera, Cleothera and

Cyra, was distinguished from Hyperaspis Redtenbacher primarily by

the presence of a basal pronotal line (Hyperaspis) or absence of a line

(Cleothera). This character was an unfortunate choice because it is

illusory or absolutely fails in many species. Mulsant himself descri-

bed taxa in Cleothera that were actually Hyperaspis, and vice versa.

CroTcH (1874) further muddled the picture by synonymizing Cleo-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 247

thera under Hyperaspis. Subsequent authors either followed Mulsant

or Crotch for generic assignments, creating a situation where correct

generic assignments can be arrived at only by examination of type

specimens.

CHAPIN (1966) was first to make sense of some generic categori-

es by using genital and antennal characters not observed by earlier

workers. EL ALI (1972) further clarified generic relationships, but it

was DUVERGER (1989) who clarified distinctions between Brachiacan-

thini Mulsant and Hyperaspidini, providing a suite of characters for

each taxon that allows accurate generic assignment. He used both

male and female genitalic structures as well as an external character

to distinguish the tribes.

Some genera of Hyperaspidini have been studied by Milléo

and colleagues. Tiphysa Mulsant was placed in Hyperaspidini by

DUVERGER (1989), but MiLLéo & ALMEIDA (2003) transferred it

to Brachiacanthini. Diazonema Weise was revised by MILLEO &

ALMEIDA (2000) and Thalassa Mulsant was revised by MILLEO et

al. (2004). Three previously unrecognized genera are described he-

rein.

BIOLOGY

Host records are not available for most South American spe-

cies of Hyperaspidini. Data for North American species of Hype-

raspis (GORDON 1985) indicate that scale insects of the Coccidae and

Pseudococcidae are common prey, although aphids and other Ho-

moptera have been recorded (http://www.sel.barc.usda.gov/scalenet/

query.htm). Biological data for members of Menoscelis did not exist

until ORIVEL et al. (2004) published on the biology of “Thalassa

saginata” (Menoscelis saginata). They discovered that the larvae of

this species develop inside colonies of the arboreal ant Dolichoderus

bidens (L.), and are obligatory myrmecophiles. They also stated that

a study of arboreal ant colonies did not reveal the presence of M.

saginata in nests of other than ant species.

The list below is taken from literature sources and from label

data. Host names have been verified or corrected using ScaleNet,

a computerized database of scale names that can be accessed at:

http://www.sel.barc.usda.gov/scalenet/query.htm.

248 R.D. GORDON & C. CANEPARI

Clypeaspis trilineata (Mulsant)

Pseudococcus calceolariae (Maskell) - ScHILDER & SCHILDER

(1928)

Saccharicoccus sacchari (Cockerell) - SCHILDER & SCHILDER

(1928)

Hyperaspis pseudodonzeli n. sp.

Orthezia insignis [Insignorthezia insignis (Browne)] - label data

Hyperaspis orthivora n. sp.

Orthezia sp. - label data

Hyperaspis onerata (Mulsant)

Orthezia sp. - label data

Phenacoccus sp. - label data

Pulvinaria sp. - label data

Hyperaspis festiva Mulsant

Sipha flava (Forbes) - SCHILDER & SCHILDER (1928)

Saccharicoccus sacchari (Cockerell) - SCHILDER & SCHILDER

(1928)

Hyperaspis connectens (‘Thunberg)

Sipha flava (Forbes) - SCHILDER & SCHILDER (1928)

Tenuisvalvae notata (Mulsant)

Phenacoccus cassava ? - label data

Phenacoccus herreni Cox and Williams - label data

Tenuisvalvae quinquepustulata (F.) = raynevali (Mulsant)

Phenacoccus sp. - label data

Planococcus citri (Risso) - label data

Tenuisvalvae bromelicola (Sicard)

Pseudococcus bromeliae = Dysmicoccus brevipes (Cockerell) -

SICARD (1925)

DIAGNOSTIC CHARACTERS

Principal morphological terms used in descriptions and keys are

briefly explained to simplify key and description usage.

Parente ta rrone and “swim cee: Culp tu nie). “Relative

size of punctures on head, thorax, and elytral and ventral surfaces

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 249

is often useful. Size and spacing of punctures on various structures

are compared. Spacing of punctures is expressed as “separated by a

3 166

diameter (of a puncture),” “separated by less than a diameter,” etc.

Color, particularly dorsal patterns, is a primary distinguishing cha-

racter. Surface appearance varies from shiny to quite dull depending

on degree of alutaceous sculpture.

Head. The clypeus is long or short, concealing the labrum

or not, apically emarginate. ‘The frons and clypeus are joined either

abruptly (Fig. 1), or smoothly, obliquely (Fig. 11), and the inner

eye margin is smoothly rounded, lacking an eye canthus projecting

from frons onto inner margin of eye.

Prosternum. Prosternal carinae usually converge at some

point from apex to base of prosternum, but usually do not reach the

base before converging, creating a “stem” that reaches the base. In a

few species the carinae are incomplete, not convergent, or absent.

Abdomen. Postcoxal line configuration is quite uniform,

but the line may or may not extend to the posterior margin of the

1° abdominal sternum; it may be flattened medially along posterior

margin or rounded; and the recurved apical portion varies in degree

of anterior extension.

Male genitalia. Genital structure is important for iden-

tification of males. GORDON (1985) illustrated and defined genitalic

parts in detail, and the most important structures are discussed here.

Basal lobe: median projection of phallobase, serving as a siphonal

guide; a simple structure, differing in length and shape. Parameres:

paired lateral projections of phallobase serving to position and hold

basal lobe in position during copulation. Phallobase: includes the

basal piece, basal lobe and paramere of male genitalia. Sipho: sclero-

tized rod inserted through the basal lobe and into the female bursa

copulatrix during copulation; corresponds to aedeagus or penis, dif-

fering in length, form of apex, and structure of basal capsule.

Female genitalia (Fig. 143). Two forms of genital pla-

tes occur in Hyperaspidini: one is short, more or less transverse, as

in Hyperaspis species, and the other long, slender, as in Tenuisvalvae

species. The spermathecae are structurally unique within the Coc-

cinellidae, making the homologies of component structures unclear.

After discussion with N. Vandenberg (pers. comm.), it seems best

to use a modified version of DOBZHANSKyY’s (1941) terminology. ‘The

250 R.D. GORDON & C. CANEPARI

spermatheca is divided into a slender basal “tube” (basal unit) and a

retort-shaped distal capsule united to the basal portion by a slender

connecting duct. The basal unit is tube shaped, lacks a visible ramus,

accessory gland is present (not shown in many illustrations), and is

rounded apically, with an apical “beak” (chitinized appendix). The

distal portion is more or less rounded, not noticeably sclerotized,

and apparently devoid of distinguishing characters (see Fig. 143).

METHODS

Morphological structures, both external and internal, were di-

scussed and illustrated by GORDON (1985). Most of the same termi-

nology is followed here, and that publication should be consulted

if clarification is needed. Genital techniques are discussed under

“Dissection” below. Lectotype and paralectotype labels were affixed

to specimens so designated throughout. Label data for all newly de-

scribed taxa are transcribed exactly as they appear on the label. All

locality records listed in the text were taken from specimens actually

examined; published records were not accepted because a defining

criteria at the species level, genitalia, were not examined by most

previous authors.

Dissection. Both sexes should be dissected when examining

Hyperaspis specimens. Specific techniques consist of softening a

specimen in hot water, removing the abdomen, placing it in a dilute

solution of potassium or sodium hydroxide until muscle and fat are

removed, rinsing abdomen and genitalic structures in clear water,

and placing cleaned structures in glycerine for examination. Genitalia

may be stored in several ways, but here they were stored in glyceri-

ne microvials or mounted on a card attached to the donor specimen.

‘Type material. Type specimens have been examined for all spe-

cific taxa unless otherwise indicated. Detailed information is inclu-

ded under “Type locality” and “Remarks”. ‘The “Methods” sections

in GORDON (1985, 1999) contain discussions of locality records and

primary type depositories that are applicable here.

Collections. The following acronyms denote depositories for spe-

cimens used in this study: (BM) Natural History Museum, London;

(CAS) California Academy of Sciences, San Francisco, California;

(CCM) Claudio Canepari, Milan; (CDA) California Department of

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 251

Food and Agriculture; (CMP) Carnegie Museum of Natural Hi-

story, Pittsburgh, Pennsylvania; (CNC) Canadian National Collec-

tion, Ottawa, Ontario; (DEI) Deutsches Entomologisches Institut,

Miuncheberg; (DZUP) Universidade Federal do Parana, Curitiba,

Brazil; (GG) Guillermo Gonzalez, Santiago, Chile; (MBR) Museo

Argentino de Ciencias Naturales “Bernardo Rivadavia”, Buenos Ai-

res, Argentina; (MHNG) Muséum d’Histoire Naturelle, Geneva,

Switzerland; (MNHL) Muséum d’Histoire Naturelle, Lyon, Fran-

ce; (MNHP) Muséum National d’Histoire Naturelle, Paris, Fran-

ce; (MHNS) Museo Nacional de Historia Natural, Santiago, Chile;

(NHMV) Naturhistorisches Museum, Vienna, Austria; (MSNG)

Museo Civico di Storia Naturale “Giacomo Doria”, Genova, Italy;

(NREA) Naturhistoriska Riksmuseum, Entomologiska Avdelningen,

Stockholm, Sweden; (SMTD) Staatliches Museum ftir Tierkun-

de, Dresden, Germany; (UMZC) Cambridge University Museum,

Cambridge, England; (USNM) U.S. National Museum of Natu-

ral History, Smithsonian Institution, Washington, DC; (ZMHB)

Zoologisches Museum, Humboldt Universitàt, Berlin, Germany;

(ZMMU) Zoological Museum of Moscow, Lomosov State Univer-

sity, Moscow, Russia; (ZMUC) Zoological Museum, University of

Copenhagen, Denmark.

DISCUSSION OF TYPE MATERIALS

Most previously described species of Hyperaspidinae were na-

med by MULSANT in a series of publications. His 1850 treatment of

the world Coccinellidae was the first, and remains the most signifi-

cant publication in the annals of lady beetle systematic. In that work

he listed material examined from many sources, both public and

private, listing the species locality followed by the collection from

which the specimens came, e.g., Collect. Dejean, Chevrolat, etc.

Most of these collections have been preserved in present museum

holdings, but a few cannot be located; hence species placements in

Hyperaspidini or Brachiacanthini are unknown or doubtful.

Types of species described by T. KIRscH (1876, 1883), mostly

from Peru, were previously presumed destroyed because an inqui-

ty concerning the types, which was directed to Dresden (SMTD)

30 years ago, elicited that response. Helmut Firsch, Passau, Ger-

SI R.D. GORDON & C. CANEPARI

many, informed us (pers. comm.) that at least some Kirsch types are

present in Dresden. Subsequent communication with Olaf Jaeger

(SMTD) confirmed this, and we have been able to examine most

of the needed specimens. MULSANT (1870) described several species

from the Bruck collection. Helmut Fursch (pers. comm.) informed

us that the Emil von Bruck collection had been in Bonn and was

destroyed by bombing in 1944. Therefore, the identity of those spe-

cies is questionable. Specimens from the Trobert collection cannot

be traced in spite of numerous inquires directed to museums. Spe-

cimens from the Melly collection also cannot be traced. In addition,

some taxa whose type material should be in existing collections, as

indicated by Mulsant, are simply not there. Following is a list of

Hyperaspidinae species for which type material has not been loca-

ted. "The Mulsant species are followed by the name of the original

collection from which his specimens came.

Cleothera adhaerens Mulsant - Sallé

Cleothera cincta Kirsch - should be in the SMTD but not located there.

Cleothera collaris Mulsant - Buquet, Melly, should be in BM, but

not located there.

Cleothera flavida Mulsant - Bruck

Cleothera florifera Vogel - should be in the ZMHB but not located there.

Cleothera gemellata Mulsant - Bruck

Cleothera gracilis Mulsant - Germar and Schaum, should be in the

ZMHB but not located there.

Cleothera graminicola Mulsant - Trobert

Cleothera laqueata Mulsant - Bruck

Cleothera matsoni Mulsant - Bruck

Cleothera millieri Mulsant - Trobert

Cleothera ormanceayi Mulsant - Germar and Schaum, Trobert,

should be in the ZMHB but not located there.

Cleothera ovatonotata Mulsant - Trobert

Cleothera pallax Mulsant - Sallé

Cleothera pedicata Mulsant - Bruck

Cleothera ponderosa Mulsant - Bruck

Cleothera prettosa Mulsant - Bruck

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 253

Cleothera proserpinae Mulsant - Bruck

Cleothera puella Mulsant - Bruck

Cleothera punctum Mulsant - Chevrolat, should be in the UMZC

but not located there.

Cleothera quadrina Mulsant - Deyrolle, should be in the UMZC but

not located there.

Cleothera septenaria Mulsant - Bruck

Cleothera subparallela Mulsant - Bruck

Cleothera turbata Mulsant - Germar and Schaum, should be in the

ZMHB but not located there.

Cleothera vaticina Mulsant - Bruck

Cleothera venalis Mulsant - Bruck

Hyperaspis vetusta Weise - Should be in the ZMHB but not located

there.

Cleothera vexata Mulsant - Should be in the UMZC but not located

there.

Lectotypes for several species are designated herein to stabilize

the current classification for future researchers.

LIST OF SOUTH AMERICAN HYPERASPIDINI

Diazonema Mulsant

1. hypocrita (Mulsant)

2. fallax Weise

. pubescens (Weise)

. cavifrons Weise (not South American)

. boothi, n. sp.

. murilloi, n. sp.

Nm DB UYU

7. eccentrica, n. Sp.

Menoscelis Mulsant

1. saginata Mulsant

2. insignis Mulsant

Thalassa Mulsant

1. pentaspilota (Chevrolat)

2. flaviceps Mulsant (not South American)

3. similaris Mulsant

4. glauca (Mulsant)

254 R.D. GORDON & C. CANEPARI

5. korschefskyi Milléo, Almeida & Gordon

6. montezumae Mulsant (not South American)

Clypeaspis, n. gen.

1. trilineata (Mulsant)

Prognataspis, n. gen.

1. surreptiva, n. sp.

Peruaspis, n. gen.

1. paprzycRii, n. sp.

2. hypocrita, n. sp.

Hyperaspis Redtenbacher

Section I

donzeli group

1. latitibia, n. sp.

2. donzeli Mulsant

3. matronata Mulsant

4. helveola, n. sp.

chapini group

5. scutifera Mulsant

6. imitatrix, n. sp.

7. chapini, n. sp.

onerata group

8. cracentis, n. Sp.

9. operaria (Mulsant)

10. limbigera (Mulsant)

11. bisignata, n. sp.

12. camargoi, n. sp.

13. biguttata Sicard

14. clerda Mulsant

15. tayronensis, n. sp.

16. pseudodonzeli, n. sp.

17. orthivora, n. sp.

18. zonula, n. sp.

19. brethesi, n. sp.

20. rosariensis, n. sp.

21. stladesma, n. sp.

22. octonotata, n. sp.

23. campbelli, n. sp.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI

24. colombiensis, n. sp.

25. dissidens, n. sp.

26. uninotata, n. sp.

27. onerata (Mulsant)

28. bicruciata (Mulsant)

joannae group

29. joannae, n. sp.

30. albopunctata Crotch

31. apicaspis, n. sp.

32. delicata Almeida & Vitorino

33. mimica, n. sp.

34. laterimacula, n. sp.

howdeni group

35. howdeni, n. sp.

36. chocoi, n. sp.

07. Gira, Niwas?

38. recordata Mulsant

Species not assigned to a group

39. eupaleoides Crotch

40. mariposa, n. sp.

41. stmlaensis, n. sp.

42. lindae, n. sp.

Species known only from females

43. c-nigrum Mulsant

44. sagittata Crotch

45. guilloryi (Mulsant)

46. pseudopavida, n. sp.

47. istmina, n. sp.

48. satipoensis, n. sp.

49. abertha, n. sp.

50. circumclusa, n. sp.

51. aemulata, n. sp.

52. dispar, n. sp.

53. histrionica (Mulsant)

Section II

vredenburgi group

54. ayacucho, n. sp.

55. herrerai, n. sp.

255

256 R.D. GORDON & C. CANEPARI

56. vredenburgi, n. sp.

57. festiva Mulsant

58. germaini Crotch

59. connectens (Thunberg)

conclusa group

60. ingrata Mulsant

61. funesta (Germain)

62. nana Mader

63. sphaeridoides Mulsant

64. conclusa Weise

65. arida, n. sp.

66. elegantissima Brèthes

67. longula Weise

68. prolata, n. sp.

Tenuisvalvae Duverger

1. unipunctata (Crotch)

. peregrina (Mulsant)

. deyrollei (Crotch)

. notata (Mulsant)

. bisquinquepustulata (F.)

. ecoffett (Mulsant)

. parenthesis, n. sp.

. caucaensis, n. Sp.

. bromelicola (Sicard)

SOON SVI BW WN

. gnoma, n. Sp.

SYSTEMATICS

Hyperaspidini Brèthes, 1923

Hyperaspidini: Brèthes, 1923: 454; Winco 1952: 17; DUvVERGER 2001: 222;

VANDENBERG 2002: 383; DUVERGER 2003: 67.

Hyperaspidina Jacobson, 1916: 969.

Hyperaspini Costa, 1849: 9; WEISE 1885: 5; CHAPIN 1966: 278; GORDON 1985: 352;

DUVERGER 1989: 146.

Description. Hyperaspidinae of small to medium size;

form ranges from elongate oval, depressed, to rounded, convex.

Dorsal surface glabrous except Blaisdelliana Gordon. Antenna short,

with 9 to 11 articles; club elongate, fusiform, apical article small, re-

SOUTH AMERCAN. COCCINELLIDAE, XI: HYPERASPIDINI 254

cessed in preceding article; antennal insertion exposed or concealed.

Eye large, entire, not emarginated by an eye canthus (Figs. 7, 11),

finely faceted, without pubescence. Maxillary palpus with apical seg-

ment securiform. Scutellum usually large. Epipleuron narrow, usu-

ally excavated for reception of femoral apices except Hyperaspidius

Crotch and Blaisdelliana. Leg short; femur grooved or flattened for

reception of tibia; protibia slender, unmodified, or expanded; tarsus

cryptotetramerous; claws toothed or not. Abdomen without gland

openings, with 6 visible sterna. Male genitalia with basal lobe asym-

metrical, toothed, lobe rooted in phallobase (Figs. 75, 80). Female

spermatheca compound, basal unit and apical portion connected by

a short or long duct (Fig. 108); genital plate basically transverse

(Fig. 108), or long, slender, stylus reduced or absent (Fig. 494).

Remarks. Eight South American genera are recognized as

belonging to Hyperaspidini: Hyperaspis Chevrolat, Tenuisvalvae Du-

verger, Thalassa Mulsant, Menoscelis Mulsant, Diazonema Mulsant,

Clypeaspis, new genus, Prognataspis, new genus, and Peruaspis, new

genus. SICARD (1912) described Helesius caseyi from Colombia, but

examination of a syntype of that species revealed that it is a member of

Cyra. Therefore, Helesius is not currently known from South America.

Of Hyperaspidine genera, Hyperaspis contains by far the greater

number of species and comprises the bulk of this study. Hypera-

spis, Tenuisvalvae, Menoscelis, and Diazonema are completely re-

vised, and Clypeaspis, Prognataspis, and Peruaspis newly described.

Thalassa was treated by MILLEO et al. (2004); therefore we provide

only species key and new information.

Diazonema and Clypeaspis have 9 or 10-articled antennae with

antennal bases concealed beneath the eye canthus, and strongly

modified protibiae. Thalassa, Menoscelis, Peruaspis, Hyperaspis, and

Tenuisvalvae have 10 or 11-articled antennae with antennal bases

partially concealed and protibia unmodified or weakly modified in

the latter three genera.

MULSANT’s original (1850) classification of these taxa was spo-

radically altered by subsequent authors such as CROTCH (1874) and

CHapUuIs (1876). Our investigations, for the most part, confirm Mul-

sant’s original arrangement of taxa conceived more than 150 years ago.

Hyperaspidini and Brachiacanthini are distinguished by both ex-

ternal and genital characters. Hyperaspidini are distinguished by the

258 R.D. GORDON & C. CANEPARI

lack of an eye canthus, inner margin of eye smoothly rounded (this is

an excellent character by which specimens are quickly distinguished

from brachiacanthines); presence of a fine, basal pronotal line, and

basomedian projection of pronotum truncate (not dependable cha-

racters). Male genitalia have parameres rooted inside basal piece.

Female spermathecae are compound, lacking infundibulum and visi-

ble ramus, with a basal unit separated from apical portion by a duct.

Brachiacanthini are distinguished by having an eye canthus of

varying size and shape extended from frons onto inner margin of

eye, emarginate eye (R. Booth, pers. comm.); without basal pronotal

line, basomedian projection of pronotum not truncate. Male genita-

lia with parameres attached to outer portion of basal piece (as typi-

cal for most Coccinellidae). Female genitalia of typical coccinellid

type with single spermathecal capsule composed of ramus, nodulus,

and cornu; with distinct infundibulum.

KEY TO GENERA OF HYPERASPIDINI

ie Postcoxal line on basal abdominal sternum incomplete, of

Scynmobiis iy pe (Pig) S7)IIAUP 100 2) Pe Menoscelis Mulsant

Postcoxal line on basal abdominal sternum incomplete, of

EVI ms CwMinus) Tye TRIO GOV oo... ee a he we es 2

2(1). Labrum extended beyond clypeal apex, wide, apically trun-

cate, dark brown, heavily sclerotized (Fig. 6); prosternal

process prominent, raised (Figs. 4, 5) ..... Peruaspis, n. gen.

Labrum extended beyond clypeal apex or not, narrow, api-

cally rounded, weakly sclerotized (Figs. 11, 12), yellow or

pale brown; prosternal process not prominent (Figs. 10, 12) 3

3(2). Protibia angulate near base, posterior margin strongly flan-

Poumon) ti. Mae BIO DI DAB Ce 4

Protibia not angulate near base, flange on posterior margin

(GE ON AMSEC o ca aha x Aa e wae LS 5

4(3). Antenna 9-articled (Fig. 25); frons on a distinctly higher

plane than clypeus (Fig.13); ground color of elytron usually

pale red (except murillot and eccentrica)....Diazonema Weise

Antenna 11-articled (Fig. 43); frons on a slightly higher

5(3).

6(5).

7(6).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 259

plane than clypeus; ground color of elytron black, with

bluish.coriereenishtlustieprnasetanetisa can Thalassa Mulsant

Eye large, approximately as wide as median frontal area

when viewed from front (Fig. 53); protibia distinctly flan-

ged; abdominal sterna 3-5 with pit on each side near lateral

Mare; ove 0 SORTE INTO Prognataspis, n. gen.

Eye small, frontal area 2 to 3 times wider than eye when

viewed from front (Fig. 1); protibia flanged or not; abdo-

minal sterna 3-5 without pit on each sided............ 6

Body elongate, nearly parallel sided, dorsoventrally flatte-

ned (Fig. 47); protibia distinctly flanged, flange narrow at

base; widened toward apex (Fig 2). 522% Clypeaspis, n. gen.

Body elongate or rounded, not parallel sided, dorsoventrally

flattened or not; protibia not flanged or feebly so (Figs. 9, 74) 7

Female genital plate short, subquadrate (Fig. 97); female

abdominal sternum 6 apically convex; antenna with 10 or 11

articles; ultimate maxillary palpal article truncate; mandi-

bulary retinaculum slightly curved, apex acute ......

PA ALOR) DURE MOTORI A): PO tk ARES Hyperaspis Chevrolat

Female genital plate long, slender (Fig. 494); female ab-

dominal sternum 6 apically triangular; antenna with 11

articles; ultimate maxillary palpal article slightly emar-

ginate; mandibulary retinaculum strongly curved, apex

POUHOCECS . ARI RR: Pr Ple eee Tenuisvalvae Duverger

Genus Diazonema Weise, 1926

Diazonema Weise, 1926: 12; KorscHersky 1931: 202; BLACKWELDER 1945: 448;

CHAPIN 1966: 279; MILLEo & ALMEIDA 2000: 66 (as synonym of Corystes).

Corystes Mulsant, 1850: 506 (not Corystes Latreille, 1802); CroTtcH 1874: 208;

CHAPUIS 1876: 249; KORSCHEFSKY 1932: 267; BLACKWELDER 1945: 448; MILLEO

& ALMEIDA 2000: 66 (not Corystes Latreille, 1802).

Corystes (Diazonema): DUVERGER 1989: 146.

Type species: of Corystes, Corystes hypocrita Mulsant, by mo-

notypy; of Diazonema, Diazonema cavifrons Weise, by subsequent

designation of KORSCHEFSKY (1932).

260 R.D. GORDON & C. CANEPARI

Description. Hyperaspidini with form rounded, convex.

Dorsal surface glabrous or pubescent. Elytral ground color variable.

Elytron maculate or not. Antenna with 9 articles, basal article wider

than long (Figs. 25, 31); antennal insertion concealed. Apical maxil-

lary article straight apically; mandibulary retinaculum strongly cur-

ved, apex rounded. Clypeus short, narrow, slightly extended beyond

frons, on lower plane than frons, abruptly joined to frons, apically

emarginate (Figs. 13, 15). Frons widely flared, concealing antennal

base. Labrum concealed beneath clypeus. Scutellum large, wider

than long. Elytral epipleuron wide, not medially grooved, descen-

ding externally, excavated for reception of middle and hind femoral

apices. Prosternal process slightly convex, with 2 carinae conver-

gent toward base. Legs with tibiae short. Protibia short, wide, flat-

tened, tibial groove strongly flanged, outer margin of flange slight-

ly sinuate, flange as wide at base as apex, angulate near base (Fig.

26); meso- and metatibia narrow, medially emarginate, not flanged,

with distinct tibial groove, angulate near base. Posterior margin of

metasternum abruptly descending between coxa and lateral margin.

Metendosternite with deep, V-shaped anterior border. T'arsal claw

with basal tooth. Postcoxal line on basal abdominal sternum incom-

plete, of Scymnus (Scymnus) type (Fig. 17). Male apical abdomi-

nal sternum weakly modified. Apex of female abdominal sternum 6

rounded. Female genital plate transverse.

Remarks. Diazonema (as Corystes) was revised by MILLEO &

ALMEIDA (2000), and that publication should be referred to for detai-

led discussions and illustrations. We do not entirely agree with their

conclusions regarding species synonymy. ‘Therefore, we have rein-

stated some species names as valid taxa. Male genitalia are extreme-

ly similar in all Diazonema species, suggesting that only a single taxa

exists, but external characters such as pubescence or lack of pubescen-

ce, and type of prosternal process, indicate there are six valid species.

A single specimen from Nicaragua, in the Natural History Mu-

seum, London collections, was examined but not identified. It is

most similar to D. fallax and D. hypocrita but differs from both. Be-

cause it is a single specimen lacking any positively definitive charac-

ter, we label it only as “Diazonema sp.” in the hope that additional

specimens will be found to clarify its taxonomic position.

Diazonema is most similar morphologically to Menoscelis (see re-

marks under that genus); it differs from all other genera of Hypera-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 261

spidinae by a combination of short clypeus, strongly expanded frons

on a higher plane than the clypeus, concealing antennal bases, 9-ar-

ticled antenna, and short, compressed protibia.

We thank Roger Booth for informing us that Corystes Mulsant,

1850 is preoccupied by Corystes Latreille, 1802. Therefore, Diazonema

Weise, 1926 becomes the next available name for this group of species.

KEY TO SPECIES OF DIAZONEMA

21).

a1).

4(2).

5(4).

6(4).

Elytron reddish yellow, immaculate, or with black border

CURUD ET PcARUIE NODE Fcc OW Bp otk eo A na Pe eee Soe per Sm She TY 3

Mistral’ eround color blaela*mmaeilate* 238 DIE eee Rai 2

Elytron with single, oval, yellow spot on apical declivity

(Dio 244 ASSALTI A LR 6. murilloi, n. sp.

Elytron with 3 red and yellow spots ..... 7. eccentrica, n. Sp.

Dorsal surface pubescent, pubescence composed of very

short Jagine SDA, TAGUUE <> ot meat Se 3. pubescens (Weise)

Porsal surfacetplabtousi tie elia lo Strole Tani bdito. 4

Head with frons and clypeus deeply dished, frons on a

‘much higher plane than clypeus, with pronounced horns;

Central America oF. Geuacor. so lu etaclas. ae ke oe 5

Head with frons and clypeus slightly depressed, frons on a

slightly higher plane than clypeus; not known from Central

FAMINE RICA ' ph i OMe NSPE E, DI Ae RI TREE es be BGP ee eee 6

Frontal horns apically angulate; ventral surface reddish

brown; Central America....... MET OU fi. 4. cavifrons Weise

Frontal horns apically acute, brown; ventral surface mostly

dark brown!) Eougdor (Pig tes, 22a be È 5. boothi, n. sp.

Prosternal process strongly narrowed basally, abruptly wi-

dened apically; elytral apex narrowly yellow; northeastern

and eastern South America.......... 1. hypocrita (Mulsant)

Prosternal process slightly narrowed basally, slightly wider

apically; elytral apex broadly yellow; Andean South Ameri-

damme, Ofte. ¢margiuuate modali not Sea 2. fallax Weise

262 R.D. GORDON & C. CANEPARI

1. Diazonema hypocrita (Mulsant, 1850), n. comb.

Corystes hypocrita Mulsant, 1850: 507; CroTcH 1874: 208; KORSCHEFSKY 1932: 267;

GORDON 1987: 26; MILLEO & ALMEIDA 2000: 71.

Corystes (Corystes) hypocrita: DUVERGER 1989: 146.

Type locality. Cayenne (French Guiana).

Description. 6, length 4.0 mm, width 3.6 mm; body

form round, convex. Dorsal surface glabrous; head dull, alutaceous;

pronotum dull, alutaceous; elytron shiny. Color yellow except pro-

~notum with 5 reddish yellow spots, median spot widely separated

from anterior margin, funnel shaped, with base narrow in front of

scutellum, apex wide, basal triangular spot laterad of scutellum, tri-

angular median spot in lateral 1/2; elytron pale red with narrow,

yellow border on lateral and apical margins (Fig. 16); pro-, meso-

and metasterna brownish yellow; abdomen dark brown except outer

1/8 yellowish brown. Head punctures fine, separated by a diameter

or less. Pronotal punctures fine, larger than on head, separated by

a diameter or less; elytral punctures slightly larger than pronotal

punctures, separated by 1 to 3 times a diameter. Metasternal punc-

tures slightly larger than on elytron, sparse, nearly absent. Punc-

tures on abdominal sterna slightly larger than on elytra, sparse me-

dially on basal sternum, dense, separated by less than a diameter on

sterna 2-6; area on 1°“ abdominal sternum inside postcoxal arc shiny,

impunctate. Clypeal apex narrow, deeply emarginate medially, not

sinuate laterally, lateral angle abruptly rounded. Prosternum narrow,

“pinched” at base, widened toward apex, sides straight, with cari-

nae narrowly separated at apex, convergent toward base, extended

1/3 distance to basal sternal margin. Protibia short, wide, deeply

grooved for reception of tarsus, posterior margin strongly flanged.

Postcoxal line on 1" abdominal sternum extended to posterior sternal

margin, flattened along margin, apex recurved 1/4 distance to basal

sternal margin. Fifth sternum with apex broadly, weakly emargin-

ate; 6 sternum weakly emarginate. Genitalia with basal lobe 7/8

length of paramere, slender, sinuate, apex with one angle rounded,

opposite angle acute; paramere slender, slightly widened to round-

ed apex in apical 1/2 (Fig. 18); sipho long, slender, evenly curved,

basal capsule with inner arm short, abruptly recurved apically, outer

arm absent, base of sipho knobbed (Figs. 19, 20).

2. None examined.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 263

Variation. Length 4.0 to 5.3 mm, width 3.6 to 4.8 mm.

Geographical distribution: Northeastern and eastern South

America.

Specimens examined: 4. Brazil, “Amazons”; Sao Paulo; Suri-

nam, “Cayenne”, Uyttenboogaart (BM) (MHNL) (USNM).

Remarks. Diazonema hypocrita seems to be an eastern South

American taxa distinguished from other Diazonema species by key

characters. It most nearly resembles the Peruvian D. fallax, but the

dorsal punctures are finer and not as closely spaced as in D. fallax.

The lectotype designated here is in the Dejean collection and is

labeled “Cayenne, Lacordaire.”

2. Diazonema fallax Weise, 1926

Diazonema fallax Weise, 1926: 13; MiLLéo & ALMEIDA 2000: 71 (as synonym of

Corystes hypocrita).

Type locality. Callanga, Peru (ZMHB), lectotype designated by

MiLLfo & ALMEIDA (2000).

Description. d, length 4.7 mm, width 4.4 mm; body

form round, convex. Dorsal surface glabrous; head dull, alutaceous;

pronotum alutaceous, shiny; elytron weakly alutaceous, shiny. Color

yellow except pronotum with 5 reddish brown spots, median spot

narrowly divided medially, forming triangular spot on each side, fee-

bly connected to base by narrow stem, triangular, basolateral spot

laterad of scutellum, indistinct median spot in lateral 1/2; elytron pale

red with narrow, yellow border on lateral margin, border abruptly

widened on apical margin; pro-, meso- and metasterna brownish yel-

low; abdomen dark brown except outer 1/8 yellowish brown. Head

punctures fine, separated by less than a diameter. Pronotal punctures

larger than on head, separated by less than a diameter; elytral punc-

tures smaller than pronotal punctures, separated by 1 to 3 times a

diameter. Metasternal punctures larger than on elytron, sparse me-

dially, becoming coarser, separated by less than a diameter laterally.

Punctures on abdominal sterna larger than on elytra, very coarse,

somewhat dense medially on basal sternum, smaller, separated by

less than a diameter on sterna 2-6; area on 1* abdominal sternum

inside postcoxal arc slightly alutaceous, shiny, impunctate. Clypeal

apex narrow, deeply emarginate medially, not sinuate laterally, lateral

204 R.D. GORDON & C. CANEPARI

angle abruptly rounded. Prosternum wide, slightly narrowed at base,

slightly widened toward apex, sides weakly rounded, with carinae

widely separated at apex, convergent toward base, extended nearly

1/2 distance to basal sternal margin. Protibia short, wide, deeply

grooved for reception of tarsus, posterior margin strongly flanged.

Postcoxal line on 1** abdominal sternum extended to posterior sternal

margin, flattened along margin, apex recurved 1/3 distance to basal

sternal margin. Fifth sternum with apex broadly, weakly emarginate;

6" sternum weakly emarginate. Genitalia with basal lobe 7/8 length

of paramere, slender, sinuate, apex with one angle rounded, opposite

angle acute; paramere slender, slightly widened to rounded apex in

apical 1/2; sipho long, slender, evenly curved, basal capsule with inner

arm short, truncate apically, outer arm long, wide, apically knobbed.

2. Similar to d except apex of 6% abdominal sternum rounded.

Genitalia with beak of basal unit apically rounded.

Variation. Length 4.5 to 4.7 mm, width 4.3 to 4.4 mm. Pronotal

maculation may be indistinct, nearly invisible; elytral color pattern

varies from that described above to forms with narrow to broadly black

lateral elytral borders and mostly black epipleurae (Tingo Maria).

Geographical distribution: Peru.

Specimens examined: 10. Peru: Callanga; Sani Beni; Huanuco;

Tingo Maria (CAS) (USNM) (ZMHB).

Remarks. Diazonema fallax was described from Peru, and

all specimens examined have been from that country. This is appar-

ently the species actually described by MILLEo & ALMEIDA (2000) as

D. hypocrita because the female genitalia agree with their illustra-

tion. It is distinguished from D. hypocrita by key characters as well

as coarser, denser, dorsal and ventral punctation.

3. Diazonema pubescens (Weise, 1926), n. comb.

Corystes pubescens Weise, 1926: 34; KorscHEFSKY 1932: 268; MiLLéo & ALMEIDA

2000: 71 (as synonym of Corystes hypocrita).

Type locality. Ecuador.

Description. &, length 5.5 mm, width 5.0 mm; body

form round, convex. Dorsal surface entirely pubescent, pubes-

cence short, golden, moderately dense; head and pronotum aluta-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 265

ceous, slightly shiny; elytron shiny. Color yellow except head and

pronotum dark yellow; elytron pale red with narrow, yellow border

on lateral and apical margins; pro-, meso- and metasterna brown-

ish yellow; abdomen reddish brown except outer 1/4 yellow. Head

punctures coarse, separated by less than a diameter. Pronotal punc-

tures smaller than on head, separated by a diameter or less; elytral

punctures larger than pronotal punctures, separated by a diameter

or less. Metasternal punctures larger than on elytron, sparse medi-

ally, becoming coarser, separated by less than a diameter laterally.

Punctures on abdominal sternum larger than on elytra, coarse, sparse

medially on basal sternum, smaller, separated by less than a diam-

eter on sterna 2-6; area on 1" abdominal sternum inside postcoxal

arc slightly alutaceous, shiny, finely, sparsely punctate. Clypeal apex

wide, shallowly emarginate medially, sinuate laterally, lateral angle

acute. Prosternum wide, slightly narrowed at base, slightly wid-

ened toward apex, sides straight, with carinae narrowly separated at

apex, convergent toward base, extended nearly 1/2 distance to basal

sternal margin. Protibia short, wide, deeply grooved for reception

of tarsus, posterior margin strongly flanged. Postcoxal line on 1%

abdominal sternum extended to posterior sternal margin, rounded

along margin, apex recurved 1/3 distance to basal sternal margin.

Fifth sternum with apex broadly, weakly emarginate; 6° sternum

weakly emarginate. Genitalia with beak of basal unit of spermatheca

having apical projection, projection apically truncate.

3. Not known.

Variation. None observed.

Geographical distribution: Ecuador and Peru.

Specimens examined: 4. Ecuador: Province Loja, Zamora; Peru:

Junin, Chanchamayo, 1300 m (BM) (USNM) (NREA: holotype).

Remarks. ‘This species is easily recognized by the unique

dorsal pubescence, laterally sinuate clypeal apex, and pronotal punc-

tures smaller than head punctures. Males were not available for exam-

ination; therefore male genitalia could not be comparatively assessed.

4. Diazonema cavifrons (Weise, 1903), n. comb.

Corystes cavifrons Weise, 1903: 208; MiLLéo & ALMEIDA 2000: 71 (as synonym of

Corystes hypocrita).

266 R.D. GORDON & C. CANEPARI

Diazonema cavifrons: WEISE 1926: 12; KorscHEFsKy 1931: 202.

Corystes (Diazonema) cavifrons: DUVERGER 1989: 146.

Type locality. Turrialba, Costa Rica (ZMHB) (lectotype desig-

nated by MILLEO & ALMEIDA 2000).

Description. 6, length 4.5 mm, width 4.0 mm; body

form rounded, widest anterior to middle of elytra, convex. Dorsal

surface glabrous; head slightly alutaceous, shiny; pronotum slightly

alutaceous, shiny; elytron shiny. Color pale yellowish brown except

head yellow with outer margins of frontal horns and clypeal apex

light brown; pronotum with lateral 1/3 yellow, median 1/3 yellow-

ish brown with oval, yellow spot near base on each side of middle;

elytron with lateral margin slightly darker brown; propleuron, legs

yellow. Head impunctate. Pronotal punctures fine, separated by less

than to 3 times a diameter; elytral punctures smaller than pronotal

punctures, separated by less than to 3 times a diameter. Metasternal

punctures slightly larger than on elytron, sparse, widely separated.

Punctures on abdominal sterna larger than on elytra, very coarse,

somewhat dense medially on basal sternum, smaller, separated by less

than a diameter on sterna 2-6; area on 1% abdominal sternum inside

postcoxal arc slightly alutaceous, shiny, impunctate. Head triangu-

larly depressed medially; clypeal apex narrow, emarginate medially,

not sinuate laterally, lateral angle abruptly rounded; frontal horns

greatly expanded, raised, tapered to acute apex. Prosternal process

wide, pinched between coxae, widened toward apex, sides weakly

rounded, with carinae widely separated at apex, convergent toward

base, extended to apical margin of coxae. Protibia short, wide, deep-

ly grooved for reception of tarsus, posterior margin strongly flanged.

Postcoxal line on 1** abdominal sternum extended to posterior ster-

nal margin, rounded along margin, apex strongly recurved toward

basal sternal margin. Fifth sternum with apex broadly, weakly emar-

ginate; 6 sternum convex. Genitalia as described for D. hypocrita.

? Similar to male. Female genitalia as described for D. pubescens.

Variation. Length 4.5 to 4.0 mm. Frontal horns may be reduced

in size, somewhat flattened; pronotum with median yellow spots

sometimes indistinct.

Geographical distribution: Costa Rica.

Specimens examined: 6. Costa Rica, “Costa Rica”; Honduras,

“Honduras” (BM) (ZMHB).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 267

Remarks. Diazonema cavifrons has the frontal area deeply

dished as shown by the excellent photograph in MILLEO & ALMEI-

DA (2000: p. 73, fig. 24), and the frons raised, forming “horns”.

This head structure differs from other Diazonema species except D.

boothi. See remarks under that species.

5. Diazonema boothi n. sp.

Type material. Holotype dé: Equador (sic), 52526, Buckley,

Corystes hypocrita (handwritten), Fry Coll. 1905.100 (BM).

Description. 6, length 5.2 mm, width 5.0 mm; body

form round, convex (Fig. 21). Dorsal surface glabrous; head dull,

alutaceous; pronotum alutaceous, dull; elytron shiny. Color yellow

except head with outer margins of frontal horns and clypeal apex

dark brown; pronotum with 5 indistinct brown spots, median spot

narrowly divided medially, forming triangular spot on each side,

feebly connected to base by narrow stem, triangular, basolateral spot

laterad of scutellum; elytron pale reddish yellow, slightly paler to-

ward lateral and apical borders (Fig. 21); pro-, meso- and metaster-

na dark brown, nearly black; mesoleg with trochanter, basal 2/3 of

femur dark brown, metaleg with trochanter, basal 3/4 of femur dark

brown; abdomen dark brown except basal sternum red medially,

outer 1/4 of sterna 1-6 yellowish brown. Head punctures fine, sepa-

rated by a diameter or less. Pronotal punctures slightly larger than

on head, separated by a diameter or less; elytral punctures larger

than pronotal punctures, separated by less than to twice a diameter.

Metasternal punctures larger than on elytron, separated by less than

to twice a diameter medially, becoming coarser, separated by less

than a diameter laterally. Punctures on abdominal sterna larger than

on elytra, very coarse, somewhat dense medially on basal sternum,

smaller, separated by less than a diameter on sterna 2-6; area on 1*

abdominal sternum inside postcoxal arc slightly alutaceous, shiny,

impunctate. Head medially depressed; clypeal apex narrow, emar-

ginate medially, not sinuate laterally, lateral angle abruptly rounded;

frontal horns greatly expanded, raised, tapered to acute apex (Fig.

14). Prosternum wide, slightly narrowed at base, slightly widened

toward apex, sides weakly rounded, with carinae widely separated at

apex, convergent toward base, extended nearly 1/2 distance to basal

268 R.D. GORDON & C. CANEPARI

sternal margin. Protibia short, wide, deeply grooved for reception

of tarsus, posterior margin strongly flanged. Postcoxal line on 1%

abdominal sternum extended to posterior sternal margin, flattened

along margin, apex slightly recurved toward basal sternal margin.

Fifth sternum with apex broadly, weakly emarginate; 6° sternum

convex. Genitalia with basal lobe 7/8 length of paramere, slender,

sinuate, apex with one angle rounded, opposite angle acute; paramere

slender, slightly widened to rounded apex in apical 1/2 (Fig. 22);

sipho long, slender, evenly curved, basal capsule with inner arm

short, truncate apically, outer arm long, wide (Fig. 23).

2. Not known.

Etymology. This species is named for Roger Booth, a colleague

who has contributed greatly to this and other coccinellid studies, both

through loan of specimens and advice on numerous technical matters.

Remarks. Diazonema booth (from Ecuador) resembles the

Central American Diazonema cavifrons, but differs by having the

head punctate; elytral punctures larger than pronotal punctures;

metasternal punctures very large, dense; ventral surface mostly dark

brown, nearly black; and postcoxal line on basal abdominal ster-

num flattened along apical sternal margin, apex slightly recurved

toward basal margin. In D. cavifrons the head is impunctate; elytral

punctures smaller than pronotal punctures; metasternal punctures

small, sparse; ventral surface yellowish brown; and postcoxal line on

basal abdominal sternum rounded along apical sternal margin, apex

strongly recurved toward basal margin.

6. Diazonema murilloi n. sp.

Type material. Holotype dé: Colombia, Val (Valle del Cauca), Rol-

danillo, 966 m, 27.V.1939, Murillo No 5001 (USNM). Paratypes 4: 3,

same data as holotype (USNM); 1, Colombia, Cnd (Cundinamarca),

nr El Colegio, alt. 1600 m, 10.III.-‘40, Murillo No 5363 (USNM).

Description. 6, length 4.0 mm, width 3.6 mm; body

round, convex. Dorsal surface with head alutaceous, shiny; prono-

tum and elytron slightly alutaceous, shiny. Color black except head

yellow; pronotum with narrow apical margin and lateral 3/8 yellow;

elytron with greenish sheen, 1 large, oval, yellow spot on apical de-

clivity (Fig. 24); ventral surface reddish yellow except median area of

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 269

pro- and mesosternum, epipleuron black. Head punctures fine, sepa-

rated by less than a diameter; pronotal punctures larger than on head,

separated by a diameter or less; elytral punctures slightly smaller

than pronotal punctures, separated by a diameter or less. Metaster-

nal punctures fine, sparse, nearly absent Punctures on median area

of basal abdominal sternum coarse, sparse, punctures on sterna 2-6

smaller than on basal sternum, separated by 1 to 3 times a diameter;

area on 1*t abdominal sternum inside postcoxal arc shiny, impunctate.

Clypeal apex weakly emarginate, slightly sinuate laterally. Antenna

with 9 articles (Fig. 25). Prosternum with intercoxal process narrow,

pinched at base, widened apically, sides straight, carinae widely sepa-

rated at extreme apex, abruptly narrowed toward base, forming long,

apical stem, extended 3/4 distance to basal sternal margin. Protibia

short, wide, deeply grooved for reception of tarsus, posterior margin

strongly flanged Postcoxal line on 1% abdominal sternum reaching

posterior sternal margin, rounded along margin, apex recurved 1/4

distance to basal sternal margin. Fifth sternum with apex broadly,

weakly emarginate; 6" sternum truncate. Genitalia with basal lobe

7/8 length of paramere, slender, sinuate, apex with one angle round-

ed, opposite angle acute, small, angulate projection on one side;

paramere slender, lateral margins parallel to apex (Fig. 27); sipho

long, robust, evenly curved, basal capsule with inner arm short,

truncate apically, outer arm absent, base of sipho knobbed (Fig. 28).

2 Not known.

Variation. Length 4.0 to 4.3 mm, width 3.6 to 3.7 mm. The

paratype from E] Colegio has the elytral spot on apical declivity

small, irregularly rounded, reddish yellow.

Etymology. This species is named for L. M. Murillo, a Co-

lombian entomologist who collected this type series as well as many

more Colombian Coccinellidae.

Remarks. This species and D. eccentrica are the only spe-

cies of Diazonema with basically black elytra. Diazonema murilloi is

the only species having prosternal carinae with a long basal stem.

7. Diazonema eccentrica. n. sp.

Type material. Holotype 2: Colombia, Vaupes, River Vaupes,

x.x11.1952, D. J. Taylor. D402 B.W/. I. C.R.S. Expedition Colom-

270 R.D. GORDON & C. CANEPARI

bia, Pred. on coccid. D. 376. on cacao, Com. Inst. Ent. Coll. No.

13110, ?Thalassa sp., Det. R.D. Pope 1953 (BM).

Description. ®, length 2.7 mm, width 2.3 mm; body

form round, convex. Dorsal surface with head, pronotum, and

elytron shiny. Color black, head with median, elongate brown spot

on vertex and frons; pronotum with narrow anterior margin and lat-

eral 1/4 yellow; elytron with 3 spots, 1 large median, pale red spot

extended from near base to apical declivity, 1 small, elongate oval,

yellow spot medially on lateral margin, 1 yellow spot on margin at

apex (elytron damaged, extent of spot not determined) (Figs. 29,

30); ventral surface, including abdomen, yellow except pro-, meso-

and metasterna yellowish brown. Head punctures fine, separated

by less than to twice a diameter. Pronotal punctures larger than on

head, separated by less than to twice a diameter; elytral punctures

slightly larger than pronotal punctures, separated by 1 to 3 times a

diameter. Metasternal punctures larger than on elytron, sparse me-

dially, becoming denser laterally, separated by less than a diameter.

Punctures on median area of basal abdominal sternum larger than

on elytra, sparse, punctures fine, dense on sterna 2-6; area on 1%

abdominal sternum inside postcoxal arc slightly alutaceous, weakly

shiny, finely, sparsely punctate. Clypeal apex wide, deeply emargin-

ate medially. Prosternum narrow, carinae narrowly separated at apex,

convergent toward base, extended 4/5 distance to basal margin, not

basally stemmed, not reaching basal margin of sternum. Postcoxal

line on 1° abdominal sternum extended to posterior margin, flat-

tened along margin, recurved 1/5 distance to basal sternal margin.

Fifth sternum with apex truncate; 6° sternum rounded. Genitalia of

the Hyperaspis type with beak of basal unit elongate, wide; connect-

ing duct long (Fig 32).

3d Not known.

Etymology. The specific name is the Latin eccentricus, meaning

different, odd, referring to the different appearance of this species

compared to other species of Diazonema.

Remarks. Like D. murilloi, this species is an atypical mem-

ber of Diazonema because of the black elytral ground color and dis-

tinct maculation. It is also the smallest species in the genus and has

female genitalia of the Hyperaspis type. The holotype lacks the left

elytron and apical portion of the right elytron.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 2710

Genus Menoscelis Mulsant, 1850

Menoscelis Mulsant, 1850: 508; CroTcH 1874: 209; CHapin 1966: 279; EL-ALI 1972:

23:

Menoscelis (Thalassa): CHAPUIS 1876: 228.

Thalassa (Menoscelis): KORSCHEFSKY 1931: 208; BLACKWELDER 1945: 449; DUVERGER

1989: 146.

Type species: Menoscelis saginata, by subsequent designation of

CROTCH (1874).

Description. Hyperaspidini with form rounded, convex.

Dorsal surface glabrous. Elytron maculate. Antenna with 11 ar-

ticles, basal article wider than long (Fig. 36); antennal insertion

concealed. Apical maxillary article straight apically; mandibulary

retinaculum strongly curved, apex rounded. Clypeus short, wide,

on lower plane than frons, abruptly joined to frons, lateral angle

abruptly rounded, apically emarginate. Frons flared, concealing

antennal base. Labrum concealed beneath clypeus. Scutellum lar-

ge, as wide as long. Elytral epipleuron wide, not medially groo-

ved, obliquely descending externally, excavated for reception of

middle and hind femoral apices. Prosternal process convex, with

2 carinae convergent toward base. Legs with tibiae long. Protibia

narrow, flattened, medially emarginate, tibial groove deep, long,

not or slightly flanged posteriorly, angulate near base (Fig. 38,

39); meso- and metatibia narrow, medially emarginate, not flan-

ged, with shallow tibial groove. Posterior margin of metasternum

abruptly descending between coxa and lateral margin. Metendo-

sternite with deep, V-shaped anterior border. Tarsal claw with

basal tooth. Postcoxal line on basal abdominal sternum incomple-

te, of Scymnobius type (Fig. 37). Male apical abdominal sternum

weakly modified. Apex of female abdominal sternum 6 rounded.

Female genital plate transverse.

Remarks. Menoscelis has been variously considered a valid

genus or subgenus. MULSANT (1850) described it as a genus; CROTCH

(1874) and CHAPIN (1966) followed suit. CHApuIs (1876) considered

it a genus with Thalassa a subgenus; KORSCHEFSKY (1931) reversed

that opinion followed by DUVERGER (1989).

It is structurally most similar to Thalassa and Diazonema, but

differs by the tibia long relative to femoral length, and a Diomus

type of postcoxal abdominal line.

272 R.D. GORDON & C. CANEPARI

KEY TO! SPECIES OF MENOSCELIS

È Length 9.0 mm; elytron light reddish brown with 5 yellow

spots Ghia, 35950 19 ERTS e./, Ps, We 2. insignis Mulsant

Length less then 6.0 mm; elytron yellow with 3 brown

SOO CS Oe ton nera 1 saginata Mulsant

1. Menoscelis saginata Mulsant, 1850

Menoscelis saginata Mulsant, 1850: 508; CroTcH 1874: 209.

Menoscelis (Thalassa) saginata: CHAPUIS 1876: 228.

Thalassa (Menoscelis) saginata: KORSCHEFSKY 1931: 208; BLACKWELDER 1945: 449.

Thalassa (Monoscelis (sic!)) saginata: ORIVEL et al. 2004: 97 (biology).

Type locality. Cayenne (French Guiana) (MNHL, lectotype

here designated).

Description. dé, length 5.8 mm, width 5.2 mm; body

form round, convex. Dorsal surface with head slightly alutaceous,

shiny; pronotum and elytron shiny. Color yellow except pronotum

with H-shaped, median dark brown macula, macula with obliquely

oval, yellow spot on each side of middle; elytron with 3 reddish

brown spots, transversely rounded, 1 discal spot on sutural margin

anterior to middle paired with spot on opposite elytron, forming

large, transverse discal macula, 1 small spot on humeral callus, 1

small spot posterior to humeral callus near lateral margin (Figs.

34, 35); pro-, meso-, and apex of metasternum black. Head punc-

tures coarse, separated by less than a diameter. Pronotal punctures

smaller than on head, separated by less than to twice a diameter;

elytral punctures slightly larger than pronotal punctures, separat-

ed by 1 to 2 times a diameter. Metasternal punctures larger than

on elytron, sparse medially, separated by 1 to 2 times a diameter,

becoming dense laterally, separated by a diameter or less. Punc-

tures on median area of basal abdominal sternum larger than on

elytra, sparse, punctures fine, dense on sterna 2-6; area on 1“ ab-

dominal sternum inside postcoxal arc slightly alutaceous, weakly

shiny, finely, sparsely punctate. Clypeal apex wide, deeply emar-

ginate medially. Prosternum narrow, carinae narrowly separated at

apex, convergent, extended 1/2 distance to basal margin, basally

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 273

stemmed, stem long, reaching basal margin of sternum. Postcoxal

line on 1% abdominal sternum joining posterior margin at lateral

1/3. Fifth sternum broadly, weakly emarginate apically; 6° ster-

num truncate. Genitalia with basal lobe 1/2 length of paramere,

wide, sinuate, apex with one angle rounded, opposite angle acute,

large, angulate projection on one side; paramere wide, slightly nar-

rowed to rounded apex, inner margin sinuate (Fig. 40); sipho long,

robust, evenly curved, basal capsule with inner arm short, truncate

apically, outer arm shorter than inner arm, base of sipho sinuate

(Fig. 41).

2. Not known.

Variation. None observed.

Geographical distribution: French Guiana and Peru.

Specimens examined: 3. French Guiana: “Cayenne” (the lecto-

type). Peru: Satipo (MNHL) (USNM).

Remarks. As indicated in the key to species, M. saginata

and M. insignis are radically different in size and dorsal coloration.

CroTcH (1874) synonymized the two names stating that “M. sagi-

nata appears to me a pale variety of imsignis, the quadrate common

spot is identical in both, and all details of structure and punctua-

tion.” From that statement it is clear that Crotch had not seen a

specimen of M. insignis nor read the original description. MUL-

SANT’s (1850) original description clearly defines the differences be-

tween M. insignis and M. saginata. KORSCHEFSKY (1931) listed M.

insignis as an aberration of M. saginata and BLACKWELDER (1945)

followed the same classification. This seemingly strange situation

is perhaps explained by CROTCH’s (1874) use of the name insignis

for a species of Hyperaspis bearing a remarkable resemblance to

M. saginata. KORSCHEFSKY (1931) inadvertently confused these two

taxa in his catalogue of the Coccinellidae, a situation perpetuated

by subsequent authors.

Only two specimens have been examined: the holotype from

French Guiana and a specimen from Peru. In spite of the widely

different geographic locations, the two specimens are identical in all

respects.

The lectotype of M. saginata is here designated to stabilize fu-

ture usage of this name. It is labeled “Cayenne, Lacordaire.”

274 R.D. GORDON & C. CANEPARI

2. Menoscelis insignis Mulsant, 1850

Menoscelis insignis Mulsant, 1850: 509; CroTcH 1874: 209.

Menoscelis (Thalassa) saginata: CHAPUIS 1876: 228.

Thalassa (Menoscelis) saginata ab. insignis: KORSCHEFSKY 1931: 208; BLACKWELDER

1945: 449.

Type locality: Cayenne (French Guiana) (UMZC).

Description. &, length 7.5 mm, width 6.5 mm; body

form round, convex. Dorsal surface with head alutaceous, dull; pro-

notum slightly alutaceous, shiny; elytron shiny. Color yellow except

pronotum with median 1/3 reddish brown, reddish brown area with

3 yellow spots, 1 small, triangular, anteromedian spot, and 2 larger,

oval basal spots; elytron reddish brown with 5 large yellow spots

(Fig. 33), scutellar spot triangular, humeral spot c-shaped around

humeral callus, discal spot obliquely transverse, posterolateral spot

oval, and apical spot irregularly rounded. Head punctures fine, sep-

arated by less than a diameter. Pronotal punctures equal in size to

head punctures, separated by a diameter or less; elytral punctures

equal in size to head punctures, separated by a diameter or less.

Metasternal punctures very coarse, much larger than on elytron,

absent medially, dense laterally, separated by less than a diameter.

Punctures on median area of basal abdominal sternum larger than

on elytron, sparse, punctures fine, dense on sterna 2-6; area on 1%

abdominal sternum inside postcoxal arc slightly alutaceous, weakly

shiny, finely, sparsely punctate. Clypeal apex wide, weakly emar-

ginate medially. Prosternum narrow, carinae narrowly separated in

basal 2/4, widened apically between coxae. Postcoxal line on 1* ab-

dominal sternum joining posterior margin at lateral 1/3. Fifth ster-

num truncate apically; 6" sternum apically arcuate.

3 Not known.

Variation. None observed.

Geographical distribution: French Guiana.

Specimens examined: 1. The holotype specimen.

Remarks. See remarks under M. saginata. The 9 type

specimen examined is considered a holotype because it is clear from

the original description that Mulsant had only one type specimen.

The holotype is damaged and has been repaired at least once in its

history; therefore, genitalia were not dissected because of possible

further damage.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 275

Genus Thalassa Mulsant, 1850

Thalassa Mulsant, 1850: 511; CroTcH 1874: 209; KorscHEFSKy 1931: 208;

BLACKWELDER 1945: 449; EL-ALI 1972: 32; MILLEo et al. 2004: 395.

Type species: Thalassa pentaspilota Chevrolat, 1835, by subse-

quent designation of CROTCH 1874.

Description. Hyperaspidini with form rounded, convex.

Dorsal surface glabrous, alutaceous, dull. Elytral ground color usu-

ally dark blue, or dark blue with greenish lustre. Sd head mostly

yellow, 2 head black or reddish yellow with black maculation.

Elytron maculate or not. Antenna with 11 articles, basal article

wider than long (Fig. 43); antennal insertion concealed. Frons on

a slightly higher plane than clypeus. Apical maxillary article slight-

ly emarginate apically; mandibulary retinaculum strongly curved,

apex rounded. Clypeus long extended beyond frons, abruptly

joined to frons, lateral angle abrupt near apex, emarginate api-

cally (Fig. 42). Labrum short, narrow, weakly sclerotized, yellow-

ish brown. Scutellum large, wider than long. Elytral epipleuron

wide, not medially grooved, descending externally, excavated for

reception of middle and hind femoral apices. Prosternal process

flat, with 2 short, parallel carinae. Protibia wide, flattened, tibial

groove strongly flanged, outer margin of flange sinuate, flange as

wide at base as apex (Fig. 44); meso- and metatibia narrow, me-

dially emarginate, not flanged (Figs. 45, 46). Posterior margin of

metasternum abruptly descending between coxa and lateral mar-

gin. Metendosternite with deep, V-shaped anterior border. ‘larsal

claw with basal tooth. Postcoxal line on basal abdominal sternum

incomplete, of Scymnus (Scymnus) type. 6 apical abdominal ster-

num weakly modified. Apex of 2 abdominal sternum 6 rounded.

2 genital plate transverse.

Remarks. Thalassa was revised by MILLEO et al. (2004).

Therefore, we provide only a modified key to species and an anno-

tated species list. See MILLEO et al. (2004) for detailed descriptions

and illustrations.

Thalassa is most similar morphologically to Diazonema but is

distinguished by the 11-articled antenna, frons only slightly raised

above clypeus, and long protibia.

276 R.D. GORDON & C. CANEPARI

KEY TO SPECIES OF THALASSA

F EEtrontaiaetdiiateS9 ret. erie IPO eel inaelpbi/i pa 2

Eytrondamaezdatehi sui Heroes VEL: Ue. Blasio 4

20). Bivironm wath siagle yellow, apical’ spot... 5... i

le a <a telat ARIE Zc’ Set ER 1 Li cali 1. pentaspilota (Chevrolat)

Pivecon wit 'Pior'2 yellow. diséal ‘spots 2 Ove Ie LE 3

3(2). Elytron with 2 yellow, discal spots.5. Rorschefskyi Milléo et al.

Elytron with single, quadrangular spot on disc..... 110)

TI ADDS UID, AIA tel Os 28, 6. montezumae Mulsant

4(1). Pronotum with anterior and lateral margin narrowly dark

Velo RUSORI, Pest, Gio Mr: Limo Verse 3. similaris Mulsant

5(4). Pronotum with lateral margin very narrowly yellow... ..

MILITO, od PE STINE B.S ara tees Di, 2. flaviceps Mulsant

Pronotum with lateral 1/4 yellow......... 4. glauca Mulsant

1. Thalassa pentaspilota (Chevrolat, 1835)

Chilocorus pentaspilotus Chevrolat, 1835: 124.

Thalassa pentastigma: MULSANT 1850: 512.

Thalassa pentaspilota: CROTCH 1874: 209; GORDON 1987: 26; MiLLéo et al. 2004:

359%;

Geographic distribution: Mexico and Venezuela. BLACKWELDER

(1945) also lists Guatemala and Cuba; Guatemala is probably cor-

rect, Cuba probably incorrect.

South American specimens examined: 9. Venezuela: Aragua,

Boca del Rio; Carabobo, Marciara; Carabobo, Yuma (USNM).

Remarks. This is an easily recognized species. because

of the single, apical yellow spot on each elytron. The distribution

(Mexico and Venezuela) is very disjunctive, but an accidental in-

troduction of live specimens from Mexico to Venezuela may be an

explanation. Lectotype designated by GORDON (1987).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI DIL

2. Thalassa flaviceps Mulsant, 1850

Thalassa flaviceps Mulsant, 1850: 513; MiLLÉéo et al. 2004: 397.

Thalassa prasina Mulsant, 1850: 516; CroTcH 1874: 209 (as synonym of T.

flaviceps).

Geographic distribution: Cuba.

South American specimens examined: 1. Cuba, Havana (CAS).

3. Thalassa similaris Mulsant, 1850

Thalassa similaris Mulsant, 1850: 514; GoRDON 1987: 26; MILLEO et al. 2004: 398.

Geographic distribution: “New Grenada” (Colombia or Venezu-

ela).

South American specimens examined: 2. The lectotype (GORDON

1987) labeled “n. gr.,” meaning Colombia or Venezuela (UMZC);

Guyana, Georgetown (BM).

Remarks. This species is easily distinguished from the

similar JT. glauca by the narrowly yellow pronotal borders.

4. Thalassa glauca (Mulsant, 1850)

Menoscelis glauca Mulsant, 1850: 510.

Thalassa glauca: CROTCH 1874: 510; MILLEO et al. 2004: 399.

Geographic distribution: Guatemala and Costa Rica south to

Colombia and Brazil.

South American specimens examined: 4. Brazil: Sào Paulo, Can-

tereira; Jaù.; Colombia: no locality (BM) (UMZC) (USNM).

Remarks. This is a widely distributed species whose distri-

bution appears disjunct. The disjunction is probably not real, but an

artifact of the scarcity of specimens in collections.

5. Thalassa korschefskyi (Milléo, Almeida & Gordon, 2004)

Hyperaspis korschefskyi Milléo, Almeida & Gordon, 2004: 399.

278 R.D. GORDON & C. CANEPARI

South American distribution: Colombia, no locality (holotype).

Remarks. This species is known only from the holotype

specimen (USNM).

6. Thalassa montezumae Mulsant, 1850

Thalassa montezumae Mulsant, 1850: 512; Gorpon 1985: 400; MILLEO et al. 2004:

398.

Geographic distribution: United States, Mexico, and Guate-

mala. =

South American specimens examined: none.

Clypeaspis n. gen.

Type species: Hyperaspis trilineata Mulsant (by monotypy and

present designation).

Description. Hyperaspidini with form elongate, paral-

lel sided, dorsum glabrous. é head mostly yellow, @ head black

or brown. Elytron maculate. Antenna with 10 articles, basal article

wider than long; antennal insertion concealed. Apical maxillary ar-

ticle slightly emarginate apically; mandibulary retinaculum strongly

curved, apex rounded. Clypeus and frons joined at abrupt angle,

clypeal apex emarginate; frons raised slightly above plane of clypeus,

partially concealing antennal base (Fig. 1). Labrum short, narrow,

weakly sclerotized, yellowish brown. Scutellum large, wider than

long. Elytral epipleuron narrow, medially grooved, excavated for re-

ception of middle and hind femoral apices. Prosternal process slight-

ly raised, with 2 carinae (Fig. 3). Protibia wide, straight, not sinuate

or emarginate, tarsal groove flanged on posterior margin (Fig. 2);

meso- and metatibia wide, straight, slightly flanged on posterior mar-

gin. Posterior margin of metasternum abruptly descending between

coxa and lateral margin. Metendosternite with anterior tendons of

fork curved toward middle. 'Tarsal claw with basal tooth. Postcoxal

line on basal abdominal sternum incomplete, of Scymnus (Scymnus)

type. Male apical abdominal sternum weakly modified. Apex of fe-

male abdominal sternum 6 rounded. Female genital plate transverse.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 279

Etymology. The generic name is a combination of the first part

of “clypeus” and the ending of Hyperaspis, referring to the modified

clypeus and relationship to Hyperaspis.

Remarks. Clypeaspis belongs to the group that includes

Diazonema, Menoscelis, and Thalassa because of the wide basal an-

tennal segment, antennal insertion concealed beneath clypeus, and

frons on a higher plane than clypeus. It is distinguished from these

genera by the protibial flange narrow at the base, widened toward

apex, tibia without basal angulation, and narrow epipleuron.

1. Clypeaspis trilineata (Mulsant, 1850), n. comb.

Hyperaspis trilineata Mulsant, 1850: 667; CrotcH 1874: 227; KORSCHEFSKY 1931:

198; GORDON 1987: 29.

Type locality. Cayenne (French Guiana).

Description. d, length 3.4 mm, width 2.2 mm; body

form elongate, parallel sided, flattened. Dorsal surface alutaceous,

dull. Color reddish yellow except vertex of head black; pronotum

with large, square, black basomedian spot in median 1/3; elytron with

black vitta on sutural margin, vitta wide at base, narrowed to apical

1/8, wide, irregular black vitta extended from basal margin across

humeral angle onto apical declivity, vitta slightly widened from base

toward apex (Figs. 47, 48); venter of head, pro-, meso-, metasterna

black; abdomen black except outer 1/3 and sterna 5-6 reddish yel-

low. Head punctures fine, distinct, separated by less than to twice

a diameter; pronotal punctures larger than on head, separated by a

diameter or less; elytral punctures larger than pronotal punctures,

separated by 1 to 2 times a diameter. Metasternal punctures slightly

larger than on elytron, nearly absent medially, larger, separated by

a diameter near lateral margin. Punctures on abdominal sterna 1-4

coarse, sparse medially, becoming dense, fine laterally; sterna 5-6

finely, densely punctured throughout; area on 1° abdominal sternum

inside postcoxal arc shiny, finely, sparsely punctured. Clypeal apex

deeply emarginate. Epipleuron oblique, grooved internally, slightly

descending externally, femoral excavations deep. Prosternum with

intercoxal carinae narrowly separated, parallel toward base, extend-

ed to basal 5/8, not stemmed, not reaching basal margin. Protibia

280 R.D. GORDON & C. CANEPARI

flanged, flange narrowest at base, becoming wider toward apex (Fig.

2). Postcoxal line on 1* abdominal sternum extended to posterior

sternal margin, flattened along margin, apex recurved 1/2 distance

to basal sternal margin. Fifth sternum with apex truncate; 6™ ster-

num truncate. Genitalia with basal lobe 1/2 length of paramere,

obliquely rounded apically, with large, angulate projection on one

side; paramere wide, tapered to rounded apex in apical 1/2 (Fig.

49); sipho short, robust, apically straight, with membranous area,

basal capsule with inner arm long, slender, outer arm wider, slightly

longer than inner arm, base emarginate (Figs. 50, 51).

2. Similar to male except head dark brown. Genitalia with beak

of basal unit long, wide; connecting duct short (Fig. 52).

Variation. Length 2.8 to 3.7 mm, width 2.0 to 2.7 mm. Dorsal

and ventral black areas often dark brown.

Geographical distribution: South America and Caribbean is-

lands.

Specimens examined: 34. Argentina: Chaco, Dep. Resistencia;

Salta, Oran. Brazil: Para, Belém. British West Indies: Barbados. Ve-

nezuela: Aragua, El Limon; Caracas Valley, Los Ruices; Rio Yara-

cuy Valley, Chivacoa; Suapure, Caura R (MBR) (USNM).

Remarks. The distribution pattern of this species, based

on specimens examined, is somewhat odd. Nearly all specimens ex-

amined were collected on Barbados, with only a few from South

America and none from Trinidad.

Type material could not be located. MULSANT (1850) stated

that his specimen(s) were from the Buquet collection. Most material

from that collection is in the BM, but specimens of C. trilineata

could not be found.

Prognataspis n. gen.

Type species: Prognataspis surreptiva n. sp.

Description. Hyperaspidini with form nearly round,

dorsum glabrous. d head yellow, 2 head unknown. Elytron macu-

late. Antenna with 9 articles, basal article wider than long (miss-

ing from illustration) (Fig. 54); antennal insertion not concealed.

Eye large, nearly as wide as median frontal area. Apical maxillary

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 281

article not emarginate apically. Frons narrow. Clypeus and frons

joined at abrupt angle, clypeus narrow, clypeal apex emarginate;

frons on same plane as clypeus, not concealing antennal base (Fig.

53). Labrum short, narrow, heavily sclerotized, brown. Scutellum

large, wider than long. Elytral epipleuron broad, medially grooved,

excavated for reception of middle and hind femoral apices. Pros-

ternal process slightly raised, with 2 carinae (Fig. 55). Protibia

wide, straight, not sinuate or emarginate, tarsal groove flanged on

posterior margin (Fig. 57); meso- and metatibia narrow, straight,

not flanged on posterior margin. Posterior margin of metasternum

gradually descending between coxa and lateral margin. Abdominal

sterna 3-5 with pit near lateral margin on each side. Tarsal claw

with basal tooth. Postcoxal line on basal abdominal sternum incom-

plete, of Scymnus (Scymnus) type (Fig. 56). Male apical abdominal

sternum 5 weakly modified.

Etymology. The generic name is a combination of the Latin

prognatus, and the ending of Hyperaspis, referring to the unusually

distinctive protruding mouthparts, and relationship to Hyperaspis.

Remarks. Prognataspis is characterized by the large eyes

nearly as wide as median frontal area (usually the frontal area is

two or three times as wide as eye); 9-articled antenna; very nar-

row clypeus and labrum; flanged protibia; and abdominal sterna 3-5

with pit on each side near lateral margin.

1. Prognataspis surreptiva n. sp.

Type material. Holotype d: Ecuador, 80 km NW Quito, 1300

m, PACTO, 26.2-2.3.2002, leg. Marek Halada (CCM).

Description. 4, length 2.6 mm, width 2.4 mm; body near-

ly round, very convex. Dorsal surface shiny. Color black except head

yellow; pronotum with lateral 1/3 and apical 1/4 yellow, elytron with

single large, yellow spot medially in apical 1/2, spot tapered at apical

and posterior ends (Figs. 61, 62); antenna, labial palpi, propleuron

yellow; labrum dark brown; legs yellowish red except femora darker;

epipleuron, pro-, meso-, and metasterna black; abdomen dark reddish

brown. Head punctures fine, separated by less than diameter; pronotal

punctures slightly larger than on head, separated by less than to twice

a diameter; elytral punctures larger than pronotal punctures, sepa-

282 R.D. GORDON & C. CANEPARI

rated by less than to twice a diameter. Metasternal punctures much

larger than on elytron, nearly absent medially, becoming dense, con-

tiguous near lateral margin. Punctures on basal abdominal sternum

much than larger than on elytra, coarse, dense medially, becoming

smaller, less dense laterally; sterna 2-6 with dense punctures becom-

ing progressively smaller and denser from sternum 2 through sternum

6; area on 1*t abdominal sternum inside postcoxal arc shiny, coarsely,

sparsely punctured. Clypeus narrow, apex deeply emarginate, clypeus

and frons joined at abrupt angle. Eye nearly as wide as median frontal

area in frontal view. Epipleuron oblique, internally grooved, descend-

ing externally, femoral excavations deep. Prosternum with intercoxal

carinae widely separated at apex, convergent from apex toward base,

basally stemmed. Protibia wide, flanged. Postcoxal line on 1" abdomi-

nal sternum reaching posterior sternal margin, rounded along mar-

gin, apex recurved 1/3 distance to basal sternal margin. Fifth sternum

with apex broadly, weakly emarginate; 6" sternum deeply emargin-

ate. Genitalia with basal lobe long, slightly shorter than paramere,

apex irregularly rounded, with truncated projection on one margin;

paramere slender, abruptly narrowed at basal 1/2, tapered from basal

1/3 to rounded apex (Fig. 58); sipho slender, long, apex sinuate, basal

capsule with inner arm irregular, apically widened, slightly knobbed,

outer arm shorter than inner, truncate apically (Figs. 59, 60).

2 Not known.

Variation. None observed.

Etymology. The species name is from the Latin surreptivus,

meaning false or fraudulent, referring to the deceptive resemblance

to many Hyperaspis species.

Remarks. Generic characters readily identify this taxon,

which otherwise looks very much like a Hyperaspis species.

Peruaspis n. gen.

Type species: Peruaspis paprzyckii n. sp.

Description. Hyperaspidini with form elongate, rounded,

dorsum glabrous. Elytron maculate. Antenna with 10 articles, basal

article wider than long (Fig. 65). Clypeus short; frons and clypeus

smoothly, obliquely joined, clypeal apex weakly emarginate; frons

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 283

on same plane as clypeus, partially concealing antennal base (Fig.

6). Labrum short, wide, strongly sclerotized (Figs. 4-6). Scutel-

lum large, as wide as long. Elytral epipleuron narrow, flat, slightly

grooved medially, excavated for reception of middle and hind femo-

ral apices. Prosternal process raised, prominent, sides ridged (Figs.

5, 6). Protibia narrow, straight, slightly flanged on posterior margin

(Fig. 70); meso- and metatibia narrow, straight, slightly flanged on

posterior margin. Posterior margin of metasternum abruptly de-

scending between coxa and lateral margin. Tarsal claw with basal

tooth. Postcoxal line on basal abdominal sternum incomplete, of

Scymnus (Scymnus) type (Fig. 66). Apex of 2 abdominal sternum 6

rounded. 9 genital plate transverse.

Etymology. The generic name is a combination of “Peru” and

the ending of Hyperaspis, referring to the country of origin of both

known species.

Remarks. ‘The labial and prosternal structures of this genus

differ widely from those of other hyperaspine genera. The labrum

is heavily sclerotized, wide, and extended beyond the clypeal apex.

The prosternal process and middle of prosternum are prominently

raised, particularly so in P. paprzycku. Size difference and dorsal

coloration between the 2 included species is extreme, but both share

the diagnostic labial and prosternal characters.

Sd are unknown for either species. Both are Peruvian species,

P. paprzycku from a high altitude Andean locality, and P. hypocrita

from low altitude on the eastern slope of the Andes.

KEY TO SPECIES OF PERUASPIS

Length less than 2.5 mm; color black, elytron with 2 yel-

low spots (Ties. 62, 640.051 eee e 1. paprzycku, n. sp.

Length more than 3.0 mm; color mostly yellow with black

maculae, (ies; 06, 69). 5. Ren 2. hypocrita, n. sp.

1. Peruaspis paprzyckii n. sp.

Type material. Holotype 9: Peru, Satipo, XI, 1942, Paprzycki

(USNM). Paratype: same data as holotype (USNM).

284 R.D. GORDON & C. CANEPARI

Description. &, length 2.1 mm, width 1.7 mm; body oval,

convex. Dorsal surface shiny except head dull, alutaceous; pronotum

and elytra shiny. Color black except head irregularly mottled with

yellow across vertex and frons, clypeus brown; pronotum with lateral

1/3 yellow, inner margin of yellow area arcuate; elytron with 2 yel-

low spots, irregularly rounded spot at middle of elytron, and irregu-

larly transverse subapical spot (Figs. 63, 64); antenna and labial palpi

brownish yellow; narrow outer border of propleuron yellow; proleg

yellowish brown, ventral surface dark brown. Head punctures dis-

tinct, separated by 1 to 2 times a diameter. Pronotal punctures larger

than on head, separated by less than to twice a diameter; elytral punc-

tures larger than pronotal punctures, separated by a diameter or less.

Metasternal punctures as large as on elytron, separated by more than

a diameter medially, dense, nearly contiguous laterally. Punctures on

abdominal sterna 1-4 coarse, sparse medially, becoming dense, very

fine laterally; sterna 5-6 finely densely punctured throughout; area

on 1* abdominal sternum inside postcoxal arc shiny, impunctate.

Clypeus short, apex slightly emarginate; labrum protruding ante-

rior to clypeal apex, labrum heavily sclerotized, dark brown, shiny,

apex slightly emarginate. Epipleuron flat, slightly grooved medially,

slightly descending externally, femoral excavations deep. Antenna

with 10 articles. Prosternum with intercoxal process raised, medially

concave, with central carina. Protibia not flanged. Postcoxal line on

1° abdominal sternum extended to posterior sternal margin, rounded

along margin, apex recurved 1/5 distance to basal sternal margin.

Fifth sternum with apex broadly, distinctly emarginate. Genitalia

with beak of basal unit large, long; connecting duct short (Fig. 67).

3. Not known.

Variation. Head of paratype entirely black.

Etymology. The species is named for the collector of the type

series.

Remarks. Peruaspis paprzyckiu is a small, 2-spotted species

similar to Hyperaspis bisignata and H. camargoi in color pattern.

2. Peruaspis hypocrita n. sp.

Type material. Holotype 2: Peru, Madre de Dios, Tambopata

Wildlife Res., 30 km SW Pto. (Puerto) Maldanado (sic !), 12 50’S,

69 20°W; 290 m, 6 XII 1982, Joseph J. Anderson (CMP).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 285

Description. ?, length 3.3 mm, width 2.7 mm; body

rounded, slightly oval, convex. Dorsal surface with head slightly

alutaceous, shiny: pronotum shiny; elytra shiny. Color yellow ex-

cept pronotum with large, black, obtriangular spot on each side of

middle in basal 1/2; elytron narrowly bordered with black, with

3 black spots, 1 spot connected to suture anterior to middle, spot

in conjunction with spot on opposite elytron forming large, cor-

date spot, 1 large, irregularly rounded spot on humeral callus, |

large, irregularly rounded spot medially on apical declivity (Figs.

68, 69); venter of head, pro-, meso- and metasterna, femoral bases

black; abdominal sterna 1-4 black except lateral 1/3 and sterna 5-6

yellow. Head punctures coarse, separated by less than a diameter;

pronotal punctures larger than on head, separated by a diameter

or less; elytral punctures larger than pronotal punctures, separated

by 1 to 2 times a diameter. Metasternal punctures larger than on

elytron, separated by a diameter medially, becoming dense, con-

tiguous laterally. Punctures on abdominal sterna larger than on

elytra, dense medially, becoming contiguous laterally, less coarse

on sterna 5-6; area on 1* abdominal sternum inside postcoxal arc

shiny, nearly impunctate. Clypeus short, apex slightly emarginate;

labrum protruding anterior to clypeal apex, labrum heavily sclero-

tized, black, shiny, apex slightly emarginate. Epipleuron oblique,

slightly grooved internally, weakly descending externally, femoral

depressions deep. Clypeal apex deeply emarginate. Protibia narrow,

not flanged, with tarsal groove deep, basal end of groove projecting.

Prosternum raised, with intercoxal carinae narrowly separated, par-

allel in apical 2/3, joined apically, stemmed, reaching basal margin.

Postcoxal line on 1*t abdominal sternum reaching posterior sternal

margin, slightly rounded along margin, apex angled 1/4 distance to

basal sternal margin. Fifth sternum with apex feebly emarginate.

Genitalia with beak of basal unit long; connecting duct long (Fig.

71) (genital plates not shown).

3 Not known.

Etymology. The specific name is the Latin hypocrita, meaning

mime, player, referring to the resemblance to Hyperaspis onerata.

Remarks. Peruaspis hypocrita is a large species whose dor-

sal color pattern is similar to that of H. onerata.

286 R.D. GORDON & C. CANEPARI

Genus Hyperaspis Chevrolat in Dejean, 1833

Hyperaspis Chevrolat in Dejean, 1833: 435; MuLsantT 1850: 649; CroTcH 1874: 224;

KORSCHEFSKY 1931: 177

Type species: Coccinella reppensis Herbst, by subsequent designation of CROTCH

1874.

Gordom Duverger, 2001: 226. (n. syn.)

Aliana Duverger, 2001: 226. (n. syn.)

Description. Hyperaspidini with form elongate, oval, or

rounded, dorsoventrally flattened or convex; dorsum glabrous. Head

usually yellow in male, at least partially brown or black in female.

Elytron with pale maculation on dark background or dark macula-

tion on pale background, rarely immaculate. Antenna with 10 or 11

articles, (Figs. 92; 103,/136,146,, 170, 179; 194, 2177, 2234228; 238,

251, 257, 269, 287, 314, 321, 334, 341, 344, 349, 359, 363, 369,

375, 381, 397, 404, 413, 424, 441, 453, 467, 476, 483), basal article

longer than wide; antennal insertion exposed. Apical maxillary ar-

ticle slightly emarginate apically; mandibulary retinaculum strongly

curved, apex rounded. Clypeus usually long, abruptly or obliquely,

smoothly joined to frons laterally (Figs. 7, 11), distinctly emarginate

apically, often concealing all or most of labrum. Labrum short, nar-

row (Figs. 11, 12), weakly sclerotized, yellow or yellowish brown.

Scutellum large, wider than long. Elytral epipleuron narrow, usually

medially grooved, excavated for reception of middle and hind femo-

ral apices. Prosternal process usually slightly convex, with 2 carinae

present (carinae completely absent in H. dissidens), carinae of vary-

ing lengths (Figs. 8, 10, 12, 77, 405) (Fig. 77, H. reppensis, type spe-

cies). Protibia usually slender, rarely slightly flanged (Figs. 74, 121,

354). Posterior margin of metasternum abruptly descending between

coxa and lateral margin. Metendosternite with anterior tendons of

fork curved toward middle or not (Figs. 93, 104, 171, 195, 274,

364, 414, 425, 442, 484). Tarsal claw with or without basal tooth.

Postcoxal line on first abdominal sternum incomplete, of Scymnus

(Scymnus) type (Figs. 78, 122) (Fig. 78, H. reppensis, type species).

Sd apical abdominal sternum weakly modified. Apex of 2 abdominal

sternum 6 rounded. 2 genitalia with coxal plate basically transverse

(Fig. 97), rather than long, narrowly triangular as in Tenuisvalvae

(Fig. 494).

Remarks. HAyperaspis is a Western Hemisphere genus with

some Old World representatives. It is particularly prolific in North

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 287

America with 103 species and subspecies present (GORDON 1985).

For South America we record 68 species of Hyperaspis, a number

dwarfed by the large brachiacanthine genus Cyra.

Asymmetry in male genitalia is expressed by the basal lobe,

which is usually apically obliquely truncate or rounded and, in ven-

tral view, the right margin of the basal lobe is sinuate with a lateral

projection. However, in H. matronata, the asymmetry is reversed;

the left margin of the basal lobe is sinuate.

GORDON (1985) divided North American Hyperaspis into two

Sections based on number of antennal articles, and also recorded

a few ancillary characters for each Section that are not useful in

the South American fauna. We divide South American Hyperaspis

into two Sections defined by clypeal and frontal structures. Section

I has the clypeus and frons joined at an abrupt angle (primitive

character state), and contains five groups characterized by number

of antennal articles and male genitalia. Section II has the clypeus

and frons smoothly, obliquely joined (derived character state), and

contains two groups characterized by male genital characters. ‘These

groupings are useful as identification aids, but also correspond with

the assumption that 10-articled antenna is a primitive character state

and 11-articled antenna a derived character state. However, H. con-

nectens, which is clearly a member of Section II, has a 10-articled

antenna. This divergence indicates that number of antennal articles

is not a useful character for distinguishing genera. Within Section

I, d genitalia of the H. onerata group are considered primitive and

those of the H. howdeni group are considered derived. In Section II

members of the H. conclusa group have genitalia with long, slender,

relatively unmodified basal lobes, which we consider a derived char-

acter state. Other derived character states of the conclusa group are

dorsoventrally flattened body, moderately deep to shallow epipleural

depressions, and elongate body.

Hyperaspis and Tenuisvalvae are distinguishable only in the

female sex. Therefore, it is possible that some Hyperaspis species

known only from males may belong in Tenuisvalvae.

Examination of the lectotype of Cleothera micilla Mulsant, 1853

(designated by GORDON 1987) revealed that it is a species of Diomus

Mulsant (Scymninae, Diomini), and a junior synonym of Diomus

pallidipennis (Mulsant, 1850).

288 R.D. GORDON & C. CANEPARI

Taxa of the brachiacanthine genus Cyra are numerous, and spe-

cies are often confused with Hyperaspis species. DUVERGER (2001)

correctly designated Cyra loricata Mulsant, 1850 the type species of

Cyra. Mulsant originally described this species as Cleothera (Cyra)

loricata. MULSANT (1850) described the genus Cleothera for a single

species, C. buqueti Mulsant, 1850, which is actually a species of the

previously described genus Hinda Mulsant, 1850, as recognized by

WEISE (1895: 127). Thus, Cleothera is a junior synonym of Hinda,

and Cyra becomes the correct generic name for those species de-

scribed as Cleothera (Cyra) by MULSANT in 1850 and in subsequent

publications.

DUVERGER (2001) described 3 American genera of Hyperas-

pidini as follows. Clavatus is a monotypic Mexican and Central

American genus based on Hyperaspis acanthicola Chapin, 1966, a

species associated with ants, which we consider valid. Gordoni is a

monotypic North American genus based on Hyperaspis significans

Casey, 1908, which we here place as a junior synonym of Hy-

peraspis because the only distinguishing character is the pubescent

head. Aliana is a monotypic genus based on Hyperaspis connectens

(Thunberg, 1808), that we also consider a junior synonym of Hy-

peraspis because the only distinguishing character is the sparsely

pubescent clypeus. Both of the latter genera also have a 10-articled

antenna, a character not considered of value at the generic level in

Hyperaspis because it occurs in obviously dissimilar groups within

the genus.

KEN iO SPECTEStOR 2Y PERASPIS

da Clypeus and frons joined at abrupt angle (Figs. 7, 8) .. 2

Clypeus and frons smoothly, obliquely joined (Figs. 11, 12) 57

2(1). Species with pale elytral maculation (usually reddish or

yellow spots) on dark background (Figs. 72, 79) ...... 3

Species with dark elytral maculation (usually brown or

black spots) on pale background (Figs. 83, 213)....... 40

3(2). Species without yellow lateral border on elytron ...... 8

Species with yellow lateral border on elytron......... 4

4(3).

5(4).

6(5).

7(5).

8(3).

9(8).

10(9).

11(10).

12(11).

13(12).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 289

Species without pale elytral spot, with yellow lateral border

on elettori (Pigs 100). 00 le, eS: 5. scutifera Mulsant

Species with 1 or more discal spots on elytron, with yellow

lateral border.) 5.4.4). nogo»sfo1g.lensgi BEER so rage Ss. 5

Species with 1 pale, discal spot on elytron and yellow late-

ral:borders!» Uusuanase. mbe pveelie why ji eee he 7

Species with 2 or more pale, discal spots and yellow lateral

BOrcder RARI peek eee en RITO a 6

Elytron with 2 pale discal spots (Fig. 134)...........

Pe er ee ee aa] re pe I ce 10. limbigera Mulsant

Elytron with 6 yellow spots (Fig. 366) 53. histrionica (Mulsant)

d pronotum with lateral border of median black spot pro-

jected onto yellow lateral margin (Fig. 204); Brazil....

La or ia ra dl ee ee e ea Dati 21. stladesma, n. sp.

d pronotum with lateral border of median black spot

straight (Fig. 100); Colombia, Curacao, Venezuela. ....

MaI REN E e cet aera a ae cI 5. scutifera Mulsant

EAS it O90 ELAN: < one taid ak cae i ce 19. brethesi, n. sp.

Elytron not vittate, with variously shaped spots....... 9

Elytronwith1 pale elvtral macula. .- ot... oe 10

Elytron with more than 1 pale, elytral macula....:... 20

Elytron with 1 yellow, humeral spot (Fig. 292) 37. atra, n. sp.

Elytron with pale macula not on humerus ............. 11

Elytron with-1' yellow, apical. soot. (Fics 273) ove; 12

Elytron with yellow spot or pale macula not apical .... 15

Length more than 3.0 mm (Fig. 347) ... 49. abertha, n. sp.

eneth less than -2,5 mundo, Abs Bi Sipe, ..... 13

Prosternal carinae nearly parallel, not reaching basal

prosternal marraige, Abr eae 33. mimica, n.Sp.

Prosternal carinae tapered from apex to base, reaching

Dasat prattetalmaneio 00 lee ete 44 14

290

14(13).

15(11).

LOS)

17(16).

18(17).

19(17).

20(9).

21(20).

DEVA ST

R*D' GORDONTE NG *CANEPARI

6 genitalia with basal lobe short, wide, with lateral an-

ouletiomGlinene264) Lh... vrs Oat 31. apicaspis, n. sp.

6 genitalia with basal lobe long, tapered from base to

apex, witnout.lateral projection (Fig. 27/O)iaed lstosel. .

TN he a a Men ds, 32. delicata Almeida & Vitorino

Riyttioniithi tewvellows,sciscalspote.,..L Asi: salma sete 16

Elytron with more than 1 yellow spot or pale macula.. 20

Length less than 2.0 mm; elytral spot irregularly rounded

(Hao:, $32), (PL). 000) toga iena sieo 45. guilloryi (Mulsant)

Length more than 2.0 mm; elytral spot irregularly

no uao ott aoe. 1800) doge iia, ded loupe hi hy

Elytron with spot large, narrow apically and widened

posenerUs cet) Gimiem letotel Wolsey. GTO bata. ... 18

Elytron with spot small, oval or irregularly rounded... 19

Elytron with anterior margin of spot reaching anteri-

or border of elytron, outer margin of spot emarginate

arouma ninmeral. cals, (Fis. 256)........ 36. chocot, n. sp.

Elytron with anterior margin of spot widely separated

from anterior border of elytron, inner margin of spot

widely emarginate on disc (Fig. 339). ... 47. istmina, n. sp.

Elytron with spot regularly oval, reddish yellow (Fig.

UE ei ln n 7. chapini, n. sp.

Elytron with spot irregularly rounded, yellow (Fig. 283);

Colonie ras "SEZ: KI. POL BIL eS 45. howdeni, n. sp.

Pili Irani 6 e ei I, erat INZIALE "ki ZI

Ritorni more than i2 pale spots tt, er UNIT... 28

Prosternal carinae completely absent (Fig. 9); discal spot

imansversely smuate (Pig. 221) 3 laed 25. dissidens, n. sp.

Prosternal carinae present; discal spot not transversely

ARS ie den init ta derrrateora è» 27

Elytron with 1 scutellar spot and 1 discal spot (Fig.

ZOU SN) Po PP) Dore, sua, 38. recordata Mulsant

23(22).

24(23).

25(24).

26(25).

27(26).

28(20).

29(28).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 291

Elytron with 1 discal and 1 apicalbspot sister rt 44 DS

Elytron with apical spot transverse, yellow; median,

black pronotal area strongly emarginated with yellow

laterally (Mies 15900 spore Cig. LOU). 13. biguttata Sicard

Elytron with apical spot rounded or slightly transverse,

or if strongly transverse, then nearly white in color; me-

dian black pronotal area not emarginated with yellow la-

feral ly, di: secreta ti pane eee! 24

Length more than 2.6 mm; anterior tibia slightly flan-

ged; antenna with 10 articles; Bolivia ....1. latitibia, n. sp.

Length less than 2.5 mm; anterior tibia not flanged; an-

tenna with UIL articles: Argentina and Brazil a a 25

Elytron with transverse, nearly white, apical spot; discal

spot small, about the diameter of an eye (Fig. 255)....

e SS cree Seer One Boe ne eee nee 30. albopunctata Crotch

Elytron with rounded, yellow apical spot; discal spot

muck larger 'thah-an eye "diameter =e, SA ees ew ee 26

Elytron with discal spot large, median; discal and apical

spots narrowly separated (Fig. 144).... 12. camargoi, n. sp.

Elytron with discal spot small anterior to middle; discal

and apical spots widely separated. +04 om) oo ee 27

Elytral surface smooth, polished; elytral punctures di-

stinctly larger than pronotal punctures (Fig. 200)......

(Fri 350 lata lione devia 20. rosariensis, n. sp.

Elytral surface weakly alutaceous, slightly shiny; elytral

punctures slightly larger than pronotal punctures (Fig. 1

SO) DN mt 1 11. bisignata, n. sp.

Elytron with 3° vellow spatelor maculae. 2.0 ere Po 29

Elytron withemore than 3 spots or maculae... ns KA 35

Pronotum with an oblique, yellow, basomedian spot on

each side of middle; apical elytral spot c-shaped (Fig. 3

$2) ee BAR Serie Hote vais 48. satipoensis, n. sp.

Pronotum without oblique basal spots; apical elytral

Spot Cos Aape ae Or Mat: E A ee ee Oe er ears he 30

292

30(29).

S130).

32(31).

34(33).

35(28).

HOC 29):

S736).

R.D. GORDON & C. CANEPARI

Elytron with yellow scutellar spot (Fig. 336) .........

Bp APEOEL LEE. EST, der cord eek soda 46. pseudopavida, n. sp.

EP lyirom wittounscutellar Spot! le, sun So ADI: Apes, I SÙ

Elytral spots small, the diameter of an eye or smaller

CRISSUDPO,IITE Roe 10 UOT FORE AGIO £ 8. cracentis, n. Sp.

Elytral spotssarcer then an eve.diametet Lt. th weit i DZ

Pronotum mostly black; median spots on elytron nar-

rowly connected (Fig. 244). ...... 28. bicruciata (Mulsant)

Pronotum mostly yellow; median spots on elytron usu-

olbsnesieoanesicd Sole ne . fiir. o:. Mer deol tien.) |. 33

Pronotum yellow with pale brown maculae, or with

large, yellow lateral area deeply emarginating median

brown spot; antenna witht Martieles 906 Pine 100%... 34

Pronotum yellow with large, rectangular, median black

area (Fig. 79); antenna with 10 articles .. 2. donzeli Mulsant

All elytral spots large, narrowly separated or contigu-

ous (Fig. 176); pronotum always yellow with pale brown

maculae; apical spot on elytron large, c-shaped, occupy-

ing most of apical declivity;.......... 17. orthivora, n. sp.

All elytral spots small, widely separated, not contiguous;

pronotum occasionally yellow with pale brown maculae,

usually with large, lateral yellow area deeply emarginat-

ing median brown spot (Fig. 167); apical spot on elytron

slightly c-shaped or not, occupying less than 1/2 of api-

call'dechivatzivo dete mn matt seoanl 16. pseudodonzeli, n. sp.

Pivivon want. vellow Spots oases ae Sean tb ee a 36

Elytron with-more than 4 yellow spots ..........%... 38

Elytron without scutellar spot (Fig. 249). 29. joannae, n. sp.

P mcroniwin seme lat. Spot. nea), spet_oeg ads BOCAS LA I

Dorsal ground color black; pronotal and elytral punc-

tures coarse; apical yellow spot on elytron transverse

e le ae ree. | 14. cleida Mulsant

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 293

38(35).

39(38).

40(2).

41(40).

42(41).

43(41).

44(43).

45(44).

Dorsal ground color dark brown; pronotal and elytral

punctures fine; apical yellow spot on elytron c-shaped

GEio 340) eA Re eee 41. simlaensis, n. sp.

Elytron with 6 yellow spots (Fig. 160) . 15. tayronensis, n. sp.

Elytron'sati S gellowtsp ts RUE R PNIEE seer) 39

Length less than 2.0 mm; median lateral spot on elytron

regularly oval (Fig. 279); antenna with 11 articles .....

eT Ar ee eee 34. laterimacula, n. sp.

Length more than 2.1 mm; median lateral spot triangu-

lar or projecting inward (Fig. 110); antenna with 10 ar-

ticles. of ws parities ratttonota-sis bersi 6. imitatrix, n. sp.

Elytron yellow with indistinct, obscure, brownish mark-

ings (Fig. 91); antenna with 10 articles .. 4. helveola, n. sp.

Elytron not as described above; antenna with 10 or 11 ar-

tiles, Liswiue .s1oce. it ect. arom siti. amonrati . . 41

Elytron with concentric arcs of varying colors ........ 42

Elytron: with’ distinct spots remain 43

Elytral background yellow, punctures pale brown, dark

brown curved vitta present medially (Fig. 184) .......

VETTA LL A RI AO AVERE, REA 18. zonula, n. sp.

Elytron with narrow, lateral, yellow band, median, oval

reddish yellow area, and broad, irregular sutural vitta

(Pig. 332). oe st ORA 50. circumclusa, n. sp.

Elytron with 1 black or dark brown spot ............ 44

Elytron with more than 1 black or dark brown spot ... 47

Elytron with black spot extended from base at scutel-

lum posteriorly to apical declivity, spot narrowed medi-

any DES A CSR Mr SETE 9. operaria (Mulsant)

Elytron with spot not as described above............ 45

Elytra with butterfly shaped median spot (Fig. 308) ...

SORE ee re elo AE 40. mariposa, n. sp.

Elytraiwithy spot! net butterfly shaped: waduneadl aetna .- 46

DIA:

R.D. GORDON & C. CANEPARI

46(45)):

47(43).

48(47).

49(48).

50(49).

51(48).

S2(51),

Elytra with large black spot surrounded by narrow, yel-

lompibandaChtels22¢)yoq2 veallev.-leorgeks 26. uninotata, n. sp.

Elytra with spot brown, c-shaped ... 43. c-nigrum Mulsant

Elytron with median, brown, discal spot, spot some-

times united with that on opposite elytron forming

single spot, and sometimes with small, obscure, lateral

spots; many brown elytral punctures presenting “freck-

led appearance” (Fig. 189); male pronotum often with 2

larve, basomedian yellow spots’ (Fre. 192). ci iui

Meh a ae RET ME pn. FR ERI Se eee ae 19. brethest, n. sp.

Elytron with varying number of spots; never with

“freckled appearance; male pronotum without large, ba-

Soiniedianespatsor.rertituanta altere risomeliun . coxph 48

Elytron with 3 dark spots, sutural margin with distinct

DET Sal O POLTI Berger Patt elles AE calle A RA a aaa me TI 49

Elytron with more than 3 dark spots, sutural margin

Wici‘ortiitfo ud oriibofder, i artteeo iu. MO NTELL pill SA

Elytron with small, discrete, brown spot anterior to

middle (Fig. 213 ); ground color of elytron mostly red-

disbimiellonialast. uan polis baro: 23. campbelli, n. sp.

Elytron with large, black, discal spot contiguous with

MMe TO OOD OSK Gucmaul Olls dg desarztal. sor .costgionua: 50

Elytra with cordate, sutural spot in anterior 1/2, spot

narrowed posteriorly, 2 large lateral spots present on

each elytron (Fig. 83); antenna with 10 articles .......

SUMORLY Uns gee Je EAM. ate 3. matronata Mulsant

Elytra with round, sutural spot in middle, 2 small later-

al spots present on each elytron (Fig. 232); antenna with

GBP (SS Ie cis aa aerate a partie 27. onerata (Mulsant)

Elytron with 4 dark spots, sutural margin narrowly or

broadly bordered with black or brown .............. 2

Elytron with 5 dark spots, sutural margin bordered or

Hor wats bla ek threw. Oe wa eA YY A bai 55

Elytron with elongate, black, humeral spot extended

from base past humeral callus, 1 small, elongate spot on

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 295

53(52).

54(53).

55(51).

56(55).

57(1).

58(57).

59(57).

60(59).

sutural margin anterior to middle, and 2 small, lateral

spots on apical declivity (Fig. 319) ...... 42. lindae, n. sp.

Elytrom notasnckesenibed falvose: ae ius rotta: DÒ

Elytron with u-shaped spot in anterior 1/2 curved from

base around humeral callus and back to base (Fig. 329)

rire be) luo, Bia: BO UU 44. sagittata Crotch

Elytron without u-shaped humeral spot ............. 54

Elytron with 3 spots in row from humerus nearly to

suture: at) ie 24. colombiensis, n. sp.

Elytron with 2 spots in anterior 1/2, and 2 spots in pos-

tenon gly? 08) cine a. 22. octonotata, n. sp.

Pronotum black medially with wide, yellow lateral mar-

SR AY Se (shee) dA 51. aemulata, n. sp.

Pronotum yellow with 4 or 5 brown maculae......... 56

Pronotum with 4 brown maculae; elytron with brown

spots separated by narrow, yellow lines (Fig. 304).....

POPES SRI FO a PEER I 39. eupaleoides Crotch

Pronotum with 5 brown maculae; elytron with 4 brown,

discrete, widely separated brown spots (Fig. 361)......

A). fon. willey, Uap ne?.,.38anols ,Sinnte 71) 52. dispar, n. sp.

Elytron pale wath dark sporta ia 58

Elyiron:dark avith pale spetsser maculae... .....+s..++: 59

Elytra with large, obverse, triangular spot on suture,

and one small humeral spot (Fig. 373) ..55. herrarai, n. sp.

Elytron with suture and 3 spots dark brown to black

(io: ASR a EO ALO be ke ee kee A 65. arida, n. sp.

Flvtron,vittate cdl ie ie lek ae ina ata 60

Elytron:'not aattatesy railed. LOO a etna aule OR ae, 61

Elytron with median yellow vitta narrow, not enlarged

basally (Fig. abd... aL) DISSI 66. elegantissima Brèthes

Elytron with median yellow vitta broad, enlarged ba-

sally (hig.d49) viel MMI Pet 64. conclusa Weise

296

61(59).

62(61).

63(62).

64(63).

65(63).

66(65).

67(66).

68(67).

R.D. GORDON & C. CANEPARI

Elytron with single reddish yellow spot on lateral mar-

ATMA 220) AR LE) virvasal 61. funesta (Germain)

Elytron without spots, or with 1 or more pale spots, if 1

spot, then -elytronywith yellow lateral border... us 62

Elytron with broad, yellow border, without spots (Fig.

oc) LA A le Pe TE IO TRA, 54. ayacucho, n. sp.

Elytron with or without yellow border, always with 1 or

MOE sPOEsm le SCMilben tia: cartd Lens Meise secre iit of 63

Body extremely elongate, flattened; apical margin of

DaSalbesbe lain tin. Shara tes. dene Weber eh ce me fly mee nea. . 64

Body not extremely elongate, flattened; apical margin of

bosalssterngimmainot Sinmaatee al l(athore bold rane: 4: 65

Pronotum with basal margin irregularly yellow; elytron

with c-shaped apical spot (Fig. 481) .... 68. prolata, n. sp.

Pronotum with basal margin black; elytron with apical

spotfiriangulare(iew472)7. eno. 04 bein 67. longula Weise

Spegiesimet Kino, Wom CMe le iero coi sla sn 66

Spec sso cene CMe ss OI 69

Elytron with single, elongate, reddish yellow spot (Fig.

FF renee ciation ot. Way ai 60. ingrata Mulsant

Elytron with more than 1 spot; French Guiana and else

Pronotum mostly black, lateral margin yellow, and pos-

terior margin with outer 1/3 narrowly yellow (Fig. 386);

elytral pattern highly variable (Figs. 386-396); widely

distributed, commonly collected species 57. festiva Mulsant

Pronotum variable, but posterior margin not narrowly

We low, 2s. Cescri Ded. above. io cMaIre tenor -« 68

Elytron with 5 yellow spots, discal and lateral spots

usually narrowly connected (Fig. 379) 56. vredenburgi, n. sp.

Elytron with reddish yellow discal and apical spots nar-

rowly connected (Fig. 410)...... 59. connectens (Thunberg)

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 297

69(65). Elytron with yellow lateral border extended across base

toyseutelhann(Fign402)a.deviytomesiva 58. germaini Crotch

Elytron with or without yellow lateral border, if border

present, then not extended across base of elytron...... 70

70(69). Elytron with small, yellow, basal spot between humeral

callus. anduscutellumeG(F 19, 430) mor AY hora 62. nana Mader

Elytron without basal spot, color pattern variable but

usually with discal and lateral spots connected (Figs.

ASTRO eee ee 63. sphaeridiodes Mulsant

Section I

Species with clypeus and frons joined at abrupt angle; protibia

slender or slightly flanged.

donzeli group

Hyperaspis species having 10-articled antennae; large body size;

clypeus and frons joined at abrupt angle; epipleura obliquely de-

scending externally; protibia apically wide, slightly flanged; and

male genitalia similar to onerata group.

1. Hyperaspis latitibia n. sp.

Type material. Holotype d: Bolivia, Santa Cruz, 4-6 k SSE Bue-

na Vista, F&F Hotel, 23-26 Oct 2000, Wappes & Morris (USNM).

Description... 3, length +3,.6.mm, width,.2.8 mm; body

oval, slightly flattened. Dorsal surface with head alutaceous, weakly

shiny: pronotum shiny; elytra shiny. Color black except head yellow

with black vertex; pronotum yellow with large, rectangular, basome-

dian spot, spot widely separated from anterior margin, lateral mar-

gin of spot deeply emarginated, elytron with 2 large yellow spots,

1 irregularly rounded, discal spot anterior to middle, 1 transversely

oval, subapical spot (Figs. 72, 73); antenna, mouthparts, propleuron

yellow; legs yellow except basal 2/3 of femora black; abdomen black

with lateral 1/4 reddish yellow. Head punctures fine, separated by

298 R.D. GORDON & C. CANEPARI

less than a diameter; pronotal punctures larger than on head, sepa-

rated by a diameter or less; elytral punctures larger than pronotal

punctures, separated by a diameter or less. Metasternal punctures

much larger than on elytron, separated by a diameter medially, dense,

contiguous laterally. Punctures on abdominal sterna 1-4 as large as

on elytra, dense medially, becoming fine, dense laterally, sterna 5-6

finely densely punctured throughout; area on 1% abdominal sternum

inside postcoxal arc shiny, finely, sparsely punctured. Clypeal apex

emarginate, clypeus and frons joined at abrupt angle. Epipleuron

oblique, internally grooved, descending externally, femoral depres-

sions moderate. Antenna with 10 articles. Protibia distinctly flanged

(Fig. 74). Prosternum with intercoxal carinae narrowly separated at

apex, convergent from apex toward base, basally stemmed, reach-

ing basal margin (Fig. 77). Postcoxal line on 1** abdominal sternum

reaching posterior sternal margin, flattened along margin, apex re-

curved 1/4 distance to basal sternal margin. Fifth sternum with apex

broadly, distinctly emarginate; 6 sternum truncate. Genitalia with

basal lobe 1/2 as long as paramere, apex narrowly rounded, with

large, angulate projection on one margin; paramere wide, tapered

from base to rounded apex; (Fig. 75); sipho robust, with apical

membranous area, basal capsule with inner arm short, wide, outer

arm wide, longer than inner arm, apically knobbed, basal border

slightly emarginate (Fig. 76).

2 Not known.

Etymology. The species name is from the Latin /atus, meaning

broad, and tibia, referring to the flanged protibia.

Remarks. This species has the same elytral color pattern

as several species of Hyperaspis, but H. latitibia is larger and the

protibia is flanged.

2. Hyperaspis donzeli (Mulsant, 1850)

Cleothera donzeli Mulsant, 1850: 638.

Hyperaspis donzelt: CROTCH 1874: 222; KORSCHEFSKY 1931: 187; BLACKWELDER 1945:

447; GORDON 1987: 28.

Type locality. Brazil. MNHP (lectotype here designated).

Description. d, length 3.0 mm, width 2.6 mm; body

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 299

form round, slightly flattened. Dorsal surface with head feebly

shiny, alutaceous; pronotum and elytron shiny. Color black except

pronotum yellow with large, black, irregularly rectangular basome-

dian spot, anterior margin of spot widely separated from anterior

pronotal margin; elytron with 3 yellow spots, 1 round spot medially

on disc, 1 triangular spot on lateral margin laterad of discal spot,

1 transversely oval, subapical spot (Fig. 79); antenna, mouthparts,

propleuron, legs yellow; abdomen black except outer 1/4 and ster-

num 5-6 reddish yellow. Head punctures fine, distinct, separated

by less than to twice a diameter. Pronotal punctures larger than on

head, separated by a diameter or less; elytral punctures larger than

pronotal punctures, separated by a diameter or less. Metasternal

punctures larger than on elytron, separated by a diameter medial-

ly, contiguous laterally. Punctures on abdominal sterna 1-4 coarse,

dense medially, becoming dense, very fine laterally; sterna 5-6 fine-

ly, densely punctured throughout; area on 1° abdominal sternum

inside postcoxal arc shiny, coarsely, sparsely punctured. Clypeal

apex deeply emarginate, clypeus and frons joined at abrupt angle.

Epipleuron oblique, internally grooved, strongly descending exter-

nally, femoral excavations deep. Antenna with 10 articles. Protibia

distinctly flanged. Prosternum with intercoxal carinae tapered to-

ward base, basally stemmed, reaching basal margin. Protibia dis-

tinctly flanged. Postcoxal line on 1% abdominal sternum extended to

posterior sternal margin, flattened along margin, apex recurved 1/4

distance to basal sternal margin. Fifth sternum with apex broadly,

distinctly emarginate; 6° sternum slightly rounded. Genitalia with

basal lobe 3/4 length of paramere, strongly, obliquely truncate api-

cally, abruptly sinuate on one side; paramere wide, nearly parallel

sided, apex rounded (Fig. 80); sipho robust, curved upward, with

membranous area, basal capsule with inner arm short, wide, outer

arm long, apically knobbed, base sinuate (Figs. 81, 82).

2 Not known.

Geographical distribution: Brazil, Paraguay.

Specimens.,, examined;...1... Paraguay: » Itapua,;. Sa... Trinidad

(USNM).

Remarks. AHyperaspis donzeli has the same elytral color pat-

tern as H. pseudodonzeli known from Trinidad and northern South

America. However, Hyperaspis donzeli differs by the 10-articled

300 R.D. GORDON & C. CANEPARI

antenna, subapical elytral spots evenly, transversely oval, and the

flanged protibia.

A lectotype is here designated to stabilize future usage of this

name.

3. Hyperaspis matronata (Mulsant, 1853)

Cleothera matronata Mulsant, 1853: 71.

Hyperaspis matronata: CROTCH 1874: 226; KoRSCHEFSKY 1931: 192; BLACKWELDER

1945: 447; GORDON 1987: 28.

Cleothera cordifera Weise, 1895: 128. (n. syn.)

Hyperaspis cordifera: KORSCHEFSKY 1931: 186; BLACKWELDER 1945: 446.

Type locality: of matronata, Brazil, UMZC (lectotype designat-

ed by GORDON 1987); of cordifera, Paraguay, ZMHB (lectotype here

designated).

Description. 6, length 2.6 mm, width 2.1 mm; body

form oval, flattened. Dorsal surface with head feebly shiny, aluta-

ceous: pronotum feebly shiny, alutaceous: elytron shiny. Color yel-

low except pronotum with large, black, basomedian spot, anterior

margin of spot widely separated from anterior pronotal margin, lat-

eral margin with earlike projections; elytron with 3 black spots, 1

obliquely oval spot connected to black sutural border on disc, spot

together with matching spot on opposite elytron forming a cordate

median macula, 1 large, black, triangular spot laterally posterior to

humerus, 1 large, more or less triangular spot medially on apical de-

clivity (Fig. 83); pro-, meso-, metasterna black; abdomen black except

outer 1/4 and sternum 5-6 reddish yellow. Head punctures fine, dis-

tinct, separated by less than to twice a diameter; pronotal punctures

larger than on head, separated by a diameter or less; elytral punc-

tures larger than pronotal punctures, separated by less than to twice

diameter. Metasternal punctures larger than on elytron, very sparse

medially, nearly contiguous laterally. Punctures on abdominal sterna

1-4 coarse, dense medially, becoming dense, fine laterally; sterna 5-6

finely, densely punctured throughout; area on 1*t abdominal sternum

inside postcoxal arc shiny, coarsely, sparsely punctured. Clypeal apex

deeply emarginate, clypeus and frons joined at abrupt angle. Epi-

pleuron oblique, feebly grooved internally, weakly descending exter-

nally, femoral excavations deep. Antenna with 10 articles. Protibia

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 301

distinctly flanged. Antenna with 11 articles (Fig. 84). Prosternum

with intercoxal carinae narrowly separated at apex, tapered toward

base, basally stemmed, reaching basal margin. Protibia not flanged.

Postcoxal line on 1% abdominal sternum extended to posterior ster-

nal margin, flattened along margin, apex recurved 1/2 distance to

basal sternal margin. Fifth sternum with apex broadly, feebly emar-

ginate; 6" sternum truncate. Genitalia with basal lobe 3/4 length of

paramere, strongly, obliquely truncate apically, abruptly sinuate on

one side (asymmetry reversed); paramere wide, tapered to rounded

apex in apical 3/4 (Fig. 85); sipho slender, apically sinuate, with mem-

branous area, basal capsule with inner arm wide, outer arm slightly

longer than inner, apically knobbed, base sinuate (Figs. 86, 87).

° Similar to d except head with vertex and triangular area on

frons black; pronotum entirely black except lateral 1/6 yellow. Geni-

talia with beak of basal unit large, hastate; connecting duct short

(Figs. 88-90).

Variation. Length 2.4 to 3.0 mm, width 1.9 to 2.5 mm. Frons

of @ varies in color from frontal area black to light brown. Size of

elytral spots varies slightly.

Geographical distribution: Southeastern South America

Specimens examined: 28. Argentina: Chaco, Charata; Corrien-

tes, Santo Tomé; Department Resistencia, Chaco; Cordoba; Misio-

nes; Salta, Guemes. Bolivia: Tiguipa; “Ost Bolivien.” Brazil: Gua-

nabara, Rio de Janeiro; Nova Teutonia. Paraguay: San Bernardino;

Amanbay; San Pedro; Cororo-Rio Yapane; Sapucay (CDA) (MBR)

(USNM) (ZMHB).

Remarks. Hyperaspis matronata has a distinct color pattern

by which it can be recognized. d genitalia differ from those of other

species because of reversed symmetry in the basal lobe.

Cleothera cordifera in no way differs from H. matronata; there-

fore it is a junior synonym. The lectotype us C. cordifera is here

designated to stabilize future usage of this name. It is labeled “Par-

aguay Drake (green paper)/Hyperaspis matronalis Muls cordifera

Ws (Weise’s writing in purple ink)/Syntypus, Cleothera cordifera

Weise, 1895, labelled by MNHB 2004 (red paper).” Two additional

specimens in the series labeled “Amer Drake (green paper),” and

“Amer.m (green paper)” are considered type material and are desig-

nated as paralectotypes.

302 R.D. GORDON & C. CANEPARI

4. Hyperaspis helveola n. sp.

Type material. Holotype d: Brazil, S. Paulo (Sao Paulo),

Cantareira, 4.XII.1958, J. Halik, 11308, Brasil, Halik Collection

1966 (USNM). Allotype. Brazil, Minas Gerais, V. Monte Verde,

14.XII.1961, J. Halik, 17761, Brasil, Halik Collection, 1966 (USNM).

Paratypes. 4. 1, Brazil, Sdo Paulo, Cantereira, 15.VIII.1963, J. Ha-

lik, 22689, Brasil, Halik Collection, 1966 (USNM); 1, Brazil S.

Paulo, horto flor. (IT G), 30.1.1959, Brasil, Halik Collectin, 1966

(USNM):; “1, Brazil, “5° Paulo; "Est. S. Paulo, 24.12, Saude 1722,

J. Melzer Brasilien, det. by F. W. Nunenmacher, H. dilatata Cr.,

Korschefsky Collection 1952 (USNM); 1, Brazil, Cipo Sao Paulo,

29 Nov 1965, V. N. Alin (BM).

Description. 6, length 2.8 mm, width 2.5 mm; body

form round, convex. Dorsal surface with head dull, strongly aluta-

ceous, pronotum less dull than head, shiny, elytra shiny. Color yel-

low except pronotum with 4 median brown spots, anterior 2 spots

triangular, basal 2 spots slightly triangular, narrowly separated an-

terior to scutellum; elytron with 2 vaguely defined brown macula, 1

short macula near suture anterior to middle, 1 long, comma- shaped

macula laterally from humeral angle to apical 1/4 (Fig. 91); antenna,

mouthparts, legs, and most of abdomen reddish yellow; pro-, meso-,

metasterna and median area of basal abdominal sternum dark brown.

Head punctures very fine, hidden in dense alutaceous sculpture. Pro-

notal punctures larger than on head, separated by less than to twice

a diameter; elytral punctures larger than pronotal punctures, sepa-

rated by 1 to 3 times a diameter. Metasternal punctures larger than

on elytron, sparse medially, nearly contiguous laterally. Punctures on

abdominal sterna 1-4 coarse, sparse medially, becoming dense, very

fine laterally; sterna 5-6 finely densely punctured throughout; area

on 1* abdominal sternum inside postcoxal arc feebly shiny, aluta-

ceous, coarsely, sparsely punctured. Clypeal apex weakly emarginate,

clypeus and clypeus and frons abruptly joined. Epipleuron oblique,

internally grooved, strongly descending externally, femoral excava-

tions deep. Antenna with 10 articles (Fig. 92). Prosternum with

intercoxal carinae convergent toward base, basally stemmed, reach-

ing basal margin. Protibia slightly flanged. Postcoxal line on 1* ab-

dominal sternum extended to posterior sternal margin, curved along

margin, apex recurved 1/2 distance to basal sternal margin. Fifth

sternum with apex broadly, distinctly emarginate; 6" sternum dis-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 303

tinctly emarginate. Genitalia with basal lobe 3/4 length of paramere,

wide, with abruptly angulate projection on one side; paramere short,

very wide, narrowed to rounded apex in apical 1/4 (Fig. 94); si-

pho robust, apex sinuate, with membranous area, basal capsule

with inner arm long, slender, outer arm short, wide (Figs. 95, 96).

2 Similar to male except vertex and upper 1/2 of frons dark

brown. Genitalia with beak of basal unit short, robust, very wide

(Figs. 97-99).

Variation. Length 2.7 to 3.2 mm, width 1.4 to 2.7 mm Elytron

with vague brown areas varying from entirely absent to expanded,

occupying most of elytron.

Etymology. The specific name is from the Latin helvus, meaning

yellow, helveolus (pale yellow), and refers to the dorsal coloration.

Remarks. ‘This is a large, pale species with elusively vague

elytral maculation. No other species of South American Hyperaspis

has the same combination of dorsal color and body shape.

chapini group

Hyperaspis species having 10-articled antenna; small body size;

clypeus and frons abruptly joined or joined at slight angle (H.

scutifera); epipleura flat, not descending externally; protibia narrow,

not flanged; and male genitalia of varying types.

Two species in this group, H. imitatrix and H. chapini, are

closely related, sharing the same type of male genitalia having an

extremely long sipho.

5. Hyperaspis scutifera (Mulsant, 1850)

Cleothera scutifera Mulsant, 1850: 565.

Hyperaspis scutifera: CROTCH 1874: 216; KorSCHEFSKY 1931: 195: BLACKWELDER

1945: 448.

Hyperapis incompleta Crotch, 1874: 225; KorsScHEFSKy 1931: 225; BLACKWELDER

1945: 445; GORDON 1987: 28. (n. syn.)

Type locality: of scutifera, Venezuela, Valencia, DEZ (lectotype

here designated); of incompleta, Colombia, UMZC (holotype).

304 R.D. GORDON & C. CANEPARI

Description. 6, length 2.4 mm, width 1.7 mm; body

oval, convex. Dorsal surface with head weakly shiny alutaceous;

pronotum and elytra shiny, slightly alutaceous. Color yellow, pro-

notum black with lateral 1/5 yellow, inner margin of yellow area

straight; elytron with irregular, black, sutural spot, spot abruptly

widened posterior to scutellum and again posterior to middle, a

small poorly defined spot on humeral callus (Figs. 100, 101); an-

tenna, mouthparts, propleuron, epipleuron and legs yellow; abdo-

men with lateral 1/8 and sterna 5-6 yellowish red. Head punc-

tures distinct, separated by less than to twice a diameter. Pronotal

punctures larger than on head, separated a diameter or less; elytral

punctures as large as on pronotum, separated by less than a di-

ameter. Metasternal punctures larger than on elytron, sparse me-

dially, dense, nearly contiguous laterally. Punctures on abdomi-

nal sterna 1-4 coarse, sparse medially, becoming dense, very fine

laterally; sterna 5-6 finely densely punctured throughout; area on

1* abominal sternum inside postcoxal arc alutaceous, coarsely,

sparsely punctured. Clypeal apex deeply emarginate, clypeus and

frons joined at slight angle. Epipleuron flat, slightly grooved medi-

ally, not descending externally, femoral excavations moderate, not

deep. Antenna with 10 articles (Fig. 103). Prosternum with inter-

coxal carinae convergent from apex toward base, basally stemmed,

reaching basal margin. Protibia narrow, not flanged. Postcoxal line

on 1* abdominal sternum extended to posterior sternal margin,

rounded along margin, apex recurved 1/2 distance to basal sternal

margin. Fifth sternum with apex broadly, weakly emarginate; 6°

sternum truncate. Genitalia with basal lobe as long as paramere,

with small projection on one margin; paramere slender, tapered

from basal 1/4 to rounded apex (Fig. 105); sipho slender, inner

arm long, outer arm shorter than inner arm, basal border widely,

shallowly emarginate (Figs. 106-107).

2 Similar to male except head black; anterolateral pronotal angle

narrowly yellow. Genitalia with beak of basal unit large, elongate,

connecting duct short (Fig. 108).

Variation. Length 2.4 to 2.6 mm, width 1.6 to 1.7 mm. Dorsal

color may be black except head yellow; elytron bordered with yel-

low except sutural margin, yellow border extended from laterad of

scutellum around elytron nearly to suture at apex, inner margin of

border sinuate, basal portion of lateral yellow border may be re-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 305

duced or absent (Fig. 102), and there is a tendency for the yellow

border to separate from an apical yellow spot. The lectotype female

has the yellow elytral border uniform, not sinuate, except for a dis-

tinct posterior extension on each side of scutellum.

Geographical distribution: Colombia, Venezuela, Curacao.

Specimens examined: 15. Colombia: Meta, Puerto Gaitàn, Cali-

magua; Valle Palmeira. Curacao: Schottgatwee; Zapateer. Venezue-

la: Aragua, Cagua; Chacas, nr. Caracas; Damacar; Tocayo; Valencia

(BM) (DEI) (USNM).

Remarks. The color pattern(s) of this species is not un-

common in South American Hyperaspis species, but unique among

those with 10-articled antennae. The dorsal color pattern is highly

variable (Figs. 100-102), but genital characters are mostly stable.

Specimens from Curacao might not belong to this species. How-

ever, the genitalic differences are not definitively distinct, and we

consider them, at least provisionally, to be MH. scutifera.

The 2 genitalia of type specimens of H. scutifera and H. in-

completa were examined and appear identical. Therefore, we con-

sider them synonyms. A & syntype of H. scutifera is here designated

to stabilize future usage of this name. It is in the DEI collection,

labeled “Venezula Schaum (handwritten)/Coll. Haag/Syntypus (pink

paper)/scutifera mls. typ.”

6. Hyperaspis imitatrix n. sp.

Type material. Holotype d: Brazil, Chapada, Acc. No. 2966,

Dec. (CMNH).

Paratype: Bolivia, Warscewski (sic !) (ZMHB).

Description. 6, length 2.3 mm, width 1.7 mm; body

oval, convex. Dorsal surface with head weakly shiny, alutaceous:

pronotum and elytra shiny, slightly alutaceous. Color black except

head yellow; pronotum with large, basomedian black spot, anterior

border of spot arcuate; elytron with 5 yellow spots, scutellar spot

at base slightly rectangular, small oval spot at middle anterior to

apical declivity, elongate humeral spot on lateral margin from base

nearly to midpoint of elytron, triangular spot on lateral margin

posterior to midpoint, and transversely oval subapical spot (Figs.

306 R.D. GORDON & C. CANEPARI

109, 110); antenna, mouthparts, propleuron, epipleuron and legs

yellow; abdomen brown with lateral 1/8 and sterna 5-6 yellow.

Head punctures distinct, separated by a diameter or less. Prono-

tal punctures slightly larger than on head, separated a diameter or

less; elytral punctures as large as pronotal punctures, separated by

less than to twice a diameter. Metasternal punctures larger than on

elytron, sparse medially, dense, nearly contiguous laterally. Punc-

tures on abdominal sterna 1-4 larger than on elytra, coarse, sparse

medially, becoming dense laterally; sterna 5-6 finely, densely punc-

tured throughout; area on 1% abdominal sternum inside postcoxal

arc alutaceous, coarsely, sparsely punctured. Clypeal apex deeply

emarginate; clypeus and frons joined at abrupt angle. Epipleuron

flat, slightly grooved medially, not descending externally, femo-

ral excavations deep. Antenna with 10 articles. Prosternum with

intercoxal carinae narrowly separated at apex, convergent from

apex toward base, not reaching basal margin, not basally stemmed.

Protibia narrow, not flanged. Postcoxal line on 1% abdominal ster-

num not reaching posterior sternal margin, rounded along margin,

apex recurved 1/2 distance to basal sternal margin. Fifth sternum

with apex broadly, weakly emarginate; 6" sternum distinctly emar-

ginate. Genitalia with basal lobe as long as paramere, with large,

rounded projection on one margin; paramere slender, tapered from

basal 1/4 to rounded apex; trabes apically curled (Fig. 111); sipho

slender, extremely long, about 5 times length of phallobase, apex

unmodified, basal capsule with inner arm long, outer arm nearly

pento hie. b2).

2 Not known.

Variation. Paratype with pronotum entirely yellow except for

narrow, black, basomedian border, border widened at each end;

elytron with subapical spot narrowly connected to median spot, ely-

tral maculation larger than in holotype.

Etymology. The specific name refers to the similarity in size,

shape, and color pattern to some variations of H. incompleta.

Remarks. ‘This species and H. scutifera are similar in ex-

ternal appearance, but H. imitatrix has male genitalia with an ex-

tremely long sipho, the longest yet observed in Hyperaspis. In addi-

tion, the apically curled trabes is shared only with H. chapini, which

has a single discal spot on each elytron.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 307

7. Hyperaspis cha pini n. sp.

Type material. Holotype 6: Trinidad, B.W.I. (British West In-

dies), 1903, G. E Bryant, G. Bryant Coll. 1919-147 (BM). Allotype

2: same data as holotype (BM). Paratypes, 2: same data as holotype

(BM).

D e's'‘etiptio nes Mena 2. anim davidii. hy tna body,

oval, convex. Dorsal surface with head slightly alutaceous, shiny;

pronotum slightly alutaceous, shiny; elytra shiny. Color black ex-

cept head yellow; pronotum yellow with large, basomedian black

spot extended 3/4 distance to anterior border, lateral margin of spot

emarginated; elytron dark brown with 1 reddish yellow, elongate

spot on disc slightly posterior to middle (Fig. 113); antenna, mouth-

parts, propleuron, legs yellow; epipleuron reddish brown; abdomen

brown with lateral 1/8 to 1/4 yellow. Head punctures distinct, sep-

arated by less than a diameter. Pronotal punctures slightly larger

than on head, separated by a diameter or less; elytral punctures as

large as pronotal punctures, separated by less than to twice a diam-

eter. Metasternal punctures larger than on elytron, separated by a

diameter medially, dense, nearly contiguous laterally. Punctures on

abdominal sterna 1-4 as large as on elytra, dense medially, becom-

ing fine laterally; sterna 5-6 finely, densely punctured throughout;

area on 1" abdominal sternum inside postcoxal arc shiny, coarsely,

densely punctured. Clypeal apex deeply emarginate, clypeus and

frons joined at abrupt angle. Epipleuron oblique, slightly grooved

internally, descending externally, femoral excavations deep. Antenna

with 10 articles. Prosternum with intercoxal carinae narrowly sepa-

rated at apex, convergent from apex toward base, basally stemmed,

reaching basal margin. Protibia narrow, not flanged. Postcoxal line

on 1* abdominal sternum not reaching posterior sternal margin,

rounded along margin, apex recurved 1/2 distance to basal ster-

nal margin. Fifth sternum with apex broadly, weakly emarginate;

6" sternum distinctly emarginate. Genitalia with basal lobe 3/4 as

long as paramere, deeply notched on one margin just before base;

paramere slender, tapered from basal 1/4 to rounded apex; trabes

apically curled (Fig. 114); sipho slender, extremely long, about 5

times length of phallobase, apex unmodified, basal capsule with in-

ner arm long, outer arm nearly absent (Figs. 115, 116).

2 Similar to male except head black; pronotum black with an-

terolateral angle and lateral border narrowly reddish yellow. Genita-

308 R.D. GORDON & C. CANEPARI

lia with beak of basal unit large, oval; connecting duct short (Figs.

117-118).

Variation. Length 2.1 to 2.4 mm, width 1.7 to 1.8 mm. Female

pronotal color varies from that described above to having lateral 1/5

bright yellow.

Etymology. This species is named for Edward A. Chapin, whose

systematic research on Coccinellidae has survived the test of time

extraordinarily well.

Remarks. ‘This species has a dorsal color pattern unique

among species having 10-articled antenna. Male genitalia share with

H. imitatrix an extremely long sipho and apically twisted trabes,

characters unknown among other Hyperaspis species.

onerata group

Hyperaspis species having 11-articled antenna; body slightly dor-

soventrally flattened or convex; clypeus and frons joined at abrupt

angle; protibia usually narrow, sometimes slightly widened apically;

male genitalia with basal lobe usually short, bent medially, obliquely

rounded or obliquely truncate apically, with a distinct lateral projec-

tion of varying size and shape, wide or oval (Figs. 123, 129, 140,

etc.). This group contains both convex and dorsoventrally flattened

species. The former are placed first on the assumption that a convex

body is a more primitive character state than a flattened body.

8. Hyperaspis cracentis n. sp.

Type material. Holotype 6: Brazil, Rio de Jan. (Rio de Janeiro),

Acc No. 2966 (CMP). Allotype 2: same data as holotype (CMP).

Paratypes, 3: same data as holotype except 2 paratypes with addi-

tional label “Oct.” (CMP) (USNM).

Description. dg, length 2.5 mm, width 1.8 mm; body

oval, somewhat elongate, convex. Dorsal surface with head aluta-

ceous, dull: pronotum slightly alutaceous, feebly shiny; elytra

shiny. Color black except head yellow; pronotum with lateral 1/5

and apical 1/4 yellow, elytron with 3 small, yellow spots, 1 spot

medially on disc irregularly round, 1 spot on medial lateral margin

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 309

medially oval, 1 transversely oval subapical spot (Figs.119, 120);

antenna, mouthparts, propleuron, epipleuron and legs yellow ex-

cept basal 1/4 of pro- and mesofemora dark brown, basal 1/2 of

metafemur dark brown; abdomen dark brown with lateral 1/8 and

sterna 5-6 brownish yellow. Head punctures fine, nearly invisible

in alutaceous sculpture, separated by less than to twice a diameter;

pronotal punctures larger than on head, separated by a diameter

or less; elytral punctures larger than pronotal punctures, separated

by a diameter or less. Metasternal punctures much larger than on

elytron, coarse, separated by a diameter medially, dense, contiguous

laterally. Punctures on abdominal sterna 1-4 larger than on elytra,

coarse, dense medially, becoming fine, dense laterally; sterna 5-6

finely densely punctured throughout; area on 1*t abdominal sternum

inside postcoxal arc shiny, coarsely, densely punctured. Clypeal

apex deeply emarginate, clypeus and frons joined at abrupt angle.

Epipleuron slightly oblique, internally grooved, weakly descending

externally, femoral excavations deep. Antenna with 11 articles. Pros-

ternum with intercoxal carinae narrowly separated at apex, conver-

gent from apex toward base, basally stemmed. Protibia narrow, not

flanged (Fig. 121). Postcoxal line on 1** abdominal sternum reach-

ing posterior sternal margin, flattened along margin, apex recurved

1/3 distance to basal sternal margin (Fig. 122). Fifth sternum with

apex broadly, weakly emarginate; 6 sternum distinctly emargin-

ate. Genitalia with basal lobe 1/2 as long as paramere, wide, apex

slightly rounded, with angulate projection on one margin; paramere

wide, tapered from apical 1/4 to rounded apex; (Fig. 123); sipho ro-

bust, short, apex sinuate with membranous area, basal capsule with

inner arm robust, outer arm shorter than inner, apically truncate

(Figs. 124, 125).

2 Similar to d except head black; pronotum black except lat-

eral 1/5 yellow. Genitalia with beak of basal unit short, curved; con-

mecting duct short (Fis. 126).

Variation. Length 2.4 to 2.7 mm, width 1.8 to 2.0 mm.

Etymology. The specific name is from the Latin cracens, mean-

ing neat, graceful, referring to the elegant and graceful dorsal ap-

pearance.

Remarks. The dorsal color pattern and very coarse ventral

punctation distinguish this species.

310 R.D. GORDON & C. CANEPARI

9. Hyperaspis operaria (Mulsant, 1853)

Cleothera operaria Mulsant, 1853: 70.

Hyperaspis operaria: CROTCH 1874: 224; KorsScHEFSKY 1931: 193; BLACKWELDER

1945: 447; GORDON 1987: 28.

Hyperaspis insignis Crotch, 1874: 226; KORSCHEFSKY 1931: 190; BLACKWELDER 1945:

447. (n. syn.)

Hyperaspis communalis Brèthes, 1925b: 4; KorscHEFsKy 1931: 186; BLACKWELDER

1945: 446. (n. syn.)

Type locality: of operaria (UMZC, lectotype designated by

GORDON 1987) and insignis (UMZC, holotype), Brazil; of communalis

(BM, holotype), Rio de Janeiro (Brazil). |

Description. 4, length 2.8 mm, width 2.2 mm; body form

oval, convex. Dorsal surface with head dull, alutaceous: pronotum

alutaceous, slightly shiny: elytron slightly alutaceous, shiny. Color

yellow except pronotum with large, black, basomedian spot, ante-

rior margin of spot broadly triangular, base of spot wide, contiguous

with black elytral spot; elytron with basal, lateral, and apical borders

very narrowly black, with 1 large black spot extended from base on

each side of scutellum posteriorly to apical declivity, spot “pinched”

medially, apically rounded (Fig. 127); pro-, meso-, metasterna black;

outer margin of propleuron yellow; outer margin of epipleuron red-

dish yellow; abdomen black except outer 1/4 and sternum 5-6 red-

dish yellow. Head punctures fine, distinct, separated by a diameter

or less; pronotal punctures larger than on head, separated by a diam-

eter or less; elytral punctures larger than pronotal punctures, sepa-

rated by less than to twice diameter. Metasternal punctures larger

than on elytron, separated by a diameter medially, nearly contiguous

laterally. Punctures on abdominal sterna 1-4 coarse, dense medially,

becoming dense, fine laterally; sterna 5-6 finely, densely punctured

throughout; area on 1% abdominal sternum inside postcoxal arc

shiny, finely, sparsely punctured. Clypeal apex emarginate, clypeus

and frons joined at abrupt angle. Epipleuron oblique, grooved inter-

nally, descending externally, femoral excavations deep. Prosternum

with intercoxal carinae narrowly separated at apex, tapered nearly

to base, basal stem short, reaching basal margin. Protibia narrow,

not flanged. Postcoxal line on 1% abdominal sternum not extended

to posterior sternal margin, slightly flattened along margin, apex

recurved 1/2 distance to basal sternal margin. Fifth sternum with

apex broadly, feebly emarginate; 6° sternum feebly emarginate.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI SI

Genitalia with basal lobe 3/4 length of paramere, sinuate, strong-

ly, obliquely truncate apically, sinuate, with angulate projection

on one side; paramere wide, tapered to rounded apex in apical 3/4

(Fig.129); sipho short, robust, apically sinuate, with membranous

area, basal capsule with inner arm wide, short, outer arm wide, lon-

ger than inner arm, apically knobbed, base sinuate (Figs.130, 131).

2 Similar to male except length 3.3 mm, width 2.7 mm; black

spot on pronotum and black spot on sutural margin of elytron

slightly smaller; elytron with small, faint, brown spot. Genitalia

with basal unit expanded, tapered on each end, beak large; connect-

ing duct short (Fig. 132, 133).

Variation. Length 2.8 to 3.3 mm, width 2.2 to 3.3 mm. Size of

black elytral spot varies from the typical to wide basally, and long,

tapered apically (Fig. 128, H. insignis), to narrow basally and short,

tapered apically; black outer elytral border absent in H. insignis.

Geographical distribution: Brazil

Specimens examined: 3 (the type specimens): 2 Brazil, 1 Brazil,

Rio de Janeiro.

Remarks. AHyperaspis operaria has a dorsal color pattern

that 1s unique within the South American fauna. The type speci-

mens of H. operaria and H. communalis have virtually identical color

patterns, but the color pattern of H. insignis varies somewhat from

the typical. After comparing these three type specimens, we con-

clude that they represent the same species; hence the synonymy

listed above. However, examination of male specimens of H. insignis

may validate that name as a distinct species.

10. Hyperaspis limbigera (Mulsant, 1853)

Cleothera limbigera Mulsant, 1853: 77.

Hyperaspis limbigera: CROTCH 1874: 225; KORSCHEFSKY 1931: 191; GORDON 1987: 28.

Hyperaspis limbifera: BLACKWELDER 1945: 447.

Type locality. Brazil, UMZC (lectotype designated by GORDON

1987).

Description. 6, length 2.8 mm, width 2.4 mm; body

form rounded, convex. Dorsal surface with head dull, alutaceous:

pronotum shiny: elytron shiny. Color yellow except pronotum with

312 R.D. GORDON & C. CANEPARI

2 large, dark brown, narrowly separated triangular spots at base,

spots widely separated from apical margin, and small, light brown,

oblong spot near lateral margin; elytron with irregular brown bor-

der on sutural margin, border slightly expanded in anterior 1/3,

and connected to large, irregular brown band on apical declivity,

brown band curved laterally and anteriorly, reaching base anterior

to humeral callus, brown sutural border feebly connected to lateral

band in basal 1/3 (Figs. 134, 135); pro-, meso-, metasterna blackish

brown; abdomen yellowish brown except outer 1/4 and sternum 5-6

reddish yellow. Head punctures fine, hidden in alutaceous sculpture;

pronotal punctures larger than on head, separated by a diameter or

less; elytral punctures smaller than pronotal punctures, separated by

1 to 2 times a diameter. Metasternal punctures coarse, much larger

than on elytron, separated by a diameter medially, nearly contigu-

ous laterally. Punctures on abdominal sterna 1-4 coarse, moderately

dense medially, becoming dense, fine laterally; sterna 5-6 finely,

densely punctured throughout; area on 1% abdominal sternum in-

side postcoxal arc alutaceous, coarsely, sparsely punctured. Clypeal

apex emarginate, clypeus and frons joined at abrupt angle. Antenna

with 11 articles (Fig. 136, basal article missing). Epipleuron oblique,

grooved internally, descending externally, femoral excavations deep.

Prosternum with intercoxal carinae widely separated at apex, conver-

gent nearly to base, basal stem short, reaching basal margin. Protibia

narrow, not flanged. Postcoxal line on 1*t abdominal sternum not ex-

tended to posterior sternal margin, slightly flattened along margin,

apex recurved 3/8 distance to basal sternal margin. Fifth sternum

with apex broadly, feebly emarginate; 6° sternum feebly emarginate.

Genitalia with basal lobe 5/8 length of paramere, sinuate, strongly,

obliquely truncate apically, sinuate, with strong, angulate projection

on one side; paramere wide, tapered to rounded apex in apical 3/4;

sipho long, robust, apically sinuate, with membranous area, basal

capsule with inner arm long, slender, apically knobbed, outer arm

short, wide, apically knobbed, base sinuate.

P Similar to male except triangular pronotal spots widely separated

from anterior margin; apex of apical tergum with row of large spines.

Genitalia with basal unit elongate; connecting duct short (Fig.137).

Variation. Length 2.7 to 3.0 mm, width 2.4 to 2.5 mm. Elytron

with connection of brown sutural border to lateral brown band fee-

ble, strong, or entirely absent.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 313

Geographical distribution: Brazil

Specimens examined: 4. Brazil: “Brazil” (lectotype); Rio de Ja-

neiro; Sào Paulo (BM) (UMZC) (USNM).

Remarks. AHAyperaspis limbigera is distinguished by a com-

bination of dorsal color pattern, elytral punctures distinctly smaller

than pronotal punctures, and row of strong spines at apex of the

female apical tergum.

11. Hyperaspis bisignata n. sp.

Type material. Holotype d: (Argentina), Est. Exp. Loreto, Dr.

A. Ogloblin, Willard H. Nutting, Jr., collection donated to the Ca-

lif. Academy of Sciences May 1990 (CAS). Allotype ®: Brazil, St

Catarina (Santa Catarina), Plaumann, Willard H. Nutting, Jr., col-

lection donated to the Calif. Academy of Sciences May 1990 (CAS).

Paratypes, 10. 1, same data as holotype (CAS); 1, same data as allo-

type (CAS); 1, (Argentina), Est. Exp. Loreto, 193..., Dr. A. Oglob-

lin, Korschefsky Collection 1952, Willard H. Nutting, Jr., collection

donated to the Calif. Academy of Sciences May 1990 (CAS); (Ar-

gentina), Est. Exp. Loreto, 193..., Dr. A. Ogloblin (CAS); 1, Brazil,

Rio (Grande) do Sul, Lagos. S. Catar (Santa Catarina), 14.1.953, W.

Wittmer (CAS); 2, Brazil, Esti.S.. Paulo! (Sao Paulo), Capital: Coll:

A. Sacca, 573, Korschefsky Collection 1952 (USNM); 1, (Brazil), Sao

Paulo, Det. by F. W. Nunenmacher, Korschefsky Collection 1952

(USNM): 1, (Brazil), ‘Sào-Paulo; “Capital, Cel. HP. Fi, 24-X-31, RE

Korschefsky cum. type comp., Hyperaspis Deyrollei Cr., Korschefsky

Collection 1952 (USNM); 1, Bolivia, Dept S. Cruz (Santa Cruz),

450 m, umg. Buenavista, Steinback coll., det. by F. W. Nunen-

macher, H. petropolitana Cr., Korschefsky Collection 1952 (USNM).

Description. 6, length 3.0 mm, width 2.4 mm; body

oval, convex. Dorsal surface with head strongly alutaceous, dull:

pronotum alutaceous, feebly shiny; elytra shiny. Color black except

head yellow; pronotum with lateral 1/4 and apical 1/4 yellow, elytron

with 2 yellow spots, median spot slightly anterior to middle, irregu-

larly rounded, subapical spot transversely oval (Figs. 138, 139); an-

tenna, labial palpi, propleuron yellow; legs reddish yellow except

basal 3/8 of femora black; abdomen black with lateral 1/8 reddish

yellow. Head punctures fine, nearly invisible in alutaceous sculpture,

314 R.D. GORDON & C. CANEPARI

separated by less than to twice a diameter; pronotal punctures larg-

er than on head, separated by a diameter or less; elytral punctures

larger than pronotal punctures, separated by less than to a diameter.

Metasternal punctures much larger than on elytron, separated by

a diameter medially, dense, contiguous laterally. Punctures on ab-

dominal sterna 1-4 larger than on elytra, dense medially, becoming

fine, dense laterally, sterna 5-6 finely densely punctured throughout;

area on 1* abdominal sternum inside postcoxal arc shiny, coarse-

ly, densely punctured. Epipleuron oblique, not internally grooved,

weakly descending externally. Clypeal apex deeply emarginate. Epi-

pleuron oblique, internally grooved, descending externally, femoral

excavations deep. Antenna with 11 articles. Prosternum with inter-

coxal carinae widely separated at apex, convergent from apex to-

ward base, basally stemmed. Protibia not flanged. Postcoxal line on

1* abdominal sternum reaching posterior sternal margin, flattened

along margin, apex recurved 1/3 distance to basal sternal margin.

Fifth sternum with apex broadly, distinctly emarginate; 6° sternum

nearly truncate. Genitalia with basal lobe 2/3 as long as paramere,

wide, apex slightly rounded, with angulate projection on one mar-

gin; paramere wide, tapered from middle to rounded apex; (Fig.

140); sipho robust, short, apex curved upward with membranous

area, basal capsule with inner arm slender, long, outer arm slender,

longer than inner, apically truncate (Fig. 141, 142).

2 Similar to d except head yellow with vertex and narrow lat-

eral border inside eye black; pronotum black except lateral 1/4 yel-

low. Genitalia with beak of basal unit short, curved; connecting duct

short (Fig. 143).

Variation. Length 2.6 to 3.3 mm, width 2.0 to 2.6 mm.

Etymology. The specific name is from the Latin bis, meaning

two, and signata, meaning mark, referring to the presence of 2 spots

on each elytron.

Remarks. This species is very similar to H. camargoi but

is distinguished from that species by the following: larger in average

size; 2 head mostly yellow; small elytral spots, median elytral spot

anterior to middle, widely separated from postapical spot; basal lobe

of 3 genitalia slightly longer and proportionately not as wide; sipho

shorter, siphonal basal capsule not sinuate, inner and outer arms not

apically knobbed.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 315

12. Hyperaspis camargoi n. sp.

Type material. Holotype 6: Brazil, Campinas, F. C. Camar-

go, Hyperaspis silvestris Ws. ab. det. R. Korschefsky 27, Korschef-

sy Collection 1952 (USNM). Allotype @: Brasil, Campinas, F. C.

Camargo, Korschefsky Collection 1952 (USNM). Paratype: Brasil,

same data as allotype (USNM).

Description. 6, length 2.4 mm, width 1.8 mm; body

oval, convex. Dorsal surface with head weakly alutaceous, dull: pro-

notum and elytron shiny. Color black except head yellow; pronotum

with lateral 1/4 and apical 1/6 yellow, elytron with 2 large yellow

spots, postmedian spot irregularly round, subapical spot transverse-

ly oval, narrowly separated from postmedian spot (Figs. 144, 145);

antenna, labial palpi, propleuron, tibiae yellow; pro-, meso-, and

metasterna brown; abdomen with median area of sterna 1-5 brown,

lateral 1/8 and 6" sternum yellow. Head punctures fine, separated

by less than to twice a diameter; pronotal punctures larger than on

head, separated by 1 to 2 times a diameter; elytral punctures larger

than pronotal punctures, separated by a diameter of less. Metaster-

nal punctures much larger than on elytron, nearly absent medial-

ly, dense, contiguous laterally. Punctures on abdominal sterna 1-4

larger than on elytra, coarse, dense medially, becoming fine, dense

laterally; sterna 5-6 finely, densely punctured throughout; area on

1° abdominal sternum inside postcoxal arc shiny, coarsely, densely

punctured. Clypeal apex deeply emarginate, clypeus and frons joined

at abrupt angle. Epipleuron oblique, internally grooved, descending

externally, femoral excavations deep. Antenna with 11 articles (Fig.

146). Prosternum with intercoxal carinae widely separated at apex,

convergent from apex toward base, basally stemmed. Protibia nar-

row, not flanged. Postcoxal line on 1" abdominal sternum reaching

posterior sternal margin, flattened along margin, apex recurved 1/2

distance to basal sternal margin. Fifth sternum with apex broadly,

distinctly emarginate; 6 sternum nearly truncate. Genitalia with

basal lobe 1/2 as long as paramere, wide, apex obliquely truncate,

with angulate projection on one margin; paramere wide, tapered

from middle to rounded apex; (Fig. 147); sipho slender, long, apex

slightly curved upward, basal capsule sinuate, with inner and outer

arms slender, long, outer arm longer than inner, both arms apically

knobbed (Figs. 148, 149).

316 R.D. GORDON & C. CANEPARI

2 Similar to male except head black. Genitalia with beak of

basal unit short, curved; connecting duct short (Figs. 150, 152).

Variation:® Wenethy 2.3 toy2-summs) width. te tot. Samm. Sizerot

elytral spots slightly variable.

Etymology. This species is named for F. C. Camargo, collector

of the type series.

Remarks. See remarks under H. bisignata.

13. Hyperaspis biguttata (Sicard, 1912)

Cleothera biguttata Sicard, 1912: 136.

Hyperaspis biguttata: KORSCHEFSKY 1931: 185; BLACKWELDER 1945: 446.

Type locality. Brazil, Santos, DEI (lectotype here designated).

Description. 6, length 2.4 mm, width 2.0 mm; body

form rounded, convex. Dorsal surface with head alutaceous, feebly

shiny; pronotum feebly alutaceous, shiny; elytron slightly aluta-

ceous, shiny. Color black except head yellow; pronotum yellow with

large, median black area, black area not extended to anterior pro-

notal margin and strongly indented laterally with yellow; elytron

with 2 yellow spots, 1 oval spot on disc, 1 transverse, apical spot

at posterolateral angle (Fig.153); mouthparts, propleuron, legs red-

dish yellow; pro-, meso-, metasterna black; abdomen dark brown

except outer 1/4 and sternum 5-6 reddish yellow. Head punctures

fine, distinct, separated by less than to twice a diameter; prono-

tal punctures larger than on head, separated by a diameter or less;

elytral punctures as large as pronotal punctures, separated by less

than to twice a diameter. Metasternal punctures larger than elytral

punctures, sparse medially, contiguous laterally. Punctures on basal

abdominal sternum coarse, sparse, punctures on sterna 2-6 finely,

densely punctured throughout; area on 1" abdominal sternum inside

postcoxal arc alutaceous, nearly impunctate. Clypeal apex weakly

emarginate, clypeus and frons joined at right angle. Epipleuron flat,

grooved internally, weakly descending externally, femoral depres-

sion deep. Prosternum with intercoxal carinae widely separated at

apex, tapered toward base, basally stemmed, reaching basal margin.

Protibia narrow, not flanged. Postcoxal line on 1* abdominal ster-

num extended to posterior sternal margin, flattened along margin,

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 317

apex recurved 1/2 distance to basal sternal margin. Fifth sternum

with apex broadly, deeply emarginate; 6 sternum distinctly emar-

ginate. Genitalia with basal lobe 3/4 length of paramere, obliquely

truncate apically, with large, obtuse projection on one side; paramere

wide, tapered to rounded apex in apical 1/2 (Fig. 154); sipho robust,

apically sinuate, with membranous area, basal capsule with inner

arm long, slender, outer arm shorter than inner arm, basal border

emarginate (Fig. 155).

2 Not known.

Variation. None observed.

Geographical distribution: Brazil.

Specimens examined: 1. Brazil (lectotype) (DEI).

Remarks. ‘This species is similar to H. bisignata in dorsal

color pattern, but the apical spot is distinctly transverse in H. big-

uttata, rounded in HM. bisignata, and H. biguttata has the median,

black pronotal area very strongly indented with yellow laterally. In

addition, the siphonal capsule differs between these species.

A d type specimen in the DEI is designated as lectotype to

stabilize future usage of this name. Lectotype labels are “Santos,

J. Metz/Coll. Kraatz Sicard det./Syntypus (pink paper) /Hyperaspis

biguttata Sic n. sp. type”.

14. Hyperaspis cleida Mulsant, 1850

Hyperaspis cleida Mulsant, 1850: 670; CroTcH 1874: 229; KORSCHEFSKY 1931: 186;

BLACKWELDER 1945: 446.

Description. ‘ length "2.6 mm, widen i.e mm. body

form elongate, slightly flattened. Dorsal surface with head aluta-

ceous, feebly shiny; pronotum alutaceous, feebly shiny; elytron

slightly alutaceous, shiny. Color black except head yellow; prono-

tum black except narrow anterior margin and lateral 1/4 yellow;

elytron with 4 yellow spots, 1 obliquely oval spot laterad of scutel-

lum, 1 round spot just anterior to apical declivity near suture, 1

small, triangular spot on lateral margin at apical declivity, 1 trans-

versely oval spot at apex near suture (Figs. 156, 157); mouthparts,

propleuron, pro- and mesolegs reddish yellow, metafemur brown;

pro-, meso-, metasterna black; abdomen dark brown except outer

318 R.D. GORDON & C. CANEPARI

1/6 of sterna 1-4 reddish yellow. Head punctures fine, distinct,

separated by less than to twice a diameter; pronotal punctures very

coarse, separated by a diameter or less; elytral punctures slightly

larger than pronotal punctures, separated by a diameter or less.

Metasternal punctures larger than elytral punctures, sparse medi-

ally, separated by less than a diameter laterally. Punctures on basal

abdominal sternum coarse, sparse, punctures on sterna 2-6 finely,

densely punctured throughout; area on 1% abdominal sternum in-

side postcoxal arc alutaceous, sparsely, coarsely punctured. Clypeal

apex strongly emarginate, clypeus and frons joined at right angle.

Epipleuron flat, grooved internally, weakly descending externally,

femoral depression deep. Prosternum with intercoxal carinae widely

separated at apex, tapered toward base, basally stemmed, reaching

basal margin. Protibia narrow, not flanged. Postcoxal line on 1*

abdominal sternum extended to posterior sternal margin, flattened

along margin, apex recurved 1/3 distance to basal sternal margin.

Fifth sternum with apex broadly, weakly emarginate; 6" sternum

weakly emarginate. Genitalia with basal lobe 1/2 length of paramere,

obliquely rounded apically, with large, angulate projection on one

side; paramere wide, tapered to rounded apex in apical 1/3 (Fig.

158); sipho robust, apically sinuate, with membranous area, basal

capsule with inner arm long, robust, outer arm shorter than inner

arm, basal border emarginate (Figs. 159).

2. Not known.

Variation. None observed.

Type locality: Brazil.

Geographical distribution: Brazil.

Specimens examined: 1. Brazil.

Remarks. ‘The species described above is recognized by

the presence of 4 yellow, elytral spots, and extremely coarse dorsal

and ventral punctation.

Hyperaspis cleida was described by Mulsant from specimen(s)

in the Chevrolat collection, which was subsequently obtained by

Crotch. A single specimen in that collection is labeled “d (green pa-

per)/187 (yellow paper) cleida m.” ‘There is no indication that this is

a type; however, it matches Mulsant’s original description extremely

well; therefore, we assume it to be an example of H. cleida, but do

not designate it as a lectotype.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 319

15. Hyperaspis tayronensis n. sp.

Type material. Holotype 6: Colombia, Magd. (Magdalena), Par-

que Tayrona, 21 mi. E. Sta. Marta, V-17-1973, Howden & Campbell

(CNC). Paratypes, 2. 1, Colombia, Magd., 12 mi. W. Sta. Marta,

V-18-1973, Campbell & Howden; 1, Colombia, Magd., 25 km. E.

Barranquilla, V-16-1973, Campbell & Howden (CNC) (USNM).

Description. dg, length 2.3 mm, width 1.7 mm; body

oval, convex. Dorsal surface with head alutaceous, dull: pronotum

and elytra slightly alutaceous, shiny. Color dark brown except head

yellow; pronotum yellow with small, round, median brown spot on

lateral 1/3, large, basomedian, anchor-shaped brown spot not ex-

tended to apical margin; elytron with 7 small, yellow spots, 1 round

scutellar spot, 1 very small, oval spot between scutellar spot and

humerus, 1 oval spot on humeral angle, 1 irregularly triangular dis-

cal spot, 1 elongate spot on lateral margin laterad of discal spot, spot

on apical declivity narrowly connected to posterior spot on lateral

angle (Figs. 160, 162); pro-, meso-, metasterna dark brown; abdo-

men brownish yellow with lateral 1/8 and sterna 5-6 yellow. Head

punctures fine, separated by less than to twice a diameter; pronotal

punctures larger than on head, separated by a diameter or less; ely-

tral punctures larger than pronotal punctures, separated by less than

to twice a diameter. Metasternal punctures larger than on elytron,

absent medially, dense, contiguous laterally. Punctures on abdomi-

nal sterna 1-4 larger than on elytra, sparse medially, becoming fine,

dense laterally, sterna 5-6 finely densely punctured throughout;

area on 1" abdominal sternum inside postcoxal arc shiny, coarsely,

sparsely punctured. Clypeal apex deeply emarginate, clypeus and

frons joined at abrupt angle. Epipleuron oblique, feebly grooved in-

ternally, descending externally, femoral excavations deep. Antenna

with 11 articles. Prosternum with intercoxal carinae narrowly sepa-

rated at apex, convergent from apex toward base, basally stemmed.

Protibia narrow, not flanged. Postcoxal line on 1% abdominal ster-

num not reaching posterior sternal margin, rounded along margin,

apex recurved 3/8 distance to basal sternal margin. Fifth sternum

with apex barely perceptibly emarginate; 6° sternum nearly truncate.

Genitalia with basal lobe 2/3 as long as paramere, apex obliquely

rounded, with strong angulate projection on one margin; paramere

wide, tapered from middle to rounded apex; (Fig. 163); sipho ro-

320 R.D. GORDON & C. CANEPARI

bust, apex sinuate, with membranous area, basal capsule with inner

arm wide, longer than outer arm, outer arm wide, short, apically

truncate (Figs. 164, 165).

2 Not known.

Variation. Length 2.3 to 2.5 mm, width 1.7 to 2.0 mm. Me-

dian pronotal spot may be divided medially into narrow, transverse

brown vitta with each end bent anteriorly, and irregularly triangular

apical spot extended laterally nearly to round lateral spot; yellow

elytral spots may be large with scutellar, discal, medial lateral, and

spot on apical declivity narrowly connected (Fig. 161).

Etymology. The species is named for Parque T'ayrona where the

holotype was collected.

Remarks. Seven spots on each elytron and the unusually

marked pronotum are characters shared only with H. histrionica.

Very few Colombian species were available to early systematists

such as Mulsant and Crotch; therefore, a high percentage of spe-

cies from this region remain undescribed. These Colombian species

tend to have very distinctive dorsal color patterns as evidenced by

H. tayronensis.

16. Hyperaspis pseudodonzeli n. sp.

Type material. Holotype d: Venezuela, Aragua, Palo Ne-

gro, 455 m, 31-VII-1959, sobre escamas en suda (sic!), E. No.

16, col. A. Notz (USNM). Allotype £: Trinidad, St. Augustine,

BWI, Feb. 52, Collector F.D. Bennett, Orthezia insignis on cro-

ton (USNM). Paratypes, 38. 8, same data as allotype (USNM); 1,

Colombia, Cali, CIAT, Nov 1977, F. D. Bennett Collector, Pred.

on Orthezia (USNM); 1, Colombia, Magd (Magdalena), Argelia,

28-IX-‘35, Murillo No 107 (USNM); 1, Curacao, Schottgatwee,

July 1-5, 62, J Maldonado C (USNM),; 3, British Guiana (Guyana),

Georgetown, August 1953, Collector F.J. Simmonds (USNM); 1,

Trinidad, Orthezia, C.1.E.16482 (BM); 7, Trinidad, St. Augustine

(CNC); 2, Trinidad W.I., S/Grande, 16-4-1961, N. Gopaul Coll.

(CNC); 4, Trinidad, Tunapuna, 2-4-I-59 (CNC); 6, Trinidad, Tu-

napuna, 6-I-1959 (CNC); 2, Trinidad, Tunapuna, Aug. 1-3,1969,

H. & A. Howden (USNM); 2, Venezuela, Aragua, El Limon, 450

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 5921!

m, 22-V-69, Predator adulto sobre escama en Croto, E. No. 16, Col.

A. Notz (USNM).

Description. 6, length 2.5 mm, width 2.0 mm; body

oval, convex. Dorsal surface with head weakly alutaceous, shiny:

pronotum and elytra shiny. Color dark brown except head yellow;

pronotum yellow with 4 pale brown, indistinct spots, 1 triangular

spot medially on each side of middle, 1 narrow, basomedian spot

on each side of middle; elytron with 3 large, yellow spots, 1 irregu-

larly rounded spot on disc near suture, 1 somewhat rectangular spot

on lateral margin laterad of discal spot, 1 comma shaped subapical

spot; antenna, mouthparts, propleuron, mesopleuron, median 1/2 of

epipleuron, legs yellow; abdomen with median area of sterna 1-4

brownish yellow, lateral 1/4 and sterna 5-6 yellow. Head punctures

fine, separated by less than a diameter; pronotal punctures larger

than on head, separated by less than to twice a diameter; elytral

punctures larger than pronotal punctures, separated by less than to

twice a diameter. Metasternal punctures much larger than on elytron,

sparse medially, dense, nearly contiguous laterally. Punctures on ab-

dominal sterna 1-4 larger than on elytra, coarse, sparse medially,

becoming fine, dense laterally; sterna 5-6 finely, densely punctured

throughout; area on 1*t abdominal sternum inside postcoxal arc shiny,

sparsely, coarsely punctured. Clypeal apex emarginate, clypeus and

frons joined at abrupt angle. Epipleuron slightly oblique, not me-

dially grooved, slightly descending externally, femoral excavations

deep. Antenna with 11 articles (Fig. 170). Prosternum with inter-

coxal carinae widely separated at apex, convergent from apex toward

middle, not basally stemmed, not reaching basal margin. Protibia

narrow, not flanged. Postcoxal line on 1** abdominal sternum reach-

ing posterior sternal margin, rounded along margin, apex recurved

2/3 distance to basal sternal margin. Fifth sternum with apex broad-

ly, weakly emarginate; 6° sternum weakly emarginate. Genitalia with

basal lobe 3/4 as long as paramere, wide, apex obliquely truncate,

with angulate projection on one margin; paramere wide, tapered from

apical 1/4 to rounded apex; (Fig. 172); sipho robust, apex sinuate,

with membranous area, basal capsule with inner arm short, wide,

outer arm wide, slightly shorter than inner arm (Figs. 173, 174).

2 Similar to male except head reddish brown; pronotum may be

immaculate, entirely yellow, or brown except lateral 1/3 yellow. Gen-

italia with beak of basal unit short; connecting duct short (Fig. 175).

BO R.D. GORDON & C. CANEPARI

Variation. Length 2.3 to 2.5 mm, width 1.8 to 2.0 mm. Male

pronotal color varies from that described above to mostly dark brown

with lateral 1/3 yellow (Figs. 167-169); elytral spots vary consider-

ably in size and the subapical spot may not be comma shaped but

irregularly rounded.

Etymology. The specific name refers to the resemblance to H.

donzeli.

Remarks. Most specimens of H. pseudodonzeli have the

comma-shaped subapical spot on each elytron, but female specimens

often have this spot rounded. This species has been identified as

H. donzeli in many collections, but H. donzeli is known only from

Brazil, and the subapical spot on each elytron is transverse rather

than comma shaped.

17. Hyperaspis orthivora n. sp.

Type material. Holotype d: Colombia, Valle (Valle del Cauca),

Palmira, ICA, 4.111.1991, J. Castillo, sp. EY-9A-91, ex. Orthezia sp.,

PIE 21639, Hyperaspis det. R.G. Booth (BM). Allotype 2; same

data as holotype (BM). Paratypes, 9. 5, same data as holotype (BM);

4, Colombia, Cali, CIAT, Nov 1977, F.D. Bennett Collector, Pred.

of Orthezia (USNM).

Description. 6, length 2.4 mm, width 2.0 mm; body

oval, convex. Dorsal surface with head strongly, feebly shiny; pro-

notum slightly alutaceous, shiny; elytron shiny. Color yellow except

pronotum with 5 pale brown spots, 2 triangular spots medially in

anterior 1/2, 1 triangular, basal spot on each side of middle, 1 baso-

median, oval spot anterior to scutellum; elytron pale mottled brown

with 3 yellow spots, 1 large, oval spot near sutural margin in anterior

portion of disc, 1 oval spot on lateral margin laterad of discal spot, 1

large, comma-shaped spot at apex, all spots feebly connected; pro-,

meso-, metasterna dark reddish brown; abdomen with median area

of sterna 1-4 dark brown, lateral 1/4 and sterna 5-6 reddish yellow.

Head punctures fine, separated by a diameter or less; pronotal punc-

tures larger than on head, separated by a diameter or less; elytral

punctures as large as pronotal punctures, separated by a diameter or

less. Metasternal punctures larger than on elytron, sparse medially,

separated by less than a diameter laterally. Punctures on abdominal

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 323

sterna 1-4 as large as on elytra, dense medially, becoming fine, dense

laterally; sterna 5-6 finely, densely punctured throughout; area on

1* abdominal sternum inside postcoxal arc shiny, sparsely, coarsely

punctured. Clypeal apex emarginate, clypeus and frons joined at

abrupt angle. Epipleuron oblique, grooved internally, weakly de-

scending externally, femoral depressions deep. Antenna with 11 arti-

cles (Fig. 179). Prosternum with intercoxal carinae widely separated

at apex, convergent toward base, basally stemmed, reaching basal

margin. Protibia narrow, not flanged. Postcoxal line on 1% abdomi-

nal sternum not quite reaching posterior sternal margin, slightly flat-

tened along margin, apex angled forward 1/3 distance to basal ster-

nal margin. Fifth sternum with apex broadly, distinctly emarginate;

6 sternum feebly emarginate. Genitalia with basal lobe 2/3 as long

as paramere, apex obliquely truncate, with large, angulate projection

on one margin; paramere wide, tapered to rounded apex in apical

1/2 (Fig. 180); sipho slender, apically sinuate, apex with membra-

nous area, basal capsule with inner arm short, slender, sinuate, outer

arm long, wide, apically knobbed, basal border sinuate (Fig. 181).

2 Similar to male except head reddish yellow; median 1/3 of

pronotum reddish yellow. Genitalia with beak of basal unit short;

connecting duct short (Figs. 182, 183).

Variation. Length 2.4 to 2.7 mm, width 2.3. Dark areas on dor-

sum and venter from light reddish brown to nearly black. Male with

dark pronotal spots from described to dark reddish brown, basal

spots joined along margin to form narrow band, anterior spots nar-

rowly connected to the basal band (Figs. 176, 177). Elytral spots

feebly connected, or strongly, definitely connected (Fig. 178).

Etymology. This specific name is a combination of the first 4

letters of the scale name Orthezia, and the Latin vorator, meaning

devourer, referring to the scale-eating habits of this species.

Remarks. The predominantly yellow color, contrasted with

a brown or black elytral ground color, in combination with 3 yellow

spots on each elytron, distinguish this species.

18. Hyperaspis zonula n. sp.

Type material. Holotype d: Brasil, Lagos, Rio do Sul, S. Cater

(Santa Catarina), 14.1.1953, W. Wittmer (CAS). Allotype ®: Bra-

324 R.D. GORDON & C. CANEPARI

zil, S.C. (Santa Catarina), Sao Joaquima, Boa Jesus Rd, 28.xii.1972,

V.F. Eastop 13,890, Brit. Mus. 1976-97, sweeping (BM). Paratype:

Brasil, (illegible word), 4496, Hist. Coll. (Coleoptera) Nr. 4496, Ge-

nus? spec. Brasil., Virmnd., Zool. Mus. Berlin (ZMHB).

Description. 6, length 3.0 mm, width 2.4 mm; body

oval, convex. Dorsal surface with head strongly alutaceous, dull;

pronotum slightly alutaceous, shiny; elytra shiny. Color yellow ex-

cept head with vertex black; pronotum with narrow, basomedian

brown band; elytron with suture narrowly brown, and brown, arcu-

ate vitta extended from base inside humeral callus nearly to sutural

margin on apical declivity (Figs. 184, 185); pro-, meso-, metasterna

dark reddish brown; abdomen with median area of sterna 1-4 dark

brown, lateral 1/4 and sterna 5-6 reddish yellow. Head punctures

fine, hidden in alutaceous sculpture; pronotal punctures larger than

on head, separated by a diameter or less; elytral punctures as large

as pronotal punctures, separated by less than to twice a diameter.

Metasternal punctures larger than on elytron, separated by a diame-

ter medially, becoming contiguous laterally. Punctures on abdominal

sterna 1-4 as large as on elytra, sparse medially, becoming fine, dense

laterally; sterna 5-6 finely, densely punctured throughout; area on 1%

abdominal sternum inside postcoxal arc shiny, sparsely punctured.

Clypeal apex emarginate, clypeus and frons joined at abrupt angle.

Epipleuron oblique, slightly grooved internally, strongly descending

externally, femoral excavations deep. Antenna with 11 articles. Pros-

ternum with intercoxal carinae widely separated at apex, convergent

to base, not basally stemmed, reaching basal margin. Protibia nar-

row, not flanged. Postcoxal line on 1% abdominal sternum reaching

posterior sternal margin, flattened along margin, apex recurved 1/2

distance to basal sternal margin. Fifth sternum with apex truncate;

6 sternum barely perceptibly emarginate. Genitalia with basal lobe

5/8 as long as paramere, apex obliquely truncate, with large, angu-

late projection on one margin; paramere wide, tapered to rounded

apex in apical 1/2 (Fig. 186); sipho long, apically sinuate, apex with

membranous area, basal capsule with inner arm long, inside margin

notched, outer arm short, wide, dark brown, apically knobbed, basal

border deeply emarginate (Fig. 187).

2 Similar to male except pronotum with 1 faint, triangular spot

on each side of middle in anterior 1/2. Genitalia with beak of basal

unit short, extremely wide, batlike; connecting duct short (Fig. 188).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 325

Variation. Length 3.0 to 3.2 mm, width 2.4 to 2.7 mm. Dark ar-

eas on dorsum and venter from light reddish brown to nearly black.

6 with dark pronotal spots from described to dark reddish brown,

basal spots joined along margin to form narrow band, anterior spots

narrowly connected basal band. Elytral spots feebly connected or

strongly, definitely connected.

Etymology. The specific name is from the Latin zona, mean-

ing belt, or girdle, referring to the curved elytral vitta and rounded

form.

Remarks. This is a distinctive species with an atypical

color pattern. The pale dorsal background with a dark curved vitta

on each elytron is similar to that of some species of the coccidu-

line genus Zenoria Mulsant, but not to any known species of South

American Hyperaspis.

19. Hyperaspis brethesi n. sp.

Type material. Holotype d: 27622, Argentina, M. Garcia (Isla

Martin Garcia), BA Aires (Buenos Aires), Korschefsky Collection

1952 (USNM). Allotype 2: Argentina, Buenos Aires, Isla Martin

Garcia, I-1938, M.J. Viana (USNM). Paratypes, 28. 19, same data

as holotype; 2, same data as allotype; 1, Argentina, Buenos Aires,

Isla Martin Garcia, 4-1937, M.J. Viana; 1. BS Aires, F. 1936 Col.M.

Viana; 3, Argentina, Camargo, Nunenmacher Collection; 1, Argen-

tina, Tantana, Chaco, 37374; 1 (Argentina) Cordoba, Alta (illegi-

ble name), III-1959, Daguerre, Argentina 1968, Colln. J. Daguerre

(BM) (CAS) (MBR) (USNM).

Other specimens: 6. 1, Argentina, Corrientes, 5. Tome; 1, Ar-

gentina, Misiones, Posadas, NLH Krauss; 1, Argentina, Salta, Oran;

3, Bolivia, Santa Cruz, 3-4 k N Agua Clara, 12 Oct. 2000, Wappes

& Dozier (MBR) (USNM).

Description. 6, length 2.1 mm, width 1.7 mm; body

oval, convex. Dorsal surface with head slightly alutaceous, shiny;

pronotum slightly alutaceous, shiny; elytron shiny. Color yellow ex-

cept pronotum with median 1/3 dark brown with yellow, eyelike

spot on each side of middle; elytron yellow, speckled with brown

dots instead of entirely dark brown above, elongate brown spot on

disc at sutural margin, another elongate brown spot near suture on

326 R.D. GORDON & C. CANEPARI

apical declivity, humeral callus with small brown spot (Fig. 189);

antenna, mouthparts, propleuron, mesopleuron, epipleuron, legs

yellow; abdomen with median area of sterna 1-4 dark brown, lat-

eral 1/4 and sterna 5-6 reddish yellow. Head punctures fine, sepa-

rated by 1 to 2 times a diameter; pronotal punctures larger than on

head, separated by a diameter or less; elytral punctures larger than

pronotal punctures, separated by less than to 2 times a diameter.

Metasternal punctures much larger than on elytron, separated by a

diameter medially, becoming dense, contiguous laterally. Punctures

on abdominal sterna 1-4 larger than on elytra, coarse, sparse medi-

ally, becoming fine, dense laterally; sterna 5-6 finely, densely punc-

tured throughout; area on 1% abdominal sternum inside postcoxal

arc shiny, sparsely, coarsely punctured. Clypeal apex emarginate,

clypeus and frons joined at abrupt angle. Epipleuron flat grooved

internally, slightly descending externally, femoral depression deep.

Antenna with 11 articles (Fig. 194). Prosternum with intercoxal

carinae narrowly separated at apex, convergent toward base, basally

stemmed, reaching basal margin. Protibia narrow, not flanged. Post-

coxal line on 1% abdominal sternum reaching posterior sternal mar-

gin, slightly flattened along margin, apex recurved 1/4 distance to

basal sternal margin. Fifth sternum with apex broadly, weakly emar-

ginate; 6 sternum slightly arcuate. Genitalia with basal lobe 3/4 as

long as paramere, apex obliquely rounded, with large, angulate pro-

jection on one margin; paramere wide, tapered to rounded apex in

apical 1/2 (Fig. 196); sipho slender, apex curved, basal capsule with

inner arm robust, apically knobbed, outer arm slightly shorter than

inner arm, basal border emarginate (Figs. 197, 198).

2 Similar to d except head brown; pronotum brown except

lateral 1/4 yellow, lacking central eyelike spots. Genitalia with beak

of basal unit short; connecting duct short (Fig. 199).

Variation. Length 2.1 to 2.4 mm, width 1.7 to 1.8 mm. Dorsal

color of pale phase varies from that described above to having either

more or fewer dark brown “speckles”; brown spots on suture may

be united with those on opposite elytron, creating 1 or 2 central

elytral spots. Dark phase brown except head yellow; pronotum yel-

low with large, brown, basomedian spot, spot widely separated from

anterior pronotal margin, apically emarginate, with 2 oblique, eye-

like yellow spots in middle; elytron laterally bordered with yellow,

curved vitta from scutellum at base to near sutural margin at apical

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI DL

declivity, vitta connected to lateral border slightly anterior to apical

declivity (Figs. 190-193).

Etymology. The species is named for Juan Bréthes, who con-

tributed much to the classification of South American Coccinellidae.

Remarks. There are two color phases of this species; the

paler being predominant among specimens examined is chosen as

typical. The remaining specimens are described under “Other speci-

mens.” Both color phases of H. brethesi are distinctive and super-

ficially very different. However, males of both have male pronota

identical in color with the central, eyelike spots and identical geni-

talia. The pale form bears a slight resemblance to H. operaria, but

the latter species is distinctly larger, more rounded, and lacks the

characteristic brown, elytral “speckles”.

Isla Martin Garcia, Buenos Aires, is the locality from which

most specimens examined were collected.

20. Hyperaspis rosariensis n. sp.

Type material. Holotype d: Argentina, Salta, Rosario de Ler-

ma, XI-10/14-1983, Malaise Trap, M. Wasbauer coll. (CDA).

Description. 6, length 2.6 mm, width 2.0 mm; body

oval, convex. Dorsal surface with head alutaceous, weakly shiny:

pronotum slightly alutaceous, shiny; elytra shiny. Color black ex-

cept head yellow; pronotum yellow with large, rectangular, baso-

median spot, spot widely separated from anterior margin, lateral

margin of spot slightly emarginated, elytron with 2 yellow spots, 1

small, round spot near suture anterior to middle of elytron, 1 small,

oval, apical spot at posterolateral angle (Fig. 200), spot at postero-

lateral angle not shown); antenna, mouthparts, propleuron yellow;

pro-, mesolegs reddish yellow except outer tibial margins brown;

metafemur brown except apical 1/8 yellow; abdomen brown with

lateral 1/8 reddish yellow. Head punctures fine, separated by less

than to twice a diameter; pronotal punctures larger than on head,

separated by less than to twice a diameter; elytral punctures slightly

larger than pronotal punctures, separated by 1 to 3 times a diameter.

Metasternal punctures much larger than on elytron, absent medi-

ally, dense, contiguous laterally. Punctures on abdominal sterna 1-4

328 R.D. GORDON & C. CANEPARI

larger than on elytra, dense medially, becoming fine, dense laterally,

sterna 5-6 finely densely punctured throughout; area on 1% abdomi-

nal sternum inside postcoxal arc shiny, coarsely, densely punctured.

Clypeal apex deeply emarginate, clypeus and frons joined at abrupt

angle. Epipleuron slightly oblique, internally grooved, slightly de-

scending externally, femoral excavations deep. Antenna with 11

articles. Protibia narrow, not flanged. Prosternum with intercoxal

carinae narrowly separated at apex, convergent from apex toward

base, basally stemmed, reaching basal margin. Postcoxal line on 1*

abdominal sternum not reaching posterior sternal margin, rounded

along margin, apex recurved 1/2 distance to basal sternal margin.

Fifth sternum with apex broadly, distinctly emarginate; 6" sternum

feebly truncate. Genitalia with basal lobe 2/3 as long as paramere,

apex narrowly rounded, with rounded projection on one margin;

paramere wide, tapered from middle to rounded apex; (Fig. 201);

sipho long, with apical membranous area, basal capsule with inner

arm wide, shorter than outer arm, apically truncate (Figs. 202, 203).

2 Not known.

Etymology. The species is named for the type locality.

Remarks. This species is similar to H. bisignata and H.

camargot, but male genitalia are quite different. In addition, dorsal

punctation differs, with H. bisignata and H. camargoi having dense

elytral punctures separated by less than to twice a diameter, dis-

tinctly larger than pronotal punctures. Hyperaspis rosariensis has ely-

tral punctures separated by 1 to 3 times a diameter and only slightly

larger than pronotal punctures.

21. Hyperaspis siladesma n. sp.

Type material. Holotype d: Brazil, Chapada, Acc. No. 2966,

Nov. (CMP).

Description. 6, length 2.1 mm, width 1.6 mm; body

oval, slightly flattened. Dorsal surface with head strongly aluta-

ceous, dull: pronotum shiny; elytra shiny. Color black except head

yellow; pronotum yellow with large, somewhat rectangular, basome-

dian spot, spot widely separated from anterior margin, with lateral

earlike projection, elytron with lateral border and discal spot yellow,

discal spot anterior to middle, irregularly rounded, lateral border

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 329

irregular, widened opposite discal spot and at apex (Fig. 204, 205);

antenna, mouthparts, propleuron yellow, pro- and mesolegs yellow;

metaleg yellow with femur brown except apical 1/8 yellow; abdo-

men brown with lateral 1/8 reddish yellow. Head punctures hid-

den in alutaceous sculpture; pronotal punctures larger than on head,

separated by less than to about a diameter; elytral punctures larger

than pronotal punctures, separated by less than to about a diameter.

Metasternal punctures much larger than on elytron, separated by a

diameter medially, dense, contiguous laterally. Punctures on abdom-

inal sterna 1-4 smaller than on elytra, sparse medially, becoming

fine, dense laterally, sterna 5-6 finely densely punctured through-

out; area on 1“ abdominal sternum inside postcoxal arc shiny, finely,

sparsely punctured. Clypeal apex deeply emarginate, clypeus and

frons joined at abrupt angle. Epipleuron slightly oblique, internally

grooved, slightly descending externally, femoral depressions moder-

ately deep. Antenna with 11 articles. Protibia narrow, not flanged.

Prosternum with intercoxal carinae widely separated at apex, con-

vergent from apex toward base, basally stemmed, reaching basal

margin. Postcoxal line on 1% abdominal sternum reaching posterior

sternal margin, flattened along margin, apex recurved 5/8 distance

to basal sternal margin. Fifth sternum with apex broadly, weakly

emarginate; 6 sternum weakly emarginate. Genitalia with basal

lobe 5/8 as long as paramere, apex narrowly rounded, with angulate

projection on one margin; paramere wide, tapered from basal 1/4 to

rounded apex (Fig. 206); (sipho lost except for basal capsule) basal

capsule with inner arm slender, shorter than outer arm, outer arm

wide, elongate, apically truncate (Fig. 207).

2 Not known.

Etymology. The species name is an arbitrary combination of

letters.

Remarks. AHyperaspis stiladesma has an elytral color pattern

similar to one of the color forms of H. scutifera, but that species occurs

in northern Colombia and Venezuela and has 10-articled antennae.

22. Hyperaspis octonotata n. sp.

Type material. Holotype d: Colombia, Magd. (Magdalena), 12

mi. W. Sta. Marta, IV-28-1973, Campbell & Howden (CNC).

330 R.D. GORDON & C. CANEPARI

Description. 6, length 2.9 mm, width 2.5 mm; body

form round, slightly flattened. Dorsal surface with head dull,

strongly alutaceous, pronotum and elytra shiny. Color yellow except

pronotum with 4 median brown spots, anterior 2 spots triangular,

basal 2 spots slightly triangular, widely separated anterior to scutel-

lum; elytron with 4 oval, brown spots and sutural margin narrowly

brown, 1 spot anterior to disc between suture and humerus, 1 spot

on humerus, 1 spot on apical declivity inside sutural margin, 1 spot

on apical declivity near lateral margin, sutural margin brown from

scutellum to apex (Fig. 208, 209); prosternum with basal border and

prosternal process pale reddish brown, pro- and mesosterna dark

reddish brown, median 2/3 of abdominal sterna 1-4 pale reddish

brown. Head punctures very fine, hidden in dense alutaceous sculp-

ture. Pronotal punctures larger than on head, separated by a diam-

eter or less; elytral punctures larger than pronotal punctures, sepa-

rated by less than to twice a diameter. Metasternal punctures larger

than on elytron, coarse, sparse medially, dense laterally. Punctures

on abdominal sterna 1-4 coarse, sparse medially, becoming dense,

fine laterally; sterna 5-6 finely, densely punctured throughout; area

on 1 abdominal sternum inside postcoxal arc feebly shiny, aluta-

ceous, coarsely, sparsely punctured. Clypeal apex deeply emargin-

ate, clypeus and frons joined at abrupt angle. Epipleuron oblique,

internally grooved, descending externally, femoral excavations deep.

Antenna with 11 articles. Prosternum with intercoxal carinae con-

vergent toward base, joined at midpoint, basally stemmed, reach-

ing basal margin. Protibia narrow, not flanged. Postcoxal line on

1** abdominal sternum extended to posterior sternal margin, slightly

flattened along margin, apex recurved 1/3 distance to basal sternal

margin. Fifth sternum with apex broadly, deeply emarginate; 6”

sternum feebly emarginate. Genitalia with basal lobe nearly as long

as paramere, wide, apex obliquely truncate, with rounded projection

on one side; paramere short, wide, rounded to apex in apical 1/4

(Fig. 210); sipho robust, apex sinuate, with membranous area, basal

capsule with inner arm long, slender, outer arm shorter and slightly

wider than inner arm (Figs. 211, 212).

2 Not known.

Etymology. The specific name is from the Latin octo, meaning

eight, and notata, meaning mark, referring to the 8 brown marks on

the elytra.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 331

Remarks. The color pattern described above, with varia-

tions, is repeated many times in the Hyperaspinae genera Hyperaspis

and Cyra, but within the onerata Group only H. octonotata and H.

campbelli have that pattern.

23. Hyperaspis campbelli n. sp.

Type material. Holotype 6: Colombia, Magd. (Magdalena), 12

mi. W. Sta. Marta, IV-29-1973, Campbell & Howden (CNC).

Description. 6, length 2.4 mm, width 2.0 mm; body

form round, slightly flattened. Dorsal surface with head dull,

strongly alutaceous, pronotum and elytra shiny. Color yellow except

pronotum with 4 median brown spots, anterior 2 spots triangular,

basal 2 spots slightly triangular, widely separated anterior to scutel-

lum; elytron with ground color in apical 2/3 reddish yellow (pink),

with 4 brown spots, 1 small, elongate spot on suture anterior to

middle, 1 narrow, elongate spot on suture in basal 1/2, 1 large, ir-

regular spot on humerus, 1 large, round spot medially on apical de-

clivity (Fig. 213); pro-, meso-, and metasterna reddish brown; basal

4 abdominal sterna pale reddish brown Head punctures very fine,

hidden in dense alutaceous sculpture. Pronotal punctures larger than

on head, separated by a diameter or less; elytral punctures larger

than pronotal punctures, separated by less than to twice a diam-

eter. Metasternal punctures larger than on elytron, sparse medial-

ly, nearly contiguous laterally. Punctures on abdominal sterna 1-4

coarse, sparse medially, becoming dense, fine laterally; sterna 5-6

finely, densely punctured throughout; area on 1% abdominal sternum

inside postcoxal arc feebly shiny, alutaceous, finely, sparsely punc-

tured. Clypeal apex deeply emarginate, clypeus and frons joined

at abrupt angle. Epipleuron oblique, internally grooved, descend-

ing externally, femoral excavations deep. Antenna with 11 articles.

Prosternum with intercoxal carinae convergent toward base, basally

stemmed, reaching basal margin. Protibia narrow, not flanged. Post-

coxal line on 1% abdominal sternum extended to posterior sternal

margin, flattened along margin, apex recurved 1/3 distance to basal

sternal margin. Fifth sternum with apex broadly, deeply emarginate;

6" sternum feebly emarginate. Genitalia with basal lobe 1/2 length

of paramere, apex obliquely rounded apically, with large, angulate

392 R.D. GORDON & C. CANEPARI

projection on one side; paramere short, wide, rounded to apex in

apical 1/2 (Fig. 214); sipho slender, apex sinuate, with membranous

area, basal capsule with inner arm long, slender, outer arm shorter,

wider than inner, apex knobbed (Fig. 215).

2 Not known.

Etymology. The species is named for one of the collectors of

the type specuinen.

Remarks. The color pattern of H. campbelli is similar to

that of H. octonotata, but the oddly “pink” elytral ground color is

unusual. In addition, the body form of H. campbelli is somewhat

flattened and the brown elytral spots are of different shapes and are

differently arranged, as illustrated.

24. Hyperaspis colombiensis n. sp.

Type material. Holotype d: Colombia, Valle (Valle del Cau-

ca), Palmira, Candelaria, alt. 1020 m., 31.141, Murillo No 5484

(USNM).

Description. 4, length 2.4 mm, width 2.0 mm; body oval,

slightly flattened. Dorsal surface with head strongly alutaceous, dull;

pronotum shiny; elytron shiny. Color yellow except pronotum with 4

brown spots, 2 triangular spots medially in anterior 1/2, 1 triangular,

basal spot on each side of middle; elytron reddish yellow (pink) with

4 brown spots and suture narrowly brown except brown area wider

on apical declivity, 1 small, elongate, oval spot near suture poste-

rior to scutellum, 1 small, elongate, oval spot between suture and

humeral callus near base, 1 large, irregular spot on humeral callus,

1, large, irregularly oval spot medially on apical declivity (Fig. 216);

mouthparts reddish yellow; pro-, meso-, metasterna black; abdomen

with median 1/3 of sterna 1-2 dark brown, remainder of abdominal

sterna yellow. Head punctures fine, separated by less than to twice

a diameter; pronotal punctures larger than on head, separated by a

diameter or less; elytral punctures as large as pronotal punctures,

separated by less than to twice a diameter. Metasternal punctures

larger than on elytron, separated by a diameter medially, nearly

contiguous laterally. Punctures on abdominal sterna 1-4 as large as

on elytra, sparse medially, becoming fine, dense laterally; sterna 5-6

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 333

finely, densely punctured throughout; area on 1% abdominal sternum

inside postcoxal arc shiny, nearly impunctate. Clypeal apex deeply

emarginate, clypeus and frons joined at abrupt angle. Epipleuron

oblique, grooved internally, weakly descending externally, femoral

depressions moderately deep. Antenna with 11 articles (Fig. 217).

Prosternum with intercoxal carinae widely separated at apex, paral-

lel, short, extended 1/2 distance to base. Clypeal apex deeply emar-

ginate, clypeus and frons joined at abrupt angle. Protibia narrow,

not flanged. Postcoxal line on 1* abdominal sternum reaching pos-

terior sternal margin, flattened along margin, recurved 1/2 distance

to basal sternal margin. Fifth sternum with apex broadly, distinctly

emarginate; 6" sternum feebly emarginate. Genitalia with basal lobe

5/8 as long as paramere, apex obliquely truncate, with small, angulate

projection on one margin; paramere wide, tapered to rounded apex

in apical 2/3 (Fig. 218); sipho slender, curved down in apical 1/4,

apex with membranous area, basal capsule heavily sclerotized, dark

brown, with inner arm long, slightly curved, outer arm longer than

inner, apically knobbed, basal border emarginate (Figs. 219, 220).

2 Not known.

Etymology. The species is named for Colombia, where the ho-

lotype was collected.

Remarks. ‘This species, known only from the holotype

male, is characterized by the “pink” ground color of the elytron,

unusual elytral spot arrangement, and short, parallel prosternal cari-

nae. In addition, three elytral spots arranged in a transverse row

across the base of each elytron is an unusual pattern in Hyperaspis.

25. Hyperaspis dissidens n. sp.

Type material. Holotype d: (Colombia), Villalobos, Barranca

de Naranjo, ex: Café Finca, Coll. DJ Calvert & I. Solis, VI-12-73

(USNM). Paratypes, 2:, same data as holotype (USNM).

Description. 6, length 2.7 mm, width 2.2 mm; body

oval, slightly flattened. Dorsal surface with head strongly alutaceous,

dull; pronotum slightly alutaceous, weakly shiny; elytron shiny. Col-

or black except head yellow with vertex narrowly black; pronotum

with lateral 1/3 yellow, yellow area with internal margin arcuate, an-

334 R.D. GORDON & C. CANEPARI

terior margin narrowly yellow; elytron with 2 yellow spots, 1 irregu-

lar, transverse spot on disc from near sutural margin to lateral mar-

gin, 1 obliquely oval, subapical spot, anterior margin of spot slightly

emarginated with black (Figs. 221, 222); antenna, propleuron yel-

low; mouthparts reddish brown; proleg reddish yellow except basal

1/3 of femur dark brown, mesoleg reddish yellow except anterior

and posterior margins of femur, outer margin of tibia dark brown,

metaleg dark brown except apex of femur, inner tibial margin and

tarsus reddish yellow; abdomen with median area of sterna 1-5 dark

brown, lateral 1/8 and sternum 6 reddish yellow. Head punctures

fine, hidden in alutaceous sculpture, separated by 1 to 2 times a

diameter; pronotal punctures larger than on head, separated by a

diameter or less; elytral punctures larger than pronotal punctures,

separated by less than to twice a diameter. Metasternal punctures

larger than on elytron, separated by a diameter medially, becom-

ing nearly contiguous laterally. Punctures on abdominal sterna 1-4

smaller than elytral punctures, sparse medially, becoming fine, dense

laterally; sterna 5-6 finely, densely punctured throughout; area on 1°

abdominal sternum inside postcoxal arc, finely, sparsely punctured.

Clypeal apex deeply emarginate, clypeus and frons joined at abrupt

angle. Epipleuron flat, slightly grooved internally, not descending

externally, femoral depressions deep. Antenna with 11 articles (Fig.

223). Prosternum without intercoxal carinae (Fig. 9). Protibia nar-

row, not flanged. Postcoxal line on 1*t abdominal sternum not reach-

ing posterior sternal margin, rounded along margin, apex recurved

1/2 distance to basal sternal margin. Fifth sternum with apex broad-

ly, weakly emarginate; 6 sternum slightly emarginate. Genitalia

with basal lobe 5/8 length of paramere, apex obliquely truncate, with

small, angulate projection on one margin; paramere wide, tapered to

rounded apex in apical 1/2 (Fig. 224); sipho robust, straight api-

cally, apex with membranous area, basal capsule heavily sclerotized,

dark brown, with inner arm long, slender, apically hooked, outer

arm as long as inner, wide, basal border sinuate (Figs. 225, 226).

2 Not known.

Etymology. This specific name is from the Latin dissidens,

meaning differing, referring to the absence of prosternal carinae.

Remarks. Prosterna without carinae are very unusual in

Hyperaspis species. This character and the median, transverse elytral

spot characterize H. dissidens.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 30D

26. Hyperaspis uninotata n. sp.

Type material. Holotype d: Brazil, Guanabara, Rio de Janeiro,

X.1963, M. Alvarenga coll. (USNM).

Description. d, length 3.0 mm, width 2.4 mm; body

rounded, slightly flattened dorsoventrally. Dorsal surface with head

alutaceous, dull: pronotum alutaceous, feebly shiny; elytra shiny.

Color yellow except pronotum with large, black, nearly rectangular

basomedian spot in median 1/2; elytra medially black with broad

yellow border, apex of black area emarginated with yellow on each

side of suture (Fig. 227); pro-, meso-, metasterna black; abdomen

reddish brown with lateral 1/8 and sterna 5-6 yellow. Head punc-

tures fine, nearly invisible in alutaceous sculpture, separated by a di-

ameter or less; pronotal punctures larger than on head, separated by

a diameter or less; elytral punctures larger than pronotal punctures,

separated by less than to twice a diameter. Metasternal punctures

much larger than on elytron, coarse, separated by a diameter medi-

ally, dense, contiguous laterally. Punctures on abdominal sterna 1-4

coarse, sparse medially, becoming fine, dense laterally, sterna 5-6

finely, densely punctured throughout; area on 1% abdominal sternum

inside postcoxal arc shiny, coarsely, sparsely punctured. Clypeal

apex deeply emarginate, clypeus and frons joined at abrupt angle.

Epipleuron flat, feebly grooved internally, not descending externally,

femoral excavations deep. Antenna with 11 articles (Fig. 228). Pros-

ternum with intercoxal carinae narrowly separated at apex, conver-

gent from apex toward base, basally stemmed. Protibia narrow, not

flanged. Postcoxal line on 1*t abdominal sternum reaching posterior

sternal margin, flattened along margin, apex recurved 1/2 distance

to basal sternal margin. Fifth sternum with apex feebly emarginate;

6 sternum feebly emarginate. Genitalia with basal lobe 3/4 as long

as paramere, apex obliquely rounded, with angulate projection on

one margin; paramere wide, tapered from middle to rounded apex

(Fig. 229); sipho slender, apex sinuate, with membranous area, basal

capsule with inner arm robust, short, outer arm wide, longer than

inner arm (Figs. 230, 231).

2 Not known.

Etymology. The specific name is from the Latin unicus, mean-

ing sole or singular, and nota, meaning mark, referring to the single

black elytral macula.

336 R.D. GORDON & C. CANEPARI

Remarks. /MHyperaspis uninotata has a unique dorsal color

pattern because the apical margin of the elytral black area is emar-

ginated with yellow on each side of the suture. Other South Ameri-

can Hyperaspis species may have a similar appearance, but the black

elytral area is differently shaped and d genitalia are different in

each instance.

27. Hyperaspis onerata (Mulsant, 1850)

Cleothera onerata Mulsant, 1850: 552.

Hyperaspis onerata: CROTCH 1874: 230; KORSCHEFSKY 1931: 193; BLACKWELDER 1945:

447; GORDON 1987: 29.

Type locality. Colombia. MNHP (lectotype here designated).

Descripiron-. .6., length 2.3 mm, .width..1.8 mm;. body

form oval, flattened. Dorsal surface with head strongly alutaceous,

dull: pronotum feebly alutaceous shiny: elytron shiny. Color yellow

except pronotum with 2 triangular, black, anteromedian spots nar-

rowly connected to transverse basomedian spot, basomedian spot

narrowly irregular (Fig. 236); elytron with basal margin narrowly

black medially, sutural margin narrowly black posterior to scutel-

lum, widely black on apical declivity, lateral margin narrowly black

from anterolateral angle nearly to suture at apex, joined to black

sutural margin, 1 large, oval, discal spot on sutural margin of each

elytron forming a single, large, round spot, 1 small black, oval spot

posterior to humeral callus, 1 large, irregularly oval spot near lat-

eral margin in apical 1/3 (Figs. 232, 233); pro-, meso-, metasterna

black; abdomen black except outer 1/4 and sternum 5-6 reddish

yellow. Head punctures fine, distinct, separated by less than to

twice a diameter; pronotal punctures larger than on head, sepa-

rated by a diameter or less; elytral punctures larger than pronotal

punctures, separated by less than to twice a diameter. Metasternal

punctures larger than on elytron, very sparse medially, nearly con-

tiguous laterally. Punctures on abdominal sterna 1-4 coarse, dense

medially, becoming dense, fine laterally; sterna 5-6 finely, densely

punctured throughout; area on 1* abdominal sternum inside post-

coxal arc shiny, coarsely, sparsely punctured. Clypeal apex deeply

emarginate, clypeus and frons joined at right angle. Epipleuron

flat, grooved internally, weakly descending externally, femoral de-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI BBY

pression deep. Antenna with 11 articles (Fig. 238). Prosternum

with intercoxal carinae widely separated at apex, tapered toward

base, basally stemmed, reaching basal margin. Protibia narrow, not

flanged. Postcoxal line on 1% abdominal sternum extended to pos-

terior sternal margin, flattened along margin, apex recurved 1/2

distance to basal sternal margin. Fifth sternum with apex broadly,

feebly emarginate; 6" sternum truncate. Genitalia with basal lobe

5/8 length of paramere, strongly, obliquely truncate apically, with

large, acutely angulate projection on one side; paramere wide, ta-

pered to rounded apex in apical 1/2 (Figs. 239, 240); sipho slender,

apically sinuate, with membranous area, basal capsule with inner

arm wide, outer arm shorter than inner arm, basal border emargin-

ate (Figs. 241, 242).

2 Similar to male except head reddish yellow with vertex black.

Beak of basal unit short; connecting duct short (Fig. 243).

Variation. Length 2.0 to 2.7 mm, width 1.9 to 2.2 mm. Dorsal

dark spots vary from black to brown; median pronotal maculation

may have anterior dark area separated from basal dark area (Fig.

237 ); elytron with tendency for lateral spots to coalesce with each

other and the apical elytral margin, size of black elytral spots highly

variable (Figs. 234, 235).

Geographical distribution: Northern South America, Caribbean

Islands.

Specimens examined: 66. Colombia: Cali; Cauca, Puerto Teja-

da; Cartagena; Cundina- marca, Fusagasuga; Cundinamarca, Mel-

gar; Huila, Campoalegre; Magdalena, Santa Marta; Tolima, Ibague;

Turbaco; Valle del Cauca, Andalucia; Valle del Cauca, Buga; Valle

del Cauca, Palmira. Venezuela: Aragua, El Limon; Caracas Valley;

Departamento Federal, Caracas (BM) (CNC) (USNM).

Remarks. AHyperaspis onerata is distinguished from other

Hyperaspis species by the dorsal color pattern. ‘The basal lobe of

the 6 genitalia with a very large, acutely rectangular lateral tooth is

a character shared with only a few other species.

A ® type specimen in the MNHP is designated the lectotype

to stabilize future usage of the name. Lectotype labels are “T'ype/

Coll. Mniszech/onerata, Cleothera, Muls., Colombie, Type”. A fe-

male specimen in the MNHP bearing identical labels is designated

a paralectotype.

338 R.D. GORDON & C. CANEPARI

28. Hyperaspis bicruciata Mulsant, 1850

Hyperaspis bicruciata Mulsant, 1850: 664; CroTcH 1874: 230; KorscHEFsKky 1931:

185; BLACKWELDER 1945: 446; GORDON 1987: 29.

Type locality. “Nouvelle-Grenade” (Colombia, Venezuela).

UMZC (lectotype designated by GORDON 1987).

Description. d, length 2.4 mm, width 1.9 mm; body,

oval, convex. Dorsal surface with head strongly alutaceous, dull;

pronotum shiny; elytron shiny. Color black except head yellow; pro-

notum yellow with basomedian brown spot consisting of narrow,

basal, transverse vitta with median, vaselike projection extended 2/3

distance to anterior pronotal margin; elytron with 3 yellow spots,

large, yellow discal spot connected to large, yellow lateral spot, and

1 large, transverse, apical spot (Figs. 244, 245); antenna, mouthparts,

propleuron, legs reddish yellow; epipleuron reddish brown. Head

punctures fine, hidden in alutaceous sculpture; pronotal punctures

larger than on head, separated by a diameter or less; elytral punc-

tures larger than pronotal punctures, separated by a diameter or less.

Metasternum with punctures larger than on elytron, separated by a

diameter medially, becoming contiguous laterally. Punctures on ab-

dominal sterna larger than on elytra, dense on sterna 1-4, becoming

fine, dense laterally and on sterna 5-6; area on 1" abdominal sternum

inside postcoxal arc shiny, coarsely, sparsely punctate. Clypeal apex

deeply emarginate, clypeus and frons joined at abrupt angle. Epipleu-

ron oblique, internally grooved, descending externally, femoral de-

pressions moderate, not deep. Antenna with 11 articles. Prosternum

with intercoxal carinae narrowly separated at apex, convergent from

apex toward base, basally stemmed, reaching basal margin. Protibia

narrow, not flanged. Postcoxal line on 1" abdominal sternum reach-

ing posterior sternal margin, flattened along margin, apex recurved

1/2 distance to basal sternal margin. Fifth sternum with apex broad-

ly, feebly emarginate; 6 sternum feebly emarginate. Genitalia with

basal lobe 2/3 as long as paramere, sinuate, apex obliquely rounded,

with small, acute projection on one margin; paramere oval, slen-

der, tapered from basal 1/4 to rounded apex; (Fig. 246); sipho long,

robust, apically sinuate with membranous area, basal capsule with

inner arm short, wide, outer arm slightly longer than inner arm,

apically knobbed, basal border weakly emarginate (Figs. 247-248).

2 Not known.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 339

Geographical distribution: Northern South America.

Specimens examined: 1 (lectotype).

Remarks. The resemblance of this species to H. connectens

was commented on by both MuLsanT (1850) and CRroTcH (1874).

Hyperaspis bicruciata is distinguished from H. connectens and other

species by the peculiar, vaselike pronotal spot; elytral color pattern

with median spots broadly connected; postcoxal line flattened along

basal margin; and long, slender basal lobe of the male genitalia.

The lectotype was the only specimen examined.

joannae group

Species with convex body; frons and clypeus joined at abrupt

angle; protibia narrow; epipleural depressions deep; d genitalia with

inner arm of siphonal basal capsule very long, outer arm very short

or nearly absent (Figs. 259, 265), paramere wide or oval.

The basal siphonal capsule with a long inner arm and short to

nonexistent outer arm is diagnostic for this group.

29. Hyperaspis joannae n. sp.

Type material. Holotype d: Colombia, Leticia, Amazonas, 700’,

July 8, 1970, H. & A. Howden (USNM).

Description. 6, length 2.5 mm, width 2.0 mm; body

oval, convex. Dorsal surface with head strongly alutaceous, dull;

pronotum slightly alutaceous, shiny; elytron shiny. Color black ex-

cept head yellow; pronotum yellow with large, black, basomedian

spot, spot widely separated from anterior pronotal margin; elytron

with 4 spots, 1 large, reddish yellow, irregularly oval discal spot,

1 small, triangular spot on anterolateral angle, 1 small, irregularly

oval spot on lateral margin medially, 1 transverse, subapical spot

(Fig. 249, 250); antenna, mouthparts, propleuron, legs yellow; epi-

pleuron reddish yellow; abdomen with median area of sterna 1-4

dark brown, lateral 1/4 and sterna 5-6 reddish yellow. Head punc-

tures not visible in alutaceous sculpture; pronotal punctures fine,

separated by less than to twice a diameter; elytral punctures larg-

er than pronotal punctures, separated by 1 to 2 times a diameter.

340 R.D. GORDON & C. CANEPARI

Metasternal punctures much larger than on elytron, sparse medially,

separated by a diameter laterally. Punctures on abdominal sterna

1-4 larger than on elytra, coarse, sparse medially, becoming fine,

dense laterally; sterna 5-6 finely, densely punctured throughout;

area on 1“ abdominal sternum inside postcoxal arc shiny, sparsely,

coarsely punctured. Clypeal apex deeply emarginate, clypeus and

frons joined at abrupt angle. Epipleuron oblique, grooved internally,

strongly descending externally, femoral excavations deep. Antenna

with 11 articles (Fig. 251, basal article missing). Prosternum with

intercoxal carinae narrowly separated at apex, convergent toward

base, basally stemmed, reaching basal margin. Protibia narrow, not

flanged. Postcoxal line on 1% abdominal sternum reaching posterior

sternal margin, flattened along margin, apex recurved 1/4 distance

to basal sternal margin. Fifth sternum with apex broadly, weakly

emarginate; 6" sternum slightly arcuate. Genitalia with basal lobe

short, 1/2 as long as paramere, wide, apex obliquely truncate, with

large, rounded projection on one margin; paramere wide, tapered to

rounded apex in apical 1/2 (Fig. 252); sipho slender, apex curved

upward, with membranous area, basal capsule, with inner arm long,

slender, apically knobbed, outer arm very short, basal margin emar-

ginate (Figs. 253, 254).

2 Not known.

Etymology. This species is named for JoAnne Gordon, without

whose influence and assistance one of us (RDG) would not be co-

authoring this revision.

Remarks. Characters distinguishing H. joannae are the 4

elytral spots whose size and arrangement are not repeated elsewhere

in this group or in the South American Hyperaspis fauna.

30. Hyperaspis albopunctata Crotch, 1874

Hyperaspis albopunctata Crotch, 1874: 229; KoRSCHEFSKY 1931: 184; BLACKWELDER

1945: 446; GoRDON 1987: 29.

Type locality Brazil, UMZC (holotype, GORDON 1987).

Description. 6, length 2.8 mm, width 2.4 mm; body

oval, convex. Dorsal surface with head strongly alutaceous, dull;

pronotum slightly alutaceous, shiny; elytron shiny. Color black ex-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 341

cept head yellow; pronotum yellow with large, black, basomedian

spot, anterior margin of spot widely separated from anterior prono-

tal margin, narrowly emarginated with yellow apically, elytron with

2 pale yellow spots, 1 very small discal spot, 1 large spot at apex,

apical spot transversely elongate, outer end tapered to acute apex

(Figs. 255, 256); antenna, mouthparts, propleuron, legs yellow; epi-

pleuron reddish brown; abdomen with median area of sterna 1-5

brown, lateral 1/8 and sterna 5-6 yellow. Head punctures fine, sepa-

rated by about a diameter; pronotal punctures larger than on head,

separated by a less than to twice a diameter; elytral punctures larger

than pronotal punctures, separated less than to twice a diameter.

Metasternal punctures much larger than on elytron, sparse medi-

ally, dense, contiguous laterally. Punctures on abdominal sterna 1-4

larger than on elytra, coarse, dense medially, becoming fine, dense

laterally; sterna 5-6 finely, densely punctured throughout; area on

1* abdominal sternum inside postcoxal arc shiny, coarsely, sparsely

punctured. Clypeal apex deeply emarginate, clypeus and frons joined

at abrupt angle. Epipleuron oblique, internally grooved, descending

externally, femoral excavations deep. Antenna with 11 articles (Fig.

257). Prosternum with intercoxal carinae widely separated at apex,

convergent from apex toward base, basally stemmed. Protibia nar-

row, not flanged. Postcoxal line on 1% abdominal sternum reaching

posterior sternal margin, flattened along margin, apex recurved 1/2

distance to basal sternal margin. Fifth sternum with apex broadly,

weakly emarginate; 6 sternum truncate. Genitalia with basal lobe

1/2 as long as paramere, apex obliquely truncate, with rounded

projection on one margin; paramere wide, tapered from middle to

rounded apex; (Fig. 258); sipho slender, long, apex sinuate, curved

upward, basal capsule with inner arm very long, apically knobbed,

outer arm nearly absent, basal margin sinuate (Figs. 259, 260).

2 Similar to 6 except head reddish brown with black vertex; pro-

notum with basomedial black spot extended to apical margin. Geni-

talia with beak of basal unit short, connecting duct short (Fig. 261).

Variation. Length 2.6 to 3.0 mm, width 2.4 to 2.5 mm. Prono-

tum of d with basomedian spot apically emarginate or not.

Geographical distribution: Brazil.

Specimens examined: 16. Brazil: Minas Gerais, Monte Verde;

Sao Paulo, Campos Jordào; Sào Paulo, Cantereira; Guarulhos, Sitio

Bananal (USNM).

342 R.D. GORDON & C. CANEPARI

Remarks. The dorsal appearance is rather elegant and dis-

tinctive within the genus.

31. Hyperaspis apicaspis n. sp.

Type material. Holotype d: Brazil, 56127 (illegible label), S.

Paulo (Sao Paulo), Fry Coll. 1905.100., Hyperaspis subapicalis Crotch

(handwritten), not subapicalis acc. to Fry coll. type. det. R.G. Booth

2003 (BM).

Description. 6, length 2.0 mm, width 1.5 mm; body

oval, convex. Dorsal surface with head strongly alutaceous, dull;

pronotum alutaceous, dull; elytron shiny. Color black except head

yellow; pronotum with lateral 1/4 yellow; elytron with 1 round spot

at apex, spot widely separated from sutural margin (Figs. 262, 263);

antenna, mouthparts, propleuron, legs reddish yellow; abdomen

with median area of sterna 1-4 dark brown, lateral 1/4 and sterna

5-6 reddish yellow. Head punctures fine, separated by 1 to 2 times

a diameter; pronotal punctures larger than on head, separated by

less than to twice a diameter; elytral punctures larger than pronotal

punctures, separated by a diameter or less. Metasternal punctures

larger than on elytron, sparse medially, separated by less than a di-

ameter laterally. Punctures on abdominal sterna 1-4 smaller than

on elytra, sparse medially, becoming fine, dense laterally; sterna 5-6

finely, densely punctured throughout; area on 1% abdominal sternum

inside postcoxal arc alutaceous, sparsely, coarsely punctured. Clypeal

apex deeply emarginate, clypeus and frons joined at abrupt angle.

Epipleuron flat, slightly grooved internally, not descending external-

ly, femoral excavations deep. Antenna with 11 articles. Prosternum

with intercoxal carinae widely separated at apex, convergent toward

base, basally stemmed, reaching basal margin. Protibia narrow, not

flanged. Postcoxal line on 1*t abdominal sternum reaching posterior

sternal margin, flattened along margin, apex recurved 1/2 distance

to basal sternal margin. Fifth sternum with apex broadly, weakly

emarginate; 6° sternum truncate. Genitalia with basal lobe short,

1/2 as long as paramere, apex obliquely rounded, with small, angu-

late projection on one margin; paramere wide, tapered to rounded

apex in apical 1/2 (Fig. 264); sipho slender, s-shaped apically, apex

with membranous area, basal capsule with inner arm long, slender,

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 343

apically knobbed, outer arm very short, apically knobbed, basal bor-

der emarginate (Figs. 265, 266).

2. Not known.

Etymology. The specific name is a combination of the Latin

apiculus, meaning apex or apical, and -aspis, the ending of Hyper-

aspis.

Remarks. AHAyperaspis apicaspis has the same dorsal color pat-

tern as 2 other species of this group. See remarks under H. delicata.

32. Hyperaspis delicata Almeida & Vitorino, 1997

Hyperaspis delicata Almeida & Vitorino, 1997: 213-216.

Type locality. Brazil, Parana, Piraquara (Universidade Federal

do Parana).

Description. 6, length 2.0 mm, width 1.4 mm; body

elongate, oval, convex. Dorsal surface with head strongly alutaceous,

dull: pronotum alutaceous, feebly shiny; elytra slightly alutaceous,

shiny. Color black except head yellow; pronotum with lateral 1/4

yellow, median black area laterally emarginated with yellow; elytron

with small, yellow, subapical spot near posterolateral angle (Fig.

267, 268); antenna, mouthparts, legs reddish yellow except metafe-

mur yellowish brown; propleuron yellow; epipleuron dark reddish

brown; abdomen brown with lateral 3/8 reddish yellow. Head punc-

tures fine, hidden in alutaceous sculpture; pronotal punctures larger

than on head, separated by less than to twice a diameter; elytral

punctures larger than pronotal punctures, separated by 1 to 2 times

a diameter. Metasternal punctures larger than on elytron, sparse

medially, separated by a diameter or less laterally. Punctures on ab-

dominal sterna much larger than on elytra, dense medially, becoming

fine, dense laterally, sterna 5-6 finely densely punctured throughout;

area on 1* abdominal sternum inside postcoxal arc alutaceous, fine-

ly, sparsely punctured. Clypeal apex deeply emarginate, clypeus and

frons joined at abrupt angle. Epipleuron flat, feebly grooved inter-

nally, not descending externally, femoral depressions deep. Antenna

with 11 articles (Fig. 269). Protibia narrow, not flanged. Prosternum

with intercoxal carinae widely separated at apex, convergent toward

base, basally stemmed, reaching basal margin. Postcoxal line on 1“

344 R.D. GORDON & C. CANEPARI

abdominal sternum not reaching posterior sternal margin, flattened

along margin, apex recurved 3/8 distance to basal sternal margin.

Fifth sternum with apex broadly, distinctly emarginate; 6 sternum

feebly emarginate. Genitalia with basal lobe 2/3 as long as paramere,

narrowed from base to apex, apex slightly, narrowly rounded, with-

out projection on one margin; paramere wide, tapered to rounded

apex in apical 1/2 (Fig. 270); sipho long, slender, apically sinuate,

with apical membranous area, basal capsule with inner arm long,

narrow, irregularly shaped, outer arm nearly absent, basal border

slightly emarginate (Figs. 271, 272).

? Similar to d except head black.

Variation. Length 2.56 to 3.20, width 1.86 to 2.44 mm.

Geographical distribution: Southeastern Brazil.

Specimens examined: 12. Brazil: Parana, Piraquara; Parana,

Colombo; Ribiero Pires; Sao Paulo, Sao Paulo (CAS) (DZUP)

(USNM).

Remarks. Of the 3 species having black elytra with a sin-

gle, yellow, subapical spot on each elytron, and male genitalia with

inner arm of basal capsule extremely long, only H. mimica is distin-

guishable by an external character. Hyperaspis apicaspis and H. deli-

cata are separable by differences in the basal lobe of the d genitalia.

The lobe is short, wide, and with a lateral angulation in the former

species, and long, tapered from base to apex, and without a lateral

projection on one side in the latter. Hyperaspis mimica also has 3

genitalia with a distinctive basal lobe, but is externally distinguished

by prosternal carinae that are narrowly separated at the apex, nearly

parallel toward base, and do not reach basal sternal margin.

33. Hyperaspis mimica n. sp.

Type material. Holotype d: Country not on labels, 1% label illeg-

ible, R. Korschefsky cum typ. comp. (pink label), Hyperaspis suba-

picalis Cr. 1926 det. R. Korschefsky, Korschefsky Collection 1952

(USNM).

Description. 6, length 2.4 mm, width 1.8 mm; body

elongate, oval, convex. Dorsal surface with head strongly alutaceous,

dull: pronotum alutaceous, feebly shiny; elytra slightly alutaceous,

shiny. Color black except head yellow; pronotum with lateral 1/4

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 345

yellow, median black area laterally emarginated with yellow; elytron

with small, yellow, subapical spot near posterolateral angle (Fig.

273); antenna, mouthparts, propleuron, proleg yellow; mesoleg red-

dish brown except apical 1/4 of femur brown; metaleg, epipleuron

brown; abdomen brown with lateral 3/8 reddish brown. Head punc-

tures fine, hidden in alutaceous sculpture; pronotal punctures larger

than on head, separated by a diameter or less; elytral punctures larger

than pronotal punctures, separated by less than to twice a diameter.

Metasternal punctures larger than on elytron, sparse medially, sepa-

rated by a diameter or less laterally. Punctures on abdominal sterna

slightly larger than on elytra, dense medially, becoming fine, dense

laterally, sterna 5-6 finely densely punctured throughout; area on 1*

abdominal sternum inside postcoxal arc alutaceous, coarsely, densely

punctured. Clypeal apex deeply emarginate, clypeus and frons joined

at abrupt angle. Epipleuron flat, feebly grooved internally, slightly

descending externally, femoral depressions deep. Antenna with 11

articles. Protibia narrow, not flanged. Prosternum with intercoxal

carinae narrowly separated at apex, convergent toward base, extend-

ed 3/4 distance to basal sternal margin, not stemmed. Postcoxal line

on 1" abdominal sternum reaching posterior sternal margin, flat-

tened along margin, apex abruptly angled 7/8 distance to basal ster-

nal margin. Fifth sternum with apex broadly, feebly emarginate; 6™

sternum distinctly emarginate. Genitalia with basal lobe about as

long as paramere, narrowed in apical 1/2 to slightly rounded apex,

with slight, angulate projection on one margin; paramere long, slen-

der, tapered to rounded apex in apical 1/2 (Fig. 275); sipho long,

slender, apically sinuate, with apical membranous area, basal capsule

with inner arm long, slender, regular in shape, outer arm very short,

basal border slightly emarginate (Fig. 276, 277).

2 Not known.

Etymology. The species name is from the Latin mimus, meaning

imitate, referring to resemblance in color pattern to H. apicalis and

H. delicata.

Remarks. In addition to characters discussed under H.

delicata, H. mimica is distinguished by the larger size; comparati-

vely small, dense, dorsal and ventral punctures; and postcoxal line

on the basal abdominal sternum apically extended nearly to basal

sternal margin.

346 R.D. GORDON & C. CANEPARI

Holotype specimen labels do not indicate country of origin,

which anyway must be in Central or South America.

34. Hyperaspis laterimacula n. sp.

Type material. Holotype d: O. Bolivien (eastern Bolivia), Prov.

Sara, J. Steinbach S.V. (ZMHB).

Description. dé, length 1.8 mm, width 1.6 mm; body

round, convex. Dorsal surface with head strongly alutaceous, dull;

pronotum slightly alutaceous, shiny; elytron shiny. Color brown ex-

cept head yellow; pronotum yellow with basomedian brown spot,

anterior margin broadly yellow, lateral 1/4 yellow, basomedian spot

broadly emarginated laterally; elytron with 5 yellow spots, 1 small,

triangular spot laterad of scutellum, 1, small, round, median discal

spot, and 3 large, oval, spots on lateral margin, 1 at humeral angle,

1 at middle, and 1 subapical (Figs. 278, 279); antenna, mouthparts,

legs reddish yellow; propleuron, epipleuron yellow; pro-, meso-,

metasterna dark reddish brown; abdomen with median area of sterna

1-4 reddish yellow, lateral 1/4 and sterna 5-6 yellow. Head punctures

fine, hidden in alutaceous sculpture; pronotal punctures larger than

on head, separated by a diameter or less; elytral punctures larger

than pronotal punctures, separated by less than to twice a diameter.

Metasternal punctures larger than on elytron, separated by a diam-

eter medially, nearly contiguous laterally. Punctures on abdominal

sterna 1-4 larger than on elytra, dense medially, becoming fine, dense

laterally; sterna 5-6 finely, densely punctured throughout; area on

1* abdominal sternum inside postcoxal arc shiny, sparsely, coarsely

punctured. Clypeal apex deeply emarginate, clypeus and frons joined

at abrupt angle. Epipleuron oblique, grooved internally, slightly de-

scending externally, femoral excavations deep. Antenna with 11 arti-

cles. Prosternum with intercoxal carinae narrowly separated at apex,

convergent toward base, basally stemmed, reaching basal margin.

Protibia narrow, not flanged. Postcoxal line on 1* abdominal ster-

num not reaching posterior sternal margin, slightly flattened along

margin, apex recurved 1/4 distance to basal sternal margin. Fifth

sternum with apex broadly, weakly emarginate; 6° sternum trun-

cate. Genitalia with basal lobe short, 1/2 as long as paramere, apex

obliquely rounded, with large, angulate projection on one margin;

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 347

paramere wide, tapered to rounded apex in apical 1/4 (Fig. 280);

sipho long, slender, curved upward apically, apex with membra-

nous area, basal capsule with inner arm long, slender, outer arm

very short, nearly absent, basal border widely emarginate (Fig. 281).

2 Not known.

Etymology. The specific name is a combination of the Latin

latus, meaning side, and macula, meaning spot, referring to the 3

large, lateral spots on each elytron.

Remarks. Hyperaspis laterimacula has a siphonal capsule of

the same type as the 3 preceding species, but is otherwise unlike

them. It is recognized by a combination of small size, round body

form, and 5 yellow spots on each elytron. In its elytral spots, H. la-

terimacula resembles H. vredenburgi, but the elytral spots of the lat-

ter species are much larger, and male genitalia differ significantly.

howdeni group

Hyperaspis species having 11-articled antennae; body flattened

or convex; frons and clypeus joined at abrupt angle; protibia nar-

row; d genitalia with paramere very slender, slightly widened in

basal 1/4, tapered to narrowly rounded apex (Fig. 283).

‘The slender, tapered paramere is diagnostic for this group.

35. Hyperaspis howdeni n. sp.

Type material. Holotype 6: Colombia, Magd. (Magdalena),

Parque Tayrona, 21 mi. E. Sta. Marta, V-13-1973, Howden &

Campbell (CNC).

Description. 6, length 2.1 mm, width 1.3 mm; body

oval, convex. Dorsal surface with head strongly alutaceous, dull;

pronotum slightly alutaceous, weakly shiny; elytron shiny. Color

black except head yellow; pronotum yellow with large, black, rect-

angular basomedian spot, spot widely separated from anterior pro-

notal margin; elytron with large, irregularly rounded, discal spot

(Fig. 282); antenna, mouthparts, propleuron yellow; legs reddish

yellow; abdomen with median area of sterna 1-5 dark brown, lat-

eral 1/4 and sternum 6 reddish yellow. Head punctures fine, hidden

348 R.D. GORDON & C. CANEPARI

in alutaceous sculpture, separated by a diameter or less; pronotal

punctures larger than on head, separated by a diameter or less; ely-

tral punctures larger than pronotal punctures, separated by less than

to twice a diameter. Metasternal punctures larger than on elytron,

sparse medially, becoming nearly contiguous laterally. Punctures on

abdominal sterna 1-4 as large as elytral punctures, dense medially,

becoming fine, dense laterally; sterna 5-6 finely, densely punctured

throughout; area on 1% abdominal sternum inside postcoxal arc

shiny, coarsely, sparsely punctured. Clypeal apex deeply emargin-

ate, clypeus and frons joined at abrupt angle. Epipleuron oblique,

grooved internally, descending externally, femoral depressions deep.

Antenna with 11 articles. Prosternum with intercoxal carinae nar-

rowly separated at apex, convergent basally, stemmed, reaching bas-

al pronotal margin. Protibia narrow, not flanged. Postcoxal line on

1" abdominal sternum not reaching posterior sternal margin, slightly

flattened along margin, apex recurved 1/2 distance to basal sternal

margin. Fifth sternum with apex broadly, weakly emarginate; 6°

sternum slightly emarginate. Genitalia with basal lobe as long as

paramere, apex obliquely truncate, with distinct, angulate projection

on one margin; paramere very slender, slightly widened in basal 1/4,

tapered to narrowly rounded apex (Fig. 283); sipho robust, straight

apically, apex with membranous area, basal capsule with inner and

outer arms short , wide (Figs. 284, 285).

2 Not known.

Etymology. The species is named for Henry Howden, one of

the type collectors, and esteemed entomological colleague.

Remarks. AHAyperaspis species with a single, discal, elytral

spot are uncommon; in addition, H. howdeni is distinguished by the

large, irregular shape of the discal spot and particularly by the very

slender paramere of the male genitalia characteristic of this group.

36. Hyperaspis chocoi n. sp.

Type material. Holotype 6: Colombia, Chocé, Istmina,

21.VIIL.40, LM Murillo No. 5342 (USNM).

Description. dé, length 2.3 mm, width 1.8 mm; body

oval, slightly flattened, widest posterior to middle. Dorsal surface

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 349

with head weakly alutaceous, shiny: pronotum and elytron shiny.

Color black except head yellow; pronotum yellow with large, baso-

median spot not extended to anterior margin, elytron with 1 large,

long, yellow spot medially from near base to apical declivity, outer

border of spot emarginate around humeral callus (Fig. 286); antenna,

mouthparts, propleuron yellow; legs yellow except basal 1/4 of pro-

and mesofemura yellowish brown, basal 2/3 of metafemur brown;

abdomen with median area of sterna 1-5 reddish brown, lateral 1/8

and 6' sternum yellow. Head punctures fine, separated by less than

a diameter; pronotal punctures larger than on head, separated by

less than to twice a diameter; elytral punctures larger than pronotal

punctures, separated by less than to twice a diameter. Metasternal

punctures much larger than on elytron, sparse medially, dense, con-

tiguous laterally. Punctures on abdominal sterna 1-4 larger than on

elytra, coarse, sparse medially, becoming fine, dense laterally; ster-

na 5-6 finely, densely punctured throughout; area on 1% abdominal

sternum inside postcoxal arc shiny, nearly impunctate. Clypeal apex

deeply emarginate, clypeus and frons joined at abrupt angle. Epi-

pleuron oblique, medially grooved, descending externally, femoral

excavations deep. Antenna with 11 articles (Fig. 287). Prosternum

with intercoxal carinae narrowly separated at apex, convergent from

apex toward base, basally stemmed. Protibia narrow, not flanged.

Postcoxal line on 1* abdominal sternum reaching posterior sternal

margin, flattened along margin, apex recurved 1/2 distance to basal

sternal margin. Fifth sternum with apex broadly, distinctly emargin-

ate; 6" sternum feebly emarginate. Genitalia with basal lobe 3/4 as

long as paramere, wide, apex truncate, with angulate projection on

one margin; paramere wide, tapered from middle to rounded apex;

(Figs. 288, 289); sipho slender, apex sinuate, with membranous area,

basal capsule, with inner arm narrow, outer arm wide, as long as

inner arm (Figs. 290, 291).

2 Not known.

Etymology. This species is named for the Department of Choco,

where the type series was collected.

Remarks. ‘The large, elongate discal spot on each elytron is

an excellent recognition character. The holotype was collected at the

same place on the same date as the female holotype of Hyperaspis

istmina, a species nearly identical in size and dorsal color pattern.

350 R.D. GORDON & C. CANEPARI

37. Hyperaspis atra n. sp.

Type material. Holotype dé: Brazil, Rio de Janeiro, Murundu

Campos, VIII, 1978, M. Alvarenga (CMP).

Description. d, length 1.8 mm, width 1.5 mm; body

oval, convex. Dorsal surface with head alutaceous, weakly shiny:

pronotum shiny; elytra shiny. Color black except head yellow; pro-

notum with apical 1/5 and lateral 1/6 yellow, elytron with 1 small,

elongate yellow spot on anterolateral angle outside of humeral

callus (Figs. 292, 293); antenna, mouthparts, propleuron yellow;

epipleuron brownish yellow; pro-, mesolegs reddish yellow, basal

1/2 of femora brown; metafemur brown except apical 1/8 yellow;

abdomen brown with lateral 1/4 reddish yellow. Head punctures

fine, separated by less than to twice a diameter; pronotal punc-

tures larger than on head, separated by less than a diameter; elytral

punctures larger than pronotal punctures, separated by less than to

a diameter. Metasternal punctures slightly larger than on elytron,

separated by a diameter medially, dense, contiguous laterally.

Punctures on abdominal sterna 1-4 larger than on elytra, dense

medially, becoming fine, dense laterally, sterna 5-6 finely, densely

punctured throughout; area on 1** abdominal sternum inside post-

coxal arc shiny, coarsely, densely punctured. Clypeal apex deeply

emarginate, clypeus and frons joined at abrupt angle. Epipleuron

slightly oblique, internally grooved, descending externally, femoral

excavations deep. Antenna with 11 articles. Protibia narrow, not

flanged. Prosternum with intercoxal carinae narrowly separated at

apex, convergent from apex toward base, basally stemmed, reaching

basal margin. Postcoxal line on 1% abdominal sternum not reaching

posterior sternal margin, rounded along margin, apex recurved 1/2

distance to basal sternal margin. Fifth sternum with apex broadly,

weakly emarginate; 6" sternum broadly, distinctly truncate. Geni-

talia with basal lobe slightly shorter than paramere, apex acute,

with distinct, angulate projection on one margin; paramere wide in

basal 1/2, slender, tapered to abruptly rounded apex in apical 1/2

(Fig. 294); sipho long, with apical membranous area, basal capsule

with inner arm slender, outer arm slender, slightly shorter than in-

ner arm, apically knobbed, basal border deeply emarginate ( Figs.

295.1290):

2 Not known.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 351

Etymology. The species name is the Latin word atra, meaning

black, referring to the mostly black dorsum.

Remarks. The black elytron with a single, small, yellow,

humeral spot will distinguish H. atra.

38. Hyperaspis recordata Mulsant, 1850

Hyperapis recordata Mulsant, 1850: 679.

Hyperaspis recordata: CROTCH 1874: 228 (as synonym of Cleothera pavida Mulsant,

1850).

Hyperaspis pavida ab. recordata: KORSCHEFSKY 1931: 193; BLACKWELDER 1945: 448.

Type locality. Brazil, UMZC (lectotype designated by GORDON

1057).

Description. ®, length 2.6 mm, width 2.0 mm; body oval,

convex. Dorsal surface with head slightly alutaceous, shiny: prono-

tum shiny; elytra shiny. Color black except pronotum with lateral

margin broadly and anterior margin narrowly yellow; elytron with 2

yellow spots, 1 small, round spot with basal margin truncated near

scutellum, 1 small, transversely oval, subapical spot at posterolat-

eral angle of elytron (Figs. 297, 298); antenna, narrow outer margin

of propleuron yellow; mouthparts dark brown; legs reddish yellow

except basal 2/3 of femur dark brown; abdominal sterna 1-5 dark

brown with lateral 1/4 and sternum 6 reddish yellow. Head punctures

fine, separated by a diameter or less; pronotal punctures larger than

on head, separated by less than a diameter; elytral punctures larger

than pronotal punctures, separated by a diameter or less. Metaster-

nal punctures larger than on elytron, separated by a diameter or less

medially, becoming dense, contiguous laterally. Punctures on ab-

dominal sterna larger than on elytra, dense medially, becoming fine,

dense laterally, sterna 5-6 finely densely punctured throughout; area

on 1* abdominal sternum inside postcoxal arc shiny, coarsely, dense-

ly punctured. Clypeal apex emarginate, clypeus and frons abruptly

Joined. Epipleuron slightly oblique, grooved internally, slightly de-

scending externally, femoral depressions deep. Antenna with 11 arti-

cles. Protibia not flanged. Prosternum with intercoxal carinae widely

separated at apex, convergent toward base, stemmed, reaching basal

margin. Postcoxal line on 1* abdominal sternum not reaching pos-

terior sternal margin, slightly flattened along margin, apex recurved

352 R.D. GORDON & C. CANEPARI

3/8 distance to basal sternal margin. Fifth sternum with apex feebly

emarginate. Genitalia with beak of basal unit longer than unit, very

wide, curved in apical 1/2; connecting duct short (Figs. 302, 302).

d: Similar to 2 except apices of abdominal sterna 5 and 6 dis-

tinctly emarginate. d genitalia with basal lobe slightly shorter than

paramere, wide, apex broadly rounded, with large, distinct, angulate

projection on one margin; paramere wide in basal 1/2, slender, ta-

pered to abruptly rounded apex in apical 1/2 (Fig. 299); sipho long,

with apical membranous area, basal capsule with inner arm slender,

outer arm slender, slightly shorter than inner arm, apically knobbed,

basal border moderately emarginate (Figs. 300, 301).

Variation. Length 2.4 to 2.6 mm, width 1.9 to 2.0 mm.

Geographical distribution: Brazil.

Specimens examined: 2. Brazil: Pernambuco (lectotype); Rio de

Janeiro (CMP).

Remarks. ‘There are several black species with two pale

spots on each elytron, but this is the only one with one of the spots

located next to the scutellum.

CROTCH (1874) considered H. recordata a synonym of the Mexi-

can H. pavida Mulsant, and his treatment was followed by both

KORSCHEFSKY (1931) and BLACKWELDER (1945). Examination of many

hyperaspidinae species indicates very little overlap between Mexican

and South American species; therefore, we consider H. recordata

a valid species, probably restricted to northeastern South America.

Species not assigned to a group

Hyperaspis species with 11-articled antennae, and male genitalia

differing from any established group and from each other. Hence,

they are placed in a “group” of their own.

39. Hyperaspis eupaleoides Crotch, 1874

Hyperaspis eupaleoides Crotch, 1874: 224; KorscHEFsky 1931: 188; BLACKWELDER

1945: 447; GoRDON 1987: 28.

Hyperaspis graphica Weise, 1902: 174; KORSCHEFSKY 1931: 190; BLACKWELDER 1945:

447. (n. syn.)

SOUTH AMERCAN-COCCINELLIDAE, XI: HYPERASPIDINI 353

Type locality: of eupaleoides, Brazil, Minas Gerais (UMZC, lec-

totype designated by GORDON 1987); of graphica, Brazil (ZMHB).

Description. 6, length 3.3 mm, width 2.7 mm; body

form oval, convex. Dorsal surface with head strongly alutaceous,

dull: pronotum shiny: elytron shiny. Color yellow except pronotum

with 4 triangular brown spots, 2 spots in anterior 1/2 narrowly con-

nected medially, 2 basal spots on each side of middle; elytron with

narrow, brown sutural margin and 4 brown spots, 1 elongate, api-

cally tapered spot in basal 1/2 near suture from near base to mid-

dle, 1 crescent shaped spot laterally from base across humeral callus

nearly to middle, narrowly connected to irregularly shaped spot in

apical 1/2, 1 small, free spot near lateral margin in apical 1/2, and

1 very small, subapical spot near sutural margin (Fig. 304); pro-,

meso-, metasterna yellowish brown; abdomen yellowish brown ex-

cept outer 1/4 and sterna 5-6 yellow. Head punctures fine, hidden in

alutaceous sculpture; pronotal punctures larger than on head, sepa-

rated by a diameter or less; elytral punctures larger than pronotal

punctures, separated by a diameter or less. Metasternal punctures

larger than on elytron, separated by a diameter medially, nearly con-

tiguous laterally. Punctures on abdominal sterna 1-4 coarse, dense

medially, becoming dense, fine laterally; sterna 5-6 finely, densely

punctured throughout; area on 1“ abdominal sternum inside post-

coxal arc shiny, coarsely, densely punctured. Clypeal apex shallowly

emarginate, clypeus and frons abruptly joined. Epipleuron oblique,

grooved internally, strongly descending externally, femoral depres-

sion deep. Protibia not flanged. Prosternum with intercoxal carinae

widely separated at apex, parallel to basal 1/4, then convergent, not

basally stemmed, reaching basal margin. Protibia widened apically.

Postcoxal line on 1%“ abdominal sternum extended to posterior ster-

nal margin, flattened along margin, apex recurved 3/8 distance to

basal sternal margin. Fifth sternum with apex broadly, feebly emar-

ginate; 6" sternum emarginate. Genitalia with basal lobe 3/4 length

of paramere, apex narrowly, obliquely rounded, with large, weakly

angulate projection on one side; paramere wide, tapered to rounded

apex in apical 1/2 (Fig. 305); sipho slender, curved upward at apex,

with membranous area, basal capsule with inner arm long, narrow,

outer arm wide, shorter than inner arm, basal border emarginate

(Figs. 306, 307).

354 R.D. GORDON & C. CANEPARI

2. Not known.

Variation. Length 3.0 to 3.3 mm, width 2.6 to 2.7 mm.

Geographical distribution: Brazil.

Specimens examined: 4. Brazil: “Brasil”; Minas Gerais; Rio de

Janeiro (BM) (ZMHB).

Remarks. AHAyperaspis eupaleoides has a dorsal color pattern

unlike that of any known Hyperaspis species, but similar to certain

species of Cyra, Eupalea Mulsant, and Psyllobora Chevrolat.

The holotype of H. graphica is identical to the lectotype of

H. eupaleoides; therefore, these are considered synonyms with MH.

graphica the junior. The holotype of the latter is labeled “Brazil

(green paper)/graphica m/Syntypus Hyperaspis graphica Weise 1902,

labelled by MNHUB 2004 (red paper)”.

40. Hyperaspis mariposa n. sp.

Type material. Ecuador (illegible name) (ZMHB).

Description. 6, length 3.0 mm, width 2.5 mm; body

form rounded, convex. Dorsal surface with head alutaceous, feebly

shiny: pronotum shiny: elytron shiny. Color yellow except pronotum

with dark brown, transverse, basomedian vitta with 2 slender, dark

brown projections, 1 on each side of middle, extended anteriorly 1/2

distance to apical margin; elytron with irregular discal spot which,

taken in conjunction with spot on opposite elytron, forms butterfly

outline (Fig. 308); pro-, meso-, metasterna brownish black; abdomen

reddish brown except outer 1/4 and sterna 4-6 yellow. Head punc-

tures fine, separated by a diameter or less; pronotal punctures larger

than on head, separated by a diameter or less; elytral punctures larger

than pronotal punctures, separated by a diameter or less. Metasternal

punctures larger than on elytron, separated by a diameter medially,

nearly contiguous laterally. Punctures on abdominal sterna 1-4 as

large as on elytron, sparse medially, becoming dense, fine laterally;

sterna 5-6 finely, densely punctured throughout; area on 1% abdomi-

nal sternum inside postcoxal arc shiny, coarsely, densely punctured.

Clypeal apex emarginate, clypeus and frons abruptly joined. Epi-

pleuron slightly oblique, weakly grooved internally, weakly descend-

ing externally, femoral depressions deep. Prosternum with intercox-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 355

al carinae narrowly separated at apex, convergent toward prosternal

base, basally stemmed, reaching basal margin. Protibia not flanged.

Postcoxal line on 1° abdominal sternum extended to posterior sternal

margin, flattened along margin, apex recurved 1/2 distance to basal

sternal margin. Fifth sternum with apex broadly, distinctly emargin-

ate; 6" sternum feebly emarginate. Genitalia with basal lobe about

as long as paramere, very slightly asymmetrical, slender, apex not

oblique, rounded, with indistinct angulate projection on one side;

paramere slender, tapered to rounded apex in apical 1/2 (Fig. 309);

sipho short, robust, straight apically, apex with membranous area,

basal capsule with inner arm short, wide, irregular in outline, outer

arm narrower and shorter than inner arm (Figs. 310, 311).

2 Not known.

Specimens examined. 1 (holotype).

Etymology. The specific name means butterfly in Spanish, re-

ferring to the butterfly-shaped elytral macula.

Remarks. ‘The large size, mostly yellow dorsum, and accu-

rate depiction of a butterfly on the elytra distinguish this species.

41. Hyperaspis simlaensis n. sp.

Type material. Holotype d: Trinidad, Pt. of Spain (Port of

Spain), from H. Caracicolo, Hyperaspis sp. (USNM). Allotype ®:

Trinidad, Montserrat, July 4.05, Aug. Busck Collector (USNM).

Paratypes, 2. Trinidad, 1, same data as allotype except date June,

30.5 (USNM); 1, Trinidad, 250 m, Simla nr. Arima, 25.XI-2

X1II.1977, W.R.M.Mason (CNC).

Other specimen: 1. Venezuela, Tiara, m 1200 (Edo. Aragua), 21

V 1967, Bordon leg. (CCM).

Description. 6, length 2.0 mm, width 1.6 mm; body

oval, convex. Dorsal surface with head alutaceous, weakly shiny:

pronotum shiny; elytra shiny. Color dark brown except head yel-

low; pronotum with narrow anterior margin and lateral 1/4 yellow,

median brown area triangularly emarginated apically; elytron with 4

yellow spots, 1 triangular scutellar spot at base, 1 triangular discal

spot near suture, 1 transversely oval spot on lateral margin laterad

of scutellar spot, 1 comma shaped, subapical spot on apical decliv-

356 R.D. GORDON & C. CANEPARI

ity (Figs. 312, 313); antenna, mouthparts, propleuron, legs yellow;

epipleuron yellowish red; prosternum mostly reddish yellow. Head

punctures fine, separated by a diameter or less; pronotal punctures

larger than on head, separated by less than to twice a diameter; ely-

tral punctures slightly larger than pronotal punctures, separated by

less than to twice a diameter. Metasternal punctures larger than on

elytron, sparse medially, dense, nearly contiguous laterally. Punc-

tures on abdominal sterna larger than on elytra, dense medially, be-

coming fine, dense laterally, sterna 5-6 finely, densely punctured

throughout; area on 1* abdominal sternum inside postcoxal arc shiny,

coarsely, sparsely punctured. Clypeal apex emarginate, clypeus and

frons abruptly joined. Epipleuron slightly oblique, grooved internal-

ly, slightly descending externally, femoral depressions deep. Antenna

with 11 articles (Fig. 314). Protibia flanged. Prosternum with inter-

coxal carinae narrowly separated at apex, convergent nearly to base,

basal stem short, reaching basal margin. Abdominal sterna 2-5 with

dense fluting in median 1/3. Postcoxal line on 1% abdominal sternum

reaching posterior sternal margin, flattened along margin, apex re-

curved 1/2 distance to basal sternal margin. Fifth sternum with apex

broadly, distinctly emarginate; 6 sternum truncate. Genitalia with

basal lobe about as long as paramere, apex narrowly rounded on one

side, with small, angulate projection on one margin; paramere oval,

tapered from basal 1/2 to rounded apex (Fig. 315); sipho slender,

apically straight, with apical membranous area, basal capsule with

inner arm long, slender, outer arm slender, as long as inner arm,

apically knobbed, basal border broadly emarginate (Figs. 316, 317).

2 Similar to d except head with vertex reddish brown, frons

and clypeus reddish yellow; apical pronotal margin entirely dark

brown, lateral 1/5 yellow. Genitalia with beak of basal unit long;

connecting duct short (Fig. 318).

Variation. Length 2.0 to 2.3, width 1.6 to 1.8. Elytron on one

paratype with trace of small, reddish yellow spot at anterolateral

angle.

Etymology. The species is named for a paratype locality.

Remarks. This is a handsome, highly polished, convex spe-

cies distinguished by the presence of 4 yellow elytral spots on a red-

dish brown background, and the unique fluting on abdominal sterna

2-5. The comma-shaped apical spot on each elytron is formed by 2

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 357

joined spots, and specimens might be found with these spots distinctly

separated. Also, the suggestion of a subhumeral spot on the anterolat-

eral angle might be counted as a spot when distinct. If so, each elytron

would have 5 spots and the species would not key out correctly.

42. Hyperaspis lindae n. sp.

Type material. Holotype d: Colombia, Cld (Caldas), Villamaria,

alt. 2000 m, 12.11.41, Murillo No 5465 (USNM).

Description. 6, length 2.6 mm, width 2.0 mm; body

oval, convex. Dorsal surface with head alutaceous, weakly shiny;

pronotum shiny; elytron shiny. Color yellow except pronotum with

a large, black, median vitta on each side of middle, vitta narrowly

separated, not reaching apical margin; elytron red with basal and

apical margins narrowly black, 1 short, wide, black vitta extended

from base past humeral callus, 1 small, round, black spot medi-

ally just anterior to apical declivity, 1 small, oval, black spot on

sutural margin in basal 1/2 (Figs. 319, 320); venter of head, most of

propleuron, pro-, meso-, metasterna black; abdomen with median

area of sterna 1-5 black, lateral 1/8 and sternum 6 reddish brown

Head punctures fine, separated by less than to twice diameter; pro-

notal punctures larger than on head, separated by less than to twice

a diameter; elytral punctures larger than pronotal punctures, sepa-

rated by less than to twice a diameter. Metasternal punctures much

larger than on pronotum, separated by about a diameter medially,

less than a diameter laterally. Punctures on abdominal sterna 1-4 as

large as pronotal punctures, sparse medially, becoming fine, dense

laterally; sterna 5-6 finely, densely punctured throughout; area on 1%

abdominal sternum inside postcoxal arc alutaceous, coarsely, sparse-

ly punctured. Clypeal apex emarginate, clypeus and frons abruptly

joined. Epipleuron oblique, internally grooved, descending exter-

nally, femoral excavations deep. Antenna with 11 articles (Fig. 321).

Protibia not flanged. Prosternum with intercoxal carinae widely sep-

arated at apex, convergent near base, basal stem short, reaching bas-

al pronotal margin. Postcoxal line on 1% abdominal sternum reach-

ing posterior sternal margin, slightly flattened along margin, apex

recurved 1/2 distance to basal sternal margin. Fifth sternum with

apex broadly, distinctly emarginate; 6" sternum slightly emarginate.

358 R.D. GORDON & C. CANEPARI

Genitalia with basal lobe 7/8 as long as paramere, apical 1/3 slender,

apex rounded, with large, angulate projection one margin; paramere

wide, tapered to rounded apex in apical 1/2 (Fig. 322); sipho robust,

long, with large membranous area in apical 1/5, basal capsule heav-

ily sclerotized, dark brown, inner arm slender, long, outer arm wide,

as long as inner arm, apically knobbed (Figs. 323, 324).

2 Not known.

Etymology. The species is named for Linda Lawrence, illustra-

tor for the Systematic Entomology Laboratory, USDA, Washing-

ton, DC, who has provided excellent illustrations of lady beetles for

many publications.

Remarks. Elytral color is striking in this species, with black

maculation on a red surface. In addition, the prosternal carinae are

somewhat unusual because they do not converge until near the basal

prosternal margin.

Species known only from females

Here we place those Hyperaspis species having 11-articled an-

tennae and known only from the @ sex.

In many instances it is nearly impossible to correctly associate

Hyperaspis 63 and ££ unless they are present in series or in large

enough numbers so the connection can be detected.

43. Hyperaspis c-nigrum Mulsant, 1850

Hyperaspis c-nigrum Mulsant, 1850: 649; CrorcH 1874: 224; KoRscHEFsKy 1931:

186; BLACKWELDER 1945: 446; GORDON 1987: 28.

Type locality. Brazil, MNHP (lectotype here designated).

Description. &, length 3.4 mm, width 2.7 mm; body

form oval, convex. Dorsal surface with head strongly alutaceous,

dull: pronotum alutaceous, shiny: elytron shiny. Color yellow ex-

cept pronotum with 4 triangular, brown spots, 2 spots in anterior

1/2 narrowly connected to 2 basal spots on each side of middle;

elytron with large, brown spot separated from all margins, spot with

median yellow area narrowly connected to basal yellow border (Fig.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 359

325, 326); pro-, meso-, metasterna reddish brown; abdomen yellow-

ish brown except outer 1/4 and sterna 5-6 yellow. Head punctures

fine, hidden in alutaceous sculpture; pronotal punctures larger than

on head, separated by a diameter or less; elytral punctures larger

than pronotal punctures, separated by less than to twice a diameter.

Metasternal punctures smaller than on elytron medially, sparse, be-

coming coarse, contiguous laterally. Punctures on abdominal sterna

1-4 coarse, dense medially, becoming dense, fine laterally; sterna 5-6

finely, densely punctured throughout; area on 1*t abdominal sternum

inside postcoxal arc shiny, coarsely, densely punctured. Clypeal apex

shallowly emarginate, clypeus and frons abruptly joined. Epipleuron

oblique, grooved internally, strongly descending externally, femoral

depression deep. Prosternum with intercoxal carinae widely separated

at apex, convergent nearly to base, basal stem short, reaching basal

margin. Protibia not flanged. Postcoxal line on 1% abdominal ster-

num not extended to posterior sternal margin, rounded along margin,

apex recurved 3/8 distance to basal sternal margin. Fifth sternum

with apex broadly, feebly emarginate; 6 sternum arcuate. Genitalia

with beak of basal unit composed of 2 large segments with earlike,

anterolateral projections, connecting duct very short (Figs. 327, 328)

d. Not examined.

Geographical distribution: Brazil.

Specimens examined: 3. Brazil: “Brazil”; Minas Gerais (MNHP)

(UMZC).

Remarks. AHyperaspis c-nigrum is a large, distinctly pat-

terned species known from 3 specimens, including the lectotype.

Similar in size and coloration only to H. eupaleoides, H. c-nigrum is

distinguished by the dorsal color pattern, as is the former species.

The 2 lectotype here designated to stabilize future usage of this

name is labeled “Muséum Paris, Bresil, Minas-Geraes a Goyaz, de

Castelnau 19 - 47/168/ Hyperaspis C-nigrum Muls., auct. det.”. An-

other specimen is designated as paralectotype and is labeled “Mu-

séum Paris, des Mines.”

44. Hyperaspis sagittata Crotch, 1874

Hyperaspis sagittata Crotch, 1874: 225; KorscHEersky 1931: 195; BLACKWELDER

1945: 448; GORDON 1987: 28.

360 R.D. GORDON & C. CANEPARI

Type locality. Bogota, Colombia, UMZC (holotype, GORDON

1987).

Description. &, length 2.6 mm, width 2.2 mm; body

form rounded, convex. Dorsal surface with head alutaceous, dull:

pronotum weakly alutaceous, slightly shiny: elytron shiny. Color

yellow except pronotum with 4 triangular, brown spots, 2 anterior

spots in anterior 1/2 weakly connected to 2 basal spots on each side of

middle; elytron with all margins narrowly dark brown, with 4 brown

spots, 1 u-shaped spot in anterior 1/2 curved from base around hu-

meral callus and back to base, 1 irregular spot on suture posterior to

middle, angled postero laterally onto apical declivity and extended

onto apical 1/4, 1 spot nearly lateral margin medially, 1 small, spot

on suture at apex (Figs. 329, 330); pro-, meso-, metasterna reddish

brown; abdomen yellowish brown except outer 1/4 yellow. Head

punctures fine, distinct, separated by a diameter or less; pronotal

punctures larger than on head, separated by a diameter or less; ely-

tral punctures larger than pronotal punctures, separated by less than

to twice a diameter. Metasternal punctures smaller than on elytron

medially, sparse, becoming coarse, contiguous laterally. Punctures

on abdominal sterna 1-4 coarse, dense medially, becoming dense,

fine laterally; sterna 5-6 finely, densely punctured throughout; area

on 1" abdominal sternum inside postcoxal arc shiny, coarsely, dense-

ly punctured. Clypeal apex deeply emarginate, clypeus and frons

abruptly joined. Epipleuron oblique, grooved internally, descend-

ing externally, femoral depression deep. Prosternum with intercoxal

carinae widely separated at apex, convergent nearly to base, basal

stem short, reaching basal margin. Protibia not flanged. Postcoxal

line on 1*t abdominal sternum not extended to posterior sternal mar-

gin, flattened along margin, apex recurved 1/2 distance to basal ster-

nal margin. Fifth sternum with apex broadly, feebly emarginate; 6™

sternum arcuate. Genitalia with basal unit large, wide medially, ta-

pered at each end, beak short, wide, connecting duct short (Fig. 331)

3 Not known.

Geographical distribution: Colombia.

Specimens examined. 1 (holotype).

Remarks. Ayperaspis sagittata is a distinctly patterned spe-

cies slightly resembling H. colombiensis and H. campbelli. It is known

only from the £ holotype.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 361

45. Hyperaspis guilloryi (Mulsant, 1850)

Cleothera guilloryi Mulsant, 1850: 1040.

Hyperaspis guilloryt: CROTCH 1874: 229; KORSCHEFSKY 1931: 190; BLACKWELDER

1945: 447; GorDON 1987: 29.

Type locality. Colombia, UMZC (lectotype designated by Gor-

DON 1987).

Description. &, length 1.6 mm, width 1.3 mm; body form

oval, rounded, convex. Dorsal surface with head slightly alutaceous,

dull; pronotum shiny; elytron shiny. Color black except elytron

dark brown with large, median, oval, yellow spot (Figs. 332, 333);

antenna, mouthparts, propleuron, epipleuron, legs reddish brown;

pro-, meso-, metasterna dark brown; abdomen brown except outer

1/4 and sterna 5-6 yellowish brown. Head punctures fine, separated

by a diameter or less; pronotal punctures larger than on head, sepa-

rated by a diameter or less; elytral punctures larger than pronotal

punctures, separated by a diameter or less. Metasternal punctures

larger than on elytron, sparse medially, becoming coarse, contiguous

laterally. Punctures on abdominal sterna 1-4 coarse, dense medially,

becoming dense, fine laterally; sterna 5-6 finely, densely punctured

throughout; area on 1% abdominal sternum inside postcoxal arc

shiny, coarsely, densely punctured. Clypeal apex shallowly emargin-

ate, clypeus and frons abruptly joined. Epipleuron oblique, grooved

internally, descending externally, femoral depression deep. Antenna

with 11 articles (Fig. 334). Prosternum with intercoxal carinae nar-

rowly separated at apex, convergent toward base, basally stemmed,

reaching basal margin. Protibia not flanged. Postcoxal line on 1* ab-

dominal sternum not extended to posterior sternal margin, rounded

along margin, apex recurved 1/3 distance to basal sternal margin.

Fifth sternum with apex truncate. Genitalia with beak of basal unit

short, connecting duct short (Fig. 335)

3. Not known.

Geographical distribution: Colombia.

Specimens examined: 2. Colombia: the lectotype; Huila, Garzon

(USNM).

Remarks. ‘This very small species with a single yellow spot

on each elytron resembles only H. howdeni. Hyperaspis howdeni is

distinctly larger with a much larger elytral spot.

362 R.D. GORDON & C. CANEPARI

The £ lectotype in the UMZC has lost an elytron, but the

remaining elytron and body size match exactly a specimen from

Colombia previously identified as H. guilloryi by E.A. Chapin in

1954.

46. Hyperaspis pseudopavida n. sp.

Type material. Holotype £: Brazil, 316, 80.23, Cleothera pavida

M. Named by Mulsant (BM).

Description. &, length 2.2 mm, width 1.8 mm; body form

oval, convex. Dorsal surface with head alutaceous, dull; pronotum

shiny; elytron shiny. Color black except elytron with 3 small, yellow

spots, 1 transversely oval spot at base next to scutellum, 1 rectangu-

lar spot on lateral margin posterior to humeral callus, 1 irregularly

rounded spot on apical declivity near apical margin, widely separated

from suture (Figs. 336, 337); antenna, mouthparts except maxillary

palpus, narrow outer margin of propleuron, tibia yellow; maxillary

palpus, epipleuron, femora dark brown; abdomen dark brown ex-

cept outer 1/4 and sterna 5-6 yellowish brown. Head punctures fine,

separated by less than to twice a diameter; pronotal punctures fine,

larger than head punctures, separated by less than to twice a diam-

eter; elytral punctures larger than pronotal punctures, separated by

1 to 2 times a diameter. Metasternal punctures fine, sparse medi-

ally, becoming very coarse, nearly contiguous laterally. Punctures on

abdominal sterna 1-4 coarse, dense medially, becoming dense, fine

laterally; sterna 5-6 finely, densely punctured throughout; area on 1%

abdominal sternum inside postcoxal arc shiny, coarsely, densely punc-

tured. Clypeal apex emarginate, clypeus and frons abruptly joined.

Epipleuron oblique, grooved internally, with coarse, cross striations,

slightly descending externally, femoral depression deep. Prosternum

with intercoxal carinae widely separated at apex, convergent toward

base, basally stemmed, reaching basal margin. Protibia not flanged.

Postcoxal line on 1% abdominal sternum not extended to posterior

sternal margin, rounded along margin, apex recurved 1/2 distance to

basal sternal margin. Fifth sternum with apex broadly emarginate.

Genitalia with beak of basal unit large, as long as unit (Fig. 338).

4. Not known.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 363

Etymology. The species name refers to the resemblance to Cleo-

thera pavida Mulsant.

Remarks. ‘There are other black species with 3 pale spots

on each elytron, but this is the only one with a single spot next to

the scutellum. Hyperaspis recordata, described as a variant of Cleo-

thera pavida, also has a scutellar spot but only 2 elytral spots.

The holotype is a single ® in the BM collection identified by

Mulsant himself as Cleothera pavida Mulsant, a species he described

from Mexican material. However, the BM specimen identified by

Mulsant as C. pavida was part of the Deyrolle collection that Mul-

sant examined in 1853; C. pavida was described by Mulsant in 1850;

therefore, the specimen described above cannot be type material of

C. pavida (R. Booth, pers. comm.).

47. Hyperaspis istmina n. sp.

Type material. Holotype ®: Colombia, Chocé, Istmina,

31.VIIL'40, LM Murillo No. 5327a (USNM).

Description. ¢, length 2.5 mm, width 2.5 mm, body

oval, rounded, convex. Dorsal surface with head slightly alutaceous,

shiny: pronotum shiny; elytra shiny. Color black except head yel-

low; pronotum black with lateral 3/8 yellow; elytron with 1 large,

yellow spot extended from humeral callus nearly to apex, spot nar-

row apically, irregularly widened to truncate apex (Fig. 339, 340);

antenna, propleuron yellow; legs reddish yellow; abdominal ster-

na 1-4 black with lateral 1/3 and sterna 5-6 reddish yellow. Head

punctures fine, separated by less than to twice a diameter; pronotal

punctures larger than on head, separated by less than to twice a di-

ameter; elytral punctures larger than pronotal punctures, separated

by 1 to 2 times a diameter. Metasternal punctures larger than on

elytron, sparse medially, dense, contiguous laterally. Punctures on

abdominal sterna larger than on elytra, dense medially, becoming

fine, dense laterally, sterna 5-6 finely densely punctured throughout;

area on 1*t abdominal sternum inside postcoxal arc slightly aluta-

ceous, finely, sparsely punctured. Clypeal apex emarginate, clypeus

and frons abruptly joined. Epipleuron flat, grooved internally, not

descending externally, femoral depressions deep. Antenna with 11

articles (Fig. 341). Protibia not flanged. Prosternum with intercox-

364 R.D. GORDON & C. CANEPARI

al carinae narrowly separated at apex, convergent nearly to basal

sternal margin, stemmed, reaching basal margin. Postcoxal line on

1* abdominal sternum reaching posterior sternal margin, rounded

along margin, apex recurved 5/8 distance to basal sternal margin.

Fifth sternum with apex weakly emarginate.

4. Not known.

Etymology. The species is named for the holotype locality, and

the name is used as a noun in apposition.

Remarks. The elytral color pattern is similar only to that of

H. chocoi, and distinguishes H. istmina from other Hyperaspis species.

48. Hyperaspis satipoensis n. sp.

Type material. Holotype ®: Peru, Satipo, XI, 1941, coll. by

Paprzycki, G.H. Dieke Coll’ n.1965 (USNM).

Description. ®, length 3.0 mm, width 2.4 mm; body

rounded, convex. Dorsal surface with head alutaceous, dull: pro-

notum shiny; elytra shiny. Color black except head yellow; prono-

tum yellow with basomedian black spot, lateral 1/4 of pronotum

yellow, basomedian spot with small, obliquely oval, yellow spot on

each side of middle at base; elytron with 3 yellow spots, discal spot

near suture anterior to middle roughly triangular, broad, spot near

lateral margin laterad of discal spot irregular in shape, large, com-

ma shaped yellow spot on apical declivity (Fig. 342, 343); antenna,

mouthparts propleuron, legs reddish yellow; abdominal sterna 1-4

black with lateral 1/4 and sterna 5-6 reddish yellow. Head punctures

fine, separated by less than to twice a diameter; pronotal punctures

larger than on head, separated by a diameter or less; elytral punc-

tures larger than pronotal punctures, separated by a diameter or less.

Metasternal punctures larger than on elytron, sparse medially, dense,

contiguous laterally. Punctures on abdominal sterna larger than on

elytra, dense medially, becoming fine, dense laterally, sterna 5-6

finely densely punctured throughout; area on 1* abdominal sternum

inside postcoxal arc shiny, coarsely, densely punctured. Clypeal apex

emarginate, clypeus and frons abruptly joined. Epipleuron oblique,

grooved internally, descending externally, femoral depressions deep.

Antenna with 11 articles (Fig. 334, outer articles lost). Protibia not

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 365

flanged. Prosternum with intercoxal carinae widely separated at

apex, convergent nearly to basal sternal margin, stemmed, reach-

ing basal margin. Postcoxal line on 1% abdominal sternum reaching

posterior sternal margin, flattened along margin, apex recurved 5/8

distance to basal sternal margin. Fifth sternum with apex broadly

arcuate. Genitalia with beak of basal unit short, back of unit heavily |

sclerotized; connecting duct short (Figs. 345, 346)

d. Not known.

Etymology. The species is named for Satipo, Peru, the holotype

locality.

Remarks. Distinguishing characters for this species are

based primarily on dorsal coloration. ‘The pronotum with 2 oblique-

ly oval, median spots at the base is unique with the genus. The 3

spotted elytron with a large, comma-shaped apical spot also differs

from that of other species.

49. Hyperaspis abertha n. sp.

Type material. Holotype 2: Brazil, Sào Paulo, Capital, 21-X-31

Col. J.P.F., Korschefsky Collection 1952 (USNM).

Description. ®, length 3.3 mm, width 2.6 mm; body

oval, convex. Dorsal surface with head alutaceous, dull: pronotum

alutaceous, slightly shiny; elytra shiny. Color black except head yel-

low with vertex and narrow border next to eye black; pronotum

black with lateral 1/4 yellow; elytron with 1 large, yellow, transverse,

apical spot near sutural margin, inner margin of spot obliquely trun-

cate (Figs. 347, 348); antenna, propleuron yellow; mouthparts dark

brown; proleg reddish yellow; meso-, metaleg reddish yellow except

basal 1/2 of femur dark brown; abdominal sterna 1-5 black with

lateral 1/4 and sternum 6 reddish yellow. Head punctures fine, sepa-

rated by a diameter or less; pronotal punctures larger than on head,

separated by a diameter or less; elytral punctures larger than pro-

notal punctures, separated by less to twice a diameter. Metasternal

punctures larger than on elytron, separated by a diameter medially,

dense, becoming contiguous laterally. Punctures on abdominal sterna

larger than on elytra, dense medially, becoming fine, dense laterally,

sterna 5-6 finely, densely punctured throughout; area on 1% abdomi-

366 R.D. GORDON & C. CANEPARI

nal sternum inside postcoxal arc shiny, coarsely, densely punctured.

Clypeal apex emarginate, clypeus and frons abruptly joined. Epi-

pleuron oblique, grooved internally, descending externally, femoral

depressions deep. Antenna with 11 articles (Fig. 349). Protibia not

flanged. Prosternum with intercoxal carinae narrowly separated at

apex, convergent nearly to basal sternal margin, stemmed, reach-

ing basal margin. Postcoxal line on 1% abdominal sternum reaching

posterior sternal margin, flattened along margin, apex recurved 3/8

distance to basal sternal margin. Fifth sternum with apex broadly

emarginate. Genitalia with beak of basal unit long; connecting duct

ei) de

d. Not known.

Etymology. The specific name is an arbitrary combination of

letters.

Remarks. Three other species - H. apicaspis, H. delicata,

and H. mimula - have black elytra with a single, yellow apical spot.

Hyperaspis abertha differs by the much larger size, and spot on

elytron large, transverse, with an obliquely truncate inner margin.

50. Hyperaspis circumclusa n. sp.

Type material. Holotype &: Brazil, Taperina, Acc. No. 2966

(CMP). Paratype ®: Brazil, Parà-Is. Marajò, Curralifo, III-VI.1988,

leg. Gallizia (CCM).

Description. ®, length 2.2 mm, width 1.8 mm; body

rounded, slightly oval, convex. Dorsal surface with head alutaceous,

dull: pronotum slightly alutaceous, shiny; elytra shiny. Color red-

dish yellow except head yellow with yellowish brown clypeus; pro-

notum dark brown with lateral 1/5 yellow; elytron reddish yellow

except lateral and apical margins with wide yellow border from base

nearly to suture at apex, sutural margin widely dark brown (Figs.

352, 353); pro-, meso- and metasterna reddish brown; basal abdomi-

nal sternum reddish brown medially, remainder of abdomen red-

dish yellow. Head punctures fine, hidden in alutaceous sculpture;

pronotal punctures larger than on head, separated by a diameter or

less; elytral punctures larger than pronotal punctures, separated by

a diameter or less. Metasternal punctures larger than on elytron,

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 367

separated by a diameter medially, dense, becoming contiguous lat-

erally. Punctures on abdominal sterna as large as on elytra, dense

medially, becoming fine, dense laterally, sterna 5-6 finely, densely

punctured throughout; area on 1% abdominal sternum inside post-

coxal arc shiny, coarsely, sparsely punctured. Clypeal apex emargin-

ate, clypeus and frons abruptly joined. Epipleuron oblique, grooved

internally, descending externally, femoral depressions deep. Antenna

with 11 articles. Protibia not flanged (Fig. 354). Prosternum with

intercoxal carinae widely separated at apex, convergent nearly to

basal sternal margin, stemmed, reaching basal margin. Postcoxal line

on 1% abdominal sternum reaching posterior sternal margin, slightly

flattened along margin, apex recurved 3/8 distance to basal sternal

margin. Fifth sternum with apex broadly emarginate. Genitalia with

beak of basal unit short, divided into 2 projections (Figs. 355, 356).

4. Not known.

Etymology. The species is named for the lateral yellow border

encircling nearly the entire body except the head.

Remarks. No other known species of South American

Hyperaspis has a similar color pattern. The lateral yellow border is the

same width throughout, enhancing the circular aspect of the color pattern.

51. Hyperaspis aemulata n. sp.

Type material. Holotype 2: Venezuela, ex Venezuela, Hoboken,

N. J. (New Jersey), on Cattleya, Hob. (Hoboken) # 934 (specimen

intercepted at customs in Hoboken, New Jersey) (USNM).

Description. 2, length 3.4 mm, width 2.7 mm; body rounded,

slightly oval, convex. Dorsal surface with head slightly alutaceous,

shiny: pronotum shiny; elytra shiny. Color yellow except pronotum

with large, black, basomedian spot extended nearly to anterior mar-

gin, lateral margins of spot deeply emarginated with yellow; elytron

with 5 dark brown spots and sutural border dark brown in posterior

5/6, sutural margin slightly widened in apical 1/3, 1 small, narrowly

oval spot between suture and humeral callus, 1 irregularly elongate

spot extended from base across humeral callus, 1 oval spot at lat-

eral margin before apical declivity, 1 elongate oval spot on apical

declivity between suture and lateral spot, 1 transversely oval api-

368 R.D. GORDON & C. CANEPARI

cal spot connected to sutural border (Figs. 357, 358); pro-, meso-,

metasterna dark brown; abdominal sterna 1-5 reddish brown except

lateral 1/4 and sternum 6 yellow. Head punctures fine, separated

by less than to twice a diameter; pronotal punctures larger than

on head, separated by a diameter or less; elytral punctures larger

than pronotal punctures, separated by less than to twice a diameter.

Metasternal punctures larger than on elytron, separated by a diam-

eter medially, becoming dense, contiguous laterally. Punctures on

abdominal sterna larger than on elytra, sparse medially, becoming

fine, dense laterally, sterna 5-6 finely, densely punctured through-

out; area on 1° abdominal sternum inside postcoxal arc shiny,

coarsely, densely punctured. Clypeal apex emarginate, clypeus and

frons abruptly joined. Epipleuron oblique, not grooved internally,

descending externally, femoral depressions deep. Antenna with 11

articles (Fig. 359). Protibia not flanged. Prosternum with intercoxal

carinae widely separated at apex, convergent 1/2 distance to base,

stem long, reaching basal margin. Postcoxal line on 1° abdominal

sternum reaching posterior sternal margin, slightly flattened along

margin, apex recurved 2/3 distance to basal sternal margin. Fifth

sternum with apex feebly emarginate. Genitalia with beak of basal

unit short, wide; connecting duct long (Fig. 360).

d. Not known.

Etymology. The specific name is from the Latin aemulor, mean-

ing to emulate, referring to the resemblance to other species of Hy-

peraspidini.

Remarks. Distinguishing characteristics of H. aemulata are

the large size, 5 dark spots on each elytron, suture with dark bor-

der, and postcoxal line on the basal abdominal sternum recurved 2/3

distance to apical sternal margin.

The dorsal coloration of this species slightly resembles that of

H. matronata, and more closely, those of Tenuisvalvae ecoffeti and

T. notata.

52. Hyperaspis dispar n. sp.

Type material. Holotype ®: Brazil, Guanabara, Rio de Janeiro,

X.1963, M. Alvarenga coll. (USNM). Paratype £: same data as ho-

lotype (USNM).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 369

Description. 2, length 3.4 mm, width 2.8 mm; body

rounded, slightly oval, convex. Dorsal surface with head alutaceous,

dull: pronotum shiny; elytra shiny. Color yellow except head with

black spot on each side of vertex, frons with poorly defined, light

brown area extended onto clypeus; pronotum with 5 dark brown

spots, a large, triangular spot on each side of middle in apical 1/2,

transverse basomedian vitta, and small, elongate, oval spot on each

side near lateral margin; elytron with 5 spots and sutural margin

dark brown, suture bordered from scutellum nearly to apex, 1 large,

elongate oval, discal spot near suture in basal 1/2, 1 irregularly oval

spot on humeral angle, 1 elongate oval spot on apical declivity near

suture, 1 narrow, somewhat triangular spot near lateral margin in

apical 1/2, 1 narrowly transverse spot at apex bordering suture (Figs.

361, 362); mouthparts, legs reddish yellow; pro-, meso-, metasterna

black; abdomen reddish yellow except basal sterna black medially.

Head punctures fine, separated by less than to twice a diameter;

pronotal punctures larger than on head, separated by a diameter or

less; elytral punctures larger than pronotal punctures, separated by 1

to 2 times a diameter. Metasternal punctures larger than on elytron,

separated by a diameter or less medially, by less than a diameter

laterally. Punctures on abdominal sterna larger than on elytra, dense

medially, becoming fine, dense laterally and on sterna 5-6; area on

1* abdominal sternum inside postcoxal arc shiny, nearly impunc-

tate. Clypeus emarginate apically, clypeus and frons abruptly joined.

Epipleuron oblique, grooved internally, descending externally, fem-

oral depressions deep. Antenna with 11 articles (Fig. 363). Protibia

not flanged. Prosternum with intercoxal carinae widely separated

at apex, convergent toward base, basally stemmed, reaching basal

margin. Postcoxal line on 1*t abdominal sternum reaching posterior

sternal margin, flattened along margin, apex recurved 1/2 distance

to basal sternal margin. Fifth sternum with apex feebly emarginate.

Genitalia with beak of basal unit absent; connecting duct long (Fig.

DOD).

3. Not known.

Etymology. The specific name is the Latin dispar, meaning un-

like or dissimilar, referring to the distinctive color pattern.

Remarks. This is a large, distinctively marked species not

resembling any other member of Hyperaspis. Rather, it has the dor-

370 R.D. GORDON & C. CANEPARI

sal aspect of several species of the tribe Brachiacanthini, and also

some species of the coccinelline tribe Synonychini.

53. Hyperaspis histrionica (Mulsant, 1850)

Cleothera histrionica Mulsant, 1850: 629.

Hyperaspis histrionica: CROTCH 1874: 222; KORSCHEFSKY 1931: 190; BLACKWELDER

1945: 447.

Type locality: “Carthagena” (Cartagena, Colombia) (MNHL,

lectotype here designated).

Description. ®, length 2.4 mm; body form elongate,

slightly convex. Color black except head dark brown; pronotum dark

yellow with black border on median part of base, with triangular black

spot fused at apex of central part of border, and small, black, round-

ed spot laterally; elytron black with yellow lateral border extended

from humeral callus posteriorly, not reaching apex, with 6 yellow

spots arranged in 3 rows, row 1 from humeral callus toward suture,

row 2 median, and row 3 preapical, each row with 2 spots sometimes

fused to each other or to lateral border (Figs. 366, 367); mouthparts,

legs dark yellow; propleuron yellow; pro-, meso-, metasterna black;

abdomen dark brown. Pronotum and elytron with equal size punc-

tures, separated by about 3 times a diameter. Clypeal apex emargin-

ate, clypeus and frons joined at right angle. Epipleuron with femo-

ral depressions. Postcoxal line on 1*t abdominal sternum incomplete

6. Not known.

Variation. None observed.

Geographical distribution: known only from the type locality.

Specimens examined: 1, the lectotype.

Remarks. ‘The species described above is recognized by the

presence of 6 yellow, elytral spots arranged in 3 transverse rows.

Hyperaspis histrionica was described by Mulsant from speci-

mens in the Dejean collection. Although Mulsant had more than

one specimen when he described this species, only one remains in

that collection. This specimen, labeled “Carthagena, Lebas” is des-

ignated as the lectotype to stabilize future usage of this name.

This species is known only from the ® lectotype; hence, it is

placed with species known only from that sex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI STI

Hyperaspis tavronensis closely resembles H. histrionica in dorsal

color pattern, but we consider them distinct species for the pres-

erie

Section II

Species with frons and clypeus smoothly, obliquely joined; pro-

tibia widened apically.

vredenburgi group

Species with body slightly flattened; frons and clypeus and

clypeus smoothly, obliquely joined; protibia widened apically, slight-

ly flanged; male genitalia with basal lobe narrowed in apical 1/4,

apex rounded, with abrupt, hooklike projection on one side (Figs.

376, 376), paramere slender, oval; protibia slightly flanged.

Hyperaspis species with either 10-articled (H. connectens) or

11-articled antennae, and smoothly, obliquely joined frons and

clypeus, the latter a character shared only with the conclusa group.

Basal lobe of the male genitalia with an abrupt, hooklike projection

on one side is diagnostic for the vredenburgi group.

54. Hyperaspis ayacucho n. sp.

Type material. Holotype dé: Venezuela, Exp. Territ. Amazo-

nas, Puerto Ayacucho, June 15, 1950, J. Maldonado Capriles Coll.

(USNM). Paratype d: Venezuela, Exp. Territ. Amazonas, Mouth

Cunucunuma, R., Apr. 2, 1950, J. Maldonado Capriles Coll.

(USNM).

Description. 6, length 2.3 mm, width 1.8 mm; body

oval, slightly flattened. Dorsal surface with head alutaceous, weakly

shiny: pronotum strongly alutaceous, dull; elytra alutaceous, weakly

shiny. Color black except head yellow; pronotum black with anterior

1/6 and lateral 1/5 yellow, basal margin with black area narrowly,

briefly emarginated with yellow laterally; elytron with broad, yellow

border on basal, lateral and apical margins, basal border enlarged be-

tween scutellum and humeral callus, lateral border irregular, apical

372 R.D. GORDON & C. CANEPARI

border not reaching suture (Fig. 368); antenna, mouthparts yellow-

ish brown; propleuron, epipleuron yellow; proleg with apical 1/3 of

tibia, and anterior 1/2 of femur yellow; abdomen black with lateral

1/4 reddish yellow. Head punctures fine, separated by a diameter or

less; pronotal punctures larger than on head, separated by less than

to twice a diameter; elytral punctures slightly larger than pronotal

punctures, separated by a diameter or less. Metasternal punctures

larger than on elytron, sparse medially, dense, contiguous laterally. —

Punctures on abdominal sterna larger than on elytra, dense medially,

becoming fine, dense laterally, sterna 5-6 finely densely punctured

throughout; area on 1*t abdominal sternum inside postcoxal arc aluta-

ceous, coarsely, sparsely punctured. Clypeal apex deeply emarginate,

clypeus and frons smoothly, obliquely joined. Epipleuron slightly

oblique, feebly grooved internally, slightly descending externally,

femoral depressions deep. Antenna with 11 articles (Fig. 369). Protibia

widened apically. Prosternum with intercoxal carinae narrowly sepa-

rated at apex, parallel 2/3 distance to anterior prosternal margin, not

stemmed, not reaching basal margin. Postcoxal line on 1*t abdominal

sternum reaching posterior sternal margin, flattened along margin,

apex recurved 1/2 distance to basal sternal margin. Fifth sternum

with apex broadly, weakly emarginate; 6" sternum truncate. Geni-

talia with basal lobe 3/4 as long as paramere, apex narrowly round-

ed, with small, angulate projection on one margin; paramere oval,

tapered from basal 1/4 to rounded apex (Fig. 370); sipho robust,

apically straight, with apical membranous area, basal capsule with

inner arm long, slender, outer arm slender, as long as inner arm,

apically knobbed, basal border slightly emarginate (Figs. 371, 372).

2 Not known.

Etymology. The species is named for the holotype locality with

the name used as a noun in apposition.

Remarks. Characters distinguishing H. ayacucho are parallel

prosternal carinae not extended to anterior prosternal margin; extreme-

ly alutaceous, dull pronotal surface; and broad, yellow elytral border.

55. Hyperaspis herrerai n. sp.

Type material. Holotype d: Colombia, Magd. (Magdalena),

Codazzi, Apr. 20, 1962, J. Herrera collector (USNM).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 373

Description. 6, length 2.4 mm, width 1.8 mm; body

oval, slightly flattened. Dorsal surface with head alutaceous, weak-

ly shiny; pronotum alutaceous, weakly shiny; elytron alutaceous,

weakly shiny. Color reddish yellow except head yellow; pronotum

with large, black, rectangular basomedian spot, spot widely sepa-

rated from anterior pronotal margin, anterolateral angle with small

projection; elytra with large, more or less triangular spot in middle,

spot narrow at scutellum, widened posteriorly onto apical decliv-

ity, abruptly narrowed to suture before apex, small, oval, black

spot on humeral callus (Fig. 373, 374); venter of head, pro-, meso-,

metasterna black; propleuron yellow; abdomen with median area of

sterna 1-5 dark brown, lateral 1/4 and sternum 6 reddish yellow.

Head punctures fine, separated by less than to twice diameter; pro-

notal punctures larger than on head, separated by 1 to 3 times diam-

eter; elytral punctures smaller than pronotal punctures, separated by

1 to 3 times a diameter. Metasternal punctures larger than on pro-

notum, sparse medially, becoming nearly contiguous laterally. Punc-

tures on abdominal sterna 1-4 as large as pronotal punctures, dense

medially, becoming fine, dense laterally; sterna 5-6 finely, densely

punctured throughout; area on 1* abdominal sternum inside post-

coxal arc shiny, coarsely, densely punctured. Clypeal apex deeply

emarginate, clypeus and frons smoothly, obliquely joined. Epipleu-

ron flat, feebly grooved internally, not descending externally, femo-

ral excavation moderate, not deep. Antenna with 11 articles (Fig.

375). Protibia widened apically. Prosternum with intercoxal carinae

narrowly separated at apex, convergent near base, basal stem short,

reaching basal pronotal margin. Postcoxal line on 1% abdominal ster-

num reaching posterior sternal margin, flattened along margin, apex

recurved 5/8 distance to basal sternal margin. Fifth sternum with

apex truncate; 6° sternum slightly arcuate. Genitalia with basal lobe

3/4 as long as paramere, apex narrowly rounded, notched in apical

1/2 on one margin; paramere oval, tapered to rounded apex in api-

cal 1/2 (Fig. 376); sipho slender, constricted before apex, apex with

membranous area, basal capsule with inner and outer arms slightly

elongate , wide (Figs. 377, 378).

2 Not known.

Etymology. The species is named for J. Herrera, collector of the

holotype specimen.

374 R.D. GORDON & C. CANEPARI

Remarks. The large, triangular elytral spot is unique to

this species, and elytral punctures smaller than pronotal punctures

are rare in Hyperaspis: the elytral punctures are nearly always larger

or at least equal in size.

56. Hyperaspis vredenburgi n. sp.

Type material. Holotype 6: Brazil, Rio Negro, Manacapruru,

July, 1935, G. V. Vredenburg, Brit. Mus. 1935-615 (BM). Allotype

2: Same data as holotype (BM). Paratypes, 12. 9, same data as ho-

lotype (BM). 3, Trinidad, Caranege, Oct. 14, 1918. A-760, Harold

Morrison (USNM).

Description. 6, length 2.2 mm, width 1.7 mm; body

oval, slightly flattened. Dorsal surface with head strongly alutaceous,

dull: pronotum alutaceous, dull; elytra alutaceous, dull. Color black

except head yellow; pronotum yellow with large, somewhat rectan-

gular, basomedian spot, spot widely separated from anterior margin,

basal margin of spot narrower than anterior margin, elytron with 3

yellow spots and lateral margin in apical 2/3 yellow, 1 large, trans-

versely oval scutellar spot, 1 large, irregularly rounded, discal spot

near suture connected to lateral yellow border, 1 large, transversely

oval spot at apex, yellow lateral border irregular, abruptly widened

near apex to join discal spot (Figs. 379, 380); antenna, propleuron,

epipleuron yellow; mouthparts reddish brown; pro-, mesolegs red-

dish yellow; metaleg with femur brown except apical 1/8 yellow,

tibia and tarsus yellow except outer margin of tibia brown; abdo-

men brown with lateral 1/8 reddish yellow. Head punctures fine,

hidden in alutaceous sculpture; pronotal punctures larger than on

head, separated by less than to 3 times a diameter; elytral punctures

slightly larger than pronotal punctures, separated by 1 to 3 times

a diameter. Metasternal punctures much larger than on elytron,

separated by about a diameter throughout. Punctures on abdomi-

nal sterna 1-4 larger than on elytra, dense medially, becoming fine,

dense laterally, sterna 5-6 finely densely punctured throughout;

area on 1°“ abdominal sternum inside postcoxal arc shiny, coarsely,

densely punctured. Clypeal apex deeply emarginate, clypeus and

frons smoothly, obliquely joined. Epipleuron nearly flat, internally

grooved, not descending externally, femoral excavations moderate,

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 375

not deep. Antenna with 11 articles (Fig. 381). Protibia widened api-

cally. Prosternum with intercoxal carinae narrowly separated, par-

allel, extended 1/2 distance to basal margin. Postcoxal line on 1%

abdominal sternum not reaching posterior sternal margin, rounded

along margin, apex recurved 5/8 distance to basal sternal margin.

Fifth sternum with apex feebly emarginate; 6" sternum slightly ar-

cuate. Genitalia with basal lobe 3/4 as long as paramere, apex nar-

rowly rounded, with angulate projection on one margin; paramere

wide, tapered from middle to rounded apex; (Fig. 382); sipho short,

with membranous area, apically sinuate, basal capsule with inner

arm slender, long, outer arm slender, as long as inner arm, apically

truncate (Figs. 383, 384).

2 Similar to male except head black; black spot on pronotum

extended to apical margin. Genitalia with beak of basal unit short,

connecting duct short (Figs. 385).

Variation. Elytron with yellow lateral border as described above

or broken into 2 spots, 1 triangular spot at humeral angle, 1 apical

spot joining discal spot.

Etymology. The species is named for the collector of the type

series.

Remarks. Elytra with scutellar spots are unusual, and the

elytral pattern combined with the short, parallel prosternal carinae

distinguishes H. vredenburgi. In addition, the consistently dull dor-

sal surface, lateral metasternal punctures separated by a diameter,

and simple protibia will distinguish H. vredenburgi from either H.

tayronensis or H. laterimaculata, each of which also has 5 spots on

each elytron.

‘This is another instance of a species described from Amazonian

Brazil occurring on Trinidad, with no other localities known. Tenuts-

valvae unipunctata shares the same distribution.

57. Hyperaspis festiva Mulsant, 1850

Hyperaspis festiva Mulsant, 1850: 659; CrorcH 1874: 230; GorHam 1895: 196;

WEISE 1922: 36; KORSCHEFSKY 1931: 188; BRETHES 1925a: 204; BLACKWELDER

1945: 447; GORDON 1987: 29.

Hyperaspis cincticollis Mulsant, 1850: 553; CROTCH 1874: 230; GORDON 1987: 29.

Cleothera cincticollis: GORHAM 1895: 195.

376 R.D. GORDON & C. CANEPARI

Hyperaspis festiva ab. cincticollis: KORSCHEFSKY 1931: 188; BLACKWELDER 1945:

447,

Hyperaspis lemniscata: Boheman, 1859: 205; CRoTcH 1874: 227; BRETHES 1925b: 203

(synonym of H. festiva).

Hyperaspis festiva ab. lemniscata: KORSCHEFSKY 1931: 188.

Hyperaspis festiva var. apicalis Weise, 1885: 167; KorsCHEFSKy 1931: 188.

Hyperaspis juniapuca Brèthes, 1925b: 5; KoRSCHEFSKY 1931: 290; BLACKWELDER

1945: 447. (n. syn.).

Type locality: of festiva, Brazil (UMZC, lectotype designa-

ted by Gorpon 1987); of cincticollis, Colombia (MNHP, lectotype

here designated); of lemniscata, unknown; of apicalis, Puerto Rico

(ZMHB, lectotype here designated); of juniapuca, Brazil, Rio Junia-

puca (Amazones) (BM, holotype).

Description. 6, length 2.0 mm, width 1.4 mm; body

elongate, oval, slightly flattened. Dorsal surface with head slightly

alutaceous, feebly shiny; pronotum alutaceous, dull; elytron aluta-

ceous, slightly shiny. Color black except head yellow with vertex

black; pronotum black with anterior border narrowly yellow, lateral

1/8 yellow, median black area deeply emarginated basally by exten-

sion of lateral yellow spot; elytron with lateral and apical margins

narrowly, irregularly bordered with yellow, yellow border apically

recurved onto apical declivity, apex enlarged, 1 triangular discal

spot anterior to middle (Fig. 388); antenna, mouthparts, legs reddish

brown; propleuron, epipleuron yellow; abdomen with median area

dark brown, lateral 1/4 reddish yellow. Head punctures fine, sepa-

rated by a diameter or less; pronotal punctures larger than on head,

separated by less than to twice a diameter; elytral punctures larger

than pronotal punctures, separated by a diameter or less. Metaster-

num with punctures larger than on elytron, punctures separated by

a diameter medially, lateral punctures separated by less than a diam-

eter. Punctures on abdomen coarse, dense on sterna 1-4, becoming

fine, dense laterally and on sterna 5-6; area on 1% abdominal ster-

num inside postcoxal arc alutaceous, nearly impunctate. Clypeal apex

deeply emarginate, clypeus and frons smoothly, obliquely joined.

Epipleuron flat, internally grooved, slightly descending externally,

femoral excavations moderate, not deep. Antenna with 11 articles

(Fig. 397). Prosternum with intercoxal carinae narrowly separated

at apex, convergent toward base, pinched medially, extended 3/4

distance to anterior prosternal margin, not stemmed, not reaching

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI Ord.

basal margin. Protibia widened apically. Basal abdominal sternum

with apical margin sinuate. Postcoxal line on 1% abdominal sternum

not reaching posterior sternal margin, flattened along margin, apex

recurved 1/2 distance to basal sternal margin. Fifth sternum with

apex truncate; 6 sternum truncate. Genitalia with basal lobe 7/8

length of paramere, tapered to narrowly rounded apex in apical 1/4,

with abrupt, angulate projection on one margin; paramere narrow,

tapered from basal 1/4 to rounded apex (Fig. 398); sipho robust,

long, apex with membranous area, basal capsule with inner arm

long, slender, outer arm shorter than inner arm, slender, curved,

basal border deeply emarginate (Figs. 399, 400).

2 Similar to male except head black. Genitalia with beak of

basal unit short, connecting duct short (Fig. 401).

Variation. Length 2.0 to 2.7 mm, width 1.4 to 2.0 mm. Ely-

tral color extremely variable, discal spot often narrowly or widely

connected to spot on apical declivity; discal spot may be enlarged

to contact lateral yellow border; lateral yellow border often broken

before apex, leaving apical spot free; size of elytral spots variable

(Figs. 386-396).

Geographical distribution: Mexico and Caribbean islands south

to Argentina and Bolivia. Recorded in the literature from the Unit-

ed States, but not present there.

Specimens examined: 321. Many localities, specimens present in

nearly every collection examined (BM) (CAS) (CDA) (CMP) (DEI)

(MBR) (SMTD) (USNM) (ZMHB).

Remarks. AHAyperaspis festiva is widely distributed through-

out South America except Chile. It is frequently found and col-

lected, and has a somewhat unstable color pattern. However, the

pronotum with the median black area deeply emarginated basally by

an extension of the lateral yellow border is an excellent, remarkably

stable one. In addition, the prosternal carinae do not reach the basal

sternal margin, and the discal spot on each elytron, when present, is

nearly always triangular.

The lectotype of H. cincticollis, here designated to stabilize fu-

ture usage of this name, bears the labels “Type/Coll. Mniszech”.

The holotype of H. juniapuca has male genitalia identical to

those of typical H. festiva, and we consider it a junior synonym of

H. festiva. Two female syntypes of H. festiva var. apicalis are in

378 R.D. GORDON & C. CANEPARI

the ZMHB collection. The first of these, labeled “Portorico/Hype-

raspis apicalis (handwritten)/ ex Coll. J. Weise”, is designated as

lectotype. The second syntype, bearing identical labels but lacking

the handwritten name, is designated as paralectotype. A syntype

of H. festiva in the DEI collection labeled “Brasil Schaum/Coll.

Haag/Syntypus (pink paper)/festzva mls. typ” is designated as pa-

ralectotype.

58. Hyperaspis germainii Crotch, 1874

Hyperaspis germaini Crotch, 1874: 228; KorSCHEFSKy 1931: 189; BLACKWELDER

1945: 447; GORDON 1987: 29.

Type locality: Chile (UMZC, holotype, GORDON 1987).

Description. 6, length 2.5 mm, width 1.8 mm; body

elongate, oval, slightly flattened. Dorsal surface with head slightly

alutaceous, shiny; pronotum alutaceous, feebly shiny; elytron slightly

alutaceous, shiny. Color black except head yellow with black vertex

and narrow border along eye black; pronotum with anterior mar-

gin narrowly yellow, anterolateral angle broadly yellow, yellow area

nearly reaching posterolateral angle; elytron with basal yellow border

extended to lateral margin and posteriorly onto apical declivity, then

inward nearly to suture, 1 small, irregularly rounded, discal spot

posterior to middle (Figs. 402, 403); antenna, mouthparts, epipleu-

ron, proleg reddish yellow; propleuron yellow; meso- and metaleg

reddish yellow with femora dark brown; abdomen with median area

dark brown, lateral 1/4 dark reddish brown. Head punctures fine,

separated by less than to twice a diameter; pronotal punctures as

large as on head, separated by 1 to 3 times a diameter; elytral punc-

tures larger than pronotal punctures, separated by 1 to 2 times a

diameter. Metasternum with punctures much larger than on elytron,

punctures separated by a diameter medially, lateral punctures sepa-

rated by less than a diameter. Punctures on abdominal sterna larger

than on elytra, sparse on sterna 1-4, becoming fine, dense later-

ally and on sterna 5-6; area on 1° abdominal sternum inside post-

coxal arc shiny, nearly impunctate. Clypeal apex deeply emarginate,

clypeus and frons smoothly, obliquely joined. Epipleuron slightly

oblique, internally grooved, descending externally, femoral excava-

tions deep. Antenna with 11 articles (Fig. 404). Prosternum with

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 379

intercoxal carinae narrowly separated at apex, convergent from apex

toward base, basally stemmed, reaching basal margin (Fig. 405).

Protibia widened apically. Postcoxal line on 15 abdominal sternum

not reaching posterior sternal margin, rounded along margin, apex

recurved 1/2 distance to basal sternal margin. Fifth sternum with

apex truncate; 6 sternum truncate. Genitalia with basal lobe nearly

as long as paramere, apex narrow, obliquely rounded, with large,

angulate projection on one margin; paramere narrow, tapered from

basal 1/4 to rounded apex (Fig. 406); sipho slender, apex sinuate,

with membranous area, basal capsule with inner arm long, slender,

outer arm as long as inner arm, widened to apex, apically knobbed,

basal border emarginate (Figs. 407, 408).

? Similar to d except head black; anterior margin of pronotum

black. Genitalia with beak of basal unit long, connecting duct short

(Fig. 409).

Variation. Length 2.4 to 3.0 mm, width 1.8 to 2.4 mm. Apex

of yellow border on elytron may be separated, forming a single spot

on apical declivity.

Geographical distribution: Argentina, Chile.

Specimens examined: 24. Chile: Angol; Cauquenes; Chillan;

Concepcion; Escuadron; Colorada; Hualqui; Curico, Peidras Negras;

El Radal, Cordillera Talca; Nuble, Estero Bulileo; Santiago (CAS)

(CCM) (CMP) (MNHS) (MSNG) (USNM) (ZMHB).

Remarks. ‘The Chilean Hyperaspis fauna is very limited,

and H. germaini has a color pattern unlike that of other Chilean

species, rendering it easily recognizable.

59. Hyperaspis connectens (‘Thunberg, 1808)

Coccinella connectens Thunberg, 1808: 157.

Hyperaspis connectens: MULSANT 1850: 662; CroTtcH 1874: 230; KoRSscHEFSKY 1931:

186; DOBZHANSKY 1941: 25; GoRDON 1985: 473; GORDON 1987: 29.

Hyperaspis lengi Schaeffer, 1905: 144; DoBZzHANSKY 1941: 25; Gorpon 1985: 473.

Description. 6, length 2.6 mm, width 1.9 mm; body

elongate, oval, slightly flattened. Dorsal surface with head slightly

alutaceous, shiny; pronotum alutaceous, feebly shiny; elytron slightly

alutaceous, shiny. Color black except head yellow; pronotum yellow

380 R.D. GORDON & C. CANEPARI

with median 1/2 black, black area extended to anterior pronotal mar-

gin, lateral margins slightly emarginated with yellow; elytron with 2

large, yellow spots connected on apical declivity, 1 spot transverse on

disc anterior to middle, 1 transverse apical spot separated from suture

(Figs. 410, 411); antenna, mouthparts yellowish brown; propleuron

yellow; legs reddish yellow except femora dark brown with apical 1/6

reddish yellow; abdomen with median area dark brown, lateral 1/4

reddish yellow. Head punctures fine, separated by less than to twice

a diameter; pronotal punctures as large as on head, separated by a

less than to twice a diameter; elytral punctures larger than prono-

tal punctures, separated by less than to twice a diameter. Metaster-

num with punctures much larger than on elytron, sparse medially,

separated by less than a diameter laterally. Punctures on abdominal

sterna larger than on elytra, sparse on sterna 1-4, becoming fine,

dense laterally and on sterna 5-6; area on 1* abdominal sternum in-

side postcoxal arc shiny, coarsely, densely punctate. Clypeal apex

deeply emarginate, clypeus and frons smoothly, obliquely joined.

Epipleuron flat, slightly grooved internally, not descending external-

ly, femoral depressions moderate, not deep. Antenna with 10 articles

(Fig. 413). Prosternum with intercoxal carinae narrowly separated at

apex, convergent from apex toward base, joined before base, basally

stemmed, not reaching basal margin. Protibia widened apically. Post-

coxal line on 1% abdominal sternum not reaching posterior sternal

margin, slightly flattened along margin, apex recurved 1/4 distance

to basal sternal margin. Fifth sternum with apex broadly, feebly

emarginate; 6 sternum nearly truncate. Genitalia with basal lobe

3/4 as long as paramere, apex obliquely rounded, with large, angulate

projection on one margin; paramere oval, tapered from basal 1/4 to

rounded apex (Fig. 415); sipho slender, apex straight with membra-

nous area, basal capsule with inner arm long, wide, outer arm as

long as inner arm, basal border deeply emarginate (Figs. 416, 417).

° Similar to d except head black, clypeus reddish yellow; an-

terior margin of pronotum black. Genitalia with beak of basal unit

long, connecting duct short (Fig. 418).

Variation. Length 2.3 to 3.0 mm, width 1.7 to 2.4 mm. Elytron

with yellow spots broadly or narrowly connected, or elytron entirely

yellow except basal, sutural, and lateral border in apical 1/2 black

(Fig. 412); 2 head varies from clypeus narrowly reddish yellow to

having reddish yellow area extended onto frons.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 381

Type locality: of connectens, “Habitat in insula St. Eustachii”

(type not examined); of lengi:, Brownsville, Texas (USNM, lectotype

designated by GORDON 1985).

Geographical distribution: Southern United States south throu-

gh Central America and the Caribbean Islands to northern South

America.

Specimens examined: 57 (including 3 from Venezuela). Vene-

zuela. Barquisemento; ‘l‘ocayo (Present in most collections).

Remarks. The typical dorsal color pattern is distinctive for

the South American fauna. Many specimens were examined, nearly

all from areas other than northern South America. It appears that

H. connectens is marginally established in coastal Venezuela.

This is the only species belonging to Section II with 10-articled

antennae.

conclusa group

Hyperaspis species having 11-articled antennae; body dors-

oventrally flattened; frons and clypeus smoothly, obliquely joined;

epipleural depressions for reception of femoral apices moderate or

shallow, never deep; protibia widened apically; basal lobe of male

genitalia usually long, weakly asymmetrical (except H. elegantissima)

(Figs. 426, 443); and most species with incomplete prosternal cari-

nae. The extremely flattened H. longula and H. prolata are the only

species observed in any group with the posterior margin of the basal

abdominal sternum sinuate.

60. Hyperaspis ingrata (Mulsant, 1850)

Cleothera ingrata Mulsant, 1850: 644.

Hyperaspis ingrata: CROTCH 1874: 223; KoRscHEFsKy 1931: 190; BLACKWELDER 1945:

447,

Type locality. “Cayenne” (French Guiana) (MHNL, lectotype

here designated).

Description. ®, length 2.4 mm; body short, oval. Color

black except anterior border of pronotum yellowish red; elytron with

382 R.D. GORDON & C. CANEPARI

oval, elongate, reddish yellow spot (Fig. 419); mouthparts, legs, yel-

lowish red; ventral surface blackish brown. Pronotum and scutellum

with deep, coarse punctures separated by 2 or 3 times a diameter;

elytron with punctures as large as on pronotum, separated by 3 to

5 times a diameter. Clypeal apex emarginate. Epipleuron flat, not

internally grooved, not descending externally, femoral excavations

moderate, not deep. Postcoxal line on 1% abdominal sternum reach-

ing posterior sternal margin, flattened along margin, apex recurved.

Genitalia with beak of basal unit long, connecting duct short (Figs.

420, 421).

3 Not known.

Variation: none observed.

Geographical distribution: known only from the type locality.

Specimens examined: 1 (lectotype).

Remarks. This species is known only from the & lecto-

type, and is readily distinguished from other members of Section II

by the elytral color pattern and distribution.

The ? lectotype here designated to stabilize future usage of the

name is labeled “Cayenne, Lacordaire.”

61. Hyperaspis funesta (Germain, 1854)

Coccinella funesta Germain, 1854: 335.

Hyperaspis funesta: BRETHES 1923: 454; KORSCHEFSKY 1931: 189; BLACKWELDER

1945: 447.

Hyperaspis chilensis Crotch, 1874: 231; BRETHES 1923: 454 (as a synonym of funesta);

KORSCHEFSKY 1931: 189; BLACKWELDER 1945: 447; GORDON 1987: 29.

Type locality: of funesta, “provincia de Santiago” (Chile)

(MHNS); of chilensis, Chile (UMZC, lectotype designated by Gor-

DON 1987).

Description. 6, length 2.3 mm, width 1.7 mm; body

elongate, oval, flattened. Dorsal surface with head slightly aluta-

ceous, shiny; pronotum shiny; elytron shiny. Color black except head

yellow; pronotum black with anterior and lateral borders narrowly

yellow; elytron with 1 irregularly transverse, yellow spot medially

on lateral margin, spot extended internally 5/8 distance to sutural

margin (Figs. 422, 423); antenna, propleuron yellow; mouthparts,

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 383

epipleuron, proleg reddish yellow; propleuron, epipleuron yellow;

pro- and mesolegs reddish yellow; metaleg with femur dark brown;

abdomen dark brown except lateral 1/8 reddish yellow. Head punc-

tures fine, separated by less than to twice a diameter; pronotal

punctures larger than on head, separated by less than to twice a

diameter; elytral punctures as large as pronotal punctures, separated

by 1 to 2 times a diameter. Metasternum with punctures larger than

on elytron, punctures absent medially, lateral punctures dense, be-

coming contiguous. Punctures on abdomen coarse, dense on sterna

1-4, becoming fine, dense laterally and on sterna 5-6; area on 1%

abdominal sternum inside postcoxal arc shiny, coarsely, densely

punctate. Clypeal apex emarginate. Epipleuron flat, not internally

grooved, not descending externally, femoral excavations moderate,

not deep. Antenna with 11 articles (Fig. 424). Prosternum with in-

tercoxal carinae widely separated at apex, convergent toward base,

basally stemmed, reaching basal sternal margin. Protibia widened

apically. Postcoxal line on 1° abdominal sternum reaching posterior

sternal margin, flattened along margin, apex recurved 5/8 distance

to basal sternal margin. Fifth sternum with apex broadly, weakly

emarginate; 6 sternum weakly emarginate. Genitalia with basal

lobe nearly as long as paramere, apex narrowly rounded on one side,

with gradual, slightly angulate projection on one margin; paramere

oval, tapered from basal 1/4 to rounded apex; (Fig. 426); sipho slen-

der, long, apex with membranous area, basal capsule with inner arm

long, slender, outer arm shorter than inner arm, narrow, basal bor-

der slightly emarginate (Figs. 427, 428).

2 Similar to d except head black; anterior pronotal margin

black. Genitalia with beak of basal unit long, connecting duct short

(Fig. 429).

Variation. Length 1.9 to 2.4 mm, width 1.5 to 1.7 mm. Male

head entirely yellow or with black vertex; elytron with yellow spot

occasionally separated from lateral margin.

Geographical distribution: Argentina, Chile.

Specimens examined: 49. Argentina: Chubut, El Hoyo; Rio

Negro, El Bolson, Rio Ternero. Chile: Aconcagua, Illapel; Aconca-

gua, Papudo; Cauquenes; Coquimbo, Rio Los Molles; Lican Ray,

Cautin; Linares; Santiago, Apoquindo; Santiago, Guyacan Santiago,

Renca; Santiago, San Jose; Santiago, Cerra Calan; Valdivia, Panqui-

384 R.D. GORDON & C. CANEPARI

pulla; Victoria. (BM) (CAS) (CCM) (CMP) (CNC) (GG) (MNHS)

(SMTD) (USNM) (ZMHB).

Remarks. This species occurs in Chile and western Argen-

tina; it is the only species of Hyperaspis with a single yellow spot on

each elytron. The d genitalia differ little from those of H. nana, and

the 2 species are otherwise very similar. Hyperaspis nana might be

a color form of H. funesta, but that cannot be stated with certainty

at present.

When CrotcH (1874) described H. chilensis, he was obviously

unaware of GERMAIN’s (1854) description of H. funesta, because he

does not mention the latter name in his monograph. In addition,

Crotch’s type specimens of H. chilensis came from Germain himself.

The BM has two specimens of this species here designated paralec-

totypes. The first specimen is labeled “37318/ Reed/ Type/ Chili/

Hyperaspis chilensis Crotch/ Chilensis) Fry Coll 1905.100/?Syntype

Hyperaspis chilensis Cr. det. R. Booth 2004.” The second specimen

is labeled “Reed/ Chili/ Fry Coll 1905.100/ ?Syntype Hyperaspis

chilensis Cr. det. R. Booth 2004.” The specimens are considered

paralectotypes because they came from the Fry collection, which

Crotch certainly had available to him as evidenced by his descrip-

tions of several other species from that collection.

62. Hyperaspis nana Mader, 1957

Hyperaspis nana Mader, 1957: 82.

Type locality: Chile, Quillota (MHNS, holotype).

Description. 6, length 2.0 mm, width 1.4 mm; body

elongate, oval, slightly flattened. Dorsal surface with head slightly

alutaceous, shiny; pronotum shiny; elytron shiny. Color black ex-

cept head yellow with vertex black; pronotum black with anterior

and lateral borders narrowly yellow; elytron with lateral and apical

margins narrowly, irregularly bordered with yellow, yellow border

apically separated from elytral margin, 1 irregularly oval scutellar

spot at base, 1 large, oval, median discal spot posterior to middle

(Figs. 430, 431); antenna, mouthparts, propleuron, epipleuron yel-

low; proleg reddish yellow; mesoleg reddish yellow except posterior

1/2 of femur dark brown; metaleg dark brown; abdomen entirely

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 385

dark brown except basal sternum bright yellow outside of postcox-

al arc, sterna 2-6 with narrow lateral margin reddish yellow. Head

punctures fine, separated by less than to twice a diameter; pronotal

punctures larger than on head, separated by less than to twice a

diameter; elytral punctures as large as pronotal punctures, separated

by less than to twice a diameter. Metasternum with punctures larg-

er than on elytron, punctures nearly absent medially, lateral punc-

tures becoming contiguous. Punctures on abdomen coarse, dense on

sterna 1-4, becoming fine, dense laterally and on sterna 5-6; area

on 1* abdominal sternum inside postcoxal arc alutaceous, coarsely,

densely punctate. Clypeal apex emarginate. Epipleuron flat, inter-

nally grooved, not descending externally, femoral excavations mod-

erate, not deep. Antenna with 11 articles. Prosternum with inter-

coxal carinae narrowly separated at apex, convergent toward base,

basally stemmed, reaching basal sternal margin. Protibia widened

apically. Postcoxal line on 1" abdominal sternum reaching posterior

sternal margin, rounded along margin, apex recurved 1/2 distance

to basal sternal margin. Fifth sternum with apex broadly, weakly

emarginate; 6 sternum weakly emarginate. Genitalia with basal

lobe as long as paramere, apex narrowly rounded on one side, with

gradual, rounded projection on one margin; paramere oval, tapered

from basal 1/4 to rounded apex; (Fig. 432); sipho slender, long, apex

with membranous area, basal capsule with inner arm short, slender,

outer arm as long as inner arm, wide, curved, apically knobbed,

basal border deeply emarginate (Figs. 433, 434).

2 Similar to d except head black; anterior pronotal margin

black. Genitalia with beak of basal unit long (Figs. 435, 436).

Variation. Length 2.0 to 2.3 mm, width 1.4 to 1.7 mm. Elytron

varies from that described above to scutellar spot connected to lateral

yellow border, or scutellar spot connected to lateral border and feebly

connected to apical yellow border, with lateral border incomplete be-

hind humeral callus, leaving small, triangular, yellow spot at antero-

lateral angle of elytron, or discal spot and lateral border forming large,

yellow spot in apical 1/2, spot sometimes with small, central, black spot.

Geographical distribution: Chile.

Specimens examined: 22. Chile: Aconcagua, Rio Blanco; Acon-

cagua, Papudo; Angol; Los Petos, (illegible name); Santiago, Cerro

Calan, Las Condes; Chillan; Santiago, San Bernardo Valparaiso, La

386 R.D. GORDON & C. CANEPARI

Cruz. (CAS) (CCM) (GG) (MNHS) (USNM).

Remarks: MHyperaspis nana is the only Chilean species with

a scutellar spot on each elytron. The dorsal color pattern otherwise

resembles that of H. funesta.

63. Hyperaspis sphaeridioides Mulsant, 1850

Hyperaspis sphaeridioides Mulsant, 1850: 665; CrorcH 1874: 231; BrÈTtHES 1923:

454; KORSCHEFSKY 1931: 197; BLACKWELDER 1945: 448; GORDON 1987: 29.

Coccinella cruciata Germain, 1854: 336.

Hyperaspis cruciata: BRETHES 1923: 454 (as synonym of sphaeridioides); KORSCHEFSKY

1931: 197; BLACKWELDER 1945: 448.

Type locality: Chile. Type depository of sphaeridioides, MHNL

(lectotype here designated); of cruciata, MHNS (lectotype here des-

ignated).

Description. dé, length 2.4 mm, width 1.7 mm; body

elongate, oval, flattened. Dorsal surface with head slightly aluta-

ceous, shiny; pronotum alutaceous, slightly shiny; elytron shiny.

Color black except head yellow with vertex black; pronotum yellow

with large, black, basomedian spot, spot widely separated from an-

terior margin, lateral margin tapered from apex to base; elytron with

discal and apical yellow spots connected to median lateral and apical

lateral spots, forming 2 irregular bands (Figs. 437, 438); antenna,

propleuron, epipleuron yellow; mouthparts, legs brown; abdomen

entirely dark brown except lateral 1/6 brownish yellow. Head punc-

tures fine, separated by 2 to 3 times a diameter; pronotal punc-

tures larger than on head, separated by a diameter or less; elytral

punctures larger than pronotal punctures, separated by a diameter

or less. Metasternum with punctures larger than on elytron, absent

medially, lateral punctures becoming contiguous. Punctures on ab-

domen coarse, dense on sterna 1-4, becoming fine, dense laterally

and on sterna 5-6; area on 1" abdominal sternum inside postcoxal

arc alutaceous, coarsely, densely punctate. Clypeal apex emarginate.

Epipleuron flat, internally grooved, not descending externally, fem-

oral excavations shallow. Antenna with 11 articles (Fig. 441). Pros-

ternum with intercoxal carinae narrowly separated at apex, parallel

toward base, reaching 2/3 distance to sternal margin, not stemmed.

Protibia widened apically. Postcoxal line on 1% abdominal sternum

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 387

reaching posterior sternal margin, slightly flattened along margin,

apex recurved 1/2 distance to basal sternal margin. Fifth sternum

with apex truncate; 6" sternum weakly emarginate. Genitalia with

basal lobe slightly shorter than paramere, apex narrowly rounded on

one side, with gradual, slightly angulate projection on one margin;

paramere oval, tapered from basal 1/2 to rounded apex; (Figs. 443,

444); sipho slender, long, apex with membranous area, basal capsule

with inner arm short, slender, outer arm longer than inner arm, en-

larged apically, basal border deeply emarginate (Figs. 445, 446).

2 Similar to d except head black; anterior pronotal margin

black. Genitalia with beak of basal unit long, connecting duct short

(Figs. 447, 448).

Variation. Length 2.3 to 2.7 mm, width 1.7 to 1.9 mm. Elytron

varies from that described above to having elytral spots narrowly

connected (Figs. 439, 440), to having spots on elytron broadly con-

nected, forming large, lateral, yellow spot medially emarginated with

black on inner border.

Geographical distribution: Chile.

Specimens examined: 247. Chile: many localities (BM) (CAS)

(CCM) (CMP) (CNC) (GG) (MBR) (MNHS) (MSNG) (USNM).

Remarks. AHAyperaspis sphaeridioides is another strictly Chil-

ean species distinguished by the dorsal color pattern. Although vari-

able, pronotal and elytral patterns do not overlap variations found in

the other Chilean species.

The lectotype of H. sphaeridioides here designated to stabilize

future usage of the name bears no labels.

64. Hyperaspis conclusa Weise, 1906

Hyperaspis conclusa Weise, 1906: 197; KORSCHEFSKY 1931: 186; BLACKWELDER 1945:

446.

Hyperaspis flavolineata Mader, 1957: 83. (n. syn.)

Type locality: of conclusa, Argentina, Tucuman (MBR, lecto-

type here designated); of flavolineata, Bolivia, Lago Titicaca, Copa-

cabana, 3850 m (MHNS).

Description. 6, length 2.2 mm, width 1.5 mm; body

elongate, oval, flattened. Dorsal surface with head slightly alutaceous,

388 R.D. GORDON & C. CANEPARI

shiny; pronotum shiny; elytron shiny. Color black except head yel-

low; pronotum yellow with large, basomedian black spot, spot wide-

ly separated from anterior and lateral angles, 1 small, irregular, free

brown spot in lateral 1/4 of pronotum; elytron vittate, lateral and

apical margins with yellow border, median vitta extended from base

inside humeral callus to apical border, vitta enlarged at base, nar-

row toward apex (Figs. 449, 450); antenna, mouthparts, propleuron,

epipleuron yellow; legs reddish yellow; abdomen dark brown except

lateral 1/5 and sterna 5-6 reddish yellow. Head punctures fine, sepa-

rated by less than to twice a diameter; pronotal punctures larger than

on head, separated by a diameter or less; elytral punctures larger than

pronotal punctures, separated by a diameter or less. Metasternum

with punctures larger than on elytron, punctures sparse medially,

becoming dense, nearly contiguous laterally. Punctures on abdomen

slightly larger than on elytron, sparse on sterna 1-4, becoming fine,

dense laterally and on sterna 5-6; area on 1% abdominal sternum

inside postcoxal arc shiny, sparsely punctate. Clypeal apex emargin-

ate. Epipleuron flat slightly grooved internally, not descending ex-

ternally, femoral excavations shallow. Antenna with 11 articles (Fig.

453). Prosternum with intercoxal carinae widely separated at apex,

convergent toward base, basally stemmed, reaching basal sternal

margin. Protibia widened apically. Postcoxal line on 1% abdominal

sternum reaching posterior sternal margin, slightly flattened along

margin, apex recurved 5/8 distance to basal sternal margin. Fifth

sternum with apex broadly, weakly emarginate; 6 sternum weakly

emarginate. Genitalia with basal lobe as long as paramere, apex nar-

rowly rounded on one side, with gradual, rounded projection on

one margin; paramere oval, tapered from basal 1/4 to rounded apex;

(Fig. 454); sipho slender, long, apex with membranous area, basal

capsule with inner arm long, slender, outer arm shorter than inner

arm, wide, basal border emarginate (Figs. 455, 456).

2 Similar to d except head black; anterior pronotal margin

black, basomedian black spot enlarged laterally to include free spot.

Genitalia with beak of basal unit short, connecting duct short (Fig.

457).

Variation. Length 1.9 to 2.2 mm, width 1.4 to 1.5 mm. Pro-

notum may have lateral free spot absorbed by basomedian spot or

be narrowly connected. Elytral ground color varies from black to

brown; central vitta and lateral yellow border vary in width (Figs.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 389

451, 452) with a single specimen from Valchete, Argentina, having

vittae so wide the elytral ground color is yellow with brown vitta.

Geographical distribution: Southern South America.

Specimens examined: 33. Argentina: Cajamarca; Chaco, Fonta-

na; Cordoba, Alta Gracia; Jujuy, San Juancito; La Rioja; Mendoza,

Poterillos; Salta, Guemes; Valchete. Bolivia: Lago Titicaca, Copa-

cabana; Saavedra, Santa Cruz. Brazil: Corumba. Uruguay: Artigas,

Colonia Palma; Carrasco (BM) (CAS) (CMP) (MBR) (MHNS)

(USNM).

Remarks. Vittate species are rare in the South American

Hyperaspis fauna, and H. conclusa resembles only H. elegantissima in

this character. The latter species has very narrow elytral vittae, and

d genitalia with a short basal lobe strongly angulate on one margin.

Hyperaspis conclusa has wide elytral vittae, and male genitalia with a

basal lobe as long as the paramere, feebly angulate on one margin.

A single 2 specimen from Venezuela appears to be this species, but

more likely represents an undescribed species.

The £ holotype of Hyperaspis flavolineata Mader appears to be

identical to typical specimens of H. conclusa.

The lectotype of H. conclusa, here designated to stabilize fu-

ture usage of the name, is labeled “Rep. Argentina, Prov. Tucuman,

5-III-1900, C. Bruch/TYPUS/Hyperaspis conclusa Weise”. A para-

lectotype in the ZMHB is bears the same locality label with addi-

tional labels “Hyperaspis conclusa W./Syntypus Hyperaspis conclusa

Weise, 1906, labeled by MNHUB 2004 (red paper)”.

65. Hyperaspis arida n. sp.

Type material. Holotype d: Peru, Lambayeque, Roadside veg.,

1 mile S.E. of town, 20.viii.1971, fertile irrigate (sic !) region in arid

coastal desert, P.S. & H.L. Broomfield, B.M. 1971-486. (BM). Al-

lotype 2: same data as holotype (BM). Paratypes, 21: same data as

holotype. (BM) (USNM).

Description. 6, length 2.4 mm, width 1.8 mm; body

form oval, slightly flattened. Dorsal surface with head alutaceous,

feebly shiny: pronotum alutaceous, slightly shiny: elytron shiny. Col-

or yellow except head with black vertex; pronotum with basomedian

390 R.D. GORDON & C. CANEPARI

black spot, spot widely separated from anterior margin, with antero-

lateral, earlike projection, narrowed from apical 1/3 to base; elytron

with base near scutellum and sutural margin black, 3 median spots

present from base to apex, black sutural margin irregular, slightly

expanded at apical declivity, 1 elongate spot from base to just across

humeral callus, large, median spot at apical declivity, and 1 small,

transverse, subapical spot joined to sutural margin (Fig. 458); pro-,

meso-, metasterna black; abdomen dark brown except outer 1/4 and

sterna 4-6 yellow. Head punctures fine, separated by less than to

twice a diameter; pronotal punctures larger than on head, separated

by a diameter or less; elytral punctures larger than pronotal punc-

tures, separated by a diameter or less. Pro- and mesosterna very

coarsely punctured medially, punctures nearly contiguous. Metaster-

nal punctures larger than on elytron, separated by a diameter me-

dially, nearly contiguous laterally. Punctures on abdominal sterna

1-4 coarse, dense medially, becoming dense, fine laterally; sterna 5-6

finely, densely punctured throughout; area on 1% abdominal sternum

inside postcoxal arc alutaceous, coarsely, densely punctured. Clypeal

apex emarginate. Epipleuron flat, grooved internally, not descending

externally, femoral depressions shallow, nearly absent. Prosternum

with intercoxal carinae narrowly separated at apex, convergent near-

ly to basal margin, with short stem reaching margin. Protibia wid-

ened apically. Postcoxal line on 1% abdominal sternum not extended

to posterior sternal margin, slightly flattened along margin, apex re-

curved 2/3 distance to basal sternal margin. Fifth sternum with apex

truncate; 6" sternum truncate. Genitalia with basal lobe 3/4 length

of paramere, apex narrowly, obliquely rounded, with large, weakly

angulate projection on one side; paramere slender, tapered to round-

ed apex in apical 3/4 (Fig. 459); sipho robust, straight apically, apex

with membranous area, basal capsule with inner arm short, wide,

apically knobbed, outer arm wide, as long as inner arm, apically

knobbed (Figs. 460, 461).

° Similar to d except head black. Genitalia with basal unit

slender, curved, beak composed of small, winglike projection on

each side (Fig. 463).

Variation. Length 2.3 to 3.6 mm, width 2.4 to 2.6 mm. Elytron

often with basal border entirely black from sutural margin to spot

at humeral callus, incorporating that spot; remaining elytral spots

vary slightly in size.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 391

Etymology. The specific name is the Latin aridus, meaning dry,

referring to the Peruvian coastal desert type locality.

Remarks. Superficially this species is a pale version of H. fe-

stiva. The pronotal maculation is particularly similar, but the shal-

low femoral depressions on the epipleuron, extremely coarse, dense

pro- and mesosternal punctation, elytra with black spots on a yellow

ground color, and basal unit of female spermathca with the peculiar

winged beak distinguish MH. arida.

66. Hyperaspis elegantissima Brèthes, 1925

Hyperaspis elegantissima Brèthes, 1925a: 206; KORSCHEFSKY 1931: 188; BLACKWELDER

1945: 447.

Type locality: Argentina, Estancia La Noria, Rio San Javier,

Santa Fé (BM, holotype).

Description. 6, length 2.0 mm, width 1.5 mm; body

elongate, oval, convex. Dorsal surface with head slightly alutaceous,

shiny; pronotum slightly alutaceous, shiny; elytron shiny. Color

black except head yellow with black vertex; pronotum with anterior

margin narrowly yellow, median black area with large, lateral ear-

like projection in lateral 1/4; elytron with lateral and apical margins

narrowly yellow, narrowly sinuate vitta on elytron from base inside

humeral callus to apical yellow border (Fig. 466); antenna, propleu-

ron, epipleuron yellow; mouthparts, pro- and mesoleg brownish

yellow; metaleg dark brown; abdomen with median area of sterna

1-5 dark brown, lateral 1/4 and sterna 5-6 dark reddish yellow.

Head punctures fine, separated by less than to twice a diameter;

pronotal punctures larger than on head, separated by a diameter or

less; elytral punctures larger than pronotal punctures, separated less

than to twice a diameter. Metasternum impunctate medially, lateral

punctures larger than on elytron, separated by less than a diameter.

Punctures on all abdominal sterna larger than on elytra, sparse on

sterna 1-2, becoming more dense on sterna 3-6; area on 1% abdomi-

nal sternum inside postcoxal arc shiny, coarsely, sparsely punctured.

Clypeal apex deeply emarginate, clypeus and frons joined at abrupt

angle. Epipleuron flat, not internally grooved, not descending exter-

nally, femoral excavations nearly absent, only slight depressions pres-

392 R.D. GORDON & C. CANEPARI

ent. Antenna with 11 articles (Fig. 467). Prosternum with intercoxal

carinae narrowly separated at apex, convergent from apex toward

base, basally stemmed, reaching basal margin. Protibia widened api-

cally. Postcoxal line on 1% abdominal sternum not reaching posterior

sternal margin, flattened along margin, apex recurved 1/2 distance

to basal sternal margin. Fifth sternum with apex truncate; 6" ster-

num truncate. Genitalia with basal lobe 3/4 as long as paramere,

apex narrow, obliquely rounded, with large, angulate projection on

one margin; paramere wide, tapered from middle to rounded apex;

(Fig. 468); sipho slender, apex straight, basal capsule with inner arm

long, slender, outer arm as long as inner arm, basal border deeply

emarginate (Figs. 469, 470).

2 Similar to 6 except head black; anterior margin of pronotum

black, lateral margin narrowly yellow, median black area without

earlike projection. Genitalia with beak of basal unit long, connecting

duct short (Fig. 471).

Variation. Length 1.8 to 2.0 mm, width 1.4 to 1.5 mm. Vitta

on elytron often broken medially, width slightly variable (Figs. 464,

465).

Geographical distribution: Argentina.

| Specimens examined: 5. Argentina: San Juan, San Juan City;

San Luis, San Juan, Jachal; San Luis City; Salta; Salta, Rosario de

Lerma. (USNM) (ZMHB).

Remarks. ‘This is one of two truly vittate species in South

America. It has typical characteristics of the conclusa group except

for male genitalia, which have a basal lobe of the type found in

the onerata Group. BRETHES (1925a) aptly named this species H.

elegantissima.

BRETHES (1925b) specifically stated that he had a single speci-

men; therefore, that specimen is the holotype. The holotype in the

BM collection is labeled “Type (orange bordered disc)/ Estancia la

Noria, Rio San Javier, Santa Fe, Argentine/ G. E. Bryant 6.1.1912/

Hyperaspis elegantissima Brèthes/ J. Brèthes det. 1924.”

67. Hyperaspis longula Weise, 1922

Hyperaspis longula Weise, 1922: 35; KorscHEersky 1931: 101; BLACKWELDER 1945:

447,

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 393

Type locality: Argentina, Buenos Aires (MBR, lectotype here

designated).

Description. d, length 2.6 mm, width 1.5 mm; body

elongate, slender, nearly parallel sided, flattened. Dorsal surface

with head alutaceous, feebly shiny; pronotum strongly alutaceous,

dull; elytron alutaceous, slightly shiny. Color black except head yel-

low; pronotum black with lateral 1/6 yellow, yellow border emar-

ginated by broad, earlike projection of black area; elytron with 3

yellow spots, 1 triangular spot on anterior portion of disc, 1 sinuate

spot on lateral margin medially, 1 transversely oval, subapical spot

(Figs. 472, 473); antenna, mouthparts, propleuron yellow; epipleu-

ron reddish yellow; abdomen with median area dark brown, lateral

1/4 reddish brown. Head punctures fine, separated by a diameter

or less; pronotal punctures larger than on head, separated by a di-

ameter or less; elytral punctures larger than pronotal punctures,

separated by a diameter or less. Metasternum with punctures larger

than on elytron, punctures separated by a diameter medially, lat-

eral punctures separated by less than a diameter. Punctures on ab-

domen coarse, dense on sterna 1-4, becoming fine, dense laterally

and on sterna 5-6; area on 1" abdominal sternum inside postcoxal

arc alutaceous, coarsely, densely punctate. Clypeal apex emarginate.

Epipleuron flat, not internally grooved, not descending externally,

femoral excavations shallow, nearly absent. Antenna with 11 articles

(Fig. 476). Prosternum with intercoxal carinae narrowly separated

at apex, parallel 5/8 distance to anterior prosternal margin, not con-

vergent, not stemmed, not reaching basal margin. Protibia widened

apically. Basal abdominal sternum with apical margin sinuate. Post-

coxal line on 1% abdominal sternum reaching posterior sternal mar-

gin, flattened along margin, apex abruptly recurved, nearly reaching

basal sternal margin. Fifth sternum with apex broadly, distinctly

emarginate; 6 sternum feebly emarginate. Genitalia with basal lobe

nearly as long as paramere, apex narrow, obliquely rounded, with

angulate projection on one margin; paramere narrow, tapered from

basal 1/4 to rounded apex; (Fig. 477); sipho slender, long, apex with

membranous area, basal capsule with inner arm long, slender, outer

arm much shorter than inner arm, widened to apex, basal border

emarginate (Figs. 478, 479).

2 Similar to d except head black. Genitalia with beak of basal

unit long, connecting duct short (Fig. 480).

394 R.D. GORDON & C. CANEPARI

Variation. Length 2.6 to 3.0 mm, width 1.5 to 1.7 mm. Elytral

color pattern varies from that described above to having the discal

and lateral spots narrowly connected, or with discal and apical spots

broadly connected (Fig. 474), and lateral margin narrowly bordered

with yellow, connected to subapical spot, or lateral margin yellow,

connected to apical spot with discal spot free (Fig. 475).

Geographical distribution: Argentina, Brazil.

Specimens examined: 22. Argentina: Buenos Aires; Buenos

Aires, Delta; Buenos Aires, San Fernando; Chaco, Dep. Resisten-

cia; Misiones; Tigre. Brazil: Corumba; Rio de Janeiro (BM) (CMP)

(MBR) (USNM). |

Remarks. AHAyperaspis longula is one of the few long, slen-

der, flattened species in South America. It is distinguished by the

body shape; short prosternal carinae; sinuate apical margin of the

basal abdominal sternum; and nearly complete postcoxal line on

basal sternum. See remarks under H. prolata.

The lectotype, here designated to stabilize usage of the name,

bears the labels “Rep. Argentina, Prov. Buenos Aires, 10-X-1906,

C. Bruch/ Typus/ Hyperaspis longula Ws.”.

68. Hyperaspis prolata n. sp.

Type material. Holotype d: Argentina, Salta, Guemes (MBR).

Allotype 2: Bolivia, Dept. S. Cruz (Santa Cruz), 450 m, Umg Bue-

navista, Steinback coll., Hyperaspis n. sp., Korschefsky Collection

1952 (USNM). Paratypes, 2. Argentina, Chaco, Dep. Resistencia,

X-XII-935 J. B. Daguerre 39749; Brazil, Chapada, Acc. No. 2966,

Augst. (CMP) (CMBR).

Description. 6, length 2.6 mm, width 1.6 mm; body

elongate, slender, nearly parallel sided, flattened. Dorsal surface with

head strongly alutaceous, dull: pronotum strongly alutaceous, dull;

elytra alutaceous, weakly shiny. Color black except head yellow with

black vertex; pronotum yellow with median, black, transverse spot,

basal margin of black spot deeply scalloped with slender, median

stem extended to margin anterior to scutellum; elytron with yellow

lateral border in basal 2/3, border enlarged at apex, 1 round discal

spot near suture anterior to middle, 1 slender, comma shaped api-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 395

cal spot at apex (Fig. 481, 482); antenna, mouthparts, legs reddish

yellow; propleuron, epipleuron yellow; abdomen with sterna 1-4

dark brown except lateral 1/4 and sterna 5-6 reddish yellow. Head

punctures fine, separated by less than to twice a diameter; pronotal

punctures larger than on head, separated by less than to twice a di-

ameter; elytral punctures larger than pronotal punctures, separated

by 1 to 2 times a diameter. Metasternal punctures larger than on

elytron, sparse medially, dense, contiguous laterally. Punctures on

abdominal sterna larger than on elytra, dense medially, becoming

fine, dense laterally, sterna 5-6 finely densely punctured through-

out; area on 1* abdominal sternum inside postcoxal arc alutaceous,

coarsely, sparsely punctured. Clypeal apex emarginate. Epipleuron

flat, feebly grooved internally, not descending externally, femoral

depressions shallow. Antenna with 11 articles (Fig. 483). Protibia

widened apically. Prosternum with intercoxal carinae narrowly sepa-

rated at apex, parallel 5/8 distance to anterior prosternal margin, not

convergent, not stemmed, not reaching basal margin. Postcoxal line

on 1* abdominal sternum reaching posterior sternal margin, flat-

tened along margin, apex abruptly recurved 3/4 distance to basal

sternal margin. Basal margin of 1* abdominal sternum medially sin-

uate. Fifth sternum with apex broadly, weakly emarginate; 6° ster-

num truncate. Genitalia with basal lobe as long as paramere, apex

truncate, without projection on one margin; paramere wide, tapered

from basal 1/2 to rounded apex (Fig. 485); sipho long, slender, api-

cally straight, with apical membranous area, basal capsule with in-

ner arm long, slender, pinched near base, outer arm shorter than

inner arm, wide, basal border slightly emarginate (Figs. 486, 487).

2 Similar to d except vertex of head broadly black, with nar-

row, median projection extended anteriorly to clypeal apex. Genita-

lia with beak of basal unit short, wide, with lateral earlike projec-

tions (Figs. 488-490)

Variation. Length 2.6 to 2.8 mm, width 1.6 to 1.8 mm. Head of

d paratype with black vertex narrowly, medially emarginated with

yellow.

Etymology. The specific name is the Latin prolatus, meaning

elongated, referring to the long, slender body form.

Remarks. Hyperaspis longula and H. prolata are morpho-

logically similar species sharing several synapomorphies such as flat,

396 R.D. GORDON & C. CANEPARI

elongate, slender body; parallel prosternal carinae not reaching basal

prosternal margin; sinuate apical margin of basal abdominal ster-

num; and postcoxal line on 1% abdominal sternum abruptly recurved

apically, extended nearly to basal sternal margin. ‘They differ from

each other in type of male genitalia, and externally by the pronotal

color pattern, with H. prolata having a central black spot and the

posterior border scalloped; H. longula having a large, basomedian

black spot at basal margin of pronotum.

Genus Tenutsvalvae Duverger, n. stat.

Hyperaspis (Tenuisvalvae) Duverger, 1989: 148.

Type species: Hyperaspis bisquinquepustulata (F.) (H. raynevali

Mulsant), by monotypy.

Description. Hyperaspini with description as for Hyper-

aspis except antenna always with 11 articles (Figs. 493, 497, 501,

509, 516, 524, 531, 538, 545); ultimate maxillary article slightly

emarginate apically; mandibulary retinaculum strongly curved, apex

rounded; clypeus and frons joined at feeble angle; protibia nar-

row, not flanged; metendosternite with anterior tendons of fork not

curved toward middle (Fig. 546); postcoxal line on 1° abdominal

sternum extended to, or nearly to, posterior sternal margin, apex re-

curved; apex of female abdominal sternum 6 triangular; 9 genitalia

with genital plates extremely long, slender (Fig. 494).

Remarks. DUVERGER (1989) listed a series of differentiat-

ing characters for Hyperaspis and Tenuisvalvae derived from a com-

parison of Hyperaspis galliae Duverger and Tenuisvalvae raynevalit

Mulsant (bisquinquepustulata F.). Some of these characters fail when

examples of a variety of species of both genera are examined; char-

acters considered to be of value are used in the above description.

No good character for separating males of these two genera has been

found. The form of the apical maxillary article may be useful but

is extremely difficult to see without dissection, and metendoster-

nite differences are not consistent for the entire fauna. Characters

used by Duverger that fail when many species are compared are the

shape of the siphonal capsule and form of the postcoxal line. ‘The

postcoxal line is said by Duverger to be of the Parascymnus Chapin

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 397

type. However, examination of the postcoxal line of T. raynevali

(type species) shows it apically recurved as in Hyperaspis, not api-

cally parallel to the hind sternal margin as in Parascymnus.

Ten species are recognized as members of this genus.

KEY TO SPECIES OF TENUISVALVAE

1(2). Elytron red with narrow black border on all margins

eXcèpi*sutura? maree atterit, dt . BION 10. gnoma, n. sp.

Elytron 2th: pale or Gark Sp Ots cos, 06h ety eu 2

2(1). Elytral color primarily black or brown with yellow or

orange, spots.(ttur cyst Gee See SIRIA RT 3

Elytral color primarily yellow with black or brown spots 8

#2. Rach elytroa witte deer die noel ee Sere ee 4

Each. elytrons ath? 2 0r more SPOSA. 4. arteria è 5

4(3). Elytron with spot at center of disc (Fig. 491).........

en e et - 1. unipunctata (Crotch)

Elytron with spot at extreme apex (Fig. 495).........

i'glanan 100% HMI 5 10090 RANROESI 2. peregrina (Mulsant)

5(3). Elytron with 2 spots, one median and one at apex (Fig.

PIO) crearti eet TTI 3. deyrollei (Crotch)

Elytron with, 3 orrmorespotssi said soy bud Sesto 6

6(5). Elytron with 5 spots (Fig. 513) .. 5. bisquinquepustulata (F.)

Elytron,avith. è SP048 matt arena lana 7

7(6). Elytron with one elongate discal spot, one triangu-

lar spot laterally at middle and one large, somewhat

rounded apical spot (Fig. 426)........ 8. caucaensis, n. Sp.

Elytron with small basoscutellar spot, one humeral spot

at base, and an elongate, curved spot at apex (Fig. 543)

ere wer a rai 9. bromelicola (Sicard)

8(2). Elytral suture black; each elytron with black, c-shaped

macula extended from center of suture posteriorly to

398 R.D. GORDON & C. CANEPARI

apical 1/4, and slender, elongate spot extended from

base inside humerus to beyond humeral callus, someti-

mes connected to median c-shaped mark (Fig. 629)....

Ra AEM SI Rari spe pià 7. parenthesis, n. sp.

Elytron variably marked but never with c-shaped macula 9

9(8). Elytron with 3 discrete black spots (Fig. 506) ........

era) dare: Of promotes: - a. 4. notata (Mulsant)

Elytron with 2 discrete black spots (Fig. 522) ........

AR RL ion, La 6. ecoffeti (Mulsant)

1. Tenuisvalvae unipunctata (Crotch, 1874), n. comb.

Hyperaspis unipunctata Crotch, 1874: 230; KorscHEFsky 1931: 199; BLACKWELDER

1945: 448; GORDON 1987: 20.

Type locality. Santarem (Brazil) (UMZC, holotype, GORDON

1987).

Description. &, length 2.3 mm, width 1.4 mm; body

form round, convex. Dorsal surface with head alutaceous, dull;

pronotum slightly alutaceous, elytron shiny. Color black except

pronotum with small, triangular, anterolateral spot nearly reaching

basal margin; elytron with large, irregularly rounded, yellow spot

just anterior to middle of elytron (Figs. 491, 492); antenna, mouth-

parts, and legs reddish brown except protibia yellow; propleuron

yellow. Head punctures fine, separated by a diameter or less. Pro-

notal punctures larger than on head, separated by a diameter or

less; elytral punctures larger than pronotal punctures, separated by

a diameter or less. Metasternal punctures larger than on elytron,

sparse medially, dense, nearly contiguous laterally. Punctures on

abdominal sterna 1-4 coarse, sparse medially, becoming dense,

very fine laterally; sterna 5-6 finely densely punctured throughout;

area on 1%“ abdominal sternum inside postcoxal arc shiny, coarsely,

sparsely punctured. Clypeal apex emarginate. Epipleuron oblique,

grooved internally, externally descending, femoral excavations

deep. Antenna with 11 articles (Fig. 493). Prosternum with inter-

coxal carinae convergent toward base, basally stemmed, reaching

basal margin. Postcoxal line on 1** abdominal sternum extended to

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 399

posterior sternal margin, flattened along margin, apex recurved 1/3

distance to basal sternal margin. Fifth sternum with apex broadly,

feebly emarginate. Genitalia with beak of basal unit short, wide,

connecting duct short (Fig. 494).

3 Not known.

Geographical distribution: Brazil, Trinidad.

Specimens examined: 6. Brazil: Santarem. Trinidad: Icacos, St.

Augustine. (UMZC) (USNM).

Remarks. In color pattern, T. unipunctata closely resem-

bles Hyperaspis binotata (Say) and similar species of the eastern

United States. he type locality is Amazonian Brazil, but all other

specimens examined were from Trinidad, a somewhat unusual com-

bination of localities. However, the type specimen and the Trinidad

specimens match in all characters, including identical 9 genitalia.

2. Tenuisvalvae peregrina (Mulsant, 1850), n. comb.

Hyperaspis peregrina Mulsant, 1850: 691; CrotcH 1874: 230; KorscHEFsKy 1931:

194; BLACKWELDER 1945: 448; GORDON 1987: 29.

Hyperaspis subapicalis Crotch, 1874: 229; KoRscHEFsKy 1931: 197; BLACKWELDER,

1945: 448; GoRDON 1987: 29. (n. syn.).

Type locality: of peregrina, Brazil (UMZC, lectotype designated

by GORDON 1987); of subapicalis, Rio de Janeiro, Brazil (UMZC,

lectotype designated by GORDON 1987).

Description. &, length 3.0 mm, width 2.5 mm; body

form round, convex. Dorsal surface with head dull, strongly aluta-

ceous, pronotum slightly less dull than head, elytra feebly shiny.

Color black except pronotum with anterolateral 1/4 yellow, inner

margin of yellow area rounded; elytron with large, round, subapical

yellow spot (Figs. 495, 496); antenna, mouthparts, tibiae and tarsi

reddish yellow; propleuron yellow; abdomen black except outer 1/4

and sternum 6 reddish yellow. Head punctures fine, nearly hidden

in alutaceous sculpture, separated by less than to 3 times a diameter.

Pronotal punctures larger than on head, separated by a diameter or

less; elytral punctures larger than pronotal punctures, separated by

less than to twice a diameter. Metasternal punctures much larger than

on elytron, dense medially, nearly contiguous laterally. Punctures on

400 R.D. GORDON & C. CANEPARI

abdominal sterna 1-4 coarse, sparse medially, becoming dense, very

fine laterally; sterna 5-6 finely densely punctured throughout; area

on 1*t abdominal sternum inside postcoxal arc strongly alutaceous,

finely punctured. Clypeal apex deeply emarginate, clypeus and frons

joined at abrupt angle. Epipleuron oblique, not internally grooved,

descending externally, femoral excavations deep. Antenna with 11

articles (Fig. 497). Prosternum with intercoxal carinae tapered to-

ward base, basally stemmed, reaching basal margin. Postcoxal line

on 1% abdominal sternum extended to posterior sternal margin,

slightly flattened along margin, apex recurved 1/2 distance to basal

sternal margin. Fifth sternum with apex broadly, distinctly emargin-

ate. Genitalia with beak of basal unit short, wide; connecting duct

short (Fig. 498).

4. Similar to 9 except head yellow; pronotum with lateral 1/4

yellow. d genitalia with basal lobe 5/8 length of paramere, apex

narrowly, obliquely rounded, with large, rounded projection on one

side; paramere wide, tapered to rounded apex in apical 1/2; sipho

long, slender, sinuate in apical 1/4, apex with membranous area,

basal capsule with inner arm long, slender, outer arm slender, short-

er than inner arm, base emarginate.

Variation. None observed.

Geographical distribution: Brazil.

Specimens examined: 3. Brazil. Rio de Janeiro; Rio de Janeiro,

Campo Grande. (USNM).

Remarks. Tenuisvalvae peregrina is characterized by a sin-

gle, yellow, subapical spot on each elytron. CROTCH himself (1874)

remarked on the similarity between H. peregrina and subapicalis. Af-

ter comparing type specimens, we conclude that they represent both

sexes of the same species.

The BM collection has two syntypes of H. subapicalis that we

designate as paralectotypes. The first syntype is labeled “TYPE/ Fry

Rio Jano/ subapicalis/ Fry Coll. 1905.100./ Type (orange bordered

circle)/ Hyperaspis subapicalis Crotch Type Brazilia/ Syntype Hy-

peraspis subapicalis Cr. det. R.G. Booth 2004.” ‘The second syntype

is labeled “TYPE/ Fry Rio Jano/ Fry Coll. 1905.100./ SYNTYPE

(blue bordered circle)/ Syntype Hyperaspis subapicalis Cr. det. R.G.

Booth 2004”.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 401

3. Tenuisvalvae deyrollei (Crotch, 1874), n. comb.

Hyperaspis deyrollei Crotch, 1874: 229; KORSCHEFSKY 1931: 187; BLACKWELDER 1945:

446; GORDON 1987: 29.

Type. locality, Teapa,..5...Paula;: (Sao, Paulo). (Brazil), (OMZC,

lectotype designated by GORDON 1987).

Description. 6, length 2.5 mm, width 1.9 mm; body

form round, convex. Dorsal surface with head dull, strongly aluta-

ceous, pronotum slightly less dull than head, elytra shiny. Color black

except head entirely yellow; pronotum mostly reddish yellow with

black basomedian macula, lateral margins of macula medially emar-

ginated by pale area; elytron with 2 large, reddish yellow spots, spot

on disc raggedly rounded, apical spot transverse (Fig. 499, 500); an-

tenna, mouthparts, legs yellow; propleuron yellow; abdomen reddish

yellow except median 1/2 of sterna 1-3 dark brown. Head punctures

fine, nearly hidden in alutaceous sculpture, separated by less than to

twice a diameter. Pronotal punctures larger than on head, separated

by less than to twice a diameter; elytral punctures larger than prono-

tal punctures, separated by less than to twice a diameter. Metaster-

nal punctures much larger than on elytron, sparse medially, dense

laterally. Punctures on abdominal sterna 1-4 coarse, sparse medially,

becoming dense, very fine laterally; sterna 5-6 finely densely punc-

tured throughout; area on 1% abdominal sternum inside postcoxal

arc slightly shiny, coarsely, densely punctured. Clypeal apex deeply

emarginate, clypeus and frons joined at abrupt angle. Epipleuron

oblique deeply grooved internally, descending externally femoral ex-

cavations deep. Antenna with 11 articles (Fig. 501). Prosternum with

intercoxal carinae convergent, basally stemmed, reaching basal mar-

gin. Postcoxal line on 1% abdominal sternum extended to posterior

sternal margin, slightly flattened along margin, apex recurved 1/2

distance to basal sternal margin. Fifth sternum with apex broadly,

feebly emarginate; 6 sternum broadly, feebly emarginate. Genitalia

with basal lobe 3/4 length of paramere, with large projection on one

side, paramere wide, feebly tapered to rounded apex (Fig. 502); sipho

slender, apex curved upward with internal membranous area, basal

capsule with inner arm broad, short, outer arm short (Figs. 503, 504).

2 Similar to d except head black; pronotum with median black

area extended nearly to apex. Genitalia with beak of basal unit short;

connecting duct long (Fig. 505).

402 R.D. GORDON & C. CANEPARI

Variation. Length 2.3 to 2.7 mm, width 1.8 to 2.1 mm. Me-

dian dark area on pronotum varies slightly in size; color of dorsal

maculation varies from very dark reddish yellow to yellow and the

lectotype has a red discal spot.

Geographical distribution: Southeastern South America.

Specimens examined: 110. Argentina: Bella Vista; Buenos Aires;

Burzaco Sur; Corrientes; Delta; Entre Rios; Haedo; Loreto; Misio-

nes; Rio Chama; Corrientes, Santo Tomé; Salto; Tigre. Uruguay:

Montevideo (CCM) (MBR) USNM).

Remarks. Tentusvalvae deyrollei is frequently collected

in Argentina where it is often found in collections under the name

“Hyperaspis muhni Brèthes”. 'This is apparently a manuscript name,

as no published record of it can be located. The 2 elytral spots and

mostly reddish yellow pronotum distinguish this species.

4. Tenuisvalvae notata (Mulsant, 1850), n. comb.

Cleothera notata Mulsant, 1850: 550.

Hyperaspis notata: CROTCH 1874: 227; KORSCHEFSKY 1931: 192; BLACKWELDER 1945:

447; GORDON 1987: 29.

Type locality: Brazil (UMZC, lectotype designated by GORDON

1987).

Description. 46, length 3.2 mm, width 2.6 mm; body

form broadly round, convex. Dorsal surface slightly shiny, aluta-

ceous. Color yellow except pronotum with 3 black maculae, 1 nar-

row, irregular basomedian macula, and 2 triangular median maculae;

elytron with 4 spots and sutural margin black, 1 oval, discal spot on

suture, 1 oval spot anterior to and laterad of discal spot, 1 comma

shaped spot medially near lateral margin, 1 irregularly oblong spot

on apical declivity, sutural margin narrowly black anterior to dis-

cal spot, and widely black posterior to discal spot; scutellum black

(Fig. 506); pro-, meso-, and metasterna black; median area of basal

3 abdominal sterna dark brown. Head punctures fine, nearly hidden

in alutaceous sculpture, separated by less than to 3 times a diameter.

Pronotal punctures larger than on head, separated by less than to

twice a diameter; elytral punctures larger than pronotal punctures,

separated by less than to twice a diameter. Metasternal punctures

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 403

fine, sparse medially, becoming much larger than on elytron, dense

laterally. Punctures on abdominal sterna 1-4 coarse, sparse medi-

ally, becoming dense, very fine laterally; sterna 5-6 finely densely

punctured throughout; area on 1° abdominal sternum inside post-

coxal arc shiny, finely, indistinctly punctured. Clypeal apex deeply

emarginate, clypeus and frons joined at abrupt angle. Epipleuron

oblique, not internally grooved, descending externally, femoral exca-

vations deep. Antenna with 11 articles (Fig. 509). Prosternum with

intercoxal carinae convergent medially, basally stemmed, reaching

basal margin. Postcoxal line on 1% abdominal sternum extended to

posterior sternal margin, flattened along margin, apex recurved 2/3

distance to basal sternal margin. Fifth sternum with apex broadly,

feebly emarginate; 6" sternum broadly, feebly emarginate. Genitalia

with basal lobe as long as paramere, slender, with feeble lateral pro-

jection on one side, paramere wide, feebly tapered to rounded apex

(Figs. 507, 508); sipho slender, straight, apex with internal mem-

branous area, basal capsule with inner arm slender, short, outer arm

wide, longer than inner arm (Figs. 510, 511).

? Similar to d except head with triangular black spot on each

side of vertex. Genitalia with beak of basal unit short; connecting

duct short (Fig. 512).

Variation. Length 3.0 to 3.4 mm, width 2.4 to 2.7 mm. Prono-

tal maculae vary somewhat in size; most Colombian specimens have

the median spots fused with basomedian macula; elytral spots vary

slightly in size, but remain quite constant in shape.

Geographical distribution: Central and northern South America.

Specimens examined: 36. Bolivia: Loma Alta. Brazil: Bahia; Rio

de Janeiro, Campo Grande; Corumba; Sta. Lucia; Sao Paulo, Cam-

pinas. Colombia: Valle del Cauca, Pto. ‘Tejada; Antioquia, Rio Ne-

gro (BM) (CMNH) (USNM).

Remarks. Pronota and elytra with yellow backgrounds and

black maculation occur frequently in species of Hyperaspidini and

especially Brachiacanthini. Spot patterns are repeated in several ge-

neric taxa, creating a confusing situation. Tenuisvalvae has 3 species

with this type of pattern, 7. parenthesis, T. notata, and T. ecoffett.

The last-named species lacks a distinct discal spot on the elytron

and the pronotum is mostly black with wide, lateral yellow areas.

Tenuisvalvae parenthesis has a c-shaped vitta on each elytron.

404 R.D. GORDON & C. CANEPARI

5. Tenuisvalvae bisquinquepustulata (Fabricius, 1801), n. comb.

Coccinella bisquinquepustulata Fabricius, 1801: 384.

Cleothera bisquinquepustulata: MULSANT 1850: 610.

Hyperaspis bisquinquepustulata: CROTCH 1874: 221; KoRSCHEFSKY 1931: 185; BLACK-

WELDER 1945: 446.

Cleothera raynevalti Mulsant, 1853: 85. (n. syn.).

Hyperaspis raynevalit: CROTCH 1874; KorscHEFSKY 1931: 195; BLACKWELDER 1945:

448; GORDON 1987: 29.

Hyperaspis (Tenuisvalvae) raynevaliu: DUVERGER 1989: 148.

Hyperaspis longicoxitis Nutting, 1980: 260. (n. syn.).

Hyperaspis (Tenuisvalvae) raynevalu longicoxitis: DUVERGER 1989: 152.

Type locality: of bisqinquepustulata, “in America meridionali”

(ZMUC, lectotype here designated); of raynevali, Cayenne (French

Guiana) (UMZC, lectotype designated by GORDON 1987); of longico-

xitis, California, San Diego Co., Jacumba (CAS, holotype).

Description. 6, length 3.2 mm, width 2.6 mm; body

form broadly round, convex. Dorsal surface with head dull, strongly

alutaceous, pronotum and elytra shiny. Color black except head en-

tirely yellow; pronotum mostly yellow with large, black, basomedian

spot, spot not extended to anterior pronotal margin, anterior margin

of spot slightly emarginate; elytron with 5 yellow spots, spot laterad

of scutellum and discal spot round, humeral spot apically tapered,

spot at middle of lateral margin elongate, apical spot transverse (Fig.

513, 514); antenna, mouthparts, legs reddish yellow; propleuron and

epipleuron yellow; abdomen reddish yellow except median 2/3 of

sterna 1-5 dark brown. Head punctures distinct, visible in aluta-

ceous sculpture, separated by less than to twice a diameter. Pronotal

punctures larger than on head, separated by less than to twice a di-

ameter; elytral punctures larger than pronotal punctures, separated

by less than to twice a diameter. Metasternal punctures fine, sparse

medially, becoming much larger, dense laterally. Punctures on ab-

dominal sterna 1-4 coarse, sparse medially, becoming dense, very

fine laterally; sterna 5-6 finely densely punctured throughout; area on

1" abdominal sternum inside postcoxal arc slightly shiny, punctures

coarse, separated by a diameter. Clypeal apex deeply emarginate,

clypeus and frons joined at abrupt angle. Epipleuron oblique, inter-

nally grooved, descending externally, femoral excavations deep. An-

tenna with 11 articles (Fig. 516). Prosternum with intercoxal carinae

convergent, basally stemmed, reaching basal margin. Postcoxal line

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 405

on 1* abdominal sternum extended to posterior sternal margin, flat-

tened along margin, apex recurved 1/2 distance to basal sternal mar-

gin. Fifth sternum with apex broadly, feebly emarginate; 6° sternum

broadly, feebly emarginate. Genitalia with basal lobe 3/4 length of

paramere, apex rounded, with large projection on one side, paramere

wide, feebly tapered to rounded apex (Fig. 517); sipho slender, apex

sinuate with internal membranous area, basal capsule with inner

arm slender, elongate, outer arm wide, elongate (Figs. 518, 519).

2 Similar to male except pronotum with median black area ex-

tended to apex. Genitalia with beak of basal unit short; connecting

duct short (Figs. 520, 521).

Variation. Length 2.7 to 3.2 mm, width 2.3 to 2.6 mm. In the

typical form described above the ground color is black, the size of

the pronotal dark area varies slightly; the scutellar and discal spots

are often narrowly connected, and the 3 lateral spots may be nar-

rowly or broadly connected. The T. longicoxitis form has the ground

color brown, median pronotal spot brown, reduced in size, and of-

ten M-shaped (Fig. 515).

Geographical distribution: Central and northern South America,

some Caribbean islands, and Central America. Introduced to Africa.

Specimens examined: 37. Brazil: Mato Grosso, Chapada; Para;

Belem, Cachimbo; Santarem; Sta. Lucia; Para, Curralino. Ecuador.

Napo, Pompeya. Guyana. Hope. Peru:. Tingo Maria. Surinam.

(No locality). Trinidad: St. Augustine; Tunapuna. Bolivia: Guanay.

(BM) (CCM) (CDA) (CMP) (CNC) (CMNH) (MSNG) (USNM)

(ZMHB).

Remarks. This is a widespread, frequently collected spe-

cies that has been the object of biological control research under the

name H. raynevalu. As a result, it was introduced into the United

States (southern California) and to central Africa (Brazzaville, Con-

go Republic).

The type specimen of Coccinella bisquinquepustulata F. proved

to be synonymous with the type of 7. raynevalit; thus, the latter

name is a junior synonym of the former.

DUVERGER (1989) placed Hyperaspis longicoxitis in his subgenus

Tenuisvalvae and, at the same time, reduced the status of 7. longi-

coxitis to subspecies. He also described a new subspecies, 7. rayne-

valu brazzavillensis, for specimens from Brazzaville.

406 R.D. GORDON & C. CANEPARI

We consider 7. longicoxitis a synonym of T. bisqinquepustulata

because the only distinguishing characters are in coloration. The

former name was applied to specimens from Trinidad and certain

Caribbean islands that have a brown dorsal ground color instead

of black, and the basomedian pronotal spot reduced and often M-

shaped. We have seen examples from Para, Brasil, that are essen-

tially identical in color to Caribbean specimens, thus making color

differences useless as distinguishing characters.

Two syntypes of Coccinella bisquinquepustulata in the ZMUC

were examined. ‘The lectotype, designated here to stabilize future

usage of the name, is labeled “Type (red label)/ C. bisquinquepustu-

lata, Autata, ex. Am. mer. Schonh.” (handwritten). The paralecto-

type designated here is labeled “Type” (red label).

6. Tenuisvalvae ecoffeti (Mulsant, 1853), n. comb.

Hyperaspis ecoffeti Mulsant, 1853: 99; CroTcH 1874: 226; KorscHEFSKY 1931: 187;

BLACKWELDER 1945: 447; GORDON 1987: 28.

Hyperaspis quadrina Mulsant, 1853: 100; KoRSCHEFSKY 1931: 187 (as a synonym of

H. ecoffett).

Type locality: of ecoffett, Brazil (UMZC, lectotype designated

by GORDON 1987); of quadrina, Sainte (sic !) Catherine, Brazil (BM,

lectotype here designated).

Description. 6, length 2.7 mm, width 2.0 mm; body

form oval, convex. Head alutaceous, dull; pronotum slightly aluta-

ceous, shiny; elytra polished, shiny. Color yellow except pronotum

black with lateral and anterior borders yellow; elytron with 2 spots

and sutural margin black, 1 irregularly rectangular spot medially in

apical 1/2, 1 irregularly round spot medially in posterior 1/2, su-

tural margin with broad, black vitta extended from just posterior to

scutellum nearly to apex, vitta slightly widened in apical 1/4, nar-

rowed nearly to apex then curved outward, lateral and apical mar-

gins of elytron narrowly black; scutellum black (Fig. 522); pro-,

meso-, and metasterna black; median area of basal 4 abdominal

sterna dark brown. Head punctures fine, distinct, separated by less

than to twice a diameter. Pronotal punctures larger than on head,

separated by less than to twice a diameter; elytral punctures larger

than pronotal punctures, separated by less than to 3 times a diam-

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 407

eter. Metasternal punctures coarse, larger than on elytron, separated

by a diameter medially, becoming contiguous laterally. Punctures

on abdominal sterna 1-4 coarse, sparse medially, becoming dense,

very fine laterally; sterna 5-6 finely densely punctured throughout;

area on 1* abdominal sternum inside postcoxal arc shiny, coarsely,

densely punctured. Clypeal apex deeply emarginate, clypeus and

frons joined at abrupt angle. Epipleuron oblique, internally grooved,

descending externally, femoral excavations deep. Antenna with 11

articles (Fig. 524). Prosternum with intercoxal carinae convergent,

basally stemmed, reaching basal margin. Postcoxal line on 1* ab-

dominal sternum extended to posterior sternal margin, slightly flat-

tened along margin, apex recurved 1/2 distance to basal sternal

margin. Fifth sternum with apex broadly, deeply emarginate; 6°

sternum broadly, feebly emarginate. Genitalia with basal lobe more

than 3/4 length of paramere, wide, with strong lateral projection on

one side, paramere wide, feebly tapered to rounded apex (Fig. 525);

sipho slender, apex slightly bent downward, with membranous area,

basal capsule with inner arm slender, short, outer arm wide, longer

than inner arm (Figs. 526, 527).

2 Similar to d except head black with median reddish yellow,

apically triangular area. Genitalia with beak of basal unit long; con-

necting duct long (Fig. 528).

Variation. Length 2.6 to 2.7 mm, width 2.2 to 2.3 mm. Pronotal

macula varies significantly from occupying median 1/2 of pronotum

to occupying most of pronotum with only narrow lateral and ante-

rior margins yellow; elytral spots vary slightly in size, sutural vitta

varies noticeably in width and shape from the typical to straight

from apex to base (quadrina) (Fig. 523).

Geographical distribution: Argentina, Brazil, Paraguay.

Specimens examined: 32. Argentina: Loreto, Misiones. Brazil:

Nova ‘Teutonia; Sao Paulo, Cantareira, Campinas; Santa Catari-

na, Florianopolis. Paraguay: Hohenau, Alto-Parana (BM) (USNM)

(ZMHB).

Remarks. This species resembles only 7. notata in color

pattern. It is distinguished by the single black pronotal macula, lack

of discal spot on the elytron, and 2 spots on each elytron.

CroTcH (1874) correctly considered Hyperaspis quadrina a ju-

nior synonym of T. ecofffeti. ‘The type specimen of T. quadrina

408 R.D. GORDON & C. CANEPARI

was from the Deyrolle collection, according to MULSANT (1853),

but was not found in the UMZC. However, the Natural History

Museum, London, collection has a single specimen of T. quadrina,

labeled as a type that bears the correct locality label recorded by

Mulsant. We consider that specimen, labeled “S. Cathar (green pa-

per)/ 261/880.23/ Type (orange bordered disc/ Hyperaspis quadrina,

M. Brasil (blue paper),” to be the lectotype of H. quadrina and so

designate it to stabilize future usage of the name. KIRSCH (1876)

described “Hyperaspis Ecoffeti var. fraudulenta,” a combination re-

peated by KoRrscHEFsKy (1931) and BLACKWELDER (1945). Examina-

tion of the H. fraudulenta type specimen (SMTD) revealed that it

is actually a valid species of Cyra (Brachiacanthini).

7. Tenuisvalvae parenthesis n. sp.

Type material: Holotype 2: Kolumb (Colombia), Cundinamarca,

Monterredondo, 1961, 1400 m, leg. Schneble (USNM). Paratypes,

5. 1, same data as holotype; 1, Colombia, Boy. (Boyaca), Guayata,

18.X.40, altitude 1720 m, Murillo No 5299 (USNM). 1, Venezue-

la, E.do Merida, La Trampa, Lagunillas 1700 m, 10.KKK.778, leg

Bordon; 2, Venezuela, Timotes, m 2200, (Edo. Merida), 23-1-1968,

leg. Bordon (CCM) (MSNG) (USNM).

Description. &, length 3.5 mm, width 2.8 mm; body

form round, convex. Dorsal surface shiny except head dull, strongly

alutaceous, pronotum shiny, slightly alutaceous. Color yellow except

head with borders black, black border on vertex medially emargin-

ated with yellow; pronotum with basal border narrowly black, 2

curved projections extended from basal border to apex; elytron with

short, apically knobbed vitta extended from base past humeral cal-

lus, suture narrowly black with parenthesis shaped, apically knobbed

vitta extended out at middle, curved posteriorly onto apical decliv-

ity, small, triangular spot near lateral margin in posterior 1/2 (Figs.

529, 530); mouthparts dark brown; venter of head, pro-, meso-,

metasterna, femora (except apices), and median area of abdominal

sterna 1-5 black. Head punctures fine, nearly hidden in alutaceous

sculpture, separated by 1 to 2 times a diameter. Pronotal punctures

larger than on head, separated by less than to 2 times a diameter;

Elytral punctures larger than pronotal punctures, separated by 1 to

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 409

2 times a diameter. Metasternal punctures larger than on elytron,

separated by a diameter medially, dense, nearly contiguous laterally.

Punctures on abdominal sterna 1-4 coarse, sparse medially, becom-

ing dense, very fine laterally; sterna 5-6 finely densely punctured

throughout; area on 1° abdominal sternum inside postcoxal arc

shiny, coarsely, sparsely punctured. Clypeal apex deeply emargin-

ate, clypeus and frons joined at abrupt angle. Epipleuron oblique,

internally grooved, descending externally, femoral excavations deep.

Antenna with 11 articles (Fig. 531). Prosternum with intercoxal

carinae tapered toward base, stemmed, reaching basal margin. Post-

coxal line on 1% abdominal sternum extended to posterior sternal

margin, slightly flattened along margin, apex recurved 1/3 distance

to basal sternal margin. Fifth sternum with apex broadly, distinctly

emarginate. Genitalia with beak of basal unit short, connecting duct

short (Fig. 535).

S Similar to 2 except d genitalia with basal lobe 5/8 length of

paramere, abruptly tapered to narrow apex in apical 1/2, with large,

acute projection on one side; paramere wide, tapered to rounded

apex in apical 1/2 (Fig. 532); sipho long, slender, sinuate in api-

cal 1/4, apex with membranous area, basal capsule with inner arm

elongate, slender, outer arm slender, shorter than inner arm, base

emarginate (Figs. 533, 534).

Variation. Length 3.2 to 3.5 mm, width 2.5 to 2.8 mm. Elytron

with triangular lateral spot sometimes connected to parenthetical

vitta; apex of parenthetical vitta may be narrowly separate from

main stem.

Etymology. The species is named for the curved elytral vitta

reminiscent of a parenthesis.

Remarks. Tenutsvalvae parenthesis bears some resemblance

to T. caucaensis, but the dorsal color pattern is thus far unique with-

in the genus.

8. Tenuisvalvae caucaensis n. sp.

Type material: Holotype 6: Colombia, Cauca, 15 mi. E. Sil-

via, 11,000’, July 16, 1970, H. & A. Howden (USNM). Allotype ®:

Colombia, Cauca;:12-mi: E. Silvia, 11,000; July 15, 1970//H. .& A.

Howden (USNM).

410 R.D. GORDON & C. CANEPARI

Description. 6, length 3.0 mm, width 2.6 mm; body

form rounded, widest behind middle of elytra, convex. Head and

pronotum slightly alutaceous, weakly shiny; elytra polished. Color

black except head with apical 3/4 yellow; pronotum with lateral 1/3

yellow, yellow area with curved inner margin; elytron with 3 large,

spots, discal spot elongate, sublateral spot oval with anteromedian

angle truncate, subapical spot more or less round (Figs. 536, 537);

antenna, propleuron yellow; tarsi reddish brown. Head punctures

distinct, separated by a diameter or less. Pronotal punctures as large

as head punctures, separated by 1 to 2 times a diameter; Elytral

punctures larger than pronotal punctures, separated by 1 to 2 times

diameter. Metasternal punctures larger than on elytra, dense, absent

medially, separated by a diameter laterally. Punctures on abdominal

sterna 1-4 coarse, sparse medially, becoming dense, fine laterally;

sterna 5-6 finely densely punctured throughout; area on 1* abdomi-

nal sternum inside postcoxal arc slightly alutaceous, coarsely, sparse-

ly punctured. Clypeal apex deeply emarginate, clypeus and frons

joined at abrupt angle. Antenna with 11 articles (Fig. 538). Epipleu-

ron oblique, internally grooved, descending externally, femoral ex-

cavations deep. Prosternum with intercoxal carinae tapered to base,

not stemmed. Postcoxal line on 1% abdominal sternum extended to

posterior sternal margin, not flattened along margin, apex recurved

2/3 distance to basal sternal margin. Fifth sternum with apex broad-

ly, distinctly emarginate; 6" sternum broadly, distinctly emarginate.

Genitalia with basal lobe 3/4 length of paramere, abruptly narrowed

apically with large lateral projection, paramere nearly parallel sided,

apex broadly rounded, trabes dark brown (Fig. 539); sipho slender,

apex straight with membranous area, capsule dark brown with inner

arm large, long, outer arm large, wide (Figs. 540, 541).

? Similar to d except head black. Genitalia with beak of basal

unit large, wide, connecting duct short (Fig. 542).

Variation. None observed.

Etymology. The species is named for the Department of Cauca

where the type specimens were collected.

Remarks. This species is differentiated from other Tenu-

svalvae species by the 3 very large, yellow elytral spots, and broad,

posteriorly widened body. ‘The dark brown genital trabes and sipho-

nal capsule are also worth noting because these structures are nearly

always yellow or at least pale in hyperaspine taxa.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 411

9. Tenuisvalvae bromelicola (Sicard, 1925), n. comb.

Cleothera bromelicola Sicard, 1925: 81.

Hyperaspis bromelicola: KORSCHEFSKY 1931: 185; BLACKWELDER 1945: 446.

Hyperaspis (Tenuisvalvae) bromelicola: PECK 2005: 202.

Type locality: Panama, Canal Zone (BM, lectotype here desi-

gnated).

Description. 6, length 2.7 mm, width 2.3 mm; body

form round, convex. Dorsal surface with head dull, strongly aluta-

ceous, pronotum shiny slightly alutaceous, elytra shiny. Color black

except head entirely yellow; pronotum mostly yellow with black ba-

somedian macula, anterior margin of macula with 2 lateral projec-

tions; elytron with 3 yellow spots, spot on base laterad of scutellum

triangular, basal humeral spot somewhat triangular, apical spot long,

curved along outer elytral margin from near suture to midpoint of

elytron (Figs. 543, 544); antenna, mouthparts, propleuron, legs yel-

low; abdomen reddish yellow except median 1/2 of sterna 1-3 dark

brown. Head punctures fine, nearly hidden in alutaceous sculpture,

separated by less than to twice a diameter. Pronotal punctures larger

than on head, separated by less than to twice a diameter; elytral

punctures larger than pronotal punctures, separated by 1 to 2 times

a diameter. Metasternal punctures much larger than on elytron,

sparse medially, dense laterally. Punctures on abdominal sterna 1-4

coarse, sparse medially, becoming dense, very fine laterally; sterna

5-6 finely densely punctured throughout; area on 1% abdominal ster-

num inside postcoxal arc shiny, finely, sparsely punctured. Clypeal

apex deeply emarginate, clypeus and frons joined at abrupt angle.

Antenna with 11 articles (Fig. 545). Epipleuron oblique, internally

grooved, descending externally, femoral excavations deep. Proster-

num with intercoxal carinae convergent, basally stemmed, reaching

basal margin. Postcoxal line on 1% abdominal sternum extended to

posterior sternal margin, flattened along margin, apex recurved 1/3

distance to basal sternal margin. Fifth sternum with apex broadly,

feebly emarginate; 6 sternum broadly, feebly emarginate. Genita-

lia with basal lobe slightly shorter than paramere, with barely per-

ceptible projection on one side, paramere wide, feebly tapered to

rounded apex (Fig. 547); sipho slender, apex curved, sinuate with

membranous area, basal capsule with inner arm broad, short, outer

arm broad, short (Figs. 548, 549).

412 R.D. GORDON & C. CANEPARI

2 Similar to d except head with vertex black on each side;

pronotum with median black area extended to apex, apex of black

area medially emarginated with clear, V-shaped area exposing me-

dian yellow area of head vertex. Genitalia with beak of basal unit

short, large; connecting duct short (Figs. 550, 551).

Variation. Length 2.6 to 3.1 mm, width 2.2 to 2.6 mm. Elytral

maculation may have the humeral spot reduced to a small, reddish

brown area, the apical spot may be expanded to cover entire apical

declivity, or elongated along elytral margin nearly to humeral spot.

Geographical distribution: Colombia, Ecuador, Panama.

Specimens examined: 26. Colombia: Choco, Istmina. Ecua-

dor: El Oro, Machala; Santo Domingo de los Colorados. Galapagos

Islands, San Cristobal. Panama: Canal Zone, Ancon. (BM) (CCM)

(USNM).

Remarks. Tenuisvalvae bromelicola is the only known spe-

cies of the genus found in both South and Central America. The

distinctive color pattern should not be confused with that of any

currently known species of Tenuisvalvae. 'The Natural History Mu-

seum, London, has three type specimens of this species, the first of

which is designated as the lectotype to stabilize future usage of this

name. It is labeled “Type (orange bordered disc)/ Panama Canal

Zone VII-1914/ Pres. by Imp. Bur. Ent. Brit. Mus. 1924-482/ Lar-

vae are predaceous on Pseudococcus bromeliae/ D.T. Fullaway Col-

lector/ Cleothera bromelicola Sic., type n. sp. (handwritten in purple

ink)”. The other two syntypes, designated as paralectotypes, are la-

beled as is the lectotype, except one is without a circular disc with a

type or syntype designation, and the other has a blue bordered disc

labeled “Syntype”.

10. Tenuisvalvae gnoma n. sp.

Type material: Holotype 2: Col. (Colombia), S.A., Apr. 22-35,

From H. Daniel, Chapin 42-31 (USNM).

Description. ®, length 3.0 mm, width 2.3 mm; body

form rounded, convex. Dorsal surface shiny with head alutaceous,

feebly shiny; pronotum alutaceous, feebly shiny, elytron shiny.

Color black except head with u-shaped yellow spot; pronotum with

lateral 1/3 yellow; elytron pale red, narrowly bordered with black

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 413

except sutural margin, basal border narrowly extended onto humeral

callus, apical border broad (Fig. 552); mouthparts dark brown; an-

tenna, protibia yellow; epipleuron red; abdomen with lateral 1/4 yel-

low. Head punctures fine, separated by a diameter or less. Pronotal

punctures larger than on head, separated by a diameter or less; Ely-

tral punctures larger than pronotal punctures, separated by less than

to twice a diameter. Metasternal punctures larger than on elytron,

separated by less than to 3 times a diameter medially, dense, contig-

uous laterally. Punctures on abdominal sterna 1-4 coarse, sparse me-

dially, becoming dense, very fine laterally; sterna 5-6 finely densely

punctured throughout; area on 1° abdominal sternum inside post-

coxal arc shiny, coarsely, sparsely punctured. Clypeal apex emargin-

ate, clypeus and frons joined at abrupt angle. Epipleuron oblique,

internally grooved, descending externally, femoral excavations deep.

Prosternum with intercoxal carinae widely separated apically, gradu-

ally tapered toward base, reaching basal margin, not stemmed. Post-

coxal line on 1% abdominal sternum extended to posterior sternal

margin, slightly flattened along margin, apex recurved 3/8 distance

to basal sternal margin. Fifth sternum with apex broadly, distinctly

emarginate. Genitalia with beak of basal unit short, connecting duct

short (Fig. 553).

4 Not known.

Etymology. ‘The species name is the Latin gnoma, meaning

dwarf, referring to the small size.

Remarks. Tenuisvalvae gnoma is the only known species of

Tenuisvalvae with mostly red elytra.

ACKNOWLEDGEMENTS

Without assistance from the following individuals this study

could not have been accomplished.

We thank Roger Booth (BM), Lothar Zerche (DEI), Lucia

Massutti de Almeida (DZUP), Axel Bachmann (MBR), Giulio

Cuccodoro (MHNG), Mario Elgueta (MNHS), Heinrich Sch6n-

mann (NHMV), Olaf Jaeger (SMTD), William Foster (UMZC),

Ole Martin (ZMUC), Bernd Jaeger and Manfred Uhlig (ZMHB),

and Nikolai Nikitskii (ZMMU) for the loan of types. We thank

Boris Korotyaev, Zoological Insitute Russian Academy of Scien-

414 R.D. GORDON & C. CANEPARI

ce, for facilitating the loan of a type specimen; L. David and Joél

Clary (MNHL) for allowing an examination of the Dejean collec-

tion; Helmut Fuùrsch, Passau, Germany, for information and advice

concerning type material; Natalia Vandenberg (USNM) for refe-

rence materials, and Gary Miller, Systematic Entomology Labora-

tory, Beltsville, Maryland, for assistance with scale names. For loan

of specimens we are indebted to Roger Booth and Max Barclay

(BM), Roberta Brett (CAS), Charles Bellamy (CDA), Robert Da-

vidson (CMP), Anthony Davies (CNC), Guillermo Gonzalez (GG),

Axel Bachmann (MBR), Mario Elgueta (MNHS), Roberto Poggi

(MSNG) and Natalia Vandenberg (USNM). Special thanks to Paul

Skelley, Florida State Collection of Arthropods, Gainesville, Flo-

rida, for providing the SEMs included here. ‘The SEMs were ta-

ken at the University of Florida, ICBR, Electron Microscopy Core

Laboratory, Gainesville, FL. For manuscript review we thank AI-

fred Wheeler, Clemson University, Clemson, South Carolina; N.

Vandenberg and A. Norrbom, both of the Systematic Entomology

Laboratory, USDA, Washington, DC. The illustrations were pre-

pared by Claudio Canepari.

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ABSTRACT

South American Hyperaspidini include 8 genera and 97 species. Genera are

Diazonema Mulsant, Menoscelis Mulsant, Thalassa Mulsant, Clypeaspis, n. gen.,

Prognataspis, n. gen., Peruaspis, n. gen., Hyperaspis Redtenbacher, and Tenuisvalvae

Duverger. The genera Gordoni Duverger and Aliana Duverger are placed as junior

synonyms of Hyperaspis. In Diazonema 7 species are recognized as valid, including

3 new species: boothi, murillot, and eccentrica. Clypeaspis is erected for Hyperaspis

trilineata Mulsant. Prognataspis contains a single new species, P. surreptiva. Peruaspis

is erected for the new species P. paprzyckii and P. hypocrita. Forty-one new species of

Hyperaspis are described: H. abertha, H. aemulata, H. apicaspis, H. arida, H. atra, H.

ayacucho, H. bisignata, H. brethesi, H. camargoi, H. campbelli, H. chapini, H. chocot,

H. circumclusa, H. colombiensis, H. cracentis, H. dispar, H. dissidens, H. helveola,

H. herrera, H. howdeni, H. imitatrix, H. istmina, H. joannae, H. laterimacula, H.

latitibia, H. lindae, H. mariposa, H. mimica, H. octonotata, H. orthivora, H. prolata,

H. pseudodonzeli, H. pseudopavida, H. rosariensis, H. satipoensis, H. siladesma, H.

simlaensis, H. tayronensis, H. uninotata, H. vredenburgi, and H. zonula. Tenuisvalvae

is recognized as a valid genus instead of a subgenus of Hyperaspis, and 3 new species

are described: T. caucaensis, T. gnoma and T. parenthesis.

The following new synonyms are stated (in each couplet the second name is

the valid one): Hyperaspis cordifera (Weise) = H. matronata Mulsant, H. incompleta

Crotch = H. scutifera Mulsant, H. insignis Crotch = H. operaria (Mulsant), H.

communalis Brèthes = H. operaria (Mulsant), H. graphica Weise = H. eupaleoides

Crotch, H. juniapuca Brèthes = H. festiva Mulsant, H. flavolineata Mader = H.

conclusa Weise, H. subapicalis Crotch = Tenuisvalvae peregrina (Mulsant), H.

raynevalu (Mulsant) = Tenuisvalvae bisquinquepustulata (F.), and H. longicoxttis

Nutting = Tenuisvalvae bisquinquepustulata (F.).

New combinations from Hyperaspis are: T. unipunctata (Crotch), T. peregrina

(Mulsant), 7. deyrollet (Crotch), T. notata (Mulsant), T. ecoffett (Mulsant), 7.

bromelicola (Sicard), and T. bisquinquepustulata (F.).

Cleothera micilla Mulsant is transferred to Diomus Mulsant and placed as a

synonym of Diomus pallidipennis (Mulsant).

418 R.D. GORDON & C. CANEPARI

RIASSUNTO

Coccinellidi del Sud America (Coleoptera). Parte XI: revisione sistematica degli

Hyperaspidini (Hyperaspidinae).

La Tribù Hyperaspidini comprende in Sud America 8 generi e 97 specie. I

generi sono: Diazonema Mulsant, Menoscelis Mulsant, Thalassa Mulsant, Clypeaspts,

n. gen., Prognataspis, n. gen., Peruaspis, n. gen., Hyperaspis Redtenbacher e

Tenuisvalvae Duverger. I generi Gordoni Duverger e Aliana Duverger sono

considerati nuovi sinonimi di Hyperaspis. In Diazonema sono riconosciute 7 specie

valide, incluse 3 nuove specie: boothi, murilloi ed eccentrica. Per Hyperaspis trilineata

Mulsant viene eretto il nuovo genere Clypeaspis. Prognataspis contiene una sola

nuova specie, P. surreptiva. Il genere Peruaspis viene istituito per P. paprzyckii e

P. hypocrita. Vengono descritte quarantuno nuove specie di Hyperaspis: H. abertha,

H. aemulata, H. apicaspis, H. arida, H. atra, H. ayacucho, H. bisignata, H. brethesi,

H. camargoi, H. campbelli, H. chapini, H. chocoi, H. circumclusa, H. colombiensis, H.

cracentis, H. dispar, H. dissidens, H. helveola, H. herrerai, H. howdeni, H. imitatrix,

H. istmina, H. joannae, H. laterimacula, H. latitibia, H. lindae, H. mariposa, H.

mimica, H. octonotata, H. orthivora, H. prolata, H. pseudodonzeli, H. pseudopavida, H.

rosariensis, H. satipoensis, H. stiladesma, H. simlaensis, H. tayronensis, H. uninotata, H.

vredenburgi e H. zonula. Tenuisvalvae viene riconosciuto come genere valido invece

di sottogenere di Hyperaspis e vengono descritte 3 nuove specie: 7. caucaensis, T.

gnoma e T. parenthests.

Vengono stabilite alcune nuove sinonimie (la specie valida è la seconda di ogni

coppia): Hyperaspis cordifera (Weise) = H. matronata Mulsant, H. incompleta Crotch

= H. scutifera Mulsant, H. insignis Crotch = H. operaria (Mulsant), H. communalis

Brèthes = H. operaria (Mulsant), H. graphica Weise = H. eupaleoides Crotch, H.

juniapuca Brèthes = H. festiva Mulsant, H. flavolineata Mader = H. conclusa Weise,

H. subapicalis Crotch = Tenuisvalvae peregrina (Mulsant), H. raynevalii (Mulsant)

= Tenuisvalvae bisquinquepustulata (F.) e H. longicoxitis Nutting = Tenuisvalvae

bisquinquepustulata (F.).

Le specie trasferite da Hyperaspis a Tenuisvalvae sono: T. unipunctata (Crotch),

T. peregrina (Mulsant), T. deyrollei (Crotch), T. notata (Mulsant), T. ecoffeti

(Mulsant), T. bromelicola (Sicard) e T. bisquinquepustulata (F.).

Cleothera micilla Mulsant è trasferita in Diomus Mulsant e considerata sinonimo

di Diomus pallidipennis (Mulsant).

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI

419

Index to scientific names

(synonyms in ttalics)

abertha, n. sp.

aemulata, n. sp.

albopunctata Crotch

apicalis Weise

Aliana Duverger

apicaspis, n. sp.

arida, n. sp.

atta in. sp.

ayacucho, n. sp.

bicruciata Mulsant

biguttata (Sicard)

bisignata, n. sp.

bisquinquepustulata (F.)

Blaisdelliana Gordon

boothi, n. sp.

Brachiacanthini Duverger

brethesi, n. sp.

bromelicola (Sicard)

c-nigrum Mulsant

camargol, n. sp.

campbelli, n. sp.

caucaensis, n. sp.

cavifrons (Weise)

chapini, n. sp.

chapini group

chilensis Crotch

chocoi, n. sp.

cincticollis Mulsant

circumclusa, n. sp.

Clavatus Duverger

cleida Mulsant

365

367

340

376

286

342

389

350

338

316

313

404

256

267

247

325

411

358

315

331

409

265

307

303

382

348

375

366

288

317

Cleothera Mulsant

Clypeaspis, n. gen.

colombiensis, n. sp.

communalis Brèthes

conclusa Weise

conclusa group

connectens (Thunberg)

cordifera Weise

Corystes Mulsant

cracentis, n. sp.

cruciata Germain

delicata Almeida & Vitorino

deyrollei (Crotch)

Diazonema Weise

dispar, n. sp.

dissidens, n. sp.

donzeli (Mulsant)

donzeli group

eccentrica, n. sp.

ecoffeti (Mulsant)

elegantissima Brèthes

eupaleoides Crotch

fallax Weise

festiva Mulsant

flaviceps Mulsant

flavolineata Mader

funesta (Germain)

germainil Crotch

glauca (Mulsant)

gnoma, n. sp.

Gordoni Duverger

246

278

332

310

387

381

379

300

259

308

386

343

401

259

368

331

298

297

269

406

391

352

263

375

244

387

382

378

277

412

286

420

graphica Weise

guilloryi (Mulsant)

Helesius Casey

helveola, n. sp.

herrerai, n. sp.

Hinda Mulsant

histrionica (Mulsant)

howdeni, n. sp.

howdeni group

Hyperaspidina

Hyperaspidini

Hyperaspidius Crotch

Hyperaspini

Hyperaspis Chevrolat

hypocrita, n. sp.

hypocrita (Mulsant)

imitatrix, n. sp.

incompleta Crotch

ingrata (Mulsant)

insignis Crotch

insignis Mulsant

istmina, n. sp.

joannae, n. sp.

joannae group

juniapuca Brèthes

korschefskyi Milléo et al.

laterimacula, n. sp.

latitibia, n. sp.

lemniscata Boheman

lengi Schaeffer

limbifera Blackwelder

limbigera (Mulsant)

lindae, n. sp.

longicoxitis Nutting

352

361

25%,

302

288

370

SET

347

256

Vest

256

286

284

202

305

303

381

310

274

363

339

389

376

277

346

29]

376

379

Sii

311

356

404

R.D. GORDON & C. CANEPARI

longula Weise

mariposa, n. sp.

matronata (Mulsant)

Menoscelis Mulsant

mimica, n. sp.

montezumae Mulsant

murilloi, n. sp.

nana Mader

notata (Mulsant)

octonotata, n. sp.

onerata group

onerata (Mulsant)

operaria (Mulsant)

orthivora, n. sp.

paprzycku, n. sp.

parenthesis, n. sp.

pavida Mulsant

pentaspilota (Chevrolat)

pentastigma Mulsant

peregrina (Mulsant)

Peruaspis, n. gen.

prasina Mulsant

Prognataspis, n. gen.

prolata, n. sp.

pseudodonzeli, n. sp.

pseudopavida, n. sp.

pubescens (Weise)

quadrina (Mulsant)

raynevalu (Mulsant)

recordata Mulsant

rosariensis, n. sp.

saginata Mulsant

sagittata Crotch

satipoensis, n. sp.

302

300

270

SARE

278

268

384

402

329

308

336

310

322

283

408

363

276

399

282

277

280

394

320

362

264

406

404

351

327

272

359

364

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI

scutifera Mulsant

siladesma, n. sp.

similaris Mulsant

simlaensis, n. sp.

sphaeridioides Mulsant

subapicalis Crotch

surreptiva, n. sp.

tayronensis, n. sp.

303

328

244

355

386

Bey

281

319

Tenuisvalvae Duverger

Thalassa Mulsant

trilineata (Mulsant)

uninotata, n. sp.

unipunctata (Crotch)

vredenburgi, n. sp.

vredenburgi group

zonula, n. sp.

421

396

275

279

398

374

371

323

422

Figs.

R.D. GORDON & C. CANEPARI

Gi,

1-6 - Hyperaspidini structures. 1-3, Clypeaspis trilineata. 1, head; 2, protibia;

3, prosternum. 4-6, Peruaspis hypocrita. 4, mouthparts and prosternum;

5, prosternum; 6, clypeus.

423

HYPERASPIDINI

ppi

SOUTH AMERCAN COCCINELLIDAE, XI

vi)

8, prosternum. 9, H.

dens, prosternum. 10, H. connectens, prosternum. 11-12, H. lon-

gula. 11, head

12, prosternum.

1SS1

Figs. 7-12 - Hyperaspis structures. 7-8, H. matronata. 7, head

424 R.D. GORDON & C. CANEPARI

Figs. 13-20 - Diazonema structures. 13-15, heads. 13, D. hypocrita. 14, D. boothi.

15, D. murillot. 16, D. hypocrita, habitus; 17, abdomen. 18-20, D.

hypocrita male genitalia. 18, phallobase; 19, sipho; 20, enlarged sipho-

nal tip.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 425

Figs. 21-23 - Diazonema boothi habitus and genitalia. 21, habitus. 22, phallobase;

23, sipho.

426 R.D. GORDON & C. CANEPARI

Figs. 24-28 - Diazonema murilloi structures. 24, habitus. 25, antenna. 26, protibia.

27-28, male genitalia. 27, phallobase; 28, sipho.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 427

Figs. 29-32 - Diazonema eccentrica structures. 29-30, habitus. 31, antenna, basal

article missing. 32, female genitalia, distal capsule omitted.

428 R.D. GORDON & C. CANEPARI

Figs. 33-39 - Menoscelis structures. 33, M. insignis, habitus. 34-39, M. saginata.

34-35, habitus. 36, antenna. 37, abdomen. 38, protibia, anterior view;

39, protibia, posterior view.

429

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI

41, sipho.

Figs. 40-41 - Menoscelis saginata genitalia. 40, phallobase

430 R.D. GORDON & C. CANEPARI

Figs. 42-46 - Thalassa montezumae structures. 42, head. 43, antenna. 44, protibia;

45, mesotibia; 46, metatibia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 431

Figs. 47-52 - Clypeaspis trilineata habitus and genitalia. 47-48, habitus. 49-51, male

genitalia. 49, phallobase; 50, sipho; 51, enlarged siphonal apex. 52,

femaie genitalia, distal capsule omitted.

432 R.D. GORDON & C. CANEPARI

Figs. 53-60 - Prognataspis surreptiva structures. 53, head. 54, antenna, basal article

missing. 55, prosternum. 56, abdomen. 57, protibia. 58-60, male gen-

italia. 58, phallobase; 59, sipho; 60, enlarged siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 433

61 62

Figs. 61-62 - Prognataspis surreptiva habitus.

434 R.D. GORDON & C. CANEPARI

Figs. 63-67 - Peruaspis paprzycku structures. 63-64, habitus. 65, antenna.

66, abdomen. 67, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 435

È È CIO

ì vendica

N A 0 if n sciare iviso dotta

POP ee

Figs. 68-71 - Peruaspis hypocrita structures. 68-69, habitus. 70, protibia. 71, female

genitalia, genital plates not shown.

436 R.D. GORDON & C. CANEPARI

Figs. 72-78 - Hyperaspis latitibia and H. reppensis structures. 72-76, H. latitibia.

72-73, habitus. 74, protibia. 75-76, male genitalia. 75, phallobase;

76, sipho. 77-78, H. reppensis. 77, prosternum. 78, abdomen.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 437

Figs. 79-82 - Hyperaspis donzeli habitus and genitalia. 79, habitus. 80-82, male

genitalia. 80, phallobase; 81, sipho; 82, enlarged siphonal apex.

438 R.D. GORDON & C. CANEPARI

Figs. 83-87 - Hyperaspis matronata structures. 83, habitus. 84, antenna. 85-87, male

genitalia. 85, phallobase; 86, sipho; 87, enlarged siphonal apex.

439

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI

eer MOR,

ie POPGNOA

A N

e SI

Beene

OR,

patata PE

nti

Nee og Yi he

DECO

NT gh =

NE ny.

Co a

i Î

/ |

}

} /

PA Î i

H }

MS:

Figs. 88-90 - Hyperaspis matronata female genitalia. 88, spermathecal capsule, fron- |

tal view; 89, genital plates; 90, genital capsule, lateral view.

440) R.D. GORDON & C. CANEPARI

a -

È,

Figs. 91-99 - Hyperaspis helveola structures. 91, habitus. 92, antenna. 93, metend-

osternite. 94-96, male genitalia. 94, phallobase; 95, sipho; 96, enlarged

siphonal apex. 97-99, female genitalia. 97, entire genitalia showing

basal portion in ventral view; 98, distal portion; 99, lateral view of

basal portion.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 441

Figs. 100-108 - Hyperaspis scutifera structures. 100-102, habitus and variations.

103, antenna. 104, metendosternite. 105-107, male genitalia. 105,

phallobase; 106, sipho; 107, enlarged siphonal apex. 108, female

genitalia.

442 R.D. GORDON & C. CANEPARI

Figs. 109-112 - Hyperaspis imitatrix habitus and male genitalia. 109-110, habitus.

111-112, male genitalia. 111, phallobase; 112, sipho.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 443

113

115

116 YY

Figs. 113-116 - Hyperaspis chapini habitus and genitalia. 113, habitus. 114-116,

male genitalia. 114, phallobase; 115, enlarged siphonal apex; 116,

sipho.

+44 R.D. GORDON & C. CANEPARI

I 18 fehl da a i, } FORSE if

ue a Li dh A ett Wi 4) hal AN?

Ui He pi aA

117

Figs. 117-118 - Hyperaspis chapini female genitalia. 117, genital plates; 118, sper-

mathecal capsule, lateral and frontal views.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 445

Figs. 119-126 - Hyperaspis cracentis structures. 119-120, habitus. 121, protibia. 122,

abdomen. 123-125, male genitalia. 123, phallobase; 124, sipho, 125,

enlarged siphonal apex. 126, female genitalia.

446

R.D. GORDON & C. CANEPARI

130

ae pit

, ty d)

MEAL!) fg ART i Hi

5 1 î

7 dI; UAL

ai 3 $

Ke 2p Hy)

i; eg ge ee È oi

ies big 3

Figs. 127-133 - Hyperaspis operaria habitus and genitalia. 127-128, habitus.

129-131, male genitalia. 129, phallobase; 130, sipho; 131, enlarged

siphonal apex. 132-133, female genitalia. 132, complete genitalia;

133, basal unit, lateral view.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 447

134 135

Figs. 134-137 - Hyperaspis limbigera structures. 134-135, habitus. 136, antenna,

basal article missing. 137, female genitalia.

448

R.D. GORDON & C. CANEPARI

KON,

i : as pranreetttties aceengs RL

n =,

(

PX {i

2 oe we y/

\ beak =e basal unit a: I \ ji

Ù = i na i J S, 4 o / /

“Ss a ; x bi ii

ZA ‘hi Oi } \ \ /;

A = ie i \

di i fe di 4

distal capsule /

| if

3 $

‘ È

P, i i

i i

\ i i

} i j si

si i Hi if

ue i i it

; 1 ii

Age H N 1

n \

accessory gland } x

bursa copulatrix

genital plate

Figs. 138-143 - Hyperaspis

bisignata habitus

and genitalia. 138-139, habitus.

140-142, male genitalia. 140, phallobase; 141, sipho; 142, enlarged

siphonal apex. 143, female genitalia.

449

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI

sta

aga

\ N

a cena

147

pow ci ent

sarti series enna i ‘ È

Mey

eo sai

IAN 0%

Pi Vai

Vea si

ee satin

eee

ar ear, ARABO Lo

149

Figs. 144-152 - Hyperaspis camargoi structures. 144-145, habitus. 146, antenna.

147-149, male genitalia. 147, phallobase; 148, sipho; 149, enlarged

siphonal apex. 150-152, female genitalia. 150, distal portion; 151,

basal portion, ventral and lateral view; 152, genital plate.

450 R.D. GORDON & C. CANEPARI

Figs. 153-159 - Hyperaspis biguttata and H. cleida habitus and male genitalia.

153-155, H. biguttata. 153, habitus. 154-155, male genitalia. 154,

phallobase; 155, sipho. 156-159, H. cleida. 156-157, habitus.

158-159, male genitalia. 158, phallobase; 159, sipho.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 451

sail at,

He pe”

ig Pes: ; A

SA fig A

dott L

iY Po #%

# i ian

pe \- |

r de

{O x For, A | i |

ANI 2 ~

of, \ j

\e ì my i

i 7 / i

À ee | |

Vea e bao ak, |

H E

" i \

j i |

163 ae REC 165

Figs. 160-165 - Hyperaspis tayronensis habitus and genitalia. 160-162, habitus and

variations. 163-165, male genitalia. 163, phallobase; 164, sipho;

165, enlarged siphonal apex.

452 R.D. GORDON & C. CANEPARI

173

Figs. 167-175 - Hyperaspis pseudodonzeli structures. 167-169, habitus and varia-

tions. 170, antenna. 171, metendosternite. 172-174, male geni-

talia. 172, phallobase; 173, sipho; 174, enlarged siphonal apex.

175, female genitalia without sperm duct.

SOUTH AMERCAN COCCINELLIDAE, XI

: HYPERASPIDINI 453

th d ire pub " th

Î | / 5 MITI pe ry paps ‘ \\

183 i i

Figs. 176-183 - Hyperaspis orthivora structures. 176-178, habitus and variations

179, antenna. 180-181, male genitalia. 180, phallobase; 181, sipho.

182-183, female genitalia. 182, genital plates; 183, basal and distal

portions.

454 R.D. GORDON & C. CANEPARI

Figs. 184-185 - Hyperaspis zonula habitus.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 455

187

Figs. 186-188 - Hyperaspis zonula genitalia. 186-187, male genitalia. 186, phal-

lobase; 187, sipho. 188, female genitalia.

456 R.D. GORDON & C. CANEPARI

SOIN

ee wed

anreetenenrtete nine

seal

Nitin coal

INI

Figs. 189-198 - Hyperaspis brethesi structures. 189-193, habitus and variations. 194,

antenna. 195, metendosternite. 196-198, male genitalia. 196, phal-

lobase; 197, sipho; 198, enlarged siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 4

Pe)

201 202 bi

Figs. 199-203 - Hyperaspis brethesi female genitalia and H. rosariensis habitus

and male genitalia. 199, H. brethesi female genitalia. 200-203, H.

rosariensis. 200, habitus. 201-203, male genitalia. 201, phallobase;

202, sipho; 203, enlarged siphonal apex.

458 R.D. GORDON & C. CANEPARI

ee E RI

Nd,

sot è

PO Oe + "OY tu

» di

: a ;

x RAI 3

he is ù

Figs. 204-212 - Hyperaspis siladesma and H. octonotata habitus and genitalia.

204-207, H. siladesma. 204-205, habitus. 206-207, male genitalia.

206, phallobase; 207, siphonal capsule, remainder of sipho lost.

208-212, H. octonotata. 208-209, habitus. 210-212, male genitalia.

210, phallobase; 211, sipho; 212, enlarged siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 459

Figs. 213-220 - Hyperaspis campbelli and H. colombiensis structures. 213-215, H.

campbelli. 213, habitus. 214-215, male genitalia. 214, phallobase;

215, sipho. 216-220, H. colombiensis. 216, habitus. 217, antenna.

218-220, male genitalia. 218, phallobase; 219, sipho; 220, enlarged

siphonal apex.

460 R.D. GORDON & C. CANEPARI

Figs. 221-226 - Hyperaspis dissidens structures. 221-222, habitus. 223, antenna.

224-226, male genitalia. 224, phallobase; 225, sipho; 226, enlarged

siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 461

Ae 230 931

Figs. 227-231 - Hyperaspis uninotata structures. 227, habitus. 228, antenna.

229-231, male genitalia. 229, phallobase; 230, sipho; 231, enlarged

siphonal apex.

462 R.D. GORDON & C. CANEPARI

Figs. 232-243 - Hyperaspis onerata structures. 232-235, habitus and variations.

236-237, pronotal variations. 238, antenna. 239-242, male genitalia.

239, phallobase; 240, enlarged basal lobe; 241, sipho; 242, enlarged

siphonal apex. 243, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 463

Figs. 244-248 - Hyperaspis bicruciata habitus and male genitalia. 244-245, habitus.

246-248, male genitalia. 246, phallobase; 247, sipho; 248, enlarged

siphonal apex.

464 R.D. GORDON & C. CANEPARI

Figs. 249-254 - Hyperaspis joannae structures. 249-250, habitus. 251, antenna, basal

article missing. 252-254, male genitalia. 252, phallobase; 253, sipho;

254, enlarged siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 465

Figs. 255-261 - Hyperaspis albopunctata structures. 255-256, habitus. 257, antenna.

258-260, male genitalia. 258, phallobase; 259, sipho; 260, enlarged

siphonal apex. 261, female genitalia.

466 R.D. GORDON & C. CANEPARI

Figs. 262-266 - Hyperaspis apicaspis habitus and male genitalia. 262-263, habitus.

264-266, male genitalia. 264, phallobase; 265, sipho; 266, enlarged

siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 467

Figs. 267-272 - Hyperaspis delicata structures. 267-268, habitus. 269, antenna.

270-272, male genitalia. 270, phallobase; 271, sipho; 272, enlarged

siphonal apex.

468 R.D. GORDON & C. CANEPARI

è, Sd

Figs. 273-277 - Hyperaspis mimica structures. 273, habitus. 274, metendosternite.

275-277, male genitalia. 275, phallobase; 276, sipho; 277, enlarged

siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 469

218

bel |

| ! È I |

283 ‘ 7 } ct /

282 39 par. | |

x, | |

284 | |

| |

Figs. 278-285 - Hyperaspis laterimacula and H. howdeni habitus and male genitalia.

278-281, H. laterimacula. 278-279, habitus. 280-281, male genita-

lia. 280, phallobase; 281, sipho. 282-285 H. howdeni. 282, habitus.

283-285, male genitalia. 283, phallobase; 284, sipho; 285, enlarged

siphonal apex.

R.D. GORDON & C. CANEPARI

470

286

fav

fur

i fe

PIE

} Î FA

(i a

i Pd

3 $ ia e;

i a Ri

| a 4

SI E Li =

\ \ ae

LX \ |

x 9 i i

Sa x f }

x N, 5 i

NS VAI:

OI E A of

289 SRE

291

Figs. 286-291 - Hyperaspis chocoi structures. 286, habitus. 287, antenna. 288-291,

male genitalia. 288, phallobase; 289, enlarged basal lobe; 290, sipho;

291, enlarged siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 471

inn

ri — ad

secenoasnannoens TI I

An Vandel i È pe a

Cranes

H ti

_ | 296 |

“ I

4 |

292 993 I t. n x)

Figs. 292-296 - Hyperaspis atra habitus and male genitalia. 292-293, habitus.

294-296, male genitalia. 294, phallobase; 295, sipho; 296, enlarged

siphonal apex.

472

R.D. GORDON & C. CANEPARI

Figs. 297-303 -

Hyperaspis recordata habitus and genitalia. 297-298, habitus.

299-301, male genitalia. 299, phallobase; 300, sipho; 301, enlarged

siphonal apex. 302-303, female genitalia. 302, complete genitalia;

303, basal unit, ventral view.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 473

307

Figs. 304-307 - Hyperaspis eupaleoides habitus and male genitalia. 304, habitus.

305-307, male genitalia. 305, phallobase; 306, sipho; 307, enlarged

siphonal apex.

474 R.D. GORDON & C. CANEPARI

311

Figs. 308-311 - Hyperaspis mariposa habitus and male genitalia. 308, habitus.

309-311, male genitalia. 309, phallobase; 310, sipho; 311, enlarged

siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 475

Figs. 312-318 - Hyperaspis simlaensis structures. 312-313, habitus. 314, antenna.

315-317, male genitalia. 315, phallobase; 316, sipho; 317, enlarged

siphonal apex. 318, female genitalia.

476 R.D. GORDON & C. CANEPARI

SORIANO

SERN

SEX

3 a

324

Figs. 319-324 - Hyperaspis lindae structures. 319-320, habitus. 321, antenna.

322-324, male genitalia. 322-phallobase; 323, sipho; 324, enlarged

siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 477

Figs. 325-328 - Hyperaspis c-nigrum habitus and female genitalia. 325-326, habitus.

327-328, female genitalia. 327, complete genitalia; 328, lateral view

of basal unit.

R.D. GORDON & C. CANEPARI

478

et +2 RSS see Ti uu

SOR ING

329

330

331, complete female genitalia and ventral view of basal unit.

Figs. 329-331 - Hyperaspis sagittata habitus and female genitalia. 329-330, habitus.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 479

332

Figs. 332-325 - Hyperaspis guilloryi structures. 332-333, habitus. 334, antenna.

335, female genitalia.

480 R.D. GORDON & C. CANEPARI

Figs. 336-338 - Hyperaspis pseudopavida habitus and female genitalia.

336-337, habitus. 338, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 481

Figs. 339-341 - Hyperaspis istmina habitus and antenna. 339-340, habitus.

341, antenna.

482 R.D. GORDON & C. CANEPARI

345

Figs. 342-346 - Hyperaspis satipoensis structures. 342-343, habitus. 344, antenna,

last 5 articles missing. 345-346, female genitalia. 345, genital plates;

346, basal and distal portions.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 483

[ \ roscoenining ssi ” y i f ap

earnest, # y f

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A ì i ‘ FA: i a al

} ‘ if A \ /

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XY 17//7 N

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350

Figs. 347-351 - Hyperaspis abertha structures. 347-348, habitus. 349, antenna.

350-351, female genitalia. 350, genital plates; 351, basal and distal

portions.

484 R.D. GORDON & C. CANEPARI

355

Figs. 352-356 - Hyperaspis circumclusa structures. 352-353, habitus. 354, protibia.

355-356, female genitalia. 355, complete genitalia with basal unit in

ventral view; 356, lateral view of basal unit.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 485

Figs. 357-360 - Hyperaspis aemulata structures. 357-358, habitus. 359, antenna.

360, female genitalia.

R.D. GORDON & C. CANEPARI

486

‘fe

a os

ao 3a

See

364

Figs. 361-365 - Hyperaspis dispar structures. 361-362, habitus. 363, antenna.

364, metendosternite. 365, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 487

Figs. 366-372 - Hyperaspis histrionica and H. ayacucho habitus and structures.

366-367, H. histrionica habitus. 368-372, H. ayacucho structures.

368, habitus. 369, antenna. 370-372, male genitalia. 370, phal-

lobase; 371, sipho; 372, enlarged siphonal apex.

488 R.D. GORDON & C. CANEPARI

Figs. 373-378 - Hyperaspis herrerai structures. 373-374, habitus. 375, antenna.

376-378, male genitalia. 376, phallobase; 377, sipho; 378, enlarged

siphonal apex.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 489

Figs. 379-385 - Hyperaspis vredenburgi structures. 379-380, habitus. 381, antenna.

382-384, male genitalia. 382, phallobase; 383, sipho; 384, enlarged

siphonal apex. 385, female genitalia.

490 R.D. GORDON & C. CANEPARI

386 388 389

394

390

395 396

Figs. 386-396 - Hyperaspis festiva habitus and variations.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 491

Figs. 397-401 - Hyperaspis festiva structures. 397, antenna. 398-400, male

genitalia. 398, phallobase; 399, sipho; 400, enlarged siphonal

apex. 401, female genitalia.

492 R.D. GORDON & C. CANEPARI

Figs. 402-409 - Hyperaspis germaini structures. 402-403, habitus. 404, antenna.

405, prosternum. 406-408, male genitalia. 406, phallobase;

407, sipho; 408, enlarged siphonal apex. 409, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 493

416 418

Figs. 410-418 - Hyperaspis connectens structures. 410-412, habitus and variations.

413, antenna. 414, metendosternite. 415-417, male genitalia. 415,

phallobase; 416, sipho; 417, enlarged siphonal apex. 418, female

genitalia.

494 R.D. GORDON & C. CANEPARI

Figs. 419-421 - Hyperaspis ingrata habitus and female genitalia. 419, habitus.

420-421, female genitalia. 420, complete genitalia; 421, ventral view

of basal unit.

495

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI

se pi

singe cl

%%,

MOTTO

en sti

susan

e ss

Figs. 422-429 - Hyperaspis funesta structures. 422-423, habitus. 424, antenna. 425,

mendosternite. 426-428, male genitalia. 426, phallobase; 427, sipho;

428, siphonal apex. 429, female genitalia.

496 R.D. GORDON & C. CANEPARI

434

Le 2

Bet FU

eh ei H $ x

(at j pi OS H n

“un j gta H 5

af (SERIE E i if

$ # 3 i IS ae ee oe SD } i Sa ae

x ¥ 1 $ TO ENTI; (I } te È $ ts i

i fmt 3 i AE: E, GEE To Agati Fist

i i #4 24 | IT £4 aetkis

13 by 4 È ij AM j i

ii ed? Gime eae a PRESS Cru |a

3531 “me + ju tS 4

Figs. 430-436 - Hyperaspis nana habitus and genitalia. 430-431, habitus. 432-434,

male genitalia. 432, phallobase; 433, sipho; 434, enlarged siphonal

apex. 435-436, female genitalia. 435, genital plates; 436, distal por-

tion, ventral view of basal portion.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 497

Figs. 437-448 - Hyperaspis sphaeridioides structures. 437-440, habitus and varia-

tions. 441, antenna. 442, metendosternite. 443-446, male genitalia.

443, phallobase, ventral view; 444, phallobase, lateral view; 445,

sipho; 446, enlarged siphonal apex; 447-446, female genitalia. 447,

complete genitalia; 448, lateral view of basal unit.

498 R.D. GORDON & C. CANEPARI

Figs. 449-457 - Hyperaspis conclusa structures. 449-452, habitus and variations.

453, antenna. 454-456, male genitalia. 454, phallobase; 455; sipho;

456, enlarged siphonal apex. 457, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 499

458

460

463

461

462

Figs. 458-463 - Hyperaspis arida habitus and genitalia. 458, habitus. 459-461, male

genitalia. 459, phallobase; 460, sipho; 461, enlarged siphonal apex.

462-463, female genitalia. 462, complete genitalia, basal unit in

ventral view; 463, basal unit in lateral view.

500 R.D. GORDON & C. CANEPARI

Figs. 464-471 - Hyperaspis elegantissima structures. 464-466, habitus and variations.

467, antenna. 468-470, male genitalia. 468, phallobase; 469, sipho;

470, enlarged siphonal apex. 471, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 501

Figs. 472-480 - Hyperaspis longula structures. 472-475, habitus and variations. 476,

antenna. 477-479, male genitalia. 477, phallobase; 478, sipho; 479,

enlarged siphonal apex. 480, female genitalia.

502 R.D. GORDON & C. CANEPARI

Lissa EZIO ue

4A di

Figs. 481-490 - Hyperaspis prolata structures. 481-482, habitus. 483, antenna. 484,

metendosternite. 485-487, male genitalia. 485, phallobase; 486,

sipho; 487, siphonal apex. 488-490, female genitalia. 488, genital

plates; 489, ventral view of basal and distal units; 490, lateral view

of basal unit.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 503

Figs. 491-494 - Tenuisvalvae unipunctata structures. 491-492, habitus. 493, antenna.

494, female genitalia.

504 R.D. GORDON & C. CANEPARI

Figs. 495-498 - Tenuisvalvae peregrina structures. 495-496, habitus. 497, antenna.

498, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 505

Figs. 499-505 - Tenuisvalvae deyrollei structures. 499-500, habitus. 501, antenna.

502-504, male genitalia. 502, phallobase; 503, sipho; 504, enlarged

siphonal apex. 505, female genitalia.

506 R.D. GORDON & C. CANEPARI

Figs. 506-512 - Tenuisvalvae notata structures. 506, habitus. 509, antenna. 507-511,

male genitalia. 507, ventral view of phallobase; 508, lateral view of

phallobase; 510, sipho; 511, enlarged siphonal apex. 512, female

genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI BOY

520 | y

Figs. 513-521 - Tenuisvalvae bisquinquepustulata structures. 513-515, habitus. 516,

antenna. 517-519, male genitalia. 517, phallobase; 518, sipho; 519,

enlarged siphonal apex. 520-521, female genitalia. 520, complete

genitalia; 521, lateral view of basal unit.

508 R.D. GORDON & C. CANEPARI

Figs. 522-528 - Tenuisvalvae ecoffeti structures. 522-523, habitus. 524, antenna.

525-527, male genitalia. 525, phallobase; 526, sipho; 527, enlarged

siphonal apex. 528, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 509

Figs. 529-535 - Tenuisvalvae parenthesis structures. 529-530, habitus. 531, antenna.

532-534, male genitalia. 532, phallobase; 533, sipho; 534, enlarged

siphonal apex. 535, female genitalia.

510 R.D. GORDON & C. CANEPARI

Figs. 536-542 - Tenuisvalvae caucaensis structures. 536-537, habitus. 538, antenna.

539-541, male genitalia. 539, phallobase; 540, sipho; 541, siphonal

apex. 542, female genitalia.

SOUTH AMERCAN COCCINELLIDAE, XI: HYPERASPIDINI 511

Figs. 543-551 - Tenuisvalvae bromelicola structures. 543-544, habitus. 545, antenna.

546, metendosternite. 547-549, male genitalia. 547, phallobase; 548,

sipho; 549, enlarged siphonal apex. 550-551, female genitalia. 550,

complete genitalia; 551, ventral view of basal unit.

512 R.D. GORDON & C. CANEPARI

Figs. 552-553 - Tenuisvalvae gnoma habitus and female genitalia. 552, habitus. 553,

female genitalia.

513

RES'LIGUSTICAE CGLIII

Luca BAGHINO*

MONITORAGGIO DELLA MIGRAZIONE

POST-RIPRODUTTIVA DEI RAPACI DIURNI NELLA

ZONA DI PROTEZIONE SPECIALE IT 1331578

“BEIGUA-TURCHINO”

(AVES, ACCIPITRIFORMES ET FALCONIFORMES)

INTRODUZIONE

Da circa dieci anni la migrazione dei rapaci rappresenta, nel

panorama ornitologico nazionale, un ambito tematico di notevole

interesse ed in continua evoluzione. Tra i siti “storici” dall’inizio

dei primi controlli sulla migrazione pre-riproduttiva risalenti alla

metà degli anni ottanta del secolo da poco passato (BAGHINO &

Leucio 1989 e 1990; BAGHINO 1996; ZALLES & BILDSTEIN 2000), i

contrafforti e le vallate appenniniche di Arenzano (GE) continuano

ad essere luogo di conteggi sistematici e standardizzati tali da offrire

una serie temporale di dati relativamente estesa.

La designazione della Zona di Protezione Speciale (ex Direttiva

79/409/CEE “Uccelli”) IT 1331578 “Beigua-Turchino” e gli obbli-

ghi di monitoraggio imposti dalla Direttiva 79/409 hanno infatti

comportato l’avvio di nuovi conteggi! finalizzati ad acquisire dati

aggiornati ed altri elementi conoscitivi intorno al fenomeno migra-

torio delle specie appartenenti agli ordini degli Accipitriformes e

Falconiformes oggetto di tutela.

* Centro Ornitologico e di E.A. di Case Vacca c/o Ente Parco del Beigua — LIPU

Liguria Via Marconi 165 16011 Arenzano (GE).

' I conteggi sono stati svolti nel 2000 e 2001 con contributi dalla Provincia di Genova

- Assessorato all Ambiente e nel 2004, 2005 e 2006 su incarico dell'Ente Parco del

Beigua nell’ambito delle azioni previste dal Progetto DOCUP Obiettivo 2 2000/2006

“Parco del Beigua: un Parco per la biodiversità: conoscenza, tutela e valorizzazione

della ZPS Beigua - Turchino”, cofinanziato dall’ Unione Europea.

514 L. BAGHINO

Essendo la migrazione post-riproduttiva dei rapaci molto meno

nota in sede locale fino al 1999, è stato avviato un periodo di cam-

pionamento standardizzato di 12 giorni, avente come specie obiet-

tivo il Biancone Circaetus gallicus (Gmelin, 1788).

METODI, TECNICHE E PROTOCOLLI DI LAVORO

Il monitoraggio è stato condotto sui contrafforti montuosi di

Arenzano (44° 25° 23” N - 8° 40° 53” E), nella parte sud-orientale

della ZPS “Beigua-Turchino”.

L’area insiste sul versante tirrenico dello spartiacque principale

appenninico, solcato da valli trasversali brevi e acclivi: il sistema

orografico del Ponente genovese, nei territori comunali di Genova

ed Arenzano, registra la minima ampiezza trasversale e 1 minimi

valori di distanza dei rilievi dalla costa.

Le osservazioni hanno avuto luogo prevalentemente in due

postazioni ubicate su culminazioni a quote tra i 300 e 1 600 m s.l.m.

lungo un crinale secondario che, dividendo le valli dei ri1 Cantarena

e Lissolo secondo un asse NNE-SSO, presenta ampie vedute pano-

ramiche con campo visivo di almeno 180 gradi in direzione est/nord-

est, cioè verso i settori di provenienza degli individui migranti. Le

postazioni sono state di giorno in giorno scelte sulla base dei venti

dominanti e dell’altitudine della copertura nuvolosa che influenzano

la dislocazione e l’altezza da terra delle rotte di volo.

Il conteggio del Biancone e delle altre specie di rapaci diurni

(Accipitriformes, Falconiformes) è stato continuativamente svolto dal

2000 al 2006 nel periodo dal 15 al 26 settembre con sessioni giorna-

liere di osservazione di otto ore (dalle 9 alle 17, ora solare) suddivise

in quattro fasce orarie di transito (9-11; 11-13; 13-15; 15-17). Le

osservazioni sono state compiute lungo l’intero arco temporale e sono

state temporaneamente sospese solo in presenza di rovesci piovosi.

Il periodo di conteggio è stato disposto attorno al picco di pas-

saggio (22 e 23 settembre) del Biancone quale era risultato in seguito

a rilievi compiuti in giornate discontinue nel 1998 e 1999, con avvio

dalla metà del mese in modo da non escludere apprezzabili movi-

menti migratori della specie già in corso dopo il 5-10 settembre.

Le osservazioni sono state compiute mediante binocoli (7x42) e

cannocchiale (28x66, 32x77) montato su treppiede.

MIGRAZIONE DI RAPACI DIURNI 515

Nelle giornate con più osservatori divisi in due squadre, si è

fatto ricorso a radio VHF per lo scambio di informazioni, finalizzato

a contenere 1 casi di mancato avvistamento di individui in migra-

zione e ad escludere nel contempo doppi conteggi.

Sono state raccolte, quando possibile, informazioni relative

all’età dei soggetti migranti di Biancone, sulla base dei caratteri e

dei criteri descritti da FORSMAN 1999 e da CLARK 2003.

I dati sono stati annotati su due tipi di schede:

* la prima, specificamente studiata per il monitoraggio del

Biancone, riportava dati relativi ai soggetti migranti della

specie con particolare riferimento all’età e ad eventuali det-

tagli sullo stato di muta;

* la seconda, utilizzata dal 1991 per i conteggi dei rapaci

migratori, riguardava il transito migratorio di tutte le altre

specie di Accipitriformes e Falconiformes, con maggiori det-

tagli riguardanti, oltre all’ora di avvistamento, al numero,

all’età e o al sesso degli individui, le variabili meteorologi-

che (forza e direzione del vento secondo la scala Beaufort,

copertura e stato del cielo, visibilità) e di passaggio (prove-

nienza, direzione di svanimento e altitudine di volo in base a

fasce altimetriche stimate di 300 m ognuna, quest’ultima dal

2005 misurata quando tecnicamente possibile con telemetro

laser).

Non sono state riportate sulle schede le osservazioni dubitativa-

mente attribuibili a individui locali (è il caso del Biancone, presente

come estivo nidificante di norma fino al 25 settembre, della Poiana,

del Gheppio, dello Sparviere); sono stati considerati in migrazione

1 soggetti osservati per almeno tre minuti senza soste e significative

deviazioni dalla direzione standard di volo.

In considerazione della distribuzione verosimilmente non nor-

male dei dati derivati da conteggi, si sono impiegati vari test stati-

stici non parametrici.

Le specie sono ordinate secondo l’assetto sistematico e la nomen-

clatura adottati da BRICHETTI & Massa (1998).

516 L. BAGHINO

RISULTATI

Sono stati complessivamente osservati 4907 individui apparte-

nenti a 12 specie di Accipitriformes e a 4 di Falconiformes (Tab. I)

Tab. I - Migrazione post-riproduttiva dei Rapaci dal 15 al 26 settembre nella ZPS

IT 1331578 “Beigua-Turchino”

Migrazione post-riproduttiva dei Rapaci dal 15 al 26 settembre nella ZPS IT

1331578 “Beigua-Turchino

2000 | 2001 | 2002 | 2003 | 2004 2006 | TOTALE

391

2005

ilvus migrans

ilvus milvus

Circaetus gallicus

Circus aeruginosus

Circus cvaneus

Buteo sp.

Aquila pomarina

teraaetus pennatus

Accipitridae indeterminato

Pandion haliaetus

alco tinnunculus

alco subbuteo

alco peregrinus

pudualy cfd Pau

Pe wi al

Pa SE

CIA ECT B24

SORATTE

deli veli, 97°

pe Odie! ddp

to aonlao ik od

cer Se ato:

Bader rie

stru ad l'alto)

lio Vila

871

MIGRAZIONE DI RAPACI DIURNI 517

Ordine ACCIPITRIFORMES

Famiglia ACCIPITRIDAE

Falco pecchiaiolo Pernis apivorus (Linnaeus, 1758)

Seconda specie in ordine di abbondanza, con indice di transito

orario di 0,68 ind/h (Fig. 1).

La data mediana di migrazione della specie è risultata il 19 set-

tembre. Nel confronto tra i contingenti annualmente campionati non

è emersa una differenza significativa (test di Kruskal-Wallis H =

3,50; g.l. = 6; P>0,05). Solo a 180 individui (46,15% del totale) si

è potuta assegnare l’età con ragionevole certezza: 162 (90,0%) sono

risultati giovani e solo 18 (10,00%) adulti. Massimo conteggio: 18

indd. il 17.IX.2006.

Pernis apivorus

giorno

Fig. 1 - Fenologia della migrazione del Falco pecchiaiolo Pernis apivorus nel peri-

odo di campionamento.

Nibbio bruno Milvus migrans (Boddaert, 1783)

Specie risultata scarsamente frequente, con indice di passaggio

pari a 0,02 ind/h. Massimo conteggio: 3 indd. il 18.IX.2004.

518 L. BAGHINO

Nibbio reale Milvus milvus (Linnaeus, 1758)

Specie di comparsa poco frequente, con indice di transito pari a

0,03 ind/h. Massimo conteggio: 3 indd. il 24.IX.2005.

Biancone Circaetus gallicus (Gmelin, 1788)

Gli aspetti legati alla definizione di uno schema migratorio

denominato a “circuito” (AGOSTINI et al. 2002a), all'utilizzo di una

strategia migratoria di tipo conservativo (AGOSTINI et al. 2002b) e di

una strategia migratoria differenziale tra giovani e adulti (AGOSTINI

et al. 2004) attraverso l’Italia sono già stati analizzati in precedenti

lavori impiegando 1 risultati dei conteggi del 2000 e del 2001.

Il Biancone costituisce il 79,93% di tutti i rapaci migratori con-

teggiati nel periodo controllato; la sua migrazione presenta due dif-

ferenti picchi secondo un andamento bimodale (Fig. 2). Gli indici

di passaggio (Fig. 3), con linea di tendenza ad andamento crescente

(u=6,19+1,57 D.S. ind/h), riflettono primariamente l’aumentata effi-

cienza del campionamento, la più efficace copertura territoriale e le

migliori conoscenze delle dinamiche del passaggio e del comporta-

mento migratorio sito-specifico acquisite con gli anni.

Circaetus gallicus

COOL LL TETTI

500

400

z 300 I

giorno

Fig. 2 - Fenologia della migrazione del Biancone Circaetus gallicus nel periodo di

campionamento.

MIGRAZIONE DI RAPACI DIURNI 519

= Indice di transito orario

Circaetus gallicus

Indici di transito orario

«==@=== Ore di effettiva osservazione

= = Lineare (Indice di transito orario)

msde hrs dey srt tn Etre nt al if Mc nee eee CRE nn bent be ane 98

2000 2001 2002 2003 2004 2005 2006

anno |

N ind ora

N ore impiegate

—_

©)

Fig. 3 - Indici di passaggio del Biancone Circaetus gallicus.

La data mediana di migrazione è risultata il 20 settembre. Nei

periodi di campionamento il numero annuale degli individui migranti

non è variato in misura statisticamente significativa (test di Kruskal-

Wallis H = 4,81; g.l. = 6; P>0,05). Solo a 1799 (45,84% del totale)

individui si è potuta assegnare l’età con ragionevole certezza: 1487

(82,65%) sono risultati adulti, 52 (2,90%) immaturi e 260 (14,45%)

giovani (Fig. 4).

Circaetus gallicus - Età |

6,6%

1,3%

El giovani

EH immaturi

O adulti

54,2%\ 37,9%

indeterminati

Fig. 4 - Classi di età del Biancone Circaetus gallicus.

520 L. BAGHINO

Delle quattro fasce orarie di due ore, il flusso migratorio risulta

maggiormente distribuito in quella dalle 11 alle 13 (x? = 249,79;

g.1 = 6; P<0,05) con il 32,8% delle osservazioni. Confrontando le

mediane degli effettivi transitati nelle giornate con venti da nord

(14 indd.) e con venti da sud (45 indd.), i bianconi migrano pre-

valentemente con venti provenienti dai quadranti meridionali (test

di Mann-Whitney U = 359, Nnord = 23, Nsud = 61, z = 3,43;

P<0,05). Massimo conteggio: 187 indd. il 25.IX.2001.

Falco di palude Circus aeruginosus (Linnaeus, 1758)

Terza specie in ordine di abbondanza con indice di passaggio di

0,34 ind/h (Fig. 5).

Circus aeruginosus

giorno

Fig. 5 - Fenologia della migrazione del Falco di palude Circus aeruginosus nel peri-

odo di campionamento.

La data mediana di migrazione della specie è risultata il 20 set-

tembre. Non è stata riscontrata una differenza statisticamente signi-

ficativa tra i contingenti in migrazione nei periodi di campionamento

(test di Kruskal-Wallis H = 12,24; g.l. = 6; P>0,05).

Solo a 114 individui (52,29% del totale) si è potuta attribuire

eta con ragionevole certezza: di questi, 55 (48,24%) sono risultati

MIGRAZIONE DI RAPACI DIURNI 521

giovani o immaturi e 57 adulti: 35 dd (30,70%) e 24 92 (21,06%).

Massimo conteggio: 23 indd. il 19.IX.2001.

Albanella reale Circus cyaneus (Linnaeus, 1766)

Una sola osservazione (1 @ adulta il 16.IX.2003) di questo

migratore atteso in date mediamente più tardive rispetto al periodo

di controllo.

Sparviere Accipiter nisus (Linnaeus, 1758)

Specie relativamente regolare e frequente (Fig. 6), con indice

di transito pari a 0,12 ind/h. Massimo conteggio: 13 indd. il

17.IX.2002.

— Acipiter nisus ™= ™ Buteo buteo

giorno

Fig. 6 - Fenologia della migrazione dello Sparviere Accipiter nisus e della Poiana

Buteo buteo nel periodo di campionamento.

Poiana Buteo b. buteo (Linnaeus, 1758)

Poiana delle steppe Buteo b. vulpinus (Gloger, 1833)

Specie di comparsa regolare e piuttosto frequente (Fig. 6), con

522 L. BAGHINO

indice di passaggio di 0,07 ind/h. Massimo conteggio (sottospecie

nominale): 8 indd. il 24.IX.2005.

La forma sottospecifica vulpinus, la cui presenza non era stata

documentata in precedenza nel sito e piu in generale in Liguria,

almeno in parte per probabili assegnazioni alla sottospecie nomi-

nale e soprattutto per la mancanza di veri controlli sistematici della

migrazione post-riproduttiva, risulta molto scarsa e irregolare. Sono

stati riconosciuti con ragionevole certezza tre soggetti con fenotipo

vulpinus e caratteri ascrivibili al morfismo rossiccio. Massimo con-

teggio: 2 indd. il 17.IX.2000.

Poiana codabianca Buteo rufinus (Cretzschmar, 1827)

Una sola osservazione (1 juv. il 26.IX.2001) che costituisce cro-

nologicamente la quinta segnalazione ligure dopo le quattro presenti

in bibliografia, riepilogate da Mincozzi (1987).

Aquila anatraia minore Aquila pomarina C. L. Brehm, 1831

Un juv. transitato il 24.IX.2005 rappresenta l’unica osservazione

ligure documentata in bibliografia dalla seconda metà del dicianno-

vesimo secolo (BAGHINO 2006): va aggiunto un maschio in pelle cat-

turato a Genova-S.Martino d’Albaro nel novembre 1917 (MSNG

33579), periodo di comparsa relativamente tardiva per I’ Italia (BRI-

CHETTI & BeNUSSI 1992).

Aquila minore Hieraaetus pennatus (Gmelin, 1788)

Specie con indice di passaggio (Fig. 7) molto basso dal 2000

al 2004 compreso (0,01 ind/h), in notevole aumento nei due anni

successivi (0,70 ind/h) per effetto di un’evidente variazione delle

abbondanze, con differenze statisticamente significative (test di Kru-

skal-Wallis H = 35,15; g.l. = 6; P<0,05). Massimo conteggio: 23

indd,.1l 23.1XX2005,

MIGRAZIONE DI RAPACI DIURNI 523

Hieraaetus pennatus

giorno

Fig. 7 - Fenologia della migrazione dell'Aquila minore Huieraaetus pennatus nel

periodo di campionamento.

Famiglia PANDIONIDAE

Falco pescatore Pandion haliaetus (Linnaeus, 1758)

Specie sempre poco frequente, con indice di passaggio pari a

0,02 ind/h. Massimo conteggio: 2 indd il 20.IX.2005.

Ordine FALCONIFORMES

Famiglia FALCONIDAE

Gheppio Falco tinnunculus (Linnaeus, 1758)

Specie di comparsa poco frequente, ma presumibilmente sotto-

stimata per confusioni con i soggetti locali; indice di transito pari a

0,02 ind/h. Massimo conteggio: 2 indd. il 20.IX.2002. Vanno inol-

tre computati 10 individui di indeterminata attribuzione specifica tra

Gheppio Falco tinnunculus e Grillaio Falco naumanni.

524 L. BAGHINO

Smeriglio Falco columbarius (Linnaeus, 1758)

Due sole osservazioni di questo migratore il cui transito è da

attendersi più tardivo rispetto al periodo di controllo: 1 ind. il

26.IX.2003 e 1 ind. il 19.IX.2004.

Lodolaio Falco subbuteo (Linnaeus, 1758)

Specie di comparsa poco frequente, con indice di transito pari a

0,03 ind/h. Massimo conteggio: 2 indd. il 21.IX.2001, il 21.IX.2005,

il 29 102005 pl 2X. 2006,

Falco pellegrino Falco peregrinus "Tunstall, 1771

Specie di comparsa molto scarsa ed irregolare con osservazioni di

individui sempre singoli: tre di questi, con età e sottospecie determi-

nate, erano adulti appartenenti alla sottospecie nominale peregrinus.

DISCUSSIONE

1. Rotte locali e origini presunte dei contingenti

La direzione generale dei rapaci migratori è orientata da Est-

Nord-Est (75°-45°) verso Sud-Ovest (240°-245°).

Tutti i bianconi ad Arenzano seguono questa rotta parallela alla

costa che, nel Ponente genovese, inizia a piegare verso Sud-Ovest.

Oltre che dal Ponente genovese in risalita dell’alto versante

ligure-tirrenico (RUGGIERI et al. 2006), 1 bianconi, almeno in parte

(6,67% del totale, N = 262; mediana = 40 indd./anno), sono visti

pervenire anche con una provenienza più settentrionale che orientale

(circa 50°), lasciando spazio all’ipotesi che dei contingenti in arrivo

da aree geografiche a nord-est della Liguria e più generalmente a

nord dell’Italia centrale (Appennino ligure orientale e tosco-emi-

liano) convergano verso la Riviera di Ponente. Tuttavia questa rotta

di ingresso sul territorio con componente settentrionale tende ad

essere controllata con maggior difficoltà, per la notevole quota di

volo (900-1200 m) e la distanza (anche fino a 3 km) dalle postazioni

più a valle: tale situazione determina il mancato avvistamento dei

MIGRAZIONE DI RAPACI DIURNI 525

soggetti in transito e la conseguente sottostima soprattutto quando

la rotta tende a coincidere con la linea dello spartiacque principale

o a situarsi al di là di questo (ad es. alta valle del Cerusa-Passo del

Faiallo-Monte Reixa, 17.IX.2004).

Nel Falco di palude, la prevalente provenienza da Nord-Est

s inquadra nella fenologia temporale, per sesso e classi di età, dei

contingenti migratori originari dei Paesi dell’Europa centro- e

nord-orientale, in particolare della Repubblica Ceca (BELAUD 1993;

VORISEK & FORMANEK 1996).

Tranne due casi registrati per il Nibbio bruno e uno per il

Nibbio reale, solo le aquile minori sono state costantemente osser-

vate in senso contrario alla migrazione prevalente, con direzione da

Ovest/Sud-Ovest verso Est. Tale dato avvalora in modo considere-

vole l'ipotesi dell’appartenenza di questi contingenti a popolazioni

dell’areale occidentale europeo di distribuzione della specie (PRE-

MUDA & BAGHINO 2004).

2. Confronti con altri siti

E° possibile confrontare almeno in parte 1 dati ottenuti con quelli

di alcuni tra i maggiori siti di migrazione, regolarmente monitorati,

in relazione geografica con la Liguria.

a) Alpi Marittime francesi-Costa Azzurra

Nella ZPS “Beigua-Turchino” la migrazione post-riprodut-

tiva del Biancone trova riscontri significativi e associazioni su base

giornaliera con il sito delle Alpi Marittime (La Turbie-Fort de la

Revère, 160 km a Sud-Ovest, percorribili ad una media di circa

5 ore di volo: BeLAUD 1993), dove tuttavia gli effettivi risultano

mediamente due volte inferiori (BELAUD 2003; JARDIN 2002 e 2003:

Tab. II); qui, al contrario, il passaggio delle altre specie, soprattutto

Falco pecchiaiolo, Falco di palude (Tab. II), Sparviere, Lodolaio e

Gheppio, viene osservato con effettivi apprezzabilmente diversi (fino

a 7-10 volte superiori) e senza significative. correlazioni con quello

della ZPS.

Tale scostamento è interpretabile nel primo caso come il man-

cato intercettamento, almeno in parte, del flusso del Biancone sui

siti coperti, nel secondo con l'apporto di contingenti migratori pro-

526 L. BAGHINO

venienti da settori più settentrionali (Pianura Padana occidentale e

Basso Piemonte) che, non seguendo la costa della Liguria centrale,

evitano di transitare sui contrafforti collinari di Arenzano.

Tab. II - Correlazioni e confronti tra gli effettivi del Falco pecchiaiolo Pernis api-

vorus, del Biancone Circaetus gallicus e del Falco di palude Circus aerugi-

nosus in migrazione nella ZPS “Beigua-Turchino” e nel sito del Fort de

la Revère (Alpi Marittime francesi-Costa Azzurra) dal 15 al 26 settembre

del 2001, 2002 e 2003 (dati: Belaud, im litt.; JARDIN 2002 e 2003; N, =

totale dei tre anni nella ZPS Beigua-Turchino, N, = totale dei tre anni

nel sito del Fort de la Revère).

Test di Spearman Test di Spearman Dei Ù Son

i= 0562; N12,

Pernis apivorus ro N.S. ro N.S. P<0,05

NA alto N 14798 Test di Wilcoxon Test di Wilcoxon Test di Wilcoxon

t = 2,9, P<0,01 t = 3,1, P<0,01 t = 2,8, P<0,01

Test di Spearman Test di Spearman Test di Spearman

TA 0:92 NE 12, i= 0185, N52, Te ,085EN=103

Circaetus gallicus P<0.01 P<0.01 P=0.01

N, = 1407; N, = 507 | Test di Wilcoxon Test di Wilcoxon Test di Wilcoxon

= DAN) t = 3,1, P<0,01 t= 3, P<0,01

Test di Spearman Test di Spearman Test di Spearman

Circus aeruginosus ro N.S. ro N.S. r, N.S.

N, = 90; N, = 626 Test di Wilcoxon Test di Wilcoxon Test di Wilcoxon

t = 2,8, P<0,01 t = 2,9, P<0,01 t = 2,8, P<0,01

b) Alpi Marittime cuneesi (Valle Stura).

La migrazione del Biancone risulta ogni anno di norma inferiore

(1129 indd. dal 1991 al 2000: BeLAUD, GrrAuDO & TOoFFoLI 2001);

molto più importante è quella del Falco pecchiaiolo che fa registrare

uno o più picchi massimi tra il 25 agosto e il 5 settembre (TOFFOLI

2000, 2001 e 2003; GirauDpo 2004; Giraupo & 'TOFFOLI 2002 e 2005)

ed una netta flessione nelle settimane successive attestandosi su valori

confrontabili con quelli di Arenzano nel periodo di riferimento.

c) Alpi Apuane (LU)

Sito in relazione migratoria ben documentata (RUGGIERI et al.

2006), anche attraverso analoghi e simultanei periodi di controllo nel

MIGRAZIONE DI RAPACI DIURNI SID

2001 (AGOSTINI et al. 2002a) e 2006 (PREMUDA 2006), relativamente

alla risalita della costa tirrenica da parte del Biancone secondo lo

schema “a circuito”.

d) Isola di Pianosa (LI):

Sito interessato da parziali controlli contemporanei (relativa-

mente agli anni 2001, 2002, 2003, 2004, 2005 con archi di copertura

temporale di diversa estensione) che mostrano flussi più intensi di

falchi di palude, falchi pecchiaioli e gheppi (PAESANI & PoLITI 2002,

2003a e 2003b, 2004, 2005).

Per i Colli Asolani (TV) (MezzaviLLA, MarTIGNAGO & SILVERI

2001; MezzaviLLa, MARTIGNAGO & SILVERI 2003; MEZZAVILLA et

al. 2002, 2003, 2004, 2005 e 2006) e le Colline moreniche garde-

sane (BS-MN-VR) (GargionI 2002, 2003, 2004, 2005 e 2006), siti

di primaria importanza per la migrazione del Falco pecchiaiolo in

Italia, non è possibile effettuare confronti in quanto i controlli sono

anticipati e conclusi entro la prima decade di settembre, mostrando

notevoli concentrazioni di flussi migratori tra il 20 e il 30 agosto.

Analoghe considerazioni valgono per l’isola d'Elba interessata da

osservazioni preliminari sulla migrazione autunnale dei rapaci con-

dotte dal 6 al 12 Settembre 2004 con prevalenza di falchi di palude

e falchi pecchiaioli, allineate in generale con le risultanze dell’isola

di Pianosa (PREMUDA, PAESANI & CoccHI 2005), ma con più signifi-

cativi contingenti di sparvieri (PAESANI & VANNI 2006).

3 Analisi per specie

La ZPS “Beigua-Turchino” risulta eminentemente qualificata

dalla concentrazione dei flussi migratori degli Accipitriformes e dei

Falconiformes (GALLI, ALUIGI & BAGHINO 2006): per alcune specie è

possibile delineare determinati aspetti riguardanti gli schemi migra-

tori.

Falco pecchiaiolo Pernis apivorus (Linnaeus, 1758)

L'attuazione del monitoraggio, centrato attorno al picco della

migrazione del Biancone, in un periodo tardivo rispetto ai mas-

528 L. BAGHINO

simi passaggi del Falco pecchiaiolo rilevati nell’ Italia settentrionale

(TOFFOLI & BELLONE 1996; MEZzzaviILLA, MARTIGNAGO & SILVERI

2003), definisce per questa specie un transito di dimensione piut-

tosto contenuta e a prevalenza di soggetti giovani, analogamente a

quanto riscontrato nell’ Italia meridionale durante la seconda metà di

settembre (AGOSTINI & Locozzo 1997; AGOSTINI et al. 1999).

Biancone Circaetus gallicus (Gmelin, 1788)

I risultati qui presentati documentano l’esistenza di un flusso

migratorio post-riproduttivo notevole per dimensione, confrontabile

come ordine di grandezza con quello primaverile (BAGHINO 2003); a

livello nazionale, il sito di Arenzano risulta il secondo in ordine di

importanza dopo il Monte Colegno (Alpi Apuane, LU) e prima delle

Alpi Marittime (Valle Stura di Demonte, CN) in un periodo di tre

giorni consecutivi dell’ultima decade di settembre disposti a ridosso

del picco migratorio autunnale (RUGGIERI et al. 2006). Questo lavoro

condotto dal 2000 al 2004 in vari siti della Penisola Italiana, oltre

a documentare lo schema della migrazione “a circuito” attuata dalla

specie in risalita della Penisola italiana, evidenzia in particolare una

più marcata proporzione dei non adulti (20% costituito da giovani e

10% da immaturi) rispetto a quella osservata ad Arenzano: la riparti-

zione delle età in un simultaneo periodo di controllo nel 2001 risulta

assai simile a quella di Monte Colegno (78% adulti-16% giovani-6%

immaturi: AGOSTINI et al. 2004).

L'indice medio di passaggio ottenuto nei sette anni ad Arenzano

è dipendente dalla maggior estensione dell’arco temporale controllato

(12 giorni contro una finestra temporale di soli 3) e dall’inclusione

in essa di periodi non coincidenti o immediatamente contigui con

il picco della migrazione post-nuziale che determina indici più alti

(RUGGIERI et al. 2006): esso risulta sensibilmente maggiore di quello

rilevato nelle Alpi Marittime dal 1991 al 2000 (0,44 ind/h: BELAUD,

GIRAUDO & TorFrFoLi 2001) e nel 2001 (ToFFoLi 2001).

Lo scarso numero di individui censiti nel 2003 rispecchia l’ano-

malo andamento generale della migrazione dei Rapaci registrato

quell’anno nella maggior parte dei siti monitorati dell’Italia set-

tentrionale (PAESANI & PoLITI 2003; TorFoLI 2003) e della vicina

MIGRAZIONE DI RAPACI DIURNI 529

Francia (JARDIN 2003) e genericamente attribuito alle eccezionali

condizioni meteo-climatiche con alte temperature.

Falco di palude Circus aeruginosus (Linnaeus, 1758)

Sparviere Accipiter nisus (Linnaeus, 1758)

Per queste specie l’inferiore consistenza numerica dei contingenti

di passo rispetto ad aree geografiche vicine (Alpi Marittime francesi:

JARDIN 2002 e 2003; Isola di Pianosa: PAESANI & POLITI 2002, 2003

e 2004) pare attribuibile ad uno schema migratorio, ancora incerto

e da definire nella sua fenologia, che escluderebbe, limitatamente al

periodo controllato, i contrafforti collinari di Arenzano dalle rotte

principali attraverso l'alto versante ligure-tirrenico; queste, presu-

mibilmente confluenti nel Ponente Ligure dal lato settentrionale

dello spartiacque principale, paiono interessare maggiormente le

Alpi Marittime, mentre, più a sud, una distinta direttrice migratoria

affluisce sull’Arcipelago Toscano verso il ponte Sardo-Corso.

Poiana delle steppe Buteo buteo vulpinus (Gloger, 1833)

La direzione comune e l’imbrancamento con piccoli gruppi di

falchi pecchiaioli di alcuni degli individui attribuiti alla sottospecie

vulpinus (da considerarsi ormai non più accidentale ma migratrice

e svernante regolare per l’Italia: Corso 1999) potrebbero riferirsi a

soggetti appartenenti a popolazioni dell Europa orientale in movi-

mento attraverso la penisola italiana verso areali di svernamento

(Corso 2002).

Aquila minore Hieraaetus pennatus (Gmelin, 1788)

Il quadro delle conoscenze sulla migrazione post-riproduttiva

della specie in Italia, recentemente aggiornato (PREMUDA & BAGHINO

2004) è stato di fatto sconvolto dallo straordinario caso di migra-

zione di massa di aquile minori avvenuto nell’Ottobre del 2004 che

ha coinvolto nell’arco di dieci giorni oltre 500 individui in ingresso

dalla Francia (BAGHINO 2005; BAGHINO & PREMUDA 2005). La feno-

530 L. BAGHINO

logia dei contingenti (Fig. 8) appare in relazione con lo straordinario

fenomeno osservato nel 2004 e con i successivi movimenti migratori

da questo derivati (BAGHINO et al. in pubbl.), facendo della ZPS

“Beigua-Turchino” uno dei siti di maggior concentrazione del flusso

migratorio finora noti in Italia.

Hieraaetus pennatus - variazioni interannuali

R? ? = 0,7736 6 |

70 paint SI — a_i Pat i |

|} -$

10 |

pete? ines (orge

sr” is mi

D00 2001 ___ 2002 __ 2003 ___ 2004 _ 2005 __ 2006

-1 _____£904

anno

Fig. 8 - Variazioni interannuali di abbondanza dell'Aquila minore Hieraaetus pen-

natus nel periodo di campionamento con linea di tendenza polinomiale.

RINGRAZIAMENTI

Si ricordano con animo grato tutti gli attivisti LIPU che negli

anni hanno preso validamente parte ai conteggi, tra i quali: Carla

Rapetti, Rosangela Pedemonte, Graziano Lovato, Mauro Giorgini,

Nicola Leugio, Sandro Divano, Mauro Silveri, Rino Bontà.

Inoltre si ringraziano la Provincia di Genova (Assessorato

all'Ambiente) e Ente Parco del Beigua per aver sostenuto per più

anni le iniziative di monitoraggio e il Corpo Forestale dello Stato -

Coordinamento Regionale della Liguria per il rilascio dei permessi.

MIGRAZIONE DI RAPACI DIURNI Dod

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(CN) - Informamigrans, Parco Naturale Alpi Marittime, Valdieri: 10.

Giraupo L., & TOFFOLI R., 2005 - La migrazione post-riproduttiva in Valle Stura

(Demonte - CN) - Info Migrans, Parco Naturale Alpi Marittime, Valdieri: 16.

JARDIN M., 2002 - La migration post-nuptiale des oiseaux au Fort de la Revére en

2002 (Eze La Turbie, Alpes-Maritimes) - Rapport LPO-PACA, Hyéres-les-

Palmiers: 77 pp.

JARDIN M., 2003 - La migration post-nuptiale des oiseaux au Fort de la Revère en

2003 (Eze La Turbie, Alpes-Maritimes) - Rapport LPO-PACA, Hyères-les-

Palmiers: 57 pp.

MEZZAVILLA F., MARTIGNAGO G., PiccoLo F. & SILVERI G., 2003 - La migrazione

post-riproduttiva dei rapaci sopra i Colli Asolani (TV) Veneto - Informami-

grans, Parco Naturale Alpi Marittime, Valdieri: 12.

MEZZAVILLA F., MARTIGNAGO G., PiccoLo F. & SILVERI G., 2004 - La migrazione

post-riproduttiva dei rapaci sopra i Colli Asolani (Maser-TV) - Info Migrans,

Parco Naturale Alpi Marittime, Valdieri: 14.

MEZZAVILLA F., MarTIGNAGO G. & SILVERI G., 2001 - La migrazione dei rapaci in

provincia di Treviso (Colli Asolani-Maser) - Informamigrans, Parco Naturale

Alpi Marittime, Valdieri: 8.

MezzaviLLa F., MaRrTIGNAGO G. & SILVERI G., 2003 - Migrazione visibile post-

riproduttiva del Falco pecchiaiolo Pernis apivorus sopra 1 Colli Asolani (NE

Italia): anni 1994-2001 - Avocetta, Pavia, 27: 52-56.

MIGRAZIONE DI RAPACI DIURNI 533

MEZZAVILLA F., MARTIGNAGO G., SILVERI G. & PiccoLo F., 2002 - La migrazione

post-riproduttiva dei rapaci sopra i Colli Asolani (Maser-Treviso) - Informami-

grans, Parco Naturale Alpi Marittime, Valdieri: 10.

MEZZAVILLA F., MARTIGNAGO G., SILVERI G. & PiccoLo F., 2005 - La migrazione

post-riproduttiva dei rapaci sopra i Colli Asolani - Info Migrans, Parco Natu-

rale Alpi Marittime, Valdieri: 16.

MEZZAVILLA F., MaRTIGNAGO G., SILVERI G. & PiccoLo F., 2006 - La migrazione

post-riproduttiva dei rapaci sui Colli Asolani (TV) - Info Migrans, Parco Natu-

rale Alpi Marittime, Valdieri: 18.

Mincozzi T., 1987 - Revisione critica delle segnalazioni italiane di Poiana coda-

bianca (Buteo rufinus CR.). In: Baccetti N. & Spagnesi M. (a cura di), Rapaci

mediterranei III - Suppl. Ric. Biol. Selv., Ozzano nell’ Emilia, 12: 155-167.

PAESANI G. & POLITI P., 2002 - Monitoraggio della migrazione autunnale dei rapaci

diurni nell’isola di Pianosa (LI). Parco Nazionale Arcipelago Toscano - Infor-

mamigrans, Parco Naturale Alpi Marittime, Valdieri: 10.

PAESANI G. & PoLITI P., 2003a - Prime osservazioni sulla migrazione autunnale dei

rapaci diurni sull’isola di Pianosa nel Parco Nazionale Arcipelago Toscano -

Campi WWF Toscana - settembre 2001 - Avocetta, Pavia, 27: 71.

PAESANI G. & Pouiti P., 2003b - La migrazione autunnale dei rapaci sull’isola di

Pianosa (LI) Parco Nazionale Arcipelago Toscano - Informamigrans, Parco

Naturale Alpi Marittime, Valdieri: 12.

PAESANI G. & PoLiti P., 2004 - La migrazione autunnale dei rapaci sull’isola di

Pianosa (LI) - Info Migrans, Parco Naturale Alpi Marittime, Valdieri:14.

PAESANI G. & POLITI P., 2005 - La migrazione post-riproduttiva dei rapaci diurni

nel Parco Nazionale dell'Arcipelago Toscano (LI) - Info Migrans, Parco Natu-

rale Alpi Marittime, Valdieri: 16.

PAESANI G. & VANNI L., 2006 - La migrazione post-riproduttiva dei rapaci sull’isola

d’Elba (LI) - Info Migrans, Parco Naturale Alpi Marittime, Valdieri: 18.

PREMUDA G., 2006 - La migrazione dei rapaci: sulle-Alpi Apuane (LU): campi

autunno 2006 - Info Migrans, Parco Naturale Alpi-Marittime, Valdieri: 18.

Premuba G. & BacHino L. 2004 - La migrazione.autunnale dell’ Aquila minore

Hieraaetus pennatus attraverso la penisola italiana - Riv. ital. Orn., Milano, 74

(2): 103-115.

PREMUDA G., PAESANI G. & CoccHi L., 2005 - Osservazioni preliminari sulla migra-

zione autunnale dei rapaci sull’isola d’Elba - Riv. ital. Orn., Milano, 75 (1):

31-36.

Rucoieri L., PREMUDA G., BAGHINO L. & Grraupo L., 2006 - Esperienza di moni-

toraggio su vasta scala della migrazione autunnale del biancone Circaetus galli-

cus in Italia e nel Mediterraneo centrale - Avocetta, Pavia, 30: 76-80.

TorFFoLI R., 2000 - Parco Naturale Alpi Marittime - Informamigrans, Parco Naturale

Alpi Marittime, Valdieri. 6.

TOFFOLI R., 2001 - Parco Naturale Alpi Marittime - [nformamigrans, Parco Naturale

Alpi Marittime, Valdieri: 8.

TorFoLI R., 2003 - La migrazione dei rapaci in Valle Stura di Demonte (CN) Pie-

monte - Informamigrans, Parco Naturale Alpi Marittime, Valdieri: 12.

TorroLI R. & BELLONE C., 1996 - Osservazioni sulla migrazione autunnale dei

rapaci diurni sulle Alpi Marittime - Avocetta, Pavia, 20: 7-11.

534 L. BAGHINO

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raptor species from the Mediterranean area (pp. 381-385) - In: Muntaner J. &

Mayol J. (Eds.): Biologia y Conservacion de las Rapaces Mediterraneas, 1994.

Monografias, n.4, SEO, Madrid, 488 pp.

ZALLES J. & BILDSTEIN K., 2000 - Raptor watch: a global directory of raptor migra-

tion sites - Cambridge UK: BirdLife International and Kempton, PA, USA:

Hawk Mountain Sanctuary, BirdLife Conservation Series No. 9, 419 pp.

RIASSUNTO

In questo lavoro sono presentati 1 risultati di un monitoraggio della migrazione

post-riproduttiva dei rapaci diurni, avente come specie target il Biancone Ctrcaetus

gallicus, ed effettuato con analoghe modalita per sette anni consecutivi (dal 2000 al

2006) in un’area del Ponente Genovese (Arenzano) compresa nel Parco Naturale

del Beigua e nella Zona di Protezione Speciale IT 1331578 (sensu Dir. 79/409/CEE

“Uccelli”) “Beigua-Turchino”.

Sebbene alcuni aspetti relativi soprattutto al comportamento e allo schema

migratorio autunnale del Biancone attraverso la penisola italiana siano gia stati

affrontati, scopo di questa indagine è documentare il volume e la fenologia della

migrazione dei Rapaci attraverso una finestra temporale di dodici giorni (15-26

Settembre) nel sito di Arenzano, già noto per la concentrazione dei flussi migratori

in stagione pre-riproduttiva. In totale sono stati censiti 4907 individui appartenenti

a 12 specie di Accipitriformes e a 4 di Falconiformes.

ABSTRACT

Survey of the post-reproductive migration of diurnal raptors in the Special

Protection Area IT 1331578 “Beigua-Turchino” (Aves, Accipitriformes et

Falconiformes).

This paper reviews the results of a raptor migration survey, setting the Short-

toed Eagle Czrcaetus gallicus as the target species, that has been carried out yearly

from 2000 to 2006 using the same standardized methods and observation protocols

at the mountain site of Arenzano (25 km west of Genoa, Liguria): this is included

in the Beigua Natural Park and in a Special Protection Area (ex Directive 79/409/

CEE “Birds”) named “Beigua-Turchino”.

Although some matters regarding mainly the Short-toed Eagle’s migrating

behaviour and strategies through the Italian Peninsula have been already dealt with,

this study aims to document both the raptor migration volume and the species

phenology across a species-specific, 12-day sampling period in the second half of

September. An overall number of 4,907 migrating raptors belonging to 16 species

were counted at this site which was already known because of its raptor migration

importance during spring.

535

RES LIGUSTICAE CCLIV

DAVIDE BADANO*

CONTRIBUTO ALLA CONOSCENZA DEI NEUROTTERI

DELLA LIGURIA

(INSECTA, NEUROPTERIDA)

Lo stato di conoscenza dei Neurotteri s. l. italiani è piuttosto

eterogeneo, specialmente per gli aspetti corologici (LETARDI 1998 e

2004). In alcune aree (quali determinati settori delle Alpi e delle Pre-

alpi, Emilia-Romagna, la Sardegna e vari parchi naturali dell’Ita-

lia centrale e meridionale) si è concentrata l’attività di ricerca dei

neurotterologi mentre altre sono poco conosciute ed oggetto solo di

segnalazioni occasionali (LETARDI 1998 e 2005).

La Liguria rientra fra le regioni in cui non sono state effet-

tuate specifiche campagne di ricerca, per cui i pochi dati disponibili

riguardano per lo più specie comuni e ad ampia distribuzione, si rife-

riscono a poche località ed in molti casi sono anche piuttosto datati.

Allo scopo di incrementare le nostre conoscenze sulla neurotte-

rofauna ligure, rendo qui noti una serie di reperti da me effettuati

durante lo svolgimento della mia tesi di laurea triennale in Scienze

Naturali, sostenuta nel 2006 presso l’Università di Genova.

AREA DI STUDIO E METODI

I campionamenti sono stati effettuati, da maggio ad agosto 2006,

nelle stazioni sotto indicate, poste nell'estremo ponente ligure, nei

territori di vari comuni, tutti facenti parte della Provincia di Impe-

ria. L’unica eccezione è costituita dalla stazione di Sassello, situata

in Provincia di Savona.

* Via Queirolo 38, 18018, Arma di Taggia (IM).

536 D. BADANO

Perinaldo (IM)

Area posta in prossimità dello spartiacque tra la Valle Verbone e

la Val Nervia. Le catture (sia diurne che notturne) sono state effet-

tuate in località “Bana-Trumè” (560 m s.l.m.) e “Monte Rebuffao”

(590 m s.l.m.), in un ambiente di tipo mediterraneo che sul versante

est è particolarmente xerico (oliveti e gariga bassa arbustivo-erbacea

con prevalenza di Cistus spp.), mentre su quello ovest presenta con-

dizioni meno severe, che consentono lo sviluppo di un bosco misto

di latifoglie e, in alcuni punti, di sclerofille.

San Biagio della Cima (IM)

Zona collinare, posta sul versante sinistro della valle del torrente

Verbone. Le catture (soltanto notturne) sono state effettuate in loca-

lità “Madonna dell’Annunciata”, a circa 220 m s.l.m.; gran parte

dell’area è occupata da oliveti; la vegetazione spontanea è costituita da

boschi misti di latifoglie con prevalenza di Quercus pubescens Willd..

Castellaro (IM)

Zona collinare, situata in Valle Argentina. L’area prescelta per

le catture (solo diurne) è ubicata in località “Santuario di Lampe-

dusa” ad un’altitudine di 370 m s.l.m., in un versante collinare di

accentuata acclività esposto prevalentemente a sud/ovest, che nella

parte inferiore presenta una vegetazione costituita da un’alternanza

di boschi misti di latifoglie termofile, mentre in quella superiore,

più esposta e xerica, predominano formazioni arbustive ed erbacee.

Val Nervia (IM)

Nell’area sono stati effettuati una serie di campionamenti diurni,

salendo progressivamente d’altitudine, fino a giungere nelle imme-

diate vicinanze del “Passo Muratone”, a ridosso del confine con la

Francia a 1.158 m slm.; tale località, caratterizzata da un’accentuata

acclività, si trova al limitare della vegetazione arborea.

Un ulteriore campionamento è stato eseguito lungo le pendici del

Monte Pietravecchia e del Monte Toraggio, percorrendo il “Sentiero

degli Alpini” (Colla Melosa 1.545 m slm. — Gola dell’Incisa 1.685

m slm. — Passo di Fonte Dragurina 1.810 m slm.), zona tipicamente

montana con versante in forte acclività e con affioramenti rocciosi.

CONTRIBUTO ALLA CONOSCENZA DEI NEUROTTERI DELLA LIGURIA 537

La vegetazione è costituita da boschi di Larix decidua Miller nel

tratto iniziale, arbustiva con alberi sparsi nel tratto mediano, per

giungere ad una prateria alpina alle pendici del Monte Toraggio.

Prov.

Savona

Monti Toraggio e Pietravecchia

Passo Muratone

Drego

Aigovo

Francia

Perinald Castellaro

erinaldo SÒ di Lampedusa

Taggia Imperia

San Biagio della Cima

Ventimiglia Bordighera

Mar Ligure

Fig. 1 - Siti di cattura in provincia di Imperia.

Sassello (SV)

Le ricerche sono state effettuate nella località “Rocche dell’ Erro”,

a circa 330 m s.l.m., in un ambiente costituito da un bosco ceduo

misto di latifoglie, con prevalenza di Quercus petrea (Mattuschka)

Liebl., con versanti circostanti a Pinus spp.; le catture, sia diurne

che notturne, sono state concentrate lungo i margini boschivi.

Infine, si sono svolte raccolte sporadiche nelle seguenti località:

Colle di Nava (941 m s.l.m.), Andagna-Drego (715-1016 m s.l.m.),

Aigovo (681 m s.l.m.) e Bordighera, Vallone del Sasso (ca. 50 m

s.l.m.), tutte in Prov. di Imperia, e alla confluenza tra il Rio Arvigo

ed il Rio Canate, Davagna, in Prov. di Genova.

Le catture diurne sono state effettuate con l’uso del retino ento-

mologico, mentre per quelle notturne si è utilizzata una trappola

luminosa, costruita secondo le modalità descritte da PARENZAN &

De Marzo (1981) e BERIO (1985); come sorgente luminosa è stato

impiegato un neon U.V.A. da 15 Watt.

538 D. BADANO

RISULTATI

Per ogni specie sono indicati i siti di cattura, la categoria coro-

logica, la distribuzione in Italia (tratta da LETARDI & PANTALEONI

1996 e LETARDI 2005) e le precedenti citazioni per la Liguria. Se non

indicato diversamente, gli esemplari sono stati raccolti dall’autore.

Megaloptera

Sialidae

Sialis fuliginosa Pictet, 1836

Confluenza tra il Rio Arvigo ed il Rio Canate, Davagna (GE), 26.V.2006 (1 larva).

Specie sibirico-europea, segnalata per gran parte delle regioni

italiane e gia nota per l’area in esame (NavAs 1928, LETARDI 1994).

Non trattandosi di una larva all'ultimo stadio, l’identificazione ha un

piccolo margine di dubbio.

Raphidioptera

Raphidiidae

Dichrostigma flavipes (Stein, 1863)

Sassello (SV), 15.VI.2006 (2 99).

Specie a distribuzione centroeuropea, presente in Italia nelle

regioni dell’arco alpino ed in alcune aree dell’ Appennino settentrio-

nale; già segnalata per la Liguria da CASTELLANI (1957).

Ornatoraphidia flavilabris (Costa, 1855)

Monte Toraggio (IM), 27.VI.2006 (1 3).

Specie a distribuzione sudeuropea; presente lungo l Appennino,

è nota anche per le Alpi Marittime francesi. Prima segnalazione per

la Liguria.

CONTRIBUTO ALLA CONOSCENZA DEI NEUROTTERI DELLA LIGURIA 539

Xanthostigma aloysiana (Costa, 1855)

Perinaldo (IM), 8.VII.2006, trappola luminosa (1 9).

Specie a distribuzione W-mediterranea, che in Italia è conosciuta

per le regioni appenniniche; prima segnalazione per la Liguria.

Inocelliidae

Parainocellia bicolor (Costa, 1855)

Valle Argentina (IM), Drego, 27.V.2006 (1 larva); Sassello (SV), 14.VI.2006 (1

2); Busalla (GE), Bastia, 18.VI.2006, M. Bertolini leg. (1 9).

Specie euriecia a distribuzione sudeuropea, segnalata per quasi

tutte le regioni italiane, sia in aree alpine che appenniniche. Pre-

cedenti segnalazioni per la Liguria si devono a NavAs (1928) e

LETARDI (1994).

Neuroptera

Coniopterygidae

Coniopteryx (Coniopteryx) tineiformis Curtis, 1834

Alta Val Nervia (IM), 20.VIII.2006 (1 d).

Specie olartica, nota per Lombardia, Emilia-Romagna, Toscana,

Lazio, Campania e Sardegna; prima segnalazione per la Liguria.

Coniopteryx (Metaconiopteryx) esbenpeterseni Tjeder, 1930

Perinaldo (IM), 9.VIII.2006, trappola luminosa (1 d, 1 9).

Specie a distribuzione europea, conosciuta per Lombardia,

Veneto, Emilia-Romagna e Toscana; prima segnalazione per la

Liguria.

540 D. BADANO

Semidalis aleyrodiformis (Stephens, 1836)

Castellaro (IM), Santuario di Lampedusa, 27.V.2006 (1 6, 2 99.

Specie paleartica, nota per quasi tutte le regioni italiane; prima

segnalazione per la Liguria.

Osmylidae

Osmylus fulvicephalus (Scopoli, 1763)

Sassello (SV), 15.VI.2006 (1 ®).

Specie a diffusione eurasiatica, presente in Italia in prevalenza

nelle regioni centro-settentrionali, ma anche in Calabria. Precedenti

segnalazioni per la Liguria: INSOM et al. (1979).

Mantispidae

Mantispa styriaca (Poda, 1761)

Perinaldo (IM), 8.VII.2006, trappola luminosa (1 6,1 9 ).

Specie a distribuzione centroasiatica-europea-mediterranea, nota

per gran parte delle regioni italiane, fra cui la Liguria (LETARDI 1994).

Hemerobiidae

Hemerobius gilvus Stein, 1863

Sassello (SV), 15.VI.2006 (1 ®).

Si tratta di una specie a distribuzione S-europea, nota per gran parte

delle regioni italiane, già segnalata per la Liguria da MONSERRAT (1991).

Hemerobius humulinus Linné, 1758

Castellaro (IM), Santuario di Lampedusa, 13.VII.2006 (1 2).

CONTRIBUTO ALLA CONOSCENZA DEI NEUROTTERI DELLA LIGURIA 541

Comune specie a distribuzione olartica, nota per quasi tutte le

regioni italiane, già citata per la Liguria (MONSERRAT 1994).

Hemerobius micans Olivier, 1792

Colle di Nava (IM), 1.VII.2006 (1 ®).

Specie a diffusione sibirico-europea, segnalata per buona parte

d’Italia, tra cui la Liguria (MONSERRAT 1994).

Wesmaelius quadrifasciatus (Reuter, 1894)

Monte Toraggio (IM), 27.VI.2006 (1 9).

Specie sibirico-europea, nota in Italia per le regioni alpine;

prima segnalazione per la Liguria.

Wesmaelius subnebulosus (Stephens, 1836)

Bordighera (IM), Vallone del Sasso, 28.V.2006 (1 3); Castellaro (IM), Santuario di

Lampedusa, 22.VI.2006 (3 99).

Specie a distribuzione olartica, conosciuta per gran parte d’Ita-

lia. Precedenti segnalazioni per la Liguria sono riportate da MoN-

SERRAT (1994).

Chrysopidae

Italochrysa italica (Rossi, 1790)

Castellaro (IM), Santuario di Lampedusa, 13.VII.2006 (1 2); San Biagio della Cima

(IM), 25.VII.2006, trappola luminosa (1 d); Perinaldo (IM) 15.VII.2006, trappola

luminosa (1 9).

Si tratta di una specie a distribuzione mediterranea, conosciuta

per buona parte delle regioni italiane, tra cui la Liguria (CAPRA

1976).

542 D. BADANO

Chrysopa formosa (Brauer, 1850)

San Biagio della Cima (IM), 25.VII.2006, trappola luminosa (2 99) e 28.VII.2006,

trappola luminosa (1 6,1 9).

Specie asiatico-europea nota per gran parte delle regioni italiane,

ma non ancora per la Liguria, di cui questa costituisce la prima

segnalazione.

Chrysopa perla (Linné) sensu Schneider, 1851

Colle di Nava (IM), 6.VII.2006 (1 3); Sassello (SV), 15.VI.2006 (1 ?).

È una comune specie a distribuzione europea, nota per buona

parte delle regioni italiane. Precedenti segnalazioni per la Liguria

sono state rese note da INSOM et al. (1985).

Chrysopa viridana Schneider, 1845

Castellaro (IM), Santuario di Lampedusa, 22.VI.2006 (2 dd, 3 99) e 13.VII.2006 (1

6); Valle Argentina (IM), Aigovo, 1.VIII.2006 (1 gd).

Si tratta di una specie a diffusione euroturanica, conosciuta per

quasi tutte le regioni italiane; già segnalata per la Liguria da NAVAS

(1929).

Chrysoperla gruppo carnea (Stephens, 1836)

Castellaro (IM), Santuario di Lampedusa, 22.VI.2006 (3 dd, 1 9) e 13.VII.2006 (1

3); Valle Argentina (IM), Aigovo, 1.VIII.2006 (1 3); San Biagio della Cima (IM),

20.VII.2006 (4 29); Val Nervia (IM), 20.VIII.2006 (5 29); Val Nervia (IM), Passo

Muratone, 20.VIII.2006 (4 dé, 1 9).

Si tratta di un complesso di specie a distribuzione olartica, dif-

ficilmente distinguibili su base morfologica, con segnalazioni liguri

ad opera di NavAs (1933) e MONSERRAT (1980).

Gli esemplari sono stati catturati sia con il retino che con la

trappola luminosa in ambienti con vegetazione arborea.

CONTRIBUTO ALLA CONOSCENZA DEI NEUROTTERI DELLA LIGURIA 543

Dichochrysa clathrata (Schneider, 1845)

Sassello (SV), 15.VI.2006 (1 d).

Specie a distribuzione E-mediterranea, nota per quasi tutta

l’Italia. Questa costituisce la prima segnalazione sicura per la Ligu-

ria, poiché in ASPOÒCK et al. 1980 l’entità è citata come presente in

Liguria ma senza una indicazione precisa di località.

Dichochrysa flavifrons (Brauer, 1850)

Castellaro (IM), Santuario di Lampedusa, 3.VI.2006 (1 ®); San Biagio della

Cima (IM), 20.VII.2006 (1 2); Valle Argentina (IM), Aigovo, 1.VIII.2006 (1 6, 1

2); Val Nervia (IM), 20.VIII.2006 (1 3, 2 99); Val Nervia (IM), Passo Muratone,

20.VIII.2006 (1 2); Sassello (SV), 15.VI.2006 (2 od, 4 29.

Specie turanico-europea-mediterranea, segnalata per quasi tutte

le regioni italiane, tra cui anche la Liguria (cfr. PrINcIPI 1956 e

MONSERRAT 1980).

Dichochrysa prasina (Burmeister, 1839)

Bordighera (IM), Vallone del Sasso, 28.V.2006 (1 d, 1 2); Colla Melosa (IM),

San Biagio della Cima (IM), 20.VII.2006 (2 29); Val Nervia (IM), Passo Muratone,

20.VIII.2006 (1 3).

Specie a distribuzione eurasiatica, diffusa e comune in tutta

Italia, già nota per la Liguria (PRINCIPI 1956).

Dichochrysa zelleri (Schneider, 1851)

Castellaro (IM), Santuario di Lampedusa, 3.VI.2006 (2 2°); San Biagio della Cima

(IM), 20.VII.2006 (1 2); Perinaldo (IM), 15.VIII.2006 (1 2).

Specie a distribuzione E-mediterranea, nota per gran parte delle

regioni italiane; prima segnalazione per la Liguria.

Myrmeleontidae

Palpares libelluloides (Linné, 1764)

Perinaldo (IM), 9.VII.2006 (1 @).

544 D. BADANO

Specie turanico-mediterranea, segnalata per quasi tutte le regioni

peninsulari italiane (incluse Liguria ed Emilia-Romagna) oltre che per

alcune stazioni xerotermiche dell’Italia settentrionale. Le precedenti

segnalazioni liguri sono state pubblicate da CASTELLANI (1957), INSoM

et al. (1985), PANTALEONI (1990) e LETARDI & PANTALEONI (1996).

Myrmeleon (Morter) inconspicuus Rambur, 1842

Sassello (SV), 15.VI.2006 (5 larve, delle quali una è stata allevata, ottenendo 1 @).

Si tratta di una specie turanico-europea-mediterranea, nota per

quasi tutta l’Italia; prima segnalazione per la Liguria.

Distoleon tetragrammicus (Fabricius, 1798)

Perinaldo (IM), 9.VII.2006 (1 2); San Biagio della Cima (IM), 25.VII.2006 (1 ®);

Valle Argentina (IM), Aigovo , 1.VIII.2006 (1 ®, con ala posteriore malformata);

Sassello, 15.VI .2006 (1 ®, 1 larva)

È una specie a distribuzione europea, comune in tutta Italia e

già segnalata in Liguria da CASTELLANI (1957).

Neuroleon nemausiensis (Borkhausen, 1791)

San Biagio della Cima (IM), 23.VII.2006, trappola luminosa in uliveto (1 9).

Specie a distribuzione circumediterranea, nota in quasi tutta

Italia, Liguria compresa (CAPRA 1976).

Neuroleon microstenus (McLachlan, 1898)

Perinaldo (IM), 15.VIII.2006 (3 dd, 1 2).

Si tratta di una specie a distribuzione E-mediterranea, nota in

Italia per Emilia-Romagna, Lazio, Puglia, Basilicata e Sicilia. Prima

segnalazione per la Liguria.

Altri esemplari liguri (dati inediti) sono peraltro da tempo conser-

vati nelle collezioni del Museo Civico di Storia Naturale “G. Doria”

di Genova: Varazze (SV), VII.1918, leg. F. Invrea e VII.1934, leg.

CONTRIBUTO ALLA CONOSCENZA DEI NEUROTTERI DELLA LIGURIA 545

Borgioli; Genova, VIII.1940; Casella (GE), VIII.1932, leg. C. Man-

cini; Cavi di Lavagna (GE), 20.VII.1931, leg. E. Berio.

Macronemurus appendiculatus (Latreille, 1807)

San Biagio della Cima (IM), 25.VII.2006 (1 3) e 20.VIII.2006 (16); Perinaldo

(IM), 9.VII.2006 (1 2) e 15.VIII.2006 (4 9%; Civezza (IM), 25.VIII.2006 (1 d).

Specie diffusa nel Mediterraneo occidentale, nota per quasi

tutte le regioni italiane e già segnalata per la Liguria da CASTELLANI

(1957), INsoM ef al. (1979, 1985).

Gli esemplari sono stati raccolti in ambienti di gariga; l’indivi-

duo del 25.VII è stato catturato con la trappola luminosa.

Ascalaphidae

Libelloides coccajus (Denis & Schiffermiller, 1775)

Valle Argentina (IM), Andagna, 27.V.2006 (1 4,1 2); Drego (IM), 27.V.2006 (1 ®);

Colla Melosa, 24.VI.2006 (1 2); Sassello (SV), 15.VI.2006 (1 ®).

Si tratta di una specie a diffusione sud-europea, comune in

tutt'Italia e già segnalata per la Liguria (AISTLEITNER 1980).

Libelloides longicornis (Linné, 1764)

Sassello (SV), 15.VI.2006 (1 6,1 2).

Questa specie presenta una distribuzione sud-europea ed è

ampiamente diffusa in Italia; già segnalata per la Liguria da LETARDI

& PANTALEONI (1996). A Sassello convive nel medesimo habitat con

la specie precedente.

CONCLUSIONI

Durante l’indagine sono state reperite 32 specie di Neuropterida,

di cui: 1 di Megaloptera, 4 di Raphidioptera e 27 di Neuroptera.

Dieci specie (2 di Raphidioptera e $ di Neuroptera) risultano

546 D. BADANO

nuove per la Liguria: Ornatoraphidia flavilabris, Xanthostigma aloy-

stana, Coniopteryx tineiformis, C. esbenpeterseni, Semidalis aleyrodi-

formis, Wesmaelius quadrifasciatus, Chrysopa formosa, Dichochrysa

zelleri, Myrmeleon inconspicuus e Neuroleon microstenus.

Ad esse si aggiunge Dichochrysa clathrata, precedentemente

frutto di una segnalazione generica, non precisata, per l’area (ASPOCK

et al. 1980).

A riprova dello scarso stato di conoscenze sulla neurotterofauna

ligure, è interessante notare che nessuna di queste specie può con-

siderarsi rara in Italia; gran parte di esse, al contrario, è nota per

quasi tutte le regioni italiane. Non è difficile ipotizzare che, prose-

guendo nelle ricerche, il numero dei Neuropterida liguri possa ulte-

riormente incrementarsi.

RINGRAZIAMENTI

L’autore ringrazia il Prof. Giorgio Troiano ed il Dott. Loris

Galli (Dip.Te.Ris, Università degli Studi di Genova), relatori della

sua tesi di laurea triennale, il Dott. Roberto Poggi (Direttore del

Museo Civico di Storia Naturale “G. Doria”, Genova), per avergli

concesso di esaminare la collezione entomologica del Museo e per la

correzione del presente lavoro, e il Dott. Agostino Letardi (ENEA,

Roma) per i preziosi consigli e per l’incoraggiamento nella stesura

di questa nota.

BIBLIOGRAFIA

AISTLEITNER. E., 1980 - Die Arten des Genus Libelloides 'Tjeder, 1972, der Iberi-

schen Halbinsel (Neuroptera, Planipennia, Ascalaphidae) - Entomofauna, Linz,

Ie 2 34-229 fe

Aspock U., Aspock H. & HòLzEL H., 1980 - Die Neuropteren Europas. Eine zusam-

menfasende Darstellung der Systematik, Okologie und Chorologie der Neuro-

pteroidea (Megaloptera, Raphidioptera, Planipennia) Europas - Ed. Goecke &

Evers, Krefeld, 2 voll., 495+355 pp.

Aspòck U., Aspòck H. & RauscH H., 1991 - Die Raphidiopteren der Erde. Eine

monographische Darstellung der Systematik, Biologie, Okologie und Chorolo-

gie der rezenten Raphidiopteren der Erde, mit einer zusammenfassende Uber-

sicht der fossilen Raphidiopteren (Insecta: Neuropteroidea) - Ed. Goecke &

Evers, Krefeld, 2 voll., 730+550 pp.

BERNARDI IoRI A., KATHIRITHAMBY J., LETARDI A., PANTALEONI R. A. & PRINCIPI M.

M., 1995 - Neuropteroidea (Megaloptera, Raphidioptera, Planipennia), Mecop-

CONTRIBUTO ALLA CONOSCENZA DEI NEUROTTERI DELLA LIGURIA 547

tera, Siphonaptera, Strepsiptera. In: Minelli A., Ruffo S. & La Posta S. (eds),

Checklist delle specie della fauna italiana - Ed. Calderini, Bologna, 62, 1-20.

BerIo E., 1985 - Fauna d’Italia XXII, Lepidoptera Noctuidae I - Ed. Calderini,

Bologna, 1030 pp.

Capra F., 1976 - Quanto si sa sugli Odonati e Neurotteri dell’ Arcipelago Toscano -

Lavori Soc. it. Biogeogr., Forlì, 5 (N. S.): 541-560.

CASTELLANI O., 1957 - Contributo alla conoscenza della fauna entomologica d’Italia.

Neuroptera - Boll. Ass. romana Ent., Roma, 11-12: 9-14.

Insom E., DEL CENTINA P. & Carri S., 1979 - Alcuni Neurotteri Planipenni italiani

(Osmylidae, Ascalaphidae, Myrmeleonidae) - Redia, Firenze, 62: 35-52.

Insom E., DEL CENTINA P. & Carri S., 1985 - II Contributo alla conoscenza della

fauna neurotterologica italiana (Neuroptera: Raphidioptera, Planipennia) -

Redia, Firenze, 68: 105-121.

Insom E., DEL CENTINA P. & Carri S., 1986 - III Contributo alla conoscenza della

fauna neurotterologica italiana: Neurotteri Planipenni della Sardegna - Redia,

Firenze, 69: 651-654.

LETARDI A., 1993 - Dati sulla distribuzione italiana di Megaloptera Sialidae,

Raphidioptera Inocelliidae e Planipennia Mantispidae con particolare riferi-

mento all’ Italia Centrale (Neuropteroidea) - Boll. Soc. ent. it., Genova, 125 (3):

199-210.

LETARDI A., 1998 - Present knowledge of Italian Neuropterida: history, distribution

and current problems - Acta zool. fenn., Helsinki, 209: 145-149.

LETARDI A., 2005 - Insecta Neuroptera, Megaloptera e Raphidioptera. In: Ruffo S.,

Stoch F. (eds) 2005. Checklist e distribuzione della fauna italiana - Mem. Mus.

civ. St. nat. Verona, (2), Sez. Scienze della Vita, 16: 235-236.

LETARDI A. & PANTALEONI R. A., 1996 - I Neurotteroidei W-paleartici della col-

lezione del Museo di Zoologia dell’ Università di Roma (Neuropteroidea) -

Fragm. ent., Roma, 28 (2): 277-305.

MonseErRRAT V. J., 1980 - Contribuciòn al conocimiento de los Neur6pteros de Italia

(Neuroptera, Planipennia) - Neur. int., Nice, 1 (2): 48-64.

MonsERRAT V. J., 1991 - Nuevos datos sobre algunas especies del género Hemerobius

L., 1758 (Insecta, Neuroptera: Hemerobiidae) - Graellsia, Madrid, 47: 61-70.

MonseERRAT V. J., 1994 - Nuevos datos sobre los Coniopterigidos de las regiones

paleartica y afrotropical (Neuroptera: Coniopterygidae) - Graellsta, Madrid, 50:

109-127.

NavAs L., 1928 - Insetti europei del Museo Civico di Genova (Megalomus andreinti

sp. n.) - Boll. Soc. ent. it., Genova, 60: 75-83.

NavAs L., 1929 - Insetti Neurotteri ed affini di Oropa (Biella) - Boll. Soc. ent. it.,

Genova, 61: 44-47.

Navas L., 1933 - Insetti Neurotteri ed affini del Piemonte - Mem. Soc. ent. it.,

Genova, 12: 150-162.

PARENZAN P. & DE Marzo L., 1981 - Una nuova trappola luminosa per la cattura di

Lepidotteri ed altri insetti ad attivita notturna - Inform. giov. Ent. (suppl. Boll.

Soc. ent. it. ), Genova; 99% 5-11.

Pocci R., 1993 - Catalogo dei tipi di Neurotteroidei del Museo Civico di Storia

Naturale “G. Doria” di Genova (Insecta) - Annali Mus. civ. St. nat. G. Doria,

Genova, 89: 415-444.

548 D. BADANO

Princip! M. M., 1956 - Contributi allo studio dei “Neurotteri” italiani. XIII. Studio

morfologico, etologico e sistematico di un gruppo omogeneo di specie del gen.

Chrysopa Leach (C. flavifrons Brauer, C. prasina Burm. e C. clathrata Schn.) -

Boll. Ist. Ent. Univ. Bologna, 21: 319-410.

RIASSUNTO

Si riportano nuovi dati riguardanti la distribuzione in Liguria di Neuropterida

(Megaloptera, Raphidioptera, Neuroptera). Sono state individuate, soprattutto nel

settore occidentale della regione, 32 specie, di cui 11 non ancora note per l’area.

ABSTRACT

Contribution to the knowledge of Neuroptera of Liguria (Italy).

The Author reports new data on the distribution in Liguria of Neuropterida

(Megaloptera, Raphidioptera, Neuroptera). The researches were carried out moreover

in the western part of the region: 32 species, 11 of which not previously found in

the area, are reported.

SER)

RES LIGUSTICAE CCLVI

PETER NICK PSOMADAKIS*, MASSIMILIANO BoTTARO**,

GIVLIANG, Depra***, FuLvwio GARIBALDE**?,

STEFANO GIUSTINO***** & MARINO VACCHI**

NOTES ON THE REGALECUS GLESNE OCCURRING IN

THE GULF OF GENOVA AND IN LIGURO-PROVENCAL

WATERS (NW MEDITERRANEAN)

(PISCES, LAMPRIDIFORMES, REGALECIDAE)

INTRODUCTION

The king of herrings or oarfish Regalecus glesne Ascanius, 1772

is an open water mesopelagic fish occurring in all oceans, except in

polar regions (OLNEY 1984). The species is characterised for having

an extremely slender, laterally compressed, and very elongated body

reaching lengths up to 17 m, and is considered the longest known

teleost (ROBINS et al. 1986).

The genus Regalecus Ascanius, 1772 is generally treated as

monotypic comprising a single biologically valid species R. glesne

(HEEMSTRA 1986; PALMER 1986; NELSON 1994). However, Japanese

and Mexican ichthyologists consistently report two other species of

Regalecus occurring in the Pacific Ocean: R. russelit (Cuvier, 1816)

and R. kinoi Castro-Aguirre et al., 1991 (see Fuji 1984; CHAVEZ et

al. 1985; CasTRO-AGUIRRE et al. 1991; RAMIREZ-MURILLO & SCHMIT-

* TCRAM, Istituto Centrale per la Ricerca scientifica e tecnologica Applicata al

Mare, Via Casalotti 300, I-00166 Roma, Italia (peterpsomadakis@inwind.it)

** ICRAM c/o MNA, Università di Genova, Viale Benedetto XV 5, I-16132

Genova, Italia

*** Museo Civico di Storia Naturale “G. Doria”, Via Brigata Liguria 9, I-16121

Genova, Italia

**** Dip.Te.Ris., Università di Genova, Corso Europa 26, I-16132 Genova, Italia

***#** Via Posillipo 85, I-80123 Napoli, Italia

550 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, EF GARIBALDI, S. GIUSTINO & M. VACCHI

TER-SOTO 1996; BALART et al. 1999; SALAZAR-HERMOSO et al. 1999).

Apparently these forms are distinguished from R. glesne on the

basis of the following features: R. Rkinoi has a lesser number of rays

either in the occipital crest or in the proper dorsal fin; it also pos-

sesses longer and typically lanceolated pectoral fins, a convex frontal

margin of the head and a proportionally higher body. Whereas, A.

russelit is principally distinguished from its congenerics for having

a larger number of gill rakers in the first gill arch (Mori 1956).

With the exception of the northern Adriatic Sea, R. glesne has

been documented throughout the entire Mediterranean basin (FISCHER

et al. 1987) and, wherever it occurs, the species is considered rare

(TORTONESE 1970). However, in the Ligurian waters, the presence of

this open water species is not exceptional and different documented

instances of stranded or moribund specimens can be found in the

scientific literature or in local journalistic articles (VINCIGUERRA 1918).

In this paper, data on the occurrence of the species in the

Liguro-Provencal waters is overviewed taking into account histori-

cal and present literature as well as specimens preserved in syste-

matic collections. "The morphometric and meristic characters of a

fresh specimen found in Arenzano harbour are compared with seven

more or less complete individuals of R. glesne housed in the Museo

Civico di Storia Naturale “G. Doria” of Genova (MSNG) and with

one head stored in the “Dipartimento per lo Studio del Territorio e

delle sue Risorse” of the University of Genova (Dip.Te.Ris.). Some

information on the diet of Mediterranean oarfishes is also presented.

MATERIALS AND METHODS

Materials used in this study came from a wide variety of sources.

Important data was obtained from extensive literature search, inclu-

ding journalistic articles and systematic collections. Records of oar-

fishes occurring in the Ligurian Sea and in the bordering localities

of the French coast are summarized in Figure 1 and Table II.

Besides the fresh specimen collected at Arenzano (GE),

14.11.2003 (assigned to R. glesne and now in MSNG with catalogue

number 53664), we disposed of seven liquid-stored examples housed

in the Museum of Genova: MSNG 12306, Borghetto Santo Spi-

rito (SV), 24.11.1906; MSNG 12307, Noli (SV), 10.V.1903; MSNG

NOTES ON REGALECUS GLESNE 551

24 LIGURIAN

SEA

BEAULIEU

SAINT-JEAN

@).A FORMIGUE

Fig. 1 - Records of Regalecus glesne in the Liguro-Provencal waters according to

Table II.

34931, Spotorno (SV), 28.VII.1950; MSNG 35367, Genova Pra

(GE), 31.XII.1953; MSNG 54166, Santa Margherita Ligure (GE),

24.XI.1980; MSNG 54167, Sanremo (IM), 8.III.1992; MSNG

54168, Sanremo (IM), 18.IV.1993. The latter is represented only by

the head and the extreme portion of the caudal end. Another indi-

vidual, MSNG 37720, Genova Pegli (GE), 15.X.1959, was useless

because cut into small pieces. Also some of the above cited speci-

mens resulted problematic; in fact, due to extensive damage on the

ventral side, the anal openings of the MSNG 35367, MSNG 54166

and MSNG 54167 specimens were not visible and parameters such

as preanal and postanal lengths were not taken. Furthermore, during

morphometric analyses on the MSNG 12306 specimen, we discove-

red that a large piece at mid-body was missing and that the anterior

and posterior portions of the fish had been sewed back together,

therefore, total length of the latter could only be estimated. Accor-

ding to body proportions measurements obtained from other sam-

ples of Regalecus examined in the present study, the postanal length

of MSNG 12306 should have been 101.9 cm, which means that the

missing piece was at least 47.7 cm; the biometric proportions were

calculated in accordance to the following adjustment of total length:

TL = preanal length (original) + postanal length (estimated), that

is to say 53 + 101.9 = 154.9 cm (see Table I). We also examined

the head of a specimen from Sanremo (IM), 19.11.1998 at present

stored in the University of Genova (Dip.Te.Ris).

552 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, FE GARIBALDI, S. GIUSTINO & M. VACCHI

Tab. I — Absolute (cm) and proportional measurements expressed as percentage (%) of total length (TL)

and head length (HL) of the Arenzano specimen (MSNG 53664) of Regalecus glesne compared

with the ones of eight individuals of the species preserved in MSNG and Dip.Te.Ris.. Some

Noli Borghetto Santo Spirito

10.V.1903 24.11.1906

MSNG 12307 MSNG 12306

absolute CANE, absolute %TL %HL

1| total length (TL) 1864% (154.9) 1702%

2 | preanal length 39.8 33.7% 629% 53.0 34.2% 582%

3 | predorsal length 1.2 1.0% 19% 2.0 1.3% 22%

4 | postanal length 78.2 66.3% 1235% (101.9) 65.8% = 1120%

5 | prepelvic length 6.5 5.5% 103% IN 7.5% 127%

6 | head length (HL) 6.3 5.4% 9.1 AA

7| head height (through center of eye) 5.0 43% 80% 8.4 5.4% 92%

8 | body height (at posterior margin of opercle) 4.8 4.0% 75% 8.1 5.2% 89%

9 | body height (at anus) 3.2 2.7% 51% al 4.6% 78%

10 | preorbital length 15 12% 17, do 1.6% 27%

11| supraorbital length iO 1.0% 19% 2.4 1.5% 26%

12 | postorbital length 117.0 99.2% 1848% (150.9) 97.4% 1638%

13 | suborbital length DS: 1.8% 33% 4.0 2.6% 44%

14 | greatest body height 5.1 44% = 81% 7.6 4.9% 84%

15 | ocular diameter vertical 1.6 1.3% 24% 189 1.2% 21%

16 | ocular diameter horizontal 107 1.4% 26% 1.9 1.2% 21%

17 | interorbital space 1 1.0% 19% 1.4 0.9% 15%

18 | internarial space 0.4 0.3% 6% NM

19| preopercular max height (vertical) DRS, 1.9% 36% 3.2 2.1% 35%

20 | preopercular max length (diagonal) 4.2 3.9% 66% STE 3.4% 58%

21 | opercular max height Da5 22% 40% 4.1 2.6% 45%

22 | maxillary length 26 2.0% 37% 3.9 2.5% 43%

23| maxillary width 0.8 0.7% 13% 1.5 1.0% 16%

24 | pectoral fin lenght 2.9 1.9% 32%

25 | pectoral fin base lenght 0.8 0.7% 12% 1 0.8% 13%

26 | max dorsal fin height (excluded occipital crest) ow 2.7% 51% 27 2.4% 41%

27 | pelvic fin lenght (including distal lobes) SPA 270% DIE 749 53.0 34.2% 582%

332 SZ ok DIO

total dorsal rays up to anal opening

a | first gill arch lenght

b | longest gill filament 1.0 Ooo, 15.2%

c | shortest gill filament 0.3 0.2% 4.1%

d |! longest gillraker 0.6 0.6% 11.4%

e | gillrakers on the first arch 8+35=43 7+32=39

f | numberof rays in occipital crest 5+7=12 5+7=12

g | total dorsal rays (occipital crest included) 164

caudal fin rays

6-7

pectoral fin rays

teeth upper jaw

barel

teeth lower jaw 6-7 me

perceivable

forehead shape slightly concave concave

caudal end intact mutilated

yes

spiny nubbins on caudal tip

pseudobranch

small rakers between larger ones yes

NOTES ON REGALECUS GLESNE 553

meristic and morphological characters are also given. pTotal length of the Arenzano and San-

remo specimens measured in the laboratory. *Capture data. Estimated values are presented in

parentheses. NM: parameter not measurable.

Spotorno Genova Prà Santa Margherita Ligure

28.VII.1950 31.XII.1953 24.X1.1980

MSNG 34931 MSNG 35367 MSNG 54166

absolute oo he %HL absolute %TL absolute % TL %HL

180.4 1611% 258.0 2043% 86.5 2109.8% 1

64.5 35.8% 576% NM NM 2

1.8 1.0% 16% Son 1.2% 24% 0.9 1.0% 22.0% 3

IS 63.9% 1029% NM NM 4

{3.5 7.5% 121% 15.0 5.8% 119% 5 5.8% 122.0% 5

12 6.2% 12.6 4.9% 100% 4.1 4.7% 6

9.4 5.2% 83% 11.8 4.6% 93% 373 3.8% 80.5% 7

FI 6.1% 99% IV 4.8% 98% 335 4.0% 85.4% 8

9.5 5.3% 85% NM NM 9

342 1.8% 29% 4.3 1.7% 34.1% 0.8 0.9% 19.5% 10

343 1.8% 30% 4.0 1.5% 31% 1 1.2% 24.4% bi

Pea 97.1% 1563% 25185 OT 36 1992% 84.5 97.7% 2061.0% | 12

4.3 2.4% 38% 5.8 2.3% 46% tS 1.7% 36.6% 13

11.0 6.1% 98% oy! 4.9% 100% 3.4 3.9% 82.9% 14

Je) 1.2% 20% 2 1.1% 23% 1 1.2% 24.4% 15

DE 1.2% 20% 90 1.1% 23% 1 1.2% 24.4% 16

2.0 1.1% 18% 2:33 0.9% 18% 0.7 0.8% 17.1% k:7

0.7 0.4% 6% 1.0 0.4% 8% 0.4 0.5% 9.8% 18

4.2 2.4% 38% Dal 2.0% 40% 9 2.2% 46.3% £9

6.9 3.8% 61% Meo 3.1% 63% 2.8 3.2% 68.3% 20

4.0 DADA, 35% 6.3 2.4% 50% 1.8 2.1% 43.9% 21

4.2 2.3% 37% 5.0 1.9% 39% 1.8 2.1% 43.9% 22

1.9 1.0% 17% Bad 0.8% 17% 0.9 1.0% 22.0% 23

3.3 1.8% 29% 4.3 1.7% 34% 0.8 0.9% 19.5% 24

1.6 0.9% 14% 1.6 0.6% 13% 0.6 0.7% 14.6% 25

NM NM 256 3.0% 63.4% 26

NM NM NM 27

5.6 3.1% 50.0% 6.4 2.5% 50.7% a

1.9 1.1% 17.0% 2.0) 0.8% 15.8% b

1.0 0.6% 8.9% 0.5 0.2% 4.0% c

1.4 0.8% £25376 1.8 0.7% 14.3% d

8+33=41 7+36=43 e

NM NM f

260 366 g

106 NM h

0 0 i

14 11-12

0 9-10

0 5

strongly concave slightly concave concave

mutilated mutilated intact

NM NM

yes yes

554 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, F GARIBALDI, S. GIUSTINO & M. VACCHI

Tab. I (cont.)

Sanremo Sanremo

8.I1II.1992 18.1V.1993

MSNG 54167 MSNG 54168

absolute %TL %HL | absolute %TL %HL

1| total length (TL) 433* 2353.3%

2| preanal length NM

3 | predorsal length NM

4 | postanal length NM

5| prepelvic length NM

6 | head length (HL) 184 4.2% 100.0%

7| head height (through center of eye) 15.5 3.6% 84.2%

8 | body height (at posterior margin of opercle) 18 4.2% 97.8%

9 | body height (at anus) NM

10 | preorbital length 4 0.9% 21.7%

11 | supraorbital length 5 1.2% 27.2%

12 | postorbital length NM

13 | suborbital length 5,5 1.3% 29.9%

14 | greatest body height NM

15 | ocular diameter vertical 4.6 1.1% 25.0%

16 | ocular diameter horizontal 46 1.1% 25.0%

17 | interorbital space B07 0.9% 20.1%

18 | internarial space 1.5 0.3% 8.2%

19 | preopercular max height (vertical) 6.4 1.5% 34.8%

20 | preopercular max length (diagonal). 11:3 2.6% 61.4%

21 | opercular max height GAI 1.4% 33.2%

22 | maxillary length 64 1.4% 33.2%

23 | maxillary width Di 0.6% 14.1%

24 | pectoral fin lenght SH 1.3% 30.4%

25 | pectoral fin base lenght DT 0.6% 14.7%

26 | max dorsal fin height (excluded occipital crest) NM

27 | pelvic fin lenght (including distal lobes) NM

a | first gill arch lenght 8.9 2.62% 9.3 2.1% 50.5%

b | longest gill filament 2.8 0.82% DA 0.6% 14.7%

c | shortest gill filament 0.4 0.12% 0.3 0.1% 1.6%

d | longest gillraker 1.8 0.53% 1.9 0.4% 10.3%

e | gillrakers on the first arch 8+29=37 9+33=42

f | numberof rays in occipital crest NM

g | total dorsal rays (occipital crest included) NM

h | total dorsal rays up to anal opening NM

i caudal fin rays NM

J pectoral fin rays 13

teeth upper jaw 0

teeth lower jaw 0

forehead shape concave straight

caudal end intact NM

spiny nubbins on caudal tip NM

pseudobranch yes

small rakers between larger ones yes

NOTES ON REGALECUS GLESNE 555

Sanremo Arenzano

19.11.1998 14.11.2003

Dip: Te. Ris. MSNG 53664

SOA absolute TL

1271.60% 2363 %

LIB ZA 36.7% 867%

DOZZA 7 2.2% 52%

20159 63.3% 1496%

7.96% 101.23% 111%

6. 7.86% 100.00%

13 6.31% 80.25% 100%

Oo OND MN BU N

87.04% 16.5 5.2% 122%

19 6.0% 141%

1.99% 25.31% 5.5 1.7% 41%

27.16% 5 1.6% 37%

306.5 96.1% 2270%

32.10% di 1.3% 30%

20.5 6.4% 152%

1.89% 24.07% dh 1.3% 30%

1.94% 24.69% 30%

1.41% 17.90%

0.44% 5.56%

3.11% 39.51%

5.34% 67.90%

3.69% 46.91%

2.86% 36.42%

1.21% 15.43%

2.235% 28.40%

13.55%

Som Sha! On o Tops

strait

mutilated

yes

a 0 0 RB SH KO:

yes

556 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, F GARIBALDI, S. GIUSTINO & M. VACCHI

VAGLIO LANTIS IONI IAA

—— TY

RETE ae

Figure 2 — Morphometric characters used in this study shown on a generalized

Regalecus silhouette depicted with a “classical” occipital crest (rays

intentionally shortened).

a) frontal view

b) lateral view

c) head measurements in detail

NOTES ON REGALECUS GLESNE 557

Morphometric and meristic analyses were carried out using 43

parameters extrapolated from different works on regalecids available

in the scientific literature. Longitudinal measurements, e.g. distan-

ces from snout tip to various points along the body, were taken

straight-line (between vertical projections to the horizontal axis of

the body), made with a metal measuring tape and recorded to the

nearest millimetre. All measurements used in the present study are

shown in Figure 2 and Table I. Immediately after its finding, the

Arenzano specimen was taken to ICRAM “Centro Raccolta Cam-

pioni Mar Ligure”, divided in two halves and frozen at -20° C.

Subsequently, in the laboratory the specimen was dissected in order

to establish sex and to gather information on stomach contents and

then deposited in MSNG.

RESULTS: AND DISSESSION

The fresh specimen of R. glesne from Arenzano was 330 cm

total length (TL). When collected, the fish was still alive but showed

visible signs of injury, probably caused by the impact with a boat

propeller. Unfortunately, neither the elongated rays of the occipital

crest nor the peculiar oar-shaped pelvic fins were intact. The Aren-

zano regalecid is described as follows.

Body long, ribbon-like covered with tubercles, which became

more elevated along the abdomen especially behind the anus. Mouth

small, vertical and protractible. Lateral line starts from above the

eye, curves downwards and runs in a strait line along the lower

quarter of the body. Four longitudinal ridges from head to tail.

Greatest body depth in the middle of its length. Anus positioned in

the anterior third of the body. Caudal extremity mutilated, showing

a healed cicatrix. Gastric caecum extends well past the anus. Eyes

rounded and fairly big. Nostrils single, on each side of the snout

just behind maxillary symphysis. Pectoral fins lanceolated, inserted

horizontally. Teeth absent. Gill rakers long with minute spines pro-

jecting on the inner and outer margins. Scales absent. Anal and

caudal fins absent. Apparently eight rays on occipital crest (No.1:

very stout; No.2: very slender; No.3 to No.6: nearly as thick as the

first; No.7: slightly less stout than the former; No.8: slender like

the second; No.9: not thicker than rays of proper dorsal fin); bases

of anterior 6 dorsal rays closely set. Fin rays simple and unseg-

558 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, EF GARIBALDI, S. GIUSTINO & M. VACCHI

mented. Coloration: body silvery-grey with irregular oblique dark

streaks and blotches especially in the first third of the body. Fore-

head, parts below edge of gill covers and sides of the mouth black.

Pupil dark and iris light. Fins crimson red. Macroscopic examina-

tion of the gonads, allowed us to identify the specimen as a male.

Stomach empty.

Measurements of total length obtained from the specimen in

fresh conditions resulted quite discordant with the ones taken after

a two year permanence in the freezing chamber of the laboratory

(see Table II). Although the Arenzano specimen did not present the

typical bluish streaks on head and body described in the literature

(HEEMSTRA 1986), an irregular blue indigo spot above the eye is vis-

ible in two individuals found in Sanremo (Figure 3).

Figure 3 — Two individuals (from Sanremo) with irregular blue indigo spot above

tiie ‘eve!

NOTES ON REGALECUS GLESNE 559

Furthermore, we examined the stomach contents of two speci-

mens (MSNG 54167 and Dip.Te.Ris. specimen); the stomach con-

tents were exclusively made by crustacean remains: more in details,

they consisted respectively of 43 heads and 7 individuals of the

Mediterranean krill, Meganyctiphanes norvegica (M. Sars, 1857),

in advanced digestive status; the identification was possible on the

basis of mandibles and other rests.

The king of herrings or oarfish is apparently very rare in the

Mediterranean (TORTONESE 1970). Its presence in this sea can be

traced back to the XVI century, when IMPERATO (1599) figured and

described a bizarre fish under the name of “spada marina” from the

Gulf of Napoli (TORTONESE 1970). Eggs and young individuals at

different stages of development have been collected in the Strait of

Messina (MAZZARELLI 1910; Sanzo 1925; SparTA 1927; BERDAR et

al. 1975; CAVALLARO et al. 1980) and Elba Island (Damiani 1913),

but adults of the species have been rarely studied. As the syn-

onymy clearly shows, various nominal species have been errone-

ously described during the XIX century by ichthyologists working

on Mediterranean specimens (see Risso 1826; CuvigeR & VALEN-

CIENNES 1835; GIGLIOLI 1880). The examples used by the above

mentioned authors were mainly obtained from Nice (France) (VIN-

CIGUERRA 1918). Also the Ligurian Sea is an extraordinary site for

oarfish findings, as proven by the following records: Noli (ARIOLA

1904); Savona (Mezzana 1909); Borghetto S. Spirito, Arenzano,

Albissola, Santa Margherita Ligure (VINCIGUERRA 1918); Spotorno

(GUIGLIA 1950). Furthermore, the following individuals, held in the

museum of Genova, are not mentioned in the literature: MSNG

35367-Genova Pra; MSNG 37720-Genova Pegli; MSNG 54166-S.

Margherita Ligure; MSNG 54167-Sanremo; MSNG 54168-San-

remo as well as two other specimens captured respectively in 1997

and 1998 off Sanremo”(see Table II).

The biometric and meristic analyses conducted on the Aren-

zano specimen and on the examples of R. glesne held at MSNG

and Dip.Te.Ris., confirmed the high individual variability in body

proportions, already described by other authors (PARKER 1883;

GoobE & BEAN 1896; HuTron 1961). Proportional data in Table I

shows anomalies essentially concentrated in two individuals: in the

Arenzano specimen four measurements related to head length (HL)

were greater than values obtained from other specimens examined

560 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, F GARIBALDI, S. GIUSTINO & M. VACCHI

Table II — Records of Regalecus glesne from the Liguro-Provengal waters (in chron-

ological order). * Data not available or not furnished by the author/s.

+ Not measurable. In parentheses, size (TL) of fresh specimen. Dip.

Te.Ris.: Dipartimento per lo Studio del Territorio e delle sue Risorse,

Università di Genova; MHNN: Muséum d’Histoire Naturelle de Nice;

MNHN: Musée National d’Histoire Naturelle de Paris; MOM: Musée

Océanographique de Monaco; MZUF: Museo di Storia Naturale, Sezi-

one di Zoologia “La Specola”, Firenze; MSNG: Museo Civico di Storia

Naturale “G. Doria”, Genova.

Date of Size TL Locality of

Reference

occurrence | specimens (cm) occurrence

Risso (1826) Drawings of caudal and ventral

fins imaginary

Cuvier & MNHN A-7125, MNHN

Valenciennes (1835)

A-7126, MNHN A-7128

MNHN 000-6479

MZUF 886

MZUF 2762

MNHN 000-4466

Probably MHNN

MSNG 12307

(near Da vo,

Nice

(near the shore)

18.VIII.1877

19.1V.1891

1896

1897

10.V.1903

Giglioli (1880)

Damiani (1913)

Vayssiére (1917)

Ariola (1904)

Beaulieu Saint-Jean

Noli

Borghetto S. Spirito

(stranded)

MSNG 12306

24.11.1906 Vinciguerra (1918)

Arenzano :

TRANI t erved

24.1.1908 Cirsidéa) Vinciguerra (1918) Specimen not pres

17.VII.1909 288, 332 SO Mezzana (1909) Specimens lost

(near the shore) MOM 91 1908

Monaco ns Probably

10.III.1910 300 (captured in ERIC Vayssiére (1917) (cast specimen)

13.IX.1913 110 Fasdeloncello Vinciguerra (1918) | Specimen not preserved

(near the shore)

Albissola

Specimen not preserved

20.VI.1915 325 (speared in Vinciguerra (1918)

shallow water)

12-11-4917 236 S. Margherita Ligure | Vinciguerra (1918) Specimen lost

Monaco

(captured in the port)

MOM 91 2349

PREIS 130 or 140

MSNG 34931

MSNG 35367

MSNG 37720

28.VII.1950

J1 ees

158%. 1959

188 Guiglia (1950)

Spotorno

Genova Pra

Genova Pegli

S. Margherita Ligure

(trawl-net)

Sanremo

(trawl-net)

MSNG 54166

24.X1.1980

MSNG 54167

8.III.1992 340 (362)

MSNG 54168

MOM 91 3743

433 Sanremo

18.1V.1993

009.111.1994

La Formigue

Sanremo

12.1X.1997 (trawl-net, Garibaldi (in verbis)| | Specimen not preserved

depth -500 m)

19.11.1998 Rapreuy Head preserved (Dip.Te.Ris)

(surface)

Arenzano

MSNG 53664

14.11.2003 319 (330) Present paper

(near the shore)

NOTES ON REGALECUS GLESNE 561

(Figure 4), and in the Sanremo Dip.Te.Ris specimen, five measure-

ments related to total length (TL) resulted as well very high (Figure

5). Moreover dorsal fin ray counts in some specimens resulted also

quite discordant (see Table I). These anomalies could be linked to a

more or less degree of mutilation of the caudal end, which in regal-

ecid fishes seems to occur quite often (VINCIGUERRA 1918). Also the

poor condition of some examined individuals (including the Aren-

zano fish) may have contributed to cause these distortions. Further-

more, it must be said that TL measurement given for the Sanremo

Dip.Te.Ris specimen was taken on the fresh animal (Garibaldi in

verbis) while the biometric data here presented derives from meas-

urements obtained from its preserved head.

Particularly interesting is the presence of teeth in the upper and

lower jaws of four preserved specimens: in MSNG 12307 they are

bigger and equally distributed around the symphysis of the dentary

and premaxillary bones; in MSNG 35367 they are much smaller

and more abundant on the premaxillary rather than on the dentary,

while in MSNG 12306 and MSNG 54166 they are barely perceiv-

able and located only around the symphysis of the dentary. The

presence of teeth seems to be a variable character among oarfishes

(TRUNOV & KUKUEV 2005). It is quite interesting to note that teeth

have rarely been described for Indo-Pacific specimens of Regalecus,

however, due to their diminutive size, teeth can be easily overlooked

in these fishes.

Furthermore, some interesting observations regarding the mor-

phology of the occipital crest in the oarfish was revealed during this

study. As indicated by different authors, the number of elongated

rays on the forehead of oarfishes, is to be considered a taxonomic

character which distinguishes different species within the genus

Regalecus (see CHAVEZ et al. 1985; CasTRO-AGUIRRE et al. 1991;

RAMIREZ-MURILLO & SCHMITTER-SOTO 1996). However such infor-

mation is not always available due to the intrinsic fragility of these

structures which, in most studied specimens, appear seriously dam-

aged. In the Arenzano fish, the precise number of elongated rays

of the occipital crest can be only tentatively assessed, but valuable

observations regarding their reciprocal thickness were made. In fact,

the “classic” occipital crest morphology described in the literature

is formed by twelve rays (HULLEY & Rau 1969), of which the first

is very stout, the second, third, fourth and fifth are very slender

2 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, F GARIBALDI, S. GIUSTINO & M. VACCHI

Predorsal lenght

0,6

0,5 ®

_, 0,4

‘6

se 0,3

0,1

: : ni 7 a 1

o 5 10 45 Da

HL in cm

Greatest body height

I pee a A

3 A

7 J or a =]

0 5 10 15 20

HL in cm

Ocular diameter vertical

0,35 ]

0,3 4 À

= ; te ni: ar 5

= gal

~ 0,15 |

< 4

0,1 4

0,05 |

O UT T ic - 7

0 5 10 15 20

HL in cm

Preopercular max height (vertical)

0,6 5

4 e

0,5 4

J +

È 1 n n 3

210,84

SI di

oO) ]

SOI

0,1 4

0 4 ec. a, LL) =n T T T T T T 1 7 r DE =

5 10 15 20

HL in cm

Figure 4 — Linear regression

of four measurements related

to head length.

@ Noli (MSNG 12307);

M Spotorno (MSNG 34931);

A Genova Prà (MSNG 35367);

@ Arenzano (MSNG 53664);

A Borghetto Santo Spirito

(MSNG 12306);

+ Santa Margherita Ligure

(MSNG 54166);

X Sanremo (MSNG _ 54168);

NOTES ON REGALECUS GLESNE 563

Ocular diameter vertical

2,0%

1,8%

1,6%

1,4% CS

ilar Sr Riina asian FTA ANIA, sees

cee

cn i

RIINA INI,

As % of TL

1,0%

0,8% e ee e e e woke:

(©)

jo}

(©)

jo}

(©)

Ocular diameter horizontal

2,1%

1,9% 2

1,7%

1,5%

1,3% lai I ANNO II TA + e

As % of TL

0,9%

0,7%

0,5% Se wo_tmtmmwl

(0) 50 100 150 200 250 300 350 400 450 50

TL em

Interorbital space

1,5%

1,4% °

1,3%

1,2%

1,1% n

1 : 0% SIMILE @e tt asia

As % of TL

0,9% I E, ; ae

0,8% ni

0,7%

O) a

0 50 100 150 200 250 300 350 400 450 50

TL cem

Preopercular max length (diagonal) |

As % of TL

Figure 5 — Linear regression 2,1%

of five measurements related to he

e bia Lee eae

@ Noli (MSNG 12307); he TL em

M Spotorno (MSNG 34931);

A Genova Pra (MSNG 35367): Ro

@ Arenzano (MSNG 53664); 3,6% °

A, Borghetto Santo Spirito 3,1%

(M SNG 1230 6); 2.6% PR al i

EA Margherita Ligure ap ks ge TA iz. veto

(MSNG 54166): ge 5

X Sanremo (MSNG 54168); SRL RE I e Li

* Sanremo (MSNG 54167); 0 50 100 150 200 250 300 350 400 450 50

Opercular max height

As % of TL

564 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, E GARIBALDI, S. GIUSTINO & M. VACCHI

and the sixth, seventh, eighth and ninth are nearly as stout as the

first, the remaining ones diminish in strength and become uniform

in diameter with the proper dorsal fin rays. This kind of crest mor-

phology was more or less observed in all liquid-stored specimens

examined during the present study (see Figure 6). However, the

Regalecus collected in Arenzano presents a different morphology of

the occipital crest, which strongly differs from the classic descrip-

tions, for having only one slender ray in position No. 2 (see Figure

7). To the best of our knowledge, a similar type of occipital crest

morphology was described only once, for a large specimen stranded

ashore in New Zealand (von Haast 1877).

Our study also revealed some incorrect biometric data furnished

by VINCIGUERRA (1918) and re-proposed by GuiGcLia (1950) regard-

ing preanal and postanal lengths of two individuals of R. glesne from

Noli and Borghetto S. Spirito. For these two specimens, the preanal

length is indicated as higher in respect to the postanal length, which

must be considered erroneous since in this species the anus is posi-

tioned on the anterior third of the body (PALMER 1986).

The biometric analyses carried out on these two specimens of

R. glesne during the present study, confirms only in part the inac-

curacies given by the aforementioned authors. Infact, if for the Noli

specimen there is an obvious mistake in the localization of the anal

opening, for the Borghetto S. Spirito one, the measurements given

by VINCIGUERRA (1918) are identical to ours and the biometric dis-

proportions are due to the fact that the specimen is missing the

central portion of its body. It is possible, that the latter individ-

ual of Regalecus was taken to the museum incomplete by its finder

and that the museum staff, unaware of this, sewed the animal back

together. Since the latter was reassembled behind the anal opening,

we were able to estimate the amount of body portion missing (see

above, in Materials and Methods).

Comparatively to gill arch morphology, it is surprising that few

or even any authors have put emphasis on the presence of very small

additional rakers between each of the larger ones. ‘These structures

were observed on the first gill arch in all of our specimens, besides

MSNG 12307 and MSNG 54166 which appeared devoid of them,

even when investigated with a binocular microscope. Since the latter

individuals are juveniles, the absence of additional rakers could be

connected with a late development of these structures within the

NOTES ON REGALECUS GLESNE 565

Figure 6 — Detail of the occipital crest in MSNG 35367 (Genova Prà) and MSNG

54168 (Sanremo) specimens.

Figure 7 — Detail of the occipital crest in MSNG 53664 (Arenzano) specimen.

566 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, F GARIBALDI, S. GIUSTINO & M. VACCHI

life cycle of regalecid fishes. 'The above mentioned individuals were

among the only ones to show an intact caudal end with 4-5 minute

spiny nubbins on the upper edge (see Figure 8). These structures

may be rudiments deriving from caudal rays which are generally

present on juvenile individuals (HEEMSTRA 1986).

Some authors have discussed whether the oarfish presence in

the Mediterranean is due to the ingression of oceanic individuals or

whether this sea possesses a stable resident population of the spe-

cies (VINCIGUERRA 1918). This is difficult to say considering that

R. glesne is an elusive mesopelagic fish whose biology and ecol-

ogy are still poorly understood. However, the finding of eggs and

young individuals at different stages of development is indicative of

a self-maintaining Mediterranean population. In addition, the Ligu-

rian Sea offers the ideal trophic conditions for a filter-feeder, as

the oarfish appears to be (HUTTON 1961; HEEMSTRA 1986). Filter-

feeding among Mediterranean oarfishes is confirmed by the stomach

contents examined in the present study, as stressed also by other

authors in the past (Orsi RELINI & RELINI 1993).

Figure 8 — MSNG 12307 (Noli) specimen: caudal end with spiny nubbins visible

on upper edge.

NOTES ON REGALECUS GLESNE 567

The Liguro-Provengal waters present important upwelling phe-

nomena (ASTRALDI et al. 1995; MiLLoT 1999). These oceanographic

features allow the creation of a highly dynamic ecosystem, in which

large masses of euphausiids and other zooplanktonic organisms play

a fundamental role in the food web. The establishment of peculiar

trophic conditions all throughout the year could explain the more

frequent occurrence of open water zooplanktivorous species like R.

glesne and of many other top predators species, just like cetaceans,

tunas, swordfish and elasmobranchs (ORsI RELINI et al. 1995; AA. VV.

2007). The general topography of this Mediterranean sector, charac-

terized by a narrow continental shelf connected to bathyal grounds

by numerous steep canyons (CORRADI et al. 1987), may also contrib-

ute to the unusual abundance of oceanic oarfishes. It is interesting

to note that oarfish findings within the Ligurian Sea have occurred

mainly in winter and spring, when planktonic production in this

area is at highest levels (FABIANO 1982).

Concerning the taxonomy of the oarfish, it is still not certain if

there is one or more species within the genus Regalecus. Therefore,

it is very Important that any specimen captured or washed ashore

should be immediately reported to the nearest scientific institution

and studied by a specialist. It is also greatly desirable that a major

revision of the genus Regalecus should be undertaken as soon as pos-

sible.

ACKNOWLEDGEMENTS

The authors of this article are thankful to R. Santoro and F.

Delfino (Porto Arenzano Spa) for making this rare fish available for

study. A special thanks goes to M. Pastorino (PiEmme foto, Aren-

zano) who provided original photographic documentation.

Staff members of the libraries of the Stazione Zoologica “A.

Dohrn”, Napoli and of the Museo Civico di Storia Naturale “G.

Doria”, Genova are acknowledged for their precious help in refe-

rence search.

Many thanks also to S. Vanni (Museo di Storia Naturale,

Sezione di Zoologia “La Specola”, Firenze) and M. Bruni (Musée

Océanographique de Monaco) for providing information on speci-

mens held in their systematic collections. We are also grateful to P.

568 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, F GARIBALDI, S. GIUSTINO & M. VACCHI

Tomassetti (ICRAM) for his help with taxonomic identification of

euphausiid remains, to P. De Matteo (Makers Associati, Napoli) for

picture preparation, to S. Lomiri (ICRAM) for the French tran-

slation of the abstract, to M. B. Invernici (MSNG) for technical

support and to D. Basili (architect, Rome) for the preparation of

graphical illustration of Regalecus figures.

Finally this article is dedicated to the everlasting memory of

“my little darling” Giulia Psomadakis.

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York, 600 pp.

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In: Moser H.G., Richards W.J., Cohen D.M., Fahay M.P., Kendall Jr. A.W.

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570 PN. PSOMADAKIS, M. BOTTARO, G. DORIA, F GARIBALDI, S. GIUSTINO & M. VACCHI

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NOTES ON REGALECUS GLESNE 571

iS) eae

The description of a regalecid fish which came ashore in the harbour of

Arenzano (Gulf of Genova) on 14 February 2003 is given. The specimen, found

moribund on the sea surface, was gathered and stored for study. Morphometric

and meristic analyses were performed on the latter and compared with those of

eight liquid-stored individuals of Regalecus glesne preserved in the Museo Civico

di Storia Naturale “G. Doria” of Genova (MSNG) and in the Dipartimento per

lo Studio del Territorio e delle sue Risorse, Universita di Genova (Dip.Te.Ris.).

Exclusive observations on the structure of the occipital crest and other interesting

morphological features, among with some dietary data, are also presented. Although

the taxonomy of the genus Regalecus is at present in a somewhat confused state, the

Arenzano fish can be provisionally assigned to Regalecus glesne Ascanius, 1772.

RIASSUNTO

Note su Regalecus glesne presente nel Golfo di Genova e nelle acque Liguro-

Provenzali (Mediterraneo nord-occidentale) (Pisces, Lampridiformes, Regalecidae).

In questo lavoro viene documentato il ritrovamento di un Regalecus nel

porticciolo di Arenzano (Golfo di Genova). Sull’esemplare, raccolto moribondo in

acque superficiali il 14.II.2003, sono state eseguite dettagliate analisi morfometriche e

meristiche. I dati ottenuti sono stati confrontati con quelli ricavati da otto esemplari

di Regalecus glesne conservati in alcool e appartenenti alle collezioni del Museo Civico

di Storia Naturale “G. Doria” di Genova (MSNG) e del Dipartimento per lo Studio

del Territorio e delle sue Risorse, Università di Genova (Dip.Te.Ris.). Vengono

presentate interessanti osservazioni morfologiche, incluse alcune inedite relative alla

struttura della cresta occipitale e sulle abitudini alimentari. Nonostante il genere

Regalecus sia attualmente contraddistinto da incertezze tassonomiche, il regaleco di

Arenzano può essere provvisoriamente attribuito a Regalecus glesne Ascanius, 1772.

RESUME

Notes sar Regalecus glesne present dans le Golfe de Genova et dans les eaux

Liguro-Provengales (Méditerranée nord-occidental) (Pisces, Lampridiformes, Rega-

lecidae).

La description d’un poisson du genre Regalecus qui est entrée dans le petit port

d’Arenzano (Golfe de Genova) le 14 Février 2003 est donnée. L’exemplaire, trouvé

moribond sur la surface de la mer, a été recueilli et conservé dans le but de | étudier.

Cet individu a été analysé et ses caractéristiques morphométriques et méristiques ont

été comparées avec celles des huit individus de Regalecus glesne, conservés en alcool

dans le Museo Civico di Storia Naturale “G. Doria” de Genova (MSNG) et dans le

Dipartimento per lo Studio del Territorio e delle sue Risorse, Università di Genova

(Dip.Te.Ris.). Le travail présent des observations morphologiques, compris des

intéressantes remarques sùr la créte occipital et sùr le régime alimentaire. Bien que

la taxonomie du genre Regalecus est a ce jour en quelque sorte confuse, le poisson de

Arenzano peut provisoirement étre attribué à Regalecus glesne Ascanius, 1772.

573

PIERFRANCO CAVAZZUTI*

NUOVI CARABUS L. E CYCHRUS F. DI CINA ED IRAN.

DESCRIZIONI E NOTE SISTEMATICHE.

(COLEOPTERA, CARABIDAE)

INTRODUZIONE

La fauna carabidologica della Cina è ancora lontana dall’es-

sere conosciuta in modo soddisfacente; lo testimonia la continua

produzione di pubblicazioni che riportano le descrizioni di nuove

specie e sottospecie provenienti da quel paese, la cui scoperta è oggi

resa possibile dall’apertura di nuove strade che rendono accessi-

bili alla ricerca località che fino a ieri erano ancora irraggiungibili.

In questo articolo vengono descritti alcuni nuovi taxa di Carabus

e Cychrus, provenienti, nella maggior parte, da zone già conosciute

e più volte indagate in passato, oppure, come nel caso di Carabus

(Cupreocarabus) lixianensis pseudomorettoi, di un taxon già raccolto

da alcuni anni e a lungo confuso con C. morettot Deuve, 1997, dal

quale è difficilmente separabile senza il ricorso all’esame degli organi

genitali maschili.

Inoltre lo studio di un importante lotto di Lamprostus, raccol-

ti in Iran da Jindrich Prochazka, ha consentito di individuare una

specie inedita proveniente dall’Iran nord-occidentale, una regione

ancora oggi quasi del tutto sconosciuta sotto l’aspetto naturalistico.

DESCRIZIONE DEI NUOVI TAXA

Carabus (Apotomopterus) mecynodes mitis n. ssp. (fig. 1)

Locus typicus: Cina occidentale. Sichuan meridionale,

Mianning Xian, Passo a Sud-Ovest di Mianning, 2650 m.

* Via Chiesa 1, 12030 Pagno (CN) Italy. E-mail: pi.cavazzuti@libero.it

574 P. CAVAZZUTI

Serie bi pica: Holoiypus <,- locus: ‘typicus, in’ coll

dell’autore. Paratypi: 49 3d, 32 99, ibidem, in coll. dell’autore, del

MSNG, e in numerose altre collezioni.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 27-32 mm. Larghezza massima delle elitre 8,4-10,8 mm.

Colorazione bicolore non molto vivace: capo e pronoto sono lucidi,

neri, con deboli sfumature blu o verdi sulle aree laterali del pronoto;

le elitre hanno un aspetto sericeo, variabili fra il rosso e il bruno

rameico più o meno scuro, talora con qualche riflesso verde nella

zona basale; margini concolori, o al più, con lievi riflessi verdi nella

zona omerale. Solo due esemplari hanno le elitre completamente

verdi. Arti e parti inferiori nere. Forma piuttosto snella.

Il capo è normale, la fronte non saliente e molto liscia, con

alcune rughe sulla sommità e sul collo. I solchi frontali sono molto

brevi e limitati alla parte anteriore e al clipeo. Le mandibole sono

robuste ma brevi. Le antenne, piuttosto sviluppate, superano la metà

delle elitre nel 6 e la raggiungono nella &.

Il pronoto è trasverso (rapp. Lp/Ip = 1,35), largo tanto ante-

riormente quanto posteriormente, con 1 lati molto arcuati e ristretti

verso la base, ma senza formare un’apprezzabile sinuosità. Gli angoli

posteriori sono ottusi, smussati e per nulla salienti. Il disco è molto

convesso: forma due lobi un poco rigonfi, incisi dalla linea mediana

e limitati poco prima della base da una depressione curvilinea. La

superficie è liscia sul disco, finemente rugosa nella parte posteriore,

sui margini e, talora, anche lungo la linea mediana.

Le elitre sono in ovale allungato, con gli omeri sfuggenti, e

mostrano una chiara sinuosità nella zona preapicale. Il disco è molto

convesso e la scultura, omodinama, è molto regolare, con solo gli

intervalli primari interrotti frequentemente da piccoli punti. Le strie

sono molto finemente punteggiate.

Edeago (figg. 12-13) caratteristico della specie, molto stretto alla

base e bruscamente flesso verso il basso, nella porzione distale. La

lama apicale è larga come nella ssp. liliana Cavazzuti, 1997 e obli-

quamente troncata. Su entrambi i lati della porzione mediana sono

presenti delle profonde rughe laterali oblique. In visione dorsale

l’apice forma un triangolo quasi retto, smussato e rivolto a destra.

Derivatio nominis. Il nome latino della nuova sot-

tospecie: muitis, dal significato di mite, mansueto, è stato suggerito

NUOVI CARABUS E CYCHRUS 575

dal suo aspetto un poco dimesso, obiettivamente meno appariscente

rispetto ai taxa conspecifici noti.

Osservazioni e note. Carabus(Apotomopterus) mecy-

nodes mitis è perfettamente caratterizzato dalle sue ridotte dimen-

sioni e dal cromatismo scuro, molto meno metallico e appariscente

delle altre sottospecie conosciute: liliana Cavazzuti, 1997, combustus

Cavazzuti & Ratti, 1999, pinchuanensis Imura, 1999 (= sericatus

Cavazzuti, 1999) ed helena Kleinfeld, 1999. La sottospecie maggior-

mente differenziata sotto ogni aspetto (cromatico, dimensionale ed

edeagico) resta comunque quella nominale (ssp. mecynodes Bates,

1890), conosciuta solo attraverso gli esemplari delle vecchie colle-

zioni, la quale, oltre ad essere molto variabile nella colorazione, ha

le elitre più dilatate posteriormente e l'estremità della lama apicale

dell’edeago ancora molto più larga.

Carabus (Apotomopterus) patroclus xinshibanus n.

ssp. (fig. 2)

Locus typicus: Cina occidentale. Sichuan meridionale,

Ganluo Xian (Xinshiba), Passo fra Ganluo e Shimian, 29°29°48° N,

10279: LI tee 2 An.

Serie. tipica: Holotypus é, locus typicus. Paratypi::4

22, ibidem, in coll. dell’autore.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 31-32 mm. Larghezza massima delle elitre 9,5-10 mm. Il

colore è uniformemente nero, opaco nella £, più lucido nel 6. La

forma è molto slanciata.

Di aspetto notevolmente più robusto nei confronti della sotto-

specie tipica, che dovrebbe corrispondere agli esemplari provenienti

dalle zone circostanti a Mianning, ha il capo più grande, ma la fronte

molto meno saliente e subpiana, con i solchi laterali debolissimi, e le

antenne non raggiungenti la metà delle elitre.

Il pronoto è decisamente più trasverso (rapp. Lp/lp = 1,26), con

i lati lungamente arcuati nel tratto anteriore e modestamente sinuati

poco prima della base. I margini sono sottilmente rilevati e recano

una setola basale e una mediana. Il disco è scarsamente convesso,

è lievemente depresso nel centro lungo la linea mediana e forma

576 P. CAVAZZUTI

una larga svasatura lungo i margini, nella quale vengono inglobate

anche le impressioni basali. Gli angoli posteriori non sono salienti

ma formano una lieve indentatura agli angoli. Tutta la superficie è

solcata da finissime e fitte rughe trasversali.

Le elitre sono molto allungate e ristrette alla base, hanno

gli omeri segnati ma sfuggenti, i lati poco dilatati e la maggiore

ampiezza situata oltre la metà. Il disco è molto convesso. La scul-

tura, eterodinama, è relativamente forte, avendo gli intervalli pri-

mari un poco più larghi e più rilevati degli altri e frequentemente e

profondamente interrotti in granuli o brevissimi segmenti. I secon-

dari e i terziari sono continui e uguali fra loro; le strie sono fine-

mente punteggiate. |

Zampe robuste e molto sviluppate.

Edeago (figg. 14-15) a lobo mediano poco curvato e con la lama

apicale alquanto allungata, pressoché diritta, non acuta all’estremita,

ma con i lati paralleli e la punta arrotondata. In visione dorsale la

lama apicale si presenta di forma triangolare, stretta e allungata.

Derivatio nominis. Dal toponimo della localita, pros-

sima al locus typicus, dal quale il nuovo taxon proviene.

Osservazioni e note. La nuova sottospecie di C.

patroclus Breuning, 1933 ha habitus molto allungato, che ricorda la

ssp. xiaoxianglingensis Cavazzuti, 1996, ma presenta elitre meno paral-

lele, piu dilatate verso il terzo distale e dotate di una scultura un poco

piu forte e aspra, tuttavia meno rilevata che nella ssp. catagraphus

Cavazzuti & Rapuzzi, 2005, infeudata molto più a ovest, presso Jiulong.

Tuttavia il carattere più peculiare e determinante per la separa-

zione del nuovo taxon è costituito dalla forma particolare dell’edeago,

che non si presenta arcuato e acuto come negli altri taxa conspeci-

fici, ma di forma più allungata, parallela e ricurva sul lato dorsale.

Carabus (Archaeocarabus) lacertosus n. sp. (fig. 3)

Diagnosi. Il pronoto insolitamente robusto e trasverso,

dotato di lobi posteriori larghi e salienti, rende immediatamente

riconoscibile la nuova specie.

Locus typicus: Cina occidentale. Sichuan meridionale,

Mianning Xian, Passo a Sud-Ovest di Mianning, 2650 m.

NUOVI CARABUS E CYCHRUS 577

3 4

Figg. 1-4 - Habitus degli Holotypi dei nuovi taxa:

1: Carabus ( Apotomopterus) mecynodes mitis n. ssp.; 2: C. (A.) patro-

clus xinshibanus n. ssp.; 3: C. (Archaeocarabus) lacertosus n. sp.; 4: C.

(Rhigocarabus) impavidus n. sp.

578 P. CAVAZZUTI

Holotypus @: locus typicus, in coll. dell’autore.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 25,7 mm. Larghezza massima delle elitre 8,9 mm. Colo-

razione perfettamente nera, relativamente lucida. Forma vigorosa.

Il capo è robusto, gli occhi molto grandi e sporgenti. La fronte,

molto rilevata e liscia, ha i solchi profondi e rugosi, con della pun-

teggiatura solo nella zona soprantennale. I margini laterali sono orlati

ma poco rilevati. Il clipeo è indistinto dalla fronte, di forma rettango-

lare e molto trasversa, profondamente solcato ai lati e dotato di due

setole. Il labbro è bilobato e abbastanza espanso lateralmente; sul

lato anteriore è lievemente incavato. Il mento ha gli epilobi subacuti,

il dente mediano molto breve e lungo meno della metà degli epilobi.

Il submento è bisetolato. Le mandibole sono robuste e arcuate: il

dente mediano del lato destro è più acuto del sinistro, i retinacoli

sono bicuspidati e circa dello stesso sviluppo, ma quello destro è

più stretto dell’altro. I palpi hanno l’estremità poco dilatata: 1 labiali

sono dicheti. Le antenne sono normali, superano la base del pronoto

con 4,5 articoli e oltrepassano di poco il primo terzo delle elitre. Lo

scapo e i primi tre articoli sono moderatamente clavati; il 4° è molto

più lungo del 2°, e, come il 3°, reca una corona di setole distali; dal

5° in poi sono tutti rivestiti di una fitta pubescenza fulva.

Il pronoto è molto grande e trasverso (rapp. Lp/lp = 1,46) e

largo quasi quanto le elitre. I lati sono arcuati nel tratto anteriore,

dove la maggiore ampiezza si trova a livello circa del terzo distale,

per poi restringersi verso la base dove formano una discreta sinuo-

sità. I lobi posteriori sono larghi e molto salienti, ma con l'estremità

perfettamente arcuata. Il lato anteriore forma una lunga arcuatura,

dall’orlo abbastanza ispessito, nella parte centrale; gli angoli ante-

riori sono chiaramente salienti. I margini sono orlati e molto rilevati,

particolarmente nel tratto posteriore dove formano una vasta doccia.

Il disco è molto convesso e finemente solcato dalla linea mediana.

Le impressioni basali sono molto grandi e profonde. La superficie è

dovunque rugosa e screpolata, ma sul disco tende a divenire quasi

liscia. È presente una sola fovea laterale mediana priva di setola.

Le elitre sono allungate, più allargate nel tratto distale e con

l'estremità un poco acuta e segnata da una lieve sinuosita latero-

distale. Gli omeri sono indicati ma perfettamente arcuati. I margini

sono sottilmente rilevati e formano una doccia abbastanza ampia. Il

disco è normalmente convesso, la scultura, subomodinama, è costi-

NUOVI CARABUS E CYCHRUS 579

tuita da intervalli primari variamente interrotti in granuli e segmenti

e da intermedi (secondari e terziari) ininterrotti e grandi quanto 1

primari. Le strie sono lisce. Mentre nell’area discale gli intervalli

intermedi sono ininterrotti ma appena segnati da una lievissima

crettatura trasversale, nelle aree laterali essi si riducono progressiva-

mente di grandezza, la spezzettatura diviene più netta e la scultura si

fa alquanto rasposa. Gli arti sono bene sviluppati e robusti.

D e'ritaluiieMicto m inis. L'aspetto tomer Mobusto della

nuova specie ha ispirato il nome che le è stato attribuito: lacertosus

= muscoloso, vigoroso.

Osservazioni e note. Nello stesso ambiente, a sud-

ovest di Mianning, C. lacertosus convive con C. (A..) dongchuanicus

mianshanicus Cavazzuti, 1996, specie dalla morfologia nettamente

diversa e dal pronoto alquanto più ridotto, meno trasverso, con lati

per nulla rilevati, appena ribordati, e con minuscoli lobi posteriori.

C. paris Breuning, 1932, che vive più a nord, nella valle di Bao-

xing, è pure caratterizzato da un pronoto molto piccolo e dotato

di angoli posteriori acuti. Nessuna relazione è pure ravvisabile nei

confronti di C. mianningensis Kleinfeld, 1999, il quale, oltre ad avere

dimensioni nettamente minori, ha una forma molto più raccolta ed il

pronoto che non ha 1 lati rilevati nella maniera sopra descritta.

Nel gruppo di C. nestor Breuning, 1934 sono presenti alcuni

taxa di forma molto più robusta e alquanto compatta, come pure nel

gruppo di C. morphocaraboides Deuve, 1989 e di bornianus Hauser,

1922, nei quali però, oltre alla forma molto compatta e robusta, si è

evoluto un cromatismo più acceso e vivace.

In conclusione ritengo che C. lacertosus possegga tutte le pre-

rogative di una buona specie autonoma, ma soltanto il ritrovamento

del maschio permetterà di far luce sulle sue reali affinità.

Carabus (Piocarabus) vladimirskyi tianzhuiensis n. nom.

pro Carabus (Piocarabus) vladimirskyi tianzhuicus Cavazzuti, 2006,

preoccupato da Carabus (Istocarabus) kiukiangensis tianzhuicus

Imura, 1996.

Come cortesemente mi ha tempestivamente comunicato il col-

lega Helmut Schutze di Gleichen (Germania), che qui ringrazio per

580 P. CAVAZZUTI

7 8

Figg. 5-8 - Habitus degli Holotypi dei nuovi taxa:

5: C. (Cupreocarabus) laevithorax arduus n. ssp.; 6: C. (C.) lixianensis

pseudomorettoi n. ssp.; 7: C. (Lamprostus) seroulikbin n. sp.; 8: C. (Cop-

tolabrus) ishizukar hubetanus n. ssp.

NUOVI CARABUS E CYCHRUS 581

la segnalazione, il nome tranzhuicus da me proposto nel 2006 per

una nuova sottospecie di Carabus ( Piocarabus) vladimirskyi Dejean,

1830 cade in omonimia con C. (Istocarabus) Riukiangensis tianzhui-

cus Imura, 1996 e deve quindi essere emendato.

Carabus (Rhigocarabus) impavidus n. sp. (fig. 4)

Diagnosi. Un Rhigocarabus di medie dimensioni, di aspetto

relativamente gracile, vicino nella forma a C. jintangicus Deuve,

2001.

L'*osciistiiito pile» Cina» Sichuan (occidentale Ganji;

Passo a Nord di Fubian, 4550 m, Buma Shan, praterie alpine,

31°33159N, 102°24679E.

Serie tipica: Holotypus dé, locus typicus; Paratypi: 2

3d, ibidem; 1 3,1 9, stessi dati, ma 4180 m.) (tutti in coll. dell’au-

tore).

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 17-19,5 mm. Larghezza massima delle elitre 6,8-7,2 mm.

Colore bruno rameico scuro, un solo paratipo è di colore rameico

più chiaro. Antenne e zampe nere, solo le tibie, le mandibole e 1

primi articoli delle antenne sono lievemente tinte di rosso. Forma

snella nei due sessi.

Il capo è normale, gli occhi sono piccoli ma molto sporgenti e

la fronte è poco rilevata e liscia. I solchi sono abbastanza profondi e

debolmente rugosi, con qualche sottile ruga presente pure sul collo; le

carene laterali sono spesse ma quasi per nulla orlate. Il clipeo è indi-

stinto e perfettamente saldato con la fronte. Il labbro è poco espanso

lateralmente e fortemente incavato sul lato anteriore. Il mento ha gli

epilobi acuti ed il dente mediano un poco più breve ma acutissimo;

il submento è dicheto. Le mandibole sono piccole e brevi, un poco

irregolarmente arcuate, ed hanno il dente mediano destro piccolo

ma ben distinto e i retinacoli bicuspidati; quello destro però è lungo

il doppio del sinistro e largo solo la metà. I palpi hanno l'ultimo arti-

colo pochissimo dilatato e uguale nei due sessi; il penultimo articolo

dei labiali reca due setole basali. Le antenne sono brevi e nodose,

superano nel d la linea basale del pronoto con 4,5 articoli, 4 nella

?. ma non raggiungono la metà delle elitre. Lo scapo è breve, tozzo

582 P. CAVAZZUTI

e un poco ricurvo, il 2° articolo è poco più breve del 4°, ma è più

clavato; il 3° e il 4° hanno una corona distale di setole, e dal 5° in poi

sono tutti uniformemente rivestiti di fitta pubescenza fulva. Gli arti-

coli dal 5° al 9° del 6 hanno un’incavatura glabra sul lato inferiore.

Il pronoto è poco trasverso e alquanto piccolo (rapp. Lp/Ip =

1,31); 1 lati sono arcuati nel tratto anteriore, dove oltre la metà è

situata la maggior ampiezza, e, verso la base, molto debolmente

sinuati e un poco ristretti. Gli angoli posteriori sono piccoli e den-

tiformi, salienti e smussati. I margini laterali sono sottilmente orlati

e rilevati; il lato anteriore, nel tratto mediano, ha una lieve orlatura.

Il disco è mediamente convesso, la linea mediana è profondamente

impressa e le fossette posteriori sono brevi ma profonde. La super-

ficie discale è liscia ma screpolata lungo la linea mediana e tende a

divenire granulosa e rugosa nelle aree laterali. Sono due le setole

di ciascun lato: una basale e una mediana. Il prosterno è liscio, le

propleure sono a lustro metallico liscio e bronzeo.

Le elitre formano un ovale molto allungato, con gli omeri sfug-

genti e pochissimo marcati. La maggior ampiezza è situata presso il

terzo distale. Il disco è molto convesso e la scultura, di tipo etero-

dinamo triploide, è costituita da intervalli primari in serie di gra-

nuli, profondamente interrotti da grosse fossette, che predominano

sul resto della scultura; questa è formata da sette serie di granuli

più o meno allineati e uguali fra loro. Gli sterniti non sono solcati.

Gli arti sono robusti e di normale sviluppo. Quattro protarso-

meri, nel maschio, sono dilatati e provvisti di fanere adesive.

Edeago (figg.16-17). Il lobo mediano è sottile nel tratto basale

e moderatamente ingrossato nella parte mediana, con la lama api-

cale allungata, digitiforme e lievemente ricurva verso l’alto in visione

laterale. In visione dorsale (fig. 16) l’apice mostra un distinto orien-

tamento a destra.

Derivatio nominis. L'ambiente dal quale proviene la

nuova specie, difficile ed in forte pendenza, gli ha valso il nome

specifico attribuito: 1mpavidus = senza paura.

Osservazioni e note. L'aspetto snello e la morfologia

generale fanno avvicinare C. impavidus a C. jintangicus (diffuso sul

massiccio dei Jia Jin Shan in ambienti analoghi), ma un esame accu-

rato dimostra che si tratta di due buone specie, nettamente separate.

In C. jintangicus la parte superiore del capo è molto e profon-

NUOVI CARABUS E CYCHRUS 583

Figg. 9-11 - Habitus degli Holotypi dei nuovi taxa:

9: Cychrus toledanoi n. sp.; 10: C. toledanoi maerkangensis n. ssp.; 11:

C. brugget lianghensis n. ssp.

584 P. CAVAZZUTI

damente rugosa ed il labbro è più dilatato lateralmente. Il pronoto,

che è moderatamente convesso e quasi liscio in ‘mpavidus, è caratte-

risticamente ammaccato, depresso e fortemente rugoso in jintangicus,

nel quale i margini del pronoto sono anche molto spessi e rilevati e

privi di setole basali. Le elitre nelle due specie sono sostanzialmente

simili, ma gli organi genitali maschili differiscono molto nella forma

della lama apicale.

Con C. xiei Deuve, 1992, diffuso nella zona di Barkam con le

ssp. barkamensis Deuve, 1994 e proximior Deuve, 1997, esistono

pure limitate affinità morfologiche, però in xzez il capo è più grande,

il pronoto più trasverso e meno ristretto posteriormente, 1 margini

molto più rilevati e le fossette basali molto più grandi; nelle elitre,

peraltro molto simili, gli intervalli quaternari sono ridotti a tracce

vestigiali. L’edeago è simile nella struttura del lobo mediano, ma la

lama apicale è più lunga e sottile. La sola femmina nota di C. impa-

vidus ha un aspetto slanciato, esattamente come i maschi della sua

specie, in contrasto con l’aspetto alquanto tarchiato delle femmine di

C. xie1, comprese le sue sottospecie.

Carabus (Cupreocarabus) laevithorax arduus n.ssp. (fig. 5)

Locus typicus: Cina, Sichuan occidentale, Xiaojin,

Passo a Nord di Fubian, 4550 m, Buma Shan, praterie alpine,

31°33159N, 102°24679E.

Serie tipica: Holotypus dg, locus typicus; Paratypi: 11

Sd, 10 P9, ibidem; 2 dd, 1 2, stessi dati eccetto 4150 m, 31°32017N,

102°26405E, conservati nella collezione del MSNG e dell’autore.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 17,8-22,4 mm. Larghezza massima delle elitre 6,7-8,2

mm. Colorazione di registro particolarmente scuro: la parte dorsale

è nera con riflessi azzurri o verdi in tutti gli esemplari della quota

più elevata, tranne uno, che è di colore bronzeo con riflesso verde,

mentre i tre esemplari provenienti dalla quota minore sono bron-

zati, e marginati di verde nella £; gli scapi e i primi quattro articoli

antennali, i femori e le articolazioni delle tibie sono di colore rosso

ocraceo. I tegumenti sono molto lucidi nel 6, un po’ meno brillanti

sulle elitre della 2. Forma elegante e slanciata nel d, più breve e

con le elitre dilatate nella @.

NUOVI CARABUS E CYCHRUS 585

Nei confronti di C. laevithorax sinuatior Deuve (2005) endemico

della valle orientale più prossima, la nuova sottospecie ha una forma

più tozza e breve nel d, e decisamente più allargata nella 9. Il capo

è proporzionalmente più macrocefalo, ha gli occhi più ridotti e meno

sporgenti, la fronte pressoché liscia e impuntata. Le antenne sono

più brevi raggiungendo appena il primo terzo delle elitre nel d, solo

il primo quarto nella £. Il pronoto è meno sinuato, ha margini più

spessi e gli angoli posteriori quasi inesistenti e arrotondati. Il disco

è più convesso e liscio. Sono presenti da 2 a 6 setole laterali distri-

buite nella metà distale, più 1 basale su ciascun lato.

Le elitre sono ovalari e decisamente meno allungate, possiedono

una scultura nettamente omodinama triploide, abbastanza forte, for-

mata da intervalli primari in granuli allungati e dominanti, interca-

lati da punti profondi e metallici; i secondari sono in cordolature

sottili e continue, grandi la metà dei primari; i terziari sono grandi

come i secondari ma meno rilevati, e sovente sono spezzettati in

granuli. Le strie sono punteggiate.

L’edeago (figg. 18-19) è poco differenziato da quello della forma

tipica, rispetto al quale ha l’apice appena un poco più troncato obli-

quamente.

Derivatio nominis. Arduus, l’aggettivo latino utiliz-

zato per nominare la nuova sottospecie, è stato utilizzato nella sua

accezione più naturale, cioè: difficile da raggiungere.

Os sett'va zioni e note Mg rearactensnche fenotipiche

della nuova sottospecie, come la colorazione nera lucente sfumata di

azzurro oO di verde, gli scapi e 1 femori rossicci, e l'elevato numero di

setole pronotali, la rendono immediatamente riconoscibile, non solo

dalle altre sottospecie di C. laevithorax Breuning, 1935, ma anche da

tutte le altre specie dell’intero sottogenere Cupreocarabus.

Carabus (Cupreocarabus) lixianensis pseudomorettoi

n. ssp. (fig. 6)

Locus typicus: Cina, Sichuan. occidentale, Xiaojin

Xian, circa 20 km N. di Lianghekou, 3500 m, foresta di conifere.

Serie tipica: Holotypus d, locus typicus, in coll. dell’au-

tore. Paratypi: 12 66, 8 29, ibidem, conservati nelle coll. Ghiretti

e dell’autore.

586 P. CAVAZZUTI

Figg. 12-19 - Lobo mediano dell’edeago degli holotypi dd in visione dorsale e laterale.

12-13: Carabus (Apotomopterus) mecynodes mitis n. ssp.; 14-15: C.

(A.) patroclus xinshibanus n. ssp.; 16-17: C. (Rhigocarabus) impavi-

dus n. sp.; 18-19: C. (Cupreocarabus) laevithorax arduus n. ssp.

NUOVI CARABUS E CYCHRUS 587

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 22-25,2 mm. Larghezza massima delle elitre 7,3-8,5 mm.

La colorazione è bronzea, discretamente metallica, con chiari riflessi

rameici sul capo e talora anche sul pronoto; tibie, palpi e antenne

sono rossicce.

La forma è prossima sia alla sottospecie tiponominale che, sep-

pure in misura molto minore, alla ssp. shuanqiaoensis Cavazzuti,

2003, ma da entrambe è distinta per i seguenti caratteri: mandibole

più sottili; fronte alquanto più fittamente rugosa e punteggiata; pro-

noto molto trasverso (rapp. Lp/lp = 1,40), più minutamente rugoso

e punteggiato, con 1 lobi posteriori più lunghi e arrotondati e dotato

di tre setole nella metà distale più una basale. Le elitre corrispon-

dono a quelle della ssp. lixzanensis Deuve, 1990, ma gli intervalli

primari sono maggiormente predominanti, i secondari continui e un

po’ meno rilevati, e 1 terziari in serie di granuli; in alcuni casi 1 gra-

nuli terziari sono ridotti e affiancati da serie disordinate di granuli

quaternari. Mentre nella forma tipica gli arti sono sovente neri o

nerastri, in pseudomorettoz le tibie sono rossicce.

Edeago (figg. 20-21) molto più vicino a quello della ssp. tipica

che a quello di shuanqiaoensis. La lama apicale è molto breve e per

nulla ricurva, in visione dorsale si presenta triangolare e brevissima,

con l'estremità che accenna appena ad una flessione a destra.

Derivatio ‘nominis: La réale*somislianza ‘con. C.

morettot Deuve, 1997, con il quale peraltro è sintopico ed era stato

confuso, ha ben meritato al nuovo taxon il nome attribuito.

Osservazioni e note. C. lixianensis pseudomorettot

è stato riscontrato sintopico, nella località tipica, con C. morettot

Deuve, 1997, dal quale è pressoché indistinguibile nella morfolo-

gia esterna, ma la diversità esistente nella struttura dei rispettivi

organi genitali maschili è talmente evidente che la separazione degli

individui di questo sesso non pone alcun problema. Ben altre pro-

blematiche s’incontrano nella separazione degli esemplari di sesso

femminile, la cui attribuzione all’una o all’altra specie, sulla scorta

dei soli caratteri esoscheletrici, è abbastanza aleatoria, com’é dimo-

strato dalla presenza, fra la serie tipica di morettoi, di esemplari

appartenenti certamente a pseudomorettot.

Più a nord, sui pascoli della zona assiale della catena dei Qion-

glai Shan, è infeudato C. brezinai Deuve, 1994, dall’aspetto molto

peculiare e assolutamente inconfondibile con lixianensis pseudomoret-

588 P. CAVAZZUTI

toi. A ovest s'incontra C. laevithorax arduus Cavazzuti, di cui si è

sopra discusso, e nella zona di Jinchuan Xian C. lixtanensis jinchu-

anicus Deuve, 2007 di recente descrizione, il quale però presenta un

edeago alquanto più lungo e affilato.

Fra le specie di Cupreocarabus presenti nella valle di Shuang-

qiao, localizzata poco più a est (laevithorax Breuning, 1935, miro-

slavi Deuve, 2000 e lixianensis shuangqiaoensis Cavazzuti), le prime

due appartengono ad altre linee filetiche, come viene suggerito dalle

diverse strutture degli organi copulatori, C. lixianensis shuangqiaoen-

sis è invece così particolare e differenziato nella morfologia edeagica,

dall’apice decisamente flesso a destra, unico nell’ambito specifico, da

mettere in dubbio la sua appartenenza a C. lixianensis, a favore della

sua elevazione a specie autonoma.

Carabus (Lamprostus) seroulikbin n. sp. (fig. 7)

Diagnosi. UnLamprostus nero, semilucido, piuttosto grande,

che ricorda indubbiamente sia C. guycolasianus Deuve, 2001 che sta-

vent Heinz, 2002, ma di forma notevolmente più allargata ed appiattita.

Locus typicus: Iranoccidentale. P. Azerbaigan-e Garbi,

4 km a est di Likbin, 1660-1700 m, 36°44 N, 45°28’ E.

Serie tipica: Holotypus d, locus typicus, 21-04/25-04-2002,

J. Prochazka legit, in coll. dell’autore; Paratypi: 15 dd, 12 9,

ibidem, nelle coll. Prochazka e dell’autore; 10 dd, 8 99, Iran occi-

dentale. P. Azerbaigan-e Garbi, 12 km a sud di Serou, 1950-2000

m, 37°38 N, 44° 43° E, 26-04/29-04-2002, J. Prochazka legit, nelle

coll. E. Hajdaj, J. Kalab, J. Prochazka e dell’autore.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 28-33 mm. Larghezza massima delle elitre 9,8-12 mm.

Colorazione perfettamente nera, semilucida, con qualche vaghissima

traccia di azzurro sui margini elitrali di alcuni esemplari. Forma

robusta e pochissimo convessa.

Il capo è di grandezza normale per il sottogenere, con gli occhi

grandi e mediamente sporgenti. Il vertice della fronte è rilevato e

liscio, 1 solchi sono poco profondi ma segnati e tendenti a congiun-

gersi nella parte posteriore, perdendosi però nell’area mediana, dove

peraltro sono presenti alcune rughe che si estendono anche al collo.

NUOVI CARABUS E CYCHRUS 589

Il clipeo è liscio, di forma chiaramente trapezoidale, e non presenta

tracce della sutura frontale. Sono presenti due setole all'estremità

anteriore dei solchi. Il labbro è piccolo, con lobi pochissimo espansi

lateralmente, e moderatamente incavato sul lato anteriore; dispone

di una setola su ciascun lobo e di una piccola frangia anteriore di

setole. Il mento ha gli epilobi eccentricamente acuti, con l’apice

situato sul margine interno, col quale forma un angolo quasi retto;

il dente mediano è molto largo e sviluppato, arrotondato, e lungo

il doppio degli epilobi. La superficie è solcata da numerose rughe

longitudinali. Il submento è privo di setole.Le mandibole sono brevi

e robuste, arcuate sul lato esterno e smussate all'estremità. Il dente

mediano destro è poco sviluppato, ma sempre più evidente del sini-

stro; i retinacoli sono larghi e bicuspidati, ma si presentano molto

consumati. I palpi sono brevi ed hanno gli articoli terminali poco

dilatati, larghi all'estremità il doppio che alla base; nei mascellari

il penultimo palpomero è sviluppato quanto l’ultimo; 1 labiali sono

dotati di 8-10 setole, distribuite sia singolarmente sia a ciuffi di 2

o 3. Le antenne sono molto brevi, non raggiungono il quarto delle

elitre nel 6 e superano la base del pronoto con appena tre articoli

nella 2, 3,5 nel gd; lo scapo è breve e tozzo e dotato di una setola; gli

articoli 2° e 4° sono debolmente clavati, il 2° è lungo circa quanto il

4°. Quest'ultimo è dotato di alcune setole disposte a corona, ma dal

5° in poi tutti gli articoli sono rivestiti di scarsa pubescenza fulva.

Il pronoto è molto trasverso (rapp. Lp/Ip = 1,396, 1,49 ?), ha

i lati arcuati, non sinuati e poco ristretti posteriormente. La base è

lievemente arcuata e continua con gli angoli posteriori, 1 quali sono

arrotondati e quasi per nulla salienti. I margini sono sottilmente

orlati e pochissimo rilevati. Il disco è molto convesso, perfettamente

liscio, con una fitta punteggiatura limitata solo alla parte basale, ma

a volte può presentarsi completamente screpolato e punteggiato nella

parte basale. La linea mediana è lieve, le impressioni deboli, punti-

formi e localizzate sugli angoli posteriori

Le elitre sono ovalari, allungate, ben allargate e arrotondate agli

omeri, ed hanno la maggiore ampiezza situata poco oltre la metà. I

margini sono sottili, rilevati, e formano una doccia abbastanza larga.

La superficie, da molto poco convessa, o addirittura subpiana sul

disco, diviene fortemente declive sui margini. La scultura è formata

da una minutissima punteggiatura, più o meno allineata in serie lon-

gitudinali parallele.

590 P. CAVAZZUTI

Le zampe sono robuste ma non molto sviluppate. Nei maschi

solo i primi tre protarsomeri sono dilatati e provvisti di spazzole di

fanere adesive.

L’edeago (figg. 22-23) presenta un lobo mediano robusto e poco

ricurvo. In visione laterale sono visibili la callosità del tratto basale,

la dilatazione del tratto mediano e la forma della lama apicale, che è

appiattita, larga e allungata, con l'estremità perfettamente arrotondata.

In visione dorsale si nota la particolare struttura, il risvolto predistale

della parte chitinizzata inferiore e la netta flessione verso destra.

Derivatio nominis. Il nome della nuova specie, su

suggerimento del suo fortunato scopritore, è formato dall’unione dei

toponimi delle due località dalle quali proviene la serie tipica: Serou

e Likbin.

Osservazioni e note. Come gia é. stato accennato

nella diagnosi, la nuova entità presenta alcune affinità con le specie

di Lamprostus iraniane di maggiori dimensioni: guycolastanus e sta-

VENI.

Ma in C. guycolasianus, compresa la ssp. colorescens Deuve, 2001,

in visione dorsale l’edeago si presenta diritto e non flesso a destra;

inoltre, sul piano morfologico, guycolastanus possiede impressioni

pronotali molto più vaste e profonde, e le elitre, molto più convesse,

hanno gli omeri completamente sfuggenti. C. staveni ha il capo più

tozzo e breve, il pronoto più squadrato, con gli angoli posteriori in

lobi arrotondati, ed il disco fortemente screpolato. Anche in C. sta-

veni le elitre sono più convesse e meno dilatate, gli omeri più sfug-

genti, e la loro maggior ampiezza è situata presso la metà.

C. calleyi Fischer, 1823, specie presente a breve distanza, poco

oltre il confine turco con la ssp. machardiellus Deuve, 1991, è un’en-

tità di forma molto diversa, che inoltre presenta misure di gran lunga

inferiori; ha il protorace molto largo e decisamente ristretto alla base

e le elitre nettamente più dilatate agli omeri. In C. calleyt machar-

diellus, inoltre, l’edeago è ben differenziato nella struttura.

Nei confronti con le specie presenti nell’area siriaco-giordana:

C. syrus Roeschke, 1898, C. hemprichi Dejean, 1826, ecc., pur in

qualche caso relativamente affini nella morfologia esoscheletrica, le

reali e fondamentali differenze presenti fra gli organi riproduttori

maschili, sono talmente evidenti che fanno escludere a priori ogni

possibilità di vicinanza specifica.

NUOVI CARABUS E CYCHRUS 591

Carabus (Coptolabrus)ishizukai hubeianus n. ssp. (fig. 8)

Locus typicus: Cina centrale. West Hubei, Muyuping

Sud, 1300 m, 31°45°N, 110°4’E.

Serie tipicac.Holetypusy ¢, lecus typicuss16-05/12-

06-2004, in coll. dell’autore. Paratypi: 1 d, 2 99, stessi dati; 1 &,

ibidem, eccetto: 20-06/12-07-2003; 3 dd, 2 22, ibidem, eccetto:

21-05706-06-20053 "3T°05°N, “110°3S°E: 7° G6,\2 99; West Hubei,

Guanmenshan, 1500 m, 16-05/14-06-2004, 31°45’N, 110°04’E; 4

Sd, 3 22, West Hubei, road Xingshan-Badong, 5 km N. Gaucho,

1500 m, 19-05/12-06-2004, 31°02’ N, 110°05’E. I paratypi sono con-

servati nelle collezioni Turna e dell’autore.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 35,3-37,6 mm. Larghezza massima delle elitre 12,2-14,4

mm. La colorazione è bruna nerastra sulle elitre, con una sottile

fascia marginale molto brillante, rameica tendente al dorato; capo e

pronoto di colore rameico abbastanza scuro, un poco più vivace sui

margini. Zampe, antenne e parti inferiori nere, ad eccezione delle

propleure ed epipleure, che sono rameiche. Forma relativamente

snella.

Il capo è praticamente identico a quello della forma tiponomi-

nale: soltanto il labbro è più profondamente incavato sul lato ante-

riore. Il pronoto (rapp. Lp/lp = 1,19) è arcuato nel tratto anteriore e

non forma quegli angoli mediani chiaramente evidenti che caratteriz-

zano la forma tipica. Gli angoli posteriori sono ottusi e poco diver-

genti. Il disco è pressoché liscio e privo della depressione trasversale

che caratterizza la sottospecie ish:zuka:r Deuve & Ohshima, 1990.

Le elitre sono ovalari molto convesse, con gli omeri quasi del

tutto sfuggenti e terminanti con due mucroni appena accennati. La

scultura è molto simile a quella della forma tipica ma meno confusa,

con gli intervalli primari e secondari in forma di goccia, ben isolati

e lucidi sul fondo lievemente bulinato e opaco. Gli intervalli terziari

sono presenti qua e la, in allineamenti di minutissimi granuli.

L’edeago (figg. 24-25) non presenta apprezzabili differenze

rispetto a quelli dei taxa conspecifici.

Derivatio nominis. La nuova sottospecie prende

il nome dalla provincia dell’Hubei, dov'è stata rinvenuta.

ishizukai proviene dal Chongqing occidentale (Wushan, Xinfeng),

592 P. CAVAZZUTI

sese pare CAS È 4) 2 9

ea Brits [eb aiae ane aa a

2233, CRI DEE,

gta,

VIRATA?

thE

ohio

Figg. 20-25 - Lobo mediano dell’edeago degli holotypi dd in visione dorsale e laterale.

20-21: C. (Cupreocarabus) lixianensis pseudomorettoi n. ssp.; 22-23:

C. (Lamprostus) seroulikbin n. sp.; 24-25: C. (Coptolabrus) tshizukat

hubeianus n. ssp.

NUOVI CARABUS E CYCHRUS 593

il nuovo taxon /lubetanus proviene dall’Hubei occidentale, presso

il massiccio del Dashennonjia. Sul piano morfologico le due sot-

tospecie si differenziano principalmente per la forma più slanciata

di huberanus, nella quale gli omeri sono più ristretti, per i mucroni

elitrali più ridotti, per il pronoto di forma più arcuata, quasi priva

delle angolosita mediane caratteristiche della ssp. zshizukaz, e per la

colorazione rameica dei margini elitrali, invece del verde brillante

della forma tiponominale.

Cychrus toledanoi n. sp. (fig. 9)

Diagnosi. Un Cychrus di ridotte dimensioni, morfologica-

mente molto vicino a C. stoetzneri Roeschke, 1923, ma con il pro-

noto molto più largo e dotato di una sola lunga setola mediana.

Locus typicus: Cina, Sichuan occidentale, Xiaojin,

valle di Fubian, 3600 m, Buma Shan.

Serie tipica: Holotypus dg, locus typicus; Paratypus: 1

6, ibidem, conservati nella collezione dell’autore.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 14,5-15 mm. Larghezza massima delle elitre 5-5,2 mm.

Colore nero lucido, ad eccezione delle mandibole e delle parti boc-

cali, della base degli antennomeri 2°-3°-4°, e dell’estremità dei palpi,

che sono rossicce.

Il capo è normale, un po’ grande, con la fronte rilevata e quasi

liscia. Il solco curvilineo interoculare è completo, benché molto atte-

nuato nel centro. Gli occhi sono grandi ma poco sporgenti, le carene

laterali sono orlate e molto rilevate, il clipeo è liscio, il labbro pos-

siede due lobi molto allungati che raggiungono la base dei primi

denti. Gli epilobi del mento sono lunghi e arrotondati. Le mandi-

bole sono sottili e di normale sviluppo, dotate di due denti interni,

fra i quali quello basale possiede un dente supplementare rudimen-

tale. I palpi sono dilatati, all’estremità i labiali sono larghi quasi il

doppio dei mascellari; il penultimo articolo dei labiali reca 3 setole:

1 basale e 2 distali gemelle. Le antenne sono abbastanza sviluppate

(purtroppo sono incomplete in entrambi gli esemplari) e superano

la base del pronoto con 5 segmenti. Lo scapo è lungo e clavato con

una setola dorsale, il 2° articolo è molto più sviluppato del 4°, ma

594 P. CAVAZZUTI

solo quest’ultimo è provvisto di alcune setole a corona; dal 5° in poi

sono tutti rivestiti di breve e scarsa pubescenza.

Il pronoto è trasverso, ampiamente arrotondato, cordiforme e

fortemente ristretto alla base, dove gli angoli sono ottusi. Il lato

anteriore è arcuato, molto più esteso di quello basale, e nella parte

centrale ha un risvolto molto largo e spesso. I lati sono appena

rilevati e recano due lunghe setole sulla metà. Il disco è convesso,

molto meno declive nella parte anteriore che in quella posteriore,

dov'è pure presente una depressione trasversale confluente con la

linea mediana. Tutta la superficie è punteggiata.

Le elitre sono ovalari e abbastanza larghe, dilatate agli omeri

e tuttavia perfettamente arcuate, con l'estremità apicale appena un

po’ appuntita. Il disco è scarsamente convesso, ma esteriormente al

terzo intervallo primario diviene improvvisamente subverticale. La

scultura è eterodinama, formata da intervalli primari in serie di gra-

nuli o segmenti, relativamente grandi e rilevati; i secondari sono in

granuli grandi quanto i primari, 0 poco meno, mentre il fondo eli-

trale è completamente ricoperto da una granulosità irregolare e scar-

samente allineata, che nel declivio anteriore viene sostituita da una

forte punteggiatura. Propleure, prosterno e mesosterno sono punteg-

giati; nelle epipleure la punteggiatura è maggiormente marcata. Gli

sterniti sono lisci.

Gli arti sono di normale sviluppo. Solo i primi due protarsomeri

sono moderatamente dilatati e provvisti di fanere adesive sul lato

inferiore.

Edeago (figg. 26-27) molto arcuato, breve, piuttosto robusto e

affilato all'estremità. In visione dorsale l’apice appare acutissimo e

dotato di margini molto sottili.

Derivatio nominis. La nuova interessante specie

è cordialmente dedicata all'amico Luca ‘Toledano, appassionato e

valente specialista del difficile gruppo dei Carabidi Bembidiini.

Osservazioni e note. Cychrus toledanoi n. sp. è

vicino al gruppo brugget/stoetzner1, con i quali condivide l'ampiezza

del pronoto e la scultura elitrale, ma C. brugge: ha il pronoto mag-

giormente allungato, più convesso e chiaramente sinuato, le elitre

sono ugualmente appiattite ma descrivono un ovale più regolare ed

hanno la scultura decisamente meno forte e priva di intervalli secon-

dari. Cychrus stoetzneri Roeschke, 1923 è di forma meno robusta,

NUOVI CARABUS E CYCHRUS 595

ma le elitre hanno una scultura sostanzialmente analoga; tuttavia sul

pronoto, meno convesso e fortemente punteggiato, sono presenti un

elevato numero di setole marginali. In C. stoetzneri, inoltre, l’edeago

è molto più sottile in visione dorsale, ma l'estremità è più larga e

arrotondata. |

Con C. thibetanus Fairmaire, 1893, che possiede un pronoto

abbastanza simile e dotato di una sola setola latero-mediana, C. tole-

danoi è distinguibile per il pronoto ancora più ampio, fortemente

ristretto alla base, ma quasi per nulla sinuato, e per le elitre squa-

drate e molto allargate agli omeri, completamente diverse dalla forma

ovalare e con gli omeri sfuggenti, tipica delle elitre di C. thibetanus.

Cychrus toledanoi maerkangensis n. ssp. (fig. 10)

Locus typicus: Cina, Sichuan occidentale, Monti a

N-Nordovest di Barkam, 4000-4600 m.

Serie tipica: Holotypus 6, locus typicus, foresta di

conifere; Paratypus: 1 9, ibidem, 4300-4600 m, praterie alpine.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 13,8-14 mm. Larghezza massima delle elitre 4,4-4,6 mm.

Colorazione nera, abbastanza lucida e appena bronzata.

Rispetto alla forma tiponominale le dimensioni sono lievemente

ridotte e l’aspetto un poco più gracile, il capo è più minuto, la fronte

meno pronunciata ed è appena più punteggiata. Anche le mandibole

sono più brevi.

Il pronoto, meno ampio e meno convesso, dispone, come in C.

toledanoi, di una sola lunga setola latero-mediana. Le elitre formano

un ovale più stretto e sono meno dilatate agli omeri. L’edeago (figg.

28-29) è pressoché indifferenziato.

Derivatio nominis. Il nome attribuito deriva dall’an-

tico nome tibetano di Barkam: Maerkang.

Osservazioni e note. Cychrus toledanoi maerkangen-

sis è distinto dalle varie sottospecie di C. stoetzneri Roeschke, 1923

che vivono nella regione, compresa la ssp. maoerganicus Deuve &

Tian, 2007, soprattutto per la forma del pronoto, più convessa e

più ristretta posteriormente, con gli angoli anteriori piccoli ma net-

tamente salienti, e per la presenza di una sola lunga setola latero-

596 P. CAVAZZUTI

mediana. In C. stoetzneri i margini del pronoto recano sempre un

numero di setole marginali molto più elevato, normalmente da 3 a

5, che di solito sono molto brevi.

Inoltre è anche distinto da C. inops Deuve, 1997 per il minore

allungamento del capo e per la forma del pronoto non ristretta nella

parte anteriore.

Cychrus bruggei lianghensis n. ssp. (fig. 11)

Locus typicus: Cina, Sichuan occidentale, Xiaojin

Xian, circa 20 km a Nord di Lianghe, foresta di conifere, 3500 m.

Serie tipica: Holotypus 6, locus typicus; Paratypi: 5

Sd, 4 99, ibidem, conservati nella collezione dell’autore.

Descrizione. Dimensioni: lunghezza totale, mandibole

comprese, 16,9-19 mm. Colorazione nera, poco lucida, solo le parti

boccali sono rossicce.

La forma è slanciata ma robusta. Il capo si presenta molto

stretto e allungato, con gli occhi lenticolari, pochissimo rilevati, la

fronte molto saliente e liscia, il solco curvilineo posteriore svanito

nella parte mediana e le mandibole sviluppatissime.

Il pronoto è allungato, poco trasverso, ristretto sia anteriormente

sia posteriormente, con 1 lati arcuati, orlati, ma pochissimo rilevati,

e dotati di 3-4 setole per lato, localizzate tutte nella metà distale. Il

disco è convesso e fittamente punteggiato.

Le elitre sono ovalari, con gli omeri sfuggenti nel 6, mentre

nella 2 sono talora molto marcati. Il disco è pochissimo convesso;

la scultura è costituita da un fondo grossolanamente granuloso, dal

quale emergono le serie dei granuli primari, normalmente poco rile-

vati, e, talvolta anche le tracce vestigiali dei secondari. I protarsi del

d non sono dilatati e sono sprovvisti di fanere adesive.

L’edeago (figg. 30-31) si presenta poco differenziato rispetto a

quello della forma tipica.

Derivatio nominis. Dal toponimo della località nei

dintorni della quale sono stati reperiti gli esemplari esaminati.

Osservazioni e note. Da C. bruggei barkamenstis

Deuve, 1991, diffuso sul versante settentrionale e sulla sommità del

Qionglai Shan (Monti a Sud Est di Barkam), la nuova ssp. lianghensts

NUOVI CARABUS E CYCHRUS 597

i &

GILL Ap,

SSN

oe”

2mm

Figg. 26-31 - Lobo mediano dell’edeago degli holotypi dd in visione dorsale e laterale.

26-27: Cychrus toledanoi n. sp.; 28-29: C. toledanot maerkangensis n.

ssp.; 30-31: C. bruggei lianghensis n. ssp.

598 P CAVAZZUTI

è separabile, a colpo d’occhio, per il pronoto più stretto, più convesso,

meno fortemente punteggiato e per la scultura meno forte delle elitre,

oltre, naturalmente, che per le dimensioni decisamente più cospicue.

Dalla ssp. microthorax Deuve, 1992, presente nella vicina valle

di Shuanggiao, è distinto pure per la forma molto più grande e

robusta, opaca e non lucidissima come in microthorax, per il pro-

noto meno convesso e per le elitre meno slanciate e dotate di una

scultura più profonda.

RINGRAZIAMENTI

Voglio cordialmente ringraziare gli amici e colleghi Jindrich

Prochazka (Hodonin, Repubblica Ceca), Jaroslav Turna (Kostelec

na Hané, Repubblica Ceca) e Daniele Ghiretti (Sermide, Italia), che

mi hanno concesso di studiare il materiale delle loro raccolte o pre-

sente nelle loro collezioni; Jaroslav Kalab (Jinacovice, Repubblica

Ceca), attento e acuto studioso del genere Carabus, che mi ha segna-

lato il problema di C. morettoi, e Thierry Deuve (Muséum National

d’Histoire Naturelle de Paris, Francia), per il prestito di esemplari

tipici, indispensabili per la determinazione di alcuni dei nuovi taxa

descritti. Infine desidero ringraziare Roberto Poggi (Museo Civico

di Storia Naturale “G. Doria” di Genova) per la benevola accetta-

zione di questo lavoro sugli “Annali”.

BIBLIOGRAFIA

BREUNING S. von, 1932 - Monographie der Gattung Carabus L. - Bestimmungs-

Tabellen der europàischen Coleopteren, Troppau, 104: 3-288.

BREUNING S. von, 1933 - Zehn neue asiatische Caraben - Entom. Nachr., Troppau,

Ver 20224)

BREUNING S. von, 1934 - Ueber Carabini - Folia zool. hydrob., Riga, 6: 29-40.

Cavazzuti P., 1996 - Quinto contributo alla conoscenza dei Carabus L. della Cina.

Su C. Apotomopterus del gruppo delavayi-patroclus, con descrizione di tre nuove

sottospecie. (Coleoptera, Carabidae) - Lambillionea, Tervuren, 96 (4): 589-596.

CavazzuT! P., 1997a - Nuovi Carabini della Cina II. Descrizione di nuovi Cychrus

Fabr. e Carabus L. del Sichuan meridionale (Coleoptera, Carabidae) - Nouv.

Rev. Ent. (N.S.), Paris, 14 (3): 217-227.

CAVAZZUTI P., 1997b - Descrizione di nuovi Cychrus F. e Carabus L. delle provin-

cie di Sichuan, Yunnan e Guizhou, Cina meridionale (Coleoptera, Carabidae)

- Coléoptères, Guyancourt, 3 (4): 47-76.

NUOVI CARABUS E CYCHRUS 599

CaAvAZzuTI P., 1999 - Sesto contributo alla conoscenza dei Carabus L. della Cina.

Descrizione di nuove specie e sottospecie dello Shaanxi, Sichuan e Yunnan

(Coleoptera, Carabidae) -Coléoptéres, Guyancourt, 5 (7): 115-137.

CAVAZZUTI P., 2003 - Sui Carabini della Cina (IV nota). Descrizione di tre nuove

sottospecie del genere Carabus delle provincie di Shaanxi, Sichuan e Tibet.

(Coleoptera, Carabidae) - Lambillionea, Tervuren, 104 (3): 413-417.

CAVAZZUTI P., 2006 - Nuovi Carabus delle regioni montagnose della Cina, province

di Gansu e Qinghai settentrionale (Coleoptera, Carabidae) - Annali Mus. civ.

St. nat.“G. Doria”, Genova, 98: 135-155.

CAVAZZuTI P. & Rapuzzi I., 1996 - Descrizione di dodici nuovi taxa del genere Cara-

bus L. originari della Cina e della Corea del Nord. Nota su C. (Pseudocoptola-

brus) taliensis firmatus Cav. (1997), con proposta della sua elevazione al rango

specifico. (Coleoptera, Carabidae) - Lambillionea, Tervuren, 105 (2): 203-220.

Cavazzuti P. & RATTI P., 1999 - Note sui Carabus della Cina. Nuove specie e sot-

tospecie di Apotomopterus Hope, Isiocarabus Reitter e Archaeocarabus Semenow

(Coleoptera, Carabidae) - Coléoptéres, Guyancourt, 5 (6): 93-114.

Deuve T., 1989 - Nouveaux Carabidae des collections de l’Institut Zoologique de

Academia Sinica de Pékin (Coleoptera) - Nouv. Rev. Ent., (N.S.), Paris, 6

(2): 159-171.

Deuve T., 1990 - Description d’un nouveau Carabus du Sichuan (Col. Carabidae) -

Bull. Soc. ent. France, Paris, 95 (5-6): 160.

Deuve T., 1991a - Nouveaux Cychrus et Carabus d’Asie - Bull. Soc. Sciences Nat,

Compiègne, 70: 17-21.

Deuve T., 1991b - Descriptions de deux Cychrini de la province chinoise du

Sichuan (Col., Carabidae) - Bull. Soc. Sciences Nat, Compiègne, 72: 19-20.

Deuve T., 1992 - Contribution a l’inventaire des Carabidae de Chine (Coleoptera)

(19° note).- Bull. Soc. ent. France, Paris, 96 (3) (1991): 223-242.

Deuve T., 1993 - Carabus Cychropsis et Cychrus nouveaux des provinces de la

Chine occidentale (Col., Carabidae) - L’Entomologiste, Paris, 48 (6) (1992):

317-322.

Deuve T., 1994 - Contribution a l’inventaire des genres Carabus et Cychrus en Chine

(Coleoptera, Carabidae) - Bull. Soc. Sciences Nat, Compiégne, 82: 23-34.

Deuve T., 1997 - Nouveaux Carabus et Cychrus du Sichuan et du Yunnan, Chine

(Coleoptera, Carabidae) - Coléoptéres, Guyancourt, 3 (1): 1-11.

Deuve T., 2001a - Note sur l’identité de Carabus (Lamprostus) cylindricior et

descriptions de nouveaux Carabus d’Iran (Coleoptera, Carabidae) - Coléoptéres,

Guyancourt, 7 (11): 147-163.

Deuve T., 2001b - Nouveaux Carabus L. et Cychrus F. et Cychropsis Boileau asiati-

ques et note sur les Megodontus Solier d’ Iran (Coleoptera, Carabidae) - Coléop-

téres, Guyancourt, 7 (7): 85-102.

Deuve T., 2005 - Trois nouveaux Carabus du Sichuan (Coleoptera, Carabidae) -

Coléoptéres, Guyancourt, 11 (1): 1-7.

Deuve T. & OusuHima S. I., 1990 - Un nouveau Coptolabrus de Chine (Coleoptera,

Carabidae) - Bull. Soc. Sciences Nat, Compiégne, 63: 8-9.

Deuve T. & Tian M.T., 2007 - Nouveaux Carabus L. et Cychrus F. du Qinghai

oriental et du Sichuan septentrional (Coleoptera, Carabidae) - Coléoptéres,

Guyancourt, 13 (2): 25-34.

600 P. CAVAZZUTI

FAIRMAIRE L., 1893 - [Communication a la Séance du 12 avril 1893] - Annales Soc.

ent. France, Paris, 62, Bull. ent.: clxxv-clxxvi.

HAuUSER G., 1922 - Beitrag zur Carabenfauna Ostasiens - Arch. Naturg., Berlin, Abt.

A, 88 (7): 103-109.

HEINZ W., 2002 - Beschreibung einer neuen Carabus (Lamprostus)-Art aus Persien

(Coleoptera, Carabidae) - Zool. Middle East, Heidelberg, 26: 151-156.

Imura Y., 1996 - Contribution to the knowledge of Carabid fauna (Coleoptera,

Carabidae) in Western Hubei and Southwestern Anhui China - Gekkan-Mushi,

Tokyo, 299: 10-14.

Imura Y., 1999 - Ten new taxa of the genera Carabus and Cychrus (Coleoptera,

Carabidae) from China - Fapan. Fourn. syst. Ent., Tokyo, 5 (1): 135-144.

KLEINFELD F., 2000 - Drei neue Carabus-Arten aus den Provinzen Sichuan und

Yunnan, China (Coleoptera: Carabidae: Carabini) - Ent. Zeitschr., Stuttgart,

110 (3): 66-70.

ROESCHKE H., 1898 - Carabologische Notizen I - Ent. Nachr., Putbus, 24:

121-126.

RIASSUNTO

Sono descritti e illustrati alcuni nuovi taxa dei generi Carabus L. e Cychrus

F. provenienti dalla Cina centro-occidentale, e dall’Iran nord-occidentale: Carabus

(Apotomopterus) mecynodes mitis n. ssp.; C. (A.) patroclus xinshibanus n. ssp.; C.

(Archaeocarabus) lacertosus n. sp.; C. (Rhigocarabus) impavidus n. sp.; C. (Cupre-

ocarabus) laevithorax arduus n. ssp.; C. (C.) lixtanensis pseudomorettoi n. ssp.; C.

(Lamprostus) seroulikbin n. sp.; C. (Coptolabrus) ishizukai huberanus n. ssp.; Cychrus

toledanoi n. sp.; C. toledanot maerRangensis n. ssp. e C. bruggei lianghensis n. ssp.

Viene infine proposto il nome nuovo: tianzhutensis per C. (Piocarabus) vladi-

mirskyi tianzhuicus Cavazzuti, 2006, preoccupato da C. (Istocarabus) kiukiangensis

tranzhuicus Imura, 1996.

RESUME

Nouveaux Carabus L. et Cychrus F. de Chine et Iran. Descriptions et notes

systématiques (Coleoptera, Carabidae).

Sont décrites et illustrée quelques nouveaux taxa des genres Carabus L. et

Cychrus F., originaires de la Chine centre-occidental, et du nord-ouest Iran: Cara-

bus ( Apotomopterus) mecynodes mitis n. ssp.; C. (A.) patroclus xinshibanus n. ssp.; C.

(Archaeocarabus) lacertosus n. sp.; C. (Rhigocarabus) impavidus n. sp.; C. (Cupre-

ocarabus) laevithorax arduus n. ssp.; C. (C.) lixtanensis pseudomorettoi n. ssp.; C.

(Lamprostus) seroulikbin n. sp.; C. (Coptolabrus) ishizukai hubetanus n. ssp.; Cychrus

toledanoi n. sp.; C. toledanot maerkangensis n. ssp. et C. bruggei lianghensis n. ssp.

Il est enfin proposé le nouveau nom: tianzhuiensis pour C. (Piocarabus) vla-

dimirskyi tianzhuicus Cavazzuti, 2006, préoccupé par C. (Istocarabus) kiukiangensis

tianzhuicus Imura, 1996.

NUOVI CARABUS E CYCHRUS 601

ABSTRACT

New Carabus L. and Cychrus F. from China and Iran. Description and syste-

matic notes (Coleoptera, Carabidae).

The Author describes and represents some news taxa of the genus Carabus L.

and Cychrus F. coming from middle-western of China, and from North-western

Iran: Carabus ( Apotomopterus) mecynodes mitis n. ssp.; C. (A.) patroclus xinshiba-

nus n. ssp.; C. (Archaeocarabus) lacertosus n. sp.; C. (Rhigocarabus) impavidus n.

sp.; C. (Cupreocarabus) laevithorax arduus n. ssp.; C. (C.) lixtanensis pseudomorettoi

n. ssp.; C. (Lamprostus) seroulikbin n. sp.; C. (Coptolabrus) ishizukai hubeianus n.

ssp.; Cychrus toledanoi n. sp.; C. toledanot maerkangensis n. ssp. and C. bruggei lian-

ghensis n. ssp.

The new name tianzhutensis is proposed for C. (Piocarabus) vladimirskyi tian-

zhuicus Cavazzuti, 2006, preoccupied by C. (Istocarabus) kiukiangensis tianzhuicus

Imura, 1996.

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eee eth Rorscuke Hu. 1898 - i " Carabologische Notizen | - «Ent. Nachr,,. |

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| RIASSUNTO |

‘Sono. itberimi € si Pliswerati alcuni nuovi taxa 'dei‘generì Car bui Lie chines i

HA avea dalla Cina centro-sccidentale, è daWlran nord-occidentale: Cai sid ar

DE, ( Apatomopterus }- metynodes mitis A. 88p-; GC. (A:) patroclus xinsliibanus n. sep; AS

È: trchaeocarabui} ‘Tacertosus Yi. #p.i Wi { Riigocarabus) - im pars ue n SD.) i f ae

Pirata ony are arden Ni sep C.C.) lixtanensis pseudomoretion n. ssp. ei ye

i va: ppi tk dal ssp E {Coptotabries ) ishizuhdi hubenanus n. =P a va ras

Blue pe howe a sp | maerkangensis th. sep, é dr ri; goa LANE ‘heseris: 11. sap.

etico Ri put ae ak oNieat jolie FRA i nome muovo: fianialusensis per L'. {Piorurabiz / < ladé-

tee ‘ mitskyi tianghuicus Cavazzuti, 2006, Lp pato da C. (fsiecarabus ). kinkiangengs

oe ee “anahaciens Inara, 1996.

AO aissumie | oe | |

i Bri | ce ui Li et È i F.:de Chine et reo. Bescriptions..et. notes x 1

nines (Coleoptera, | Carabidae). | ee oe i i sal Se

_ Sent dferites et iMastrée quelques “nave ut toxo, des genres Corabii Li et

Le originaires: de Ja Chine Getitre-ciccidiess tal, 6t du notd-ovest Fran: Cara: 4

i mecymodes vebtas ni sp €14.) j-perrorius s enna ti: ssp (Ca

combat. bacertosta ne wp; oC. sbigacarabn + diapacidus n AO Cp a

TIT ris de sep C/C) bian patent) n.569 CE

na i einen n sp C7 Coptolabries) ishizihai huberanus De s8p.; A ve hr 4

WOE si, aes € caledumoa maerkangentis Ti aap. et:€. brugge! lidaghensis b. sip. Ji

Bai sof a le nouveau nom: hanazhuiensis pour C. {Piocarabus) olan’

Crui, 2006, sid par Cc. (sicari hiukiangemars

603

Piero LEO*

OSSERVAZIONI SU DICHILLUS CORSICUS E

DESCRIZIONE.DI TRE. NUOVE SPECIE DEL

MEDITERRANEO OCCIDENTALE

(COLEOPTERA, TENEBRIONIDAE)

INTRODUZIONE

La sistematica e la nomenclatura delle specie mediterranee del

genere Dichillus Jacquelin du Val, 1861 risultano ancor oggi pro-

blematiche. L’ultimo lavoro che tratta contemporaneamente tutte le

specie del genere risale ormai a più di novanta anni fa (REITTER

1916) e purtroppo, data la scarsa utilità diagnostica di alcuni dei

caratteri utilizzati, unitamente alla notevole variabilità morfologica

intraspecifica di certe specie e ad alcuni problemi nomenclatoriali

mai definitivamente risolti, non consente una sicura identificazione

di parte dei taxa noti.

Dichillus corsicus (Solier, 1838) è stato oggetto negli ultimi anni

di alcuni lavori, le cui conclusioni risultano discordanti. ARDOIN

(1973), basandosi sull’esame di materiale tipico, ipotizza la sinoni-

mia di Dichillus pumilus (Solier, 1838) con D. corsicus; ritiene però

che gli esemplari di Sardegna da lui esaminati possano appartenere

ad una specie nuova che indica provvisoriamente come “Dichillus (s.

str.) corsicus (Solier) subsp. pumilus (Solier)”. GARDINI (1976) osserva

giustamente che “le specie di Dichillus del gruppo corsicus-pumilus-

subcostatus sono particolarmente bisognose di revisione”; inoltre cita

D. pumilus, precisando che si tratta del taxon interpretato nel senso

di REITTER (1916), per alcune isole dell’ Arcipelago ‘Toscano, per la

Corsica e per la Sardegna, fornendone una breve descrizione e il

disegno dell’organo copulatore maschile. Quasi contemporaneamente,

* Via Tola 21, I-09128 Cagliari, Italia. E-mail: piero.leo@tiscali.it

604 P. LEO

CANZONERI (1977) cita di Calabria alcuni esemplari come “Dichillus

sp.? prope corsicus Sol.”, a suo parere morfologicamente intermedi

tra due “forme” (“a” e “b”) presenti in Corsardinia, che descrive

brevemente attribuendo loro rispettivamente i nomi di “corsicus Sol.

(= pumilus Sol.)” e “pumilus di Baudi (nec Solier)”.

Successivamente questa interpretazione viene seguita da BoN-

NEAU (1988), GARDINI (1995) e SOLDATI & SOLDATI (1998). Questi

ultimi però, solo pochi anni più tardi (SOLDATI & SOLDATI 2002),

affermano che in Corsica e Sardegna è presente solo D. corsicus;

elencano i caratteri che avrebbero indotto gli autori precedenti

(BaupI 1874, Porta 1934, ARDOIN 1973, GARDINI 1976 e CANZo-

NERI 1977) a differenziare corsicus e pumilus (sensu Auctorum) (in

realtà aggregano, con una interpretazione personale e qualche errore

di traduzione, i diversi caratteri discriminanti indicati dai singoli

autori citati) e ne contestano la validità, sostenendo che “en réalité,

il existe deux formes extrémes, et toute une gamme d’intermédiai-

res. Cette variabilité existe entre les différentes populations mais

parfois aussi au sein méme d’une population”; infine, sull'esame del

materiale tipico di Solier, ripropongono la sinonimia tra corsicus e

pumilus.

Nel presente lavoro, sulla base di un cospicuo materiale (circa

5.300 exx., provenienti da oltre 180 località, in gran parte della Sar-

degna ma anche di altre regioni italiane e della Corsica) e sull'esame

di nuovi caratteri diagnostici e di altri già utilizzati da alcuni autori

precedenti, riesamino D. corsicus (Solier, 1838) e D. pumilus (sensu

ReITTER 1916, nec SoLIER 1838), descrivendo quest’ultimo come

nuova specie, essendo il nome pumzlus preoccupato dalla specie di

Solier. Con l’occasione descrivo altre due specie inedite recente-

mente scoperte, rispettivamente, in Sardegna e nelle Baleari.

Abbreviazioni. MSNG: Museo Civico di Storia Natu-

rale “G. Doria” di Genova; MSNM: Museo Civico di Storia Natu-

rale di Milano; MZUR: Museo di Zoologia dell’Università degli

Studi di Roma “La Sapienza”; CA: Coll. V. Aliquò, Palermo; CF:

Coll. L. Fancello, Cagliari; CFA: Coll. S. Fattorini, Roma; CG:

Coll. G. Gardini, Genova; CL: Coll. P. Leo, Cagliari; CLI: Coll.

A. Liberto, Roma; CM: Coll. C. Meloni, Cagliari; CMO: Coll. A.

Molinu, Sassari; CS: Coll. D. Sechi, Quartu Sant’Elena (Cagliari);

CSO: Coll. F. Soldati, Montpellier.

OSSERVAZIONI SU DICHILLUS CORSICUS 605

Dichillus (Dichillus) corsicus (Solier, 1838) (figg. 1, 3, 5, 7, 9)

Tagenia corsica Solier, 1838: 33, loc. typ.: Corsica

Dichillus (Dichillus) corsicus: REITTER 1916: 159; Porta 1934: 112

Dichillus corsicus: SOLDATI & SOLDATI 2002: 167 (pars)

= Tagenia pumila Solier, 1838: 33, loc. typ.: Sardegna

D.es.crizione,.Lunghezza..totale 2,/0=3,/8.mm.. Corpo

e antenne generalmente di colore bruno scuro, zampe rossicce; gli

esemplari più o meno immaturi possono essere uniformemente

bruno rossicci o anche bicolori: capo e pronoto bruno rossicci ed

elitre bruno scure. T'egumenti lucidi.

Capo un po’ più lungo che largo (rapporto lunghezza/larghezza:

1,1-1,2), con una fitta e robusta punteggiatura ben impressa sulla

Fig. 1 - Habitus del d di Dichillus corsicus (Corsica, Serra di Pigno) (foto A. Liberto).

606 P. LEO

fronte e sul vertice, più rada e più sottile sul clipeo. Guance poco

dilatate in avanti, tempie debolmente convergenti in addietro, bru-

scamente ristrette all’altezza del collo. Cresta sopraoculare debole.

Antenne con lunghe e fitte setole giallastre; 1° antennomero un

po’ più lungo che largo, 2°-10° trasversi, 11° subquadrato; 2° anten-

nomero più breve del 3°; 3° circa 1,2 volte più largo che lungo.

Pronoto piuttosto variabile nella forma, sempre più stretto del

capo, un po’ più lungo che largo (rapporto lunghezza/larghezza:

1,1-1,2), con la massima larghezza nella metà anteriore. Lati del

pronoto brevemente ristretti in avanti e più lungamente in addietro,

talvolta debolmente arrotondati, raramente sinuati avanti alla base.

Angoli anteriori ottusi, non sporgenti in avanti; angoli posteriori

debolmente ottusi, raramente quasi retti; ribordo laterale indistinto,

Fig. 2 - Habitus del d di Dichillus tyrrhenicus n. sp. (Sardegna, Marrubiu: paraty-

pus) (foto A. Liberto).

OSSERVAZIONI SU DICHILLUS CORSICUS 607

talvolta appena accennato nel quarto anteriore. Punteggiatura robu-

sta e abbastanza fitta, dello stesso diametro di quella della fronte.

Pleure protoraciche con punti fitti, simili per diametro a quelli del

pronoto; prosterno e metasterno con punteggiatura molto robusta e

leggermente diradata.

Elitre convesse, oblunghe, snelle, più di due volte più lunghe

che larghe (rapporto lunghezza/larghezza: 2,05-2,25), con la massima

larghezza poco dietro la metà, attenuate in punta all’apice. Scultura

elitrale molto variabile: ciascun’elitra porta otto serie longitudinali di

punti ben impressi, più o meno robusti, più o meno ravvicinati tra

loro; nella gran parte delle popolazioni gli intervalli terzo, quinto e

settimo sono più elevati dei rimanenti: il terzo e il quinto almeno nel

tratto basale, il settimo, per lo più careniforme, per gran parte della

sua lunghezza; in alcune popolazioni di Corsica e Sardegna tutti gli

intervalli sono quasi piani, o solo il settimo è appena più elevato

nella metà basale. Intervalli elitrali con una serie un po’ irregolare di

micropunti, visibili a forte ingrandimento. Orlo laterale delle elitre

ben rilevato, subcareniforme; epipleure ben delimitate dalla base

all’apice elitrale, prive di serie regolare di punti.

Sterniti addominali punteggiati; 1 punti sono più robusti sul

primo sternite visibile (con un diametro leggermente minore di

quelli del metasterno), più piccoli e fitti sul quarto e quinto. Nei dd

i primi due urosterniti sono leggermente spianati nella zona centrale.

Zampe (figg. 3, 5, 7) snelle; femori leggermente clavati e sinuosi,

tibie subrette e relativamente lunghe (il rapporto lunghezza/larghezza

è di 4,5 nelle tibie anteriori e mediane, di 5,5 nelle posteriori), tarsi

molto allungati, unghie lunghe e robuste. Nei dd tutte le tibie pre-

sentano, al lato interno, una serie di piccoli tubercoletti e un dentino

aguzzo apicale, quasi perpendicolare all’asse longitudinale delle tibie

stesse.

Organo copulatore maschile (fig. 9) relativamente slanciato, con

il tegmen lungo come la capsula paramerica; questa è 3,9 volte più

lunga che larga, con apice piuttosto attenuato all’apice.

Moa toe:robra laesnnehsmarmmaaaiat thos mershideicsmtianr Shi aa

zione. Dichillus corsicus è presente in Corsica, Sardegna e Italia

meridionale (Campania, Basilicata, Calabria) (cfr. fig. 11). Personal-

mente ho esaminato circa 3.200 esemplari provenienti dalle seguenti

località:

608 P. LEO

Corsica: Serra di Pigno, Bastia, Saint Florent, L’Ile-Rousse,

Ponte Leccia (Haute-Corse); Bocognano, Ajaccio, Col de Bavella,

‘Tassinca, Sartene-foce Ortolo (Corse-du-Sud).

Sardegna: Chiaramonti, Cargeghe, Alà dei Sardi, Ardara,

Alghero-Torre Porticciolo, Bultei, Padria, Semestene (prov. Sas-

sari); Onani, M. Tuttavista, Marina di Orosei, Dorgali, Badde Sali-

ghes, Sindia, Macomer, Silanus, Baunei, Urzulei, Supramonte di

Orgosolo, M. Novo San Giovanni, Arcu Correboi, Villanova Stri-

saili, Arzana, Sorgono, Desulo, Aritzo, Bruncu Spina, M. Spada,

Laconi, Nuragus, Isili, M. Tonneri, Gairo Taquisara, Sarule (prov.

Nuoro); Cuglieri, Arborea, M. Arci, Sini, Gonnostramatza, Masul-

las (prov. Oristano); Flumentorgiu, Giara di Gesturi, Tuili, Turri,

Ussaramanna, Siddi, Las Plassas, Sardara, Pabillonis, Guspini, M.

Arcuentu, Arbus-Montevecchio, Arbus-Costa Verde, M. Linas,

Suelli, Fluminimaggiore-Tempio di Antas, Buggerru, Iglesias, Gon-

nesa, Domusnovas-Sa Duchessa, Furtei, Senorbì, Serrenti, Pimen-

tel, Villagreca, Nuraminis, Serramanna, Soleminis, M. Sette Fratelli,

Decimomannu, Uta, M. Arcosu, Assemini, Elmas, Cagliari, Perda-

xius, Santadi, Domusdemaria, Isola S. Antioco (prov. Cagliari).

Campania: S. Maria di Castellabate, Laurino, Cilento-M. Sacro

(prov. Salerno).

Basilicata: Senise, M. Pollino (prov. Potenza).

Calabria: Orsomarso-fiume Argentino, Lungro, Bocchigliero-

Bosco Basilicò, S. Pietro in Guarano (prov. Cosenza); Cotronei

(prov. Crotone); Passo del Mercante, Passo di Pietraspada (prov.

Reggio Calabria).

Sono sicuramente da attribuirsi a D. corsicus le citazioni di

D. subcostatus (Solier, 1838) per la Calabria (REITTER 1916) e di

Dichillus sp. per la Campania (Ascea Marina, Scavi di Velia e Vallo

Lucano, Laurito: GARDINI 1976) e la Calabria (Montalto: CANZO-

NERI 1977); anche la segnalazione di D. minutus (Solier, 1838) per

Senise (Potenza) (LEO 1985) è in realtà da riferire a D. corsicus.

Restano dubbie le località citate da SoLDATI & SOLDATI (2002),

poiché appare evidente che i due autori francesi associano sotto D.

corsicus anche D. tyrrhenicus n. sp., descritto nel presente lavoro. E’

di dubbia interpretazione, ma probabilmente errata, la segnalazione

di D. corsicus per Abruzzo (ANDREETTI et al. 1997), mentre sono

certamente da riferire a D. tyrrhenicus n. sp. le citazioni (sempre

OSSERVAZIONI SU DICHILLUS CORSICUS 609

sotto D. corsicus) di Marcuzzi (1985) per l’Isola d’Elba e di Lo

Cascio (2001) per l’ Arcipelago Toscano (da notare però che la foto

riprodotta in quest'ultimo lavoro rappresenta realmente un esem-

plare di D. corsicus, evidentemente di altra provenienza).

Osservazioni. Come altre specie congeneri, D. corsi-

cus è morfologicamente piuttosto variabile, soprattutto per quanto

riguarda dimensioni e scultura elitrale. Anche se le singole popola-

zioni si presentano generalmente stabili morfologicamente, la varia-

bilità non sembra grandemente influenzata da fattori geografici: in

Sardegna e in Corsica gli esemplari di popolazioni limitrofe possono

presentare una facies più o meno differente, ma del tutto coinci-

dente con esemplari di provenienza geografica anche molto lontana.

Nell’Italia meridionale la variabilità sembra essere minore, tuttavia

la morfologia degli esemplari appartenenti a tali popolazioni non è

affatto dissimile da quella riscontrabile in numerose stazioni sardo-

corse. Nel complesso, ritengo senz’altro che la specie non possa

essere suddivisibile in razze geografiche.

Per quanto riguarda la sinonimia con Tagenia pumila Solier,

1838, pur non avendo potuto esaminare materiale tipico, ritengo che

non sussistano dubbi in proposito; questa sinonimia, già ipotizzata

da ARDOIN (1973) sull’esame dei tipi di Solier, è stata ufficializzata

da CANZONERI (1977) e ribadita nei lavori di BONNEAU (1988), GAR-

DINI (1995) e SoLDATI & SOLDATI (1998). Successivamente, SOL-

DATI & SOLDATI (2002), ristudiando il materiale tipico, confermano

ancora la sinonimia suddetta; va osservato però che i due autori non

considerano D. tyrrhenicus n. sp., descritto nel presente lavoro, un

taxona ben differenziato da D. corsicus; in ogni caso è sufficiente leg-

gere le descrizioni di SOLIER (1838: 33-34) per appurare che la sua

Tagenia pumila non può corrispondere a Dichillus tyrrhenicus n. sp.:

“Nigra vel rufo-obscura, angustior ovalis...” e ancora “Roussastre

ou obscure et presque noire, trés-étroite...” “Antennes...couvertes

de très petits poils d’un jaunàtre un peu doré qui les font paraître de

cette couleur”. Dall'esame comparativo delle descrizioni originali di

Tagenia corsica e T. pumila, considerando anche la variabilità morfo-

logica cui si è accennato, non emergono inoltre caratteri morfologici

tali da poter considerare distinti i due taxa.

Dichillus corsicus sembra essere affine soprattutto a D. subco-

status (Solier, 1838), della Penisola Iberica e del Marocco setten-

trionale; quest’ultimo si differenzia però abbastanza agevolmente da

610 P. LEO

D. corsicus per le dimensioni (minime, medie e massime) maggiori

(3-4,5 mm), per il pronoto più allungato (rapporto lunghezza/lar-

ghezza: 1,2-1,35) e con gli angoli anteriori leggermente protratti in

avanti, per il ribordo laterale dello stesso pronoto nettamente più

marcato e per il terzo antennomero non trasverso. Più numerose e

più nette appaiono le differenze con D. tyrrhenicus n. sp., per le

quali si rimanda alla discussione di quest’ultima specie.

Note ecologiche. Dichillus corsicus è presente in vari

ambienti, dal livello del mare a circa 1800 m di quota. Sembra pre-

ferire la macchia mediterranea degradata, 1 pascoli incolti e le fasce

di separazione tra i coltivi, mentre rifugge le zone fittamente fore-

state e gli ambienti costieri più xerici. Presente tutto l’anno, si rin-

viene più facilmente dall’autunno alla primavera, in genere sotto i

sassi o vagliando il terriccio alla base di vari arbusti, taora nei nidi

di formiche; nei terreni pianeggianti sottoposti a periodici allaga-

menti è talvolta possibile osservarne un gran numero di esemplari

rifugiati sotto corteccia di Eucalyptus.

Dichillus (Dichillus) tyrrhenicus n.sp. (figg. 2, 4,6,8,10)

Dichillus (Dichillus) pumilus: BauDI 1874: 297; REITTER 1916: 159; Porta 1934: 112;

GARDINI 1976 : 651; CANZONERI 1977: 261; BONNEAU 1988: 134 (nec Tagenia pumila

Solier, 1838)

Dichillus (Dichillus) corsicus ssp. pumilus: ARDOIN 1973: 258 (nec Tagenia pumila

Solier, 1838)

Dichillus corsicus: SOLDATI & SOLDATI 2002: 167 (pars)

Diagnosi. Un Dichillus s. str. di medie dimensioni, subo-

paco, con corpo, zampe e antenne di colore uniformemente bruno

rossiccio. Punteggiatura dorsale robusta. Pronoto all’incirca così

lungo che largo, non ribordato ai lati. Elitre con la massima lar-

ghezza dietro la metà, arrotondate all’apice, con otto strie di robusti

punti ben impressi, intervalli elitrali subpiani; ribordo elitrale indi-

stinto, epipleure delimitate da un intervallo marginale appena con-

vesso, non careniforme. Antenne robuste, con antennomeri 2°-10°

fortemente trasversi; setole delle antenne molto corte e rade, poco

visibili nei tre antennomeri basali. 'Tibie corte, nel maschio con

robusto dente apicale al lato interno; tarsi brevi, pretarsi piuttosto

tozzi. Organo copulatore maschile relativamente corto e robusto.

OSSERVAZIONI SU DICHILLUS CORSICUS 611

Figg. 3-8 - Profilo schematico delle zampe sinistre anteriori (3 e 4), mediane (5 e 6)

e posteriori (7 e 8) dei Sé di Dichillus corsicus (3, 5, 7) e D. tyrrhenicus

n. sp. (4, 6, 8). Scala 0,5 mm.

612 P. LEO

Serie tipica. Holotypus d: Sardegna (prov. Oristano),

Marrubiu, 21.III.2005, leg. P. Leo. Paratypi dd e 99 (stessa località

dell’holotypus): 25.X1.1979, leg. C. Meloni, 39 exx.; 21.III.2005,

legni, eo, 2660exx:!

Holotypus e 10 paratypi depositati presso 11 Museo Civico di

Storia Naturale “G. Doria” di Genova (MSNG); altri paratypi nelle

seguenti collezioni: MSNM, MZUR, CA, CF, CFA, CG, CL, CLI,

CM, CMO, CS, CSO.

Dreher: zione RBunghezza” totale’ 23703752 rim. YCorpo

subopaco, interamente bruno rossiccio, zampe e antenne concolori.

Capo subquadrato o appena più lungo che largo (rapporto lun-

ghezza/larghezza: 1,0-1,1), con una robusta e densa punteggiatura,

ben impressa sulla fronte e sul vertice, più rada e superficiale sul

clipeo. Guance piuttosto dilatate, tempie appena convergenti in

addietro, bruscamente ristrette all’altezza del collo. Cresta soprao-

culare debolmente ingrossata.

Antenne grosse con setole giallastre molto corte e rade, soprat-

tutto sul primi tre articoli che spesso appaiono quasi glabri; 1° anten-

nomero un po’ più lungo che largo, 2°-10° nettamente trasversi, 11°

subquadrato; 2° antennomero più breve del 3°; 3° 1,4-1,5 volte più

largo che lungo;

Pronoto più stretto del capo, appena più lungo che largo o

subquadrato (rapporto lunghezza/larghezza: 1,0-1,1), con la mas-

sima larghezza nella metà anteriore. Lati del pronoto arrotondati,

brevemente ristretti in avanti e più o meno lungamente in addietro,

talvolta leggermente sinuati avanti alla base; angoli anteriori ottusi,

non sporgenti in avanti; angoli posteriori un poco ottusi, a volte

quasi retti; ribordo laterale indistinto. Punteggiatura dorsale densa

e robusta, dello stesso diametro di quella della fronte. Pleure pro-

toraciche con punti fitti, simili per diametro a quelli del pronoto;

prosterno e metasterno con punteggiatura ancora più robusta, dira-

data.

Elitre ovali, poco convesse, spesso un po’ appiattite sul disco, rela-

tivamente poco allungate (rapporto lunghezza/larghezza: 1,85-2,10),

con la massima larghezza nettamente dietro la metà, piuttosto arro-

tondate all’apice. Scultura elitrale variabile: ciascun’elitra porta otto

serie longitudinali di punti ben impressi e robusti, più o meno rav-

vicinati tra loro; 1 punti delle strie, più robusti e marcati sulle strie

OSSERVAZIONI SU DICHILLUS CORSICUS 613

più interne e nella metà basale, tendono a ridursi come diametro e

profondità sulle strie esterne e nella zona apicale dell’elitra; intervalli

elitrali quasi piani, talvolta il terzo, quinto e settimo sono un po’

più convessi nella zona basale dell’elitra; intervalli elitrali con radi

micropunti, visibili solo a forte ingrandimento. L'orlo laterale delle

elitre è debolmente rilevato, non careniforme, e delimita appena le

epipleure; epipleure elitrali prive di serie regolare di punti.

Sterniti addominali punteggiati; 1 punti sono più robusti e radi

sul primo e secondo sternite visibile, più fini e densi sul terzo,

quarto e quinto. Nei dd i primi due urosterniti sono nettamente

spianati nella zona centrale.

Zampe (figg. 4, 6, 8) con femori debolmente clavati e sinuosi,

tibie subrette e molto tozze (il rapporto lunghezza/larghezza è di 3,6

nelle tibie anteriori e mediane, di 4,0 nelle posteriori), tarsi corti,

unghie piccole. Nei Sd tutte le tibie presentano, al lato interno, una

serie di piccoli tubercoletti e un dente apicale relativamente robusto

e un poco pendente distalmente.

Organo copulatore maschile (fig. 10) relativamente tozzo, con il

tegmen appena più corto della capsula paramerica; questa è 3,4 volte

più lunga che larga, con apice robusto.

Mita ae FU es #0 COSTO ped RT

zione. Dichillus tyrrhenicus n. sp. è diffuso in Sardegna (comprese

numerose isole circumsarde), in Corsica, in alcune isole dell’Arcipe-

lago Toscano e sul promontorio dell’ Argentario (Toscana) (cfr. fig.

12). Sono senz’altro da riferire alla nuova specie le segnalazioni di

ARDOIN (1973) per la Sardegna (Cussedu presso Tempio Pausania,

Abbasanta e Domusnovas, sub D. corsicus ssp. pumilus) e di GAR-

DINI (1976) per l’Arcipelago Toscano (Isola d’Elba, Giglio, Capraia

e Montecristo, sub D. pumilus); anche le citazioni di MARCUZZI

(1985) per l'Isola d’Elba e di Lo Cascio (2001) per l’ Arcipelago

Toscano (entrambe sub D. corsicus) vanno riferite a D. tyrrhenicus

n. Sp.

Personalmente, oltre alla serie tipica, ho potuto esaminare circa

1.800 esemplari, provenienti dalle seguenti località:

Corsica: Cervione, Vaccaja, Aleria (Haute-Corse); Porto-Vec-

chio, Tonnara Plage, Bonifacio (Corse-du-Sud).

Sardegna: Isola Razzoli, Isola Spargiotto, Isola delle Bisce, Isola

Soffi, Arzachena, Badesi, Isola Asinara, Stintino, Golfo Aranci,

614 P. LEO

Olbia, Chiaramonti, Osilo, Sassari, Cargeghe, Alà dei Sardi, Chili-

vani, Capo Caccia, Alghero, Torralba, Giave, Bonorva, Padria, Seme-

stene, Burgos (prov. Sassari); Loculi, Onanì, M. Tuttavista, Marina

di Orosei, Monte Albo, Lula, Campeda, Bosa, Magomadas, Sindia,

Dorgali, Oniferi, Silanus, Macomer, Birori, Dualchi, Borore, Orani,

Ottana, Mamoiada, Baunei, M. Novo San Giovanni, Arcu Corre-

boi, Villanova Strisaili, Arzana, Sorgono, Aritzo, Ortuabis, Laconi,

M. Tonneri, Seui, Esterzili, Isili, Orroli, Tertenia (prov. Nuoro);

Sedilo, Abbasanta, Paulilatino, S. Caterina di Pittinuri, Riola Sardo,

Isola Mal di Ventre, Cabras, Stagno di Mistras, Oristano, San Gio-

vanni di Sinis, Asuni, M. Arci, Arborea, Uras, Mogoro, San Nicolò

Arcidano (prov. Oristano); Flumentorgiu, Giara di Gesturi, Col-

linas, Sardara, Pabillonis, Guspini, Arbus, Arbus-Montevecchio,

Arbus-Costa Verde, Gonnosfanadiga, M. Linas, Villacidro, Flu-

minimaggiore, Buggerru, Nebida, Iglesias, Gonnesa, Domusnovas,

Senorbì, San Nicolò Gerrei, Quirra, Villasalto, Serrenti, Villagreca,

Villaputzu, Siliqua, Uta, Burcei, M. Sette Fratelli, Elmas, Stagno

Simbirizzi, S. Isidoro, Cagliari, Carbonia, Capoterra, Capitana, Cala

Regina, Santadi, Perdaxius, Villasimius, Isola Serpentara, Isola dei

Cavoli, Sarroch, Pula, Isola San Pietro, Isola S. Antioco, Isola Vacca,

S. Anna Arresi, Domusdemaria, Porto Teulada (prov. Cagliari).

Toscana: Isola Capraia, Isola d’Elba (prov. Livorno); Monte

Argentario (prov. Grosseto).

Oss'ervazioni > inote Comparative. Dichillus

tyrrhenicus n. sp. corrisponde sicuramente a D. pumilus sensu BAUDI

(1874) e REITTER (1916); quest’ultimo Autore indica, in contrappo-

sizione con D. corsicus, caratteri differenziali nella colorazione, nella

conformazione del margine elitrale e nella chetotassi delle antenne;

anche D. pumilus citato e brevemente descritto da GARDINI (1976)

(che specifica chiaramente “...l’entità che chiamo Dichillus pumilus

nel senso di Reitter...”) è da riferire senza dubbio alla nuova specie

qui descritta.

D. tyrrhenicus n. sp. non può invece essere identificato con la Tage-

nia pumila Soler, 1838 la cui descrizione originale è in contrasto con

la morfologia della nuova specie qui descritta: su quest’ultimo punto

e sulla sinonimia tra D. pumilus (Solier, 1838) e D. corsicus (Solier,

1838) si vedano le osservazioni a proposito di quest’ultima specie.

Nonostante la confusione cui ha dato luogo in passato, D. tyrrhe-

OSSERVAZIONI SU DICHILLUS CORSICUS 615

nicus n. sp. appare ben differenziato dalle congeneri; l’unica specie con

cui si possa confrontare è D. corsicus, rispetto alla quale, però, pre-

senta numerosi e costanti caratteri differenziali: la nuova specie pre-

senta costantemente i tegumenti dorsali più opachi (soprattutto capo

e pronoto), l’apice elitrale più arrotondato, l’orlo laterale delle elitre

non carenato, le antenne molto meno pubescenti (soprattutto i primi

articoli) con il terzo antennomero più trasverso, tutte le tibie e 1 tarsi

nettamente più tozzi (cfr. figg. 3-8), organo copulatore maschile piu

corto e diversamente conformato (cfr. figg. 9-10); inoltre in D. corsi-

cus le dimensioni massime sono maggiori e, nella media degli esem-

plari, il colore è più scuro e capo, pronoto ed elitre sono più slanciati.

Dichillus tyrrhenicus n. sp. e D. corsicus convivono strettamente

Fig. 9 - Organo copulatore maschile, in visione ventrale, di Dichillus corsicus (Cor-

sica, Serra di Pigno); fig. 10: idem di D. tyrrhenicus n. sp. (Sardegna, Mar-

rubiu: paratypus). Scala 0,2 mm.

616 P. LEO

in numerose località della Sardegna; in alcune occasioni mi è capi-

tato anche di raccogliere esemplari delle due specie insieme sotto lo

Stesso sasso, seppure aggregati in gruppetti monospecifici distinti.

Tuttavia, benché abbia esaminato più di cinquemila individui delle

due specie, non ho mai incontrato esemplari che potessero far pen-

sare ad ibridi interspecifici; evidentemente la distanza genetica tra

le due specie è tale da non permettere, nonostante la stretta sinto-

pia, fenomeni di ibridazione. L'affermazione di SOLDATI & SOLDATI

(2002), secondo i quali “...il existe deux formes extrémes, et toute

une gamme d’intermédiaires”, è dovuta evidentemente ad una errata

interpretazione dei caratteri differenziali.

Note ecologiche. Dichillus tyrrhenicus n. sp. colonizza

vari ambienti, dal livello del mare fino ad almeno 1300 m di quota.

Manifesta una netta preferenza per gli ambienti xerici con suoli com-

Fig. 11 - Distribuzione geografica di Dichillus corsicus (località limitrofe sono se-

gnate sotto lo stesso simbolo).

OSSERVAZIONI SU DICHILLUS CORSICUS 617

patti e affioramenti rocciosi ed è frequente nelle garighe costiere e di

media montagna, nelle zone a macchia mediterranea più degradata

e nelle aree sottoposte a pascolo intensivo e ad incendi periodici.

Presente tutto l’anno, è molto più frequente in autunno-inverno ed

è rinvenibile, spesso in serie numerose di esemplari, sotto i sassi o

vagliando il terriccio alla base dei pulvini, talvolta associato a varie

specie di formiche.

Derivatio nominis. Il nome della nuova specie deriva

dalla sua distribuzione geografica.

Fig. 12 - Distribuzione geografica di Dichillus tyrrhenicus n. sp. (località limitrofe

sono segnate sotto lo stesso simbolo).

618 P. LEO

Dichillus (Dichillus) tapinomae n. sp. (figg. 13, 15)

Diagnosi. Un Dichillus s. str. di medie dimensioni, con

tegumenti molto opachi, corpo di colore bruno scuro, zampe e antenne

bruno rossicce. Punteggiatura dorsale poco robusta. Pronoto subcilin-

drico, più lungo che largo, non ribordato ai lati. Elitre slanciate, con

otto serie di punti relativamente piccoli e ravvicinati; settimo intervallo

elitrale molto convesso, subcarenato; ribordo elitrale distinto, epipleure

ben delimitate. Antenne sottili, con setole corte ma fitte, ben visibili.

Tibie e tarsi slanciati. Dimorfismo sessuale secondario attenuato.

Serie tipica. Holotypus d: Sardegna (prov. Oristano),

San Giovanni di Sinis, 7.IV.1995, leg. P. Leo. Paratypi dd e P9

(stessa località dell’holotypus): 5.X.1979, leg. C. Meloni 1 ex.;

Fig. 13 - Habitus del dé di Dichillus tapinomae n. sp. (Sardegna, San Giovanni di

Sinis: paratypus) (foto A. Liberto).

OSSERVAZIONI SU DICHILLUS CORSICUS 619

10:VIE1980y leg. <Fancello ol ex.j1731V21998; leenePoritbeo? 1

ex; (7.1V.1995, lege Pi Leo renyg 2211997016 Pireo text:

14.X.1998, leg. D. Sechi 34 exx.; 19.III.2007, leg. P. Leo 9 ex.

Holotypus e due paratypi depositati presso il Museo Civico di

Storia Naturale “G. Doria” di Genova (MSNG); altri paratypi nelle

seguenti collezioni: CG, CL, CLI, CM, CS, CSO.

Descrizione. Lunghezza totale 3,0-3,83 mm. Tegumenti

estremamente opachi, alutacei, bruno scuri, zampe e antenne bruno

rossicce.

Capo più lungo che largo (rapporto lunghezza/larghezza:

1,10-1,15), con punteggiatura mediocremente robusta, fitta e ben

impressa sulla fronte e sul vertice, più rada e superficiale sul clipeo.

Guance poco dilatate in avanti, tempie debolmente convergenti in

addietro, cresta sopraoculare debolmente ingrossata.

Antenne con setole giallastre piuttosto corte ma abbastanza fitte,

ben visibili su tutti gli antennomeri. Primo articolo antennale un

po’ più lungo che largo; 2° e 3° subeguali, 1,1 volte più larghi che

lunghi; 4°-10° trasversi, 11° subquadrato.

Pronoto più stretto del capo, subcilindrico, più lungo che largo

(rapporto lunghezza/larghezza: 1,15-1,2), con la massima larghezza

intorno alla metà Lati del pronoto assai poco arrotondati, appena

ristretti in avanti e in addietro, non sinuati avanti alla base; angoli

anteriori e posteriori ottusi; ribordo laterale indistinto, appena

accennato nel quarto anteriore. Punteggiatura dorsale mediocre-

mente robusta, dello stesso diametro di quella della fronte, piuttosto

fitta. Pleure protoraciche con punti simili per diametro a quelli del

pronoto, ma più radi; prosterno e metasterno con punteggiatura un

poco più robusta, diradata;.

Elitre poco convesse, oblunghe, snelle, più di due volte più

lunghe che larghe (rapporto lunghezza/larghezza: 2,12-2,22), poco

arrotondate ai lati, con la massima larghezza appena dietro la metà,

attenuate in punta all'apice. Elitre con otto serie longitudinali di

punti relativamente piccoli ma ben impressi, soprattutto sulle strie

interne e nella metà basale dell’elitra, piuttosto ravvicinati tra loro;

intervalli elitrali subpiani, ad eccezione del settimo che è molto con-

vesso e quasi careniforme, e del terzo e quinto che sono un po’ con-

vessi nella metà basale dell’elitra; intervalli elitrali con microscultura

molto forte, quasi granulosa. Orlo laterale delle elitre ben rilevato,

620 P. LEO

subcareniforme; epipleure ben delimitate dalla base all’apice elitrale,

prive di serie ben definite di punti ma con una scultura granulosa e

plissettata e qualche confuso punto irregolare.

Sterniti addominali mediocremente punteggiati; i punti sono più

robusti sul primo e secondo sternite visibile (con un diametro simile

a quelli del metasterno), più fitti sul quarto e quinto. Nei dd i primi

due urosterniti sono appena spianati nella zona centrale.

Zampe snelle; femori leggermente clavati e sinuosi, tibie subrette

e relativamente lunghe (il rapporto lunghezza/larghezza è di 4,5

nelle tibie anteriori e mediane, di 5,5 nelle posteriori), tarsi molto

allungati, unghie lunghe e robuste. Nei dd tutte le tibie presentano,

al lato interno, una serie di minuscoli tubercoletti, appena apprezza-

bili a forte ingrandimento, e un piccolissimo dentino aguzzo apicale,

quasi perpendicolare all’asse longitudinale delle tibie stesse.

Organo copulatore maschile (fig. 15) poco sclerificato, con il

tegmen un po’ più corto della capsula paramerica; questa è 3,8 volte

più lunga che larga, subparallela, bruscamente attenuata all’apice;

pezzo mediano relativamente breve.

OSS €fy,.a 71.0 11. 1¢. "mote CO mipa.r.a.ti ve... Dichillus

tapinomae n. sp. sembra essere un endemita della Sardegna ed é

attualmente noto di un’unica localita costiera della penisola del Sinis

(Sardegna occidentale).

La nuova specie appare nettamente differenziata da tutte le con-

generi per i caratteri elencati nella diagnosi; in particolare 1 tegumenti

estremamente opachi ed alutacei la distinguono a colpo d’occhio da

tutte le specie occidentali del sottogenere Dichillus; rispetto a D. cor-

sicus e D. tyrrhenicus n. sp., le altre specie presenti in Sardegna,

presenta inoltre una punteggiatura dorsale e sternale più minuta, il

pronoto più cilindrico, i caratteri sessuali secondari delle tibie e degli

sterniti addominali attenuati, organo copulatore maschile diversa-

mente conformato (cfr. figg. 9, 10, 15); da D. tyrrhenicus n. sp. si

differenzia ulteriormente per la colorazione più scura, la conforma-

zione più slanciata di capo, pronoto ed elitre, il margine elitrale ben

delimitato, gli articoli antennali meno trasversi e più villosi, le tibie

più strette e 1 tarsi più allungati.

Note ecologiche. Dichillus tapinomae n. sp. è stato

raccolto nelle dune e nel retroduna sabbioso della spiaggia di San

Giovanni di Sinis, esclusivamente nei nidi della formica Tapinoma

OSSERVAZIONI SU DICHILLUS CORSICUS 621

simrothi Krausse, 1909 (det. M. Mei), specie della sottofamiglia

Dolichoderinae ampiamente distribuita nella parte meridionale del

bacino mediterraneo. Vari tentativi di raccolta effettuati vagliando la

sabbia all’esterno dei formicai sono sempre risultati infruttuosi.

Derivatio nominis. La nuova specie prende il nome

dalla formica ospite.

Dichillus (Dichillus) maioricensis n. sp. (figg. 14, 16)

Diagnosi. Un Dichillus s. str. di dimensioni medio-grandi,

con tegumenti lucidi di colore bruno. Punteggiatura dorsale robusta.

Pronoto più lungo che largo, ribordato ai lati. Elitre appiattite sul disco,

slanciate, acuminate all’apice; strie elitrali formate da punti molto

robusti; terzo, quinto e settimo intervallo molto elevati, careniformi.

Epipleure elitrali con una distinta serie di punti. Antenne robuste

con lunghe e fittissime setole. Tibie e tarsi slanciati. Organo copula-

tore maschile molto slanciato, con capsula paramerica lunga e stretta.

Serie tipica. Holotypus 6: Isole Baleari, Mallorca,

Sierra de Torrellas, 20.IV.1993, leg. H. Pierotti. Paratypi: stessa

località dell’holotypus, 20.IV.1993, leg. H. Pierotti 1 6 e 1 9; Mal-

lorca, Llucmajor, Son Sunyer, 24.I1V.2000, leg. C. Bellò 1 d e 2 99;

Mallorca, Palma, Son Muntaner, 24.1V.2000, leg. C. Bello 1 9.

Holotypus depositato presso il Museo Civico di Storia Naturale

“G. Doria” di Genova (MSNG); paratypi: 2 dd e 2 22 (CL), 1 2

(CT) 1° CREDI

Descrizione. Lunghezza totale 3,12-4,25 mm. Corpo,

antenne e zampe di colore rosso bruno, talvolta le elitre più scure.

Tegumenti lucidi.

Capo più lungo che largo (rapporto lunghezza/larghezza:

1,17-1,22) con punteggiatura fitta e robusta sulla fronte e sul ver-

tice, più rada e più sottile sul clipeo. Guance poco dilatate in avanti,

tempie debolmente convergenti in addietro, bruscamente ristrette

all’altezza del collo; cresta sopraoculare ben tracciata.

Antenne robuste con setole giallastre lunghe ed estremamente

fitte, compatte; 1° antennomero allungato, 2° un po’ trasverso,

distintamente più piccolo del 3° che è all’incirca così lungo che

largo; antennomeri dal 4° al 10° trasversi, 11° subquadrato.

622 P LEO

Pronoto più stretto del capo, subcilindrico, circa 1,2 volte

più lungo che largo, con la massima larghezza intorno alla metà o

appena spostata in avanti. Lati del pronoto poco arrotondati; angoli

anteriori con vertice leggermente protratto in avanti; ribordo laterale

sottile ma distinto e completo. Punteggiatura del pronoto robusta e

abbastanza fitta, 1 punti di un diametro un poco maggiore di quelli

della fronte. Pleure protoraciche e prosterno con punteggiatura più

robusta e meno fitta di quella del pronoto; metasterno con punti

ancora più grossi e radi.

Elitre molto appiattite sul disco, oblunghe, più di due volte

più lunghe che larghe (rapporto lunghezza/larghezza: 2,10-2,23),

con la massima larghezza appena dietro la metà, attenuate in punta

Fig. 14 - Habitus del d di Dichillus maioricensis n. sp. (Baleari, Is. Maiorca, Sierra

de Torrellas: paratypus) (foto A. Liberto).

OSSERVAZIONI SU DICHILLUS CORSICUS 623

all’apice. Elitre con otto serie longitudinali di punti, molto grossi e

ben impressi. terzo, quinto e settimo intervallo elitrale fortemente

elevati, e costiformi dalla base al declivio apicale. Orlo laterale

delle elitre rilevato, subcareniforme; epipleure delimitate dalla base

all’apice elitrale, con una serie longitudinale di punti, più piccoli di

quelli delle serie dorsali ma ben distinti.

Sterniti addominali punteggiati; 1 punti sono più robusti e radi

sul primo e secondo sternite visibile, progressivamente più fini e

densi sul terzo, quarto e quinto. Nei dé i primi due urosterniti sono

appena spianati nella zona centrale.

Zampe snelle; femori leggermente clavati, tibie appena sinuose e

relativamente lunghe (il rapporto lunghezza/larghezza è di 4,5 nelle

tibie anteriori e mediane, di 5,5 nelle posteriori), tarsi allungati,

unghie lunghe e robuste. Nei dd tutte le tibie presentano, al lato

interno, una serie di piccoli tubercoletti e un dentino aguzzo apicale,

quasi perpendicolare all’asse longitudinale delle tibie stesse.

Organo copulatore maschile (fig. 16) molto slanciato, con ca-

psula paramerica appena più lunga del tegmen, 4,5 volte più lunga

che larga, subparallela, bruscamente attenuata all’apice.

Osservazionlii e note compateacive. Dichil-

lus matoricensis n. sp. sembra essere un raro endemita dell’isola

di Maiorca ed è la sola specie del sottogenere nominale presente

nell’ Arcipelago delle Baleari. Dichillus (Dichillinus) laeviusculus

Kraatz, 1862, specie iberico-magrebina citata di Maiorca, Ibiza e

Formentera (EspaNoL 1954, EspaNoL & Comas 1981), si distingue

a colpo d’occhio dalla nuova specie per numerosi caratteri (dimen-

sioni minori, aspetto d’insieme molto tozzo, punteggiatura dorsale

finissima, pronoto ed elitre privi di orlo laterale, intervalli elitrali

piani, antenne molto più grosse e con setole cortissime e rade, ecc.).

Per vari caratteri, quali dimensioni, forma generale, pronoto distin-

tamente orlato ai lati e con gli angoli anteriori un poco protratti

in avanti e terzo antennomero non trasverso, D. matoricensis n. sp.

mostra evidenti affinità con D. subcostatus (Solier, 1838), specie

ampiamente diffusa nella Penisola Iberica e nel nord del Marocco,

morfologicamente molto variabile e di cui andrebbe approfondita la

sistematica a livello subspecifico. Infatti, nonostante le sinonimie

proposte da VINOLAS & CARTAGENA 2005, è probabile che le diverse

“Varietà” descritte per questa entità vadano almeno elevate al rango

di razza geografica (cfr. anche ESPANOL 1947, 1956).

624 P LEO

La nuova specie si differenzia però nettamente da qualsiasi

forma di D. subcostatus sensu lato per i seguenti caratteri: setole delle

antenne molto più fitte, soprattutto sugli articoli 2°-5°; disco elitrale

più appiattito; terzo, quinto e settimo intervallo elitrale molto ele-

vati (molto più che in D. subcostatus “var.” elevatus Reitter, 1886),

careniformi dalla base al declivio apicale; epipleure elitrali con una

distinta serie di punti; organo copulatore maschile molto più slan-

ciato, con capsula paramerica molto lunga e stretta (cfr. h. o.: fig.

16, EspaNoL 1947: fig. 2 e VitoLas & CARTAGENA 2005: fig. 133).

Note ecologiche. Dichillus matoricensis n. sp. è stato

raccolto al vaglio nella macchia mediterranea.

Fig. 15 - Organo copulatore maschile, in visione ventrale, di Dichillus tapinomae

n. sp. (Sardegna, San Giovanni di Sinis: paratypus); fig. 16: idem di D.

matoricensis n. sp. (Baleari, Is. Maiorca, Sierra de Torrellas: holotypus).

Scala 0,2 mm.

OSSERVAZIONI SU DICHILLUS CORSICUS 625

Derivatio nominis. Dal latino Maiorica-ae, antico

nome dell’isola di Maiorca.

RINGRAZIAMENTI

Ringrazio sentitamente i seguenti amici e colleghi che, favoren-

domi in vario modo, hanno contribuito alla realizzazione di questa

nota: Luca Fancello (Cagliari), Dr. Giulio Gardini (Genova), Andrea

Liberto (Roma), Maurizio Mei (Dipartimento di Biologia Animale

e dell Uomo, Università degli Studi di Roma “La Sapienza”), Dr.

Roberto Poggi (Direttore del Museo Civico di Storia Naturale “G.

Doria” di Genova), Dr. Daniele Sechi (Assessorato dell’Agricol-

tura e Riforma agro-pastorale, Regione Autonoma della Sardegna,

Cagliari), Dr. Fabien Soldati (Montpellier). Desidero infine riservare

un ringraziamento particolarmente caloroso e riconoscente all'amico

Carlo Meloni di Cagliari, compagno di numerose escursioni e al

quale debbo la comunicazione di un grandissimo numero di dati e

di materiali, fra i quali la serie tipica di Dichillus matoricensis n. sp.

BIBLIOGRAFIA

ANDREETTI A., D1 GAETANO B., D1 Marco C., OsELLA G. & RITI M., 1997 - 23.

Coleoptera Tenebrionidae (Insecta) (pp. 425-443) - In: Osella G., Biondi M..,

Di Marco C. & Riti M. - Ricerche sulla Valle Peligna (Italia centrale, Abruzzo).

Quaderni di Provincia oggi, L'Aquila, 23 (2): 419-814.

ARDOIN P., 1973 - Contribution a l’étude des Tenebrionidae (Coleoptera) de Sardai-

gne - Annales Soc. ent. Fr., (n. s.), Paris, 9 (2): 257-307.

BauDI F., 1874 - Coleotteri Tenebrioniti delle collezioni italiane. Parte prima - Bull.

Soc. ent. ital., Firenze, 6: 183-200, 275-301.

BONNEAU P., 1988 - Contribution a la rédaction d’un catalogue des Tenebrioni-

dae (Coleoptera) de France. Première partie - L’Entomologiste, Paris, 44 (3):

133-145.

CANZONERI S., 1977 - Contributo alla conoscenza dei Tenebrionidi appenninici

(XXXI Contributo allo studio dei Tenebrionidi) - Boll. Mus. civ. St. nat.

Verona, 4: 227-285.

EspANoL F., 1947 - Algunos Stenosinae y Opatrinae recogidos durante una breve

campana entomologica por la provincia de Castellon (Col. Tenebrionidae) -

Eos, Madrid, 23 (3): 185-200.

626 P. LEO

EspaNoL F., 1954 - Los Tenebrionidos (Col.) de Baleares - Trab. Mus. Cien. nat.

Barcelona, (n. s., Zool.), 1 (5): 1-96.

EspANOL F., 1956 - Contribucién al estudio de los tenebrionidos del noroeste de

Espanfia (Col. Heteromeros) - Publ. Inst. Biol. apl., Barcelona, 24: 5-71.

EspaNoL F. & Comas J., 1981 - Sobre algunos Tenebrionidos (Col.) del Mediterra-

neo occidental - Eos, Madrid, 57: 91-96.

GARDINI G., 1976 - Materiali per lo studio dei Tenebrionidi dell’ Arcipelago

Toscano (Col. Heteromera) - Lavori Soc. ital. Biogeogr., (n.s.), Forlì, 5 (1974):

637-723.

GARDINI G., 1995 - Coleoptera Polyphaga XIII (Lagriidae, Alleculidae, ‘Tenebrio-

nidae), 17 pp. - In: Minelli A., Ruffo S. & La Posta S. (eds.). Checklist delle

specie della fauna italiana, 58, Calderini, Bologna.

Leo P., 1985 - Segnalazioni faunistiche italiane. 54. Dichillus (s. str.) minutus RR,

- Boll. Soc. ent. ital., Genova, 117 (1-3): 64.

Lo Cascio P., 2001 - Tenebrionidae (pp. 219-228) - In: A. Sforzi & L. Bartolozzi

(eds). Libro Rosso degli insetti della Toscana. Arsia, Firenze, 376 pp.

Marcuzzi G., 1985 - I Tenebrionidi (Coleoptera Heteromera) conservati presso il

Museo di Scienze Naturali di Bergamo - Rivista Mus. Sc. nat. Bergamo, 9:

117-125

PORTA A., 1934 - Fauna Coleopterorum Italica, 4. Heteromera-Phytophaga - Stab.

Tip. Piacentino, Piacenza, 415 pp.

REITTER E., 1916 - Bestimmungs-Tabellen der palaearktischer Coleopteren. 79. Heft.

Tenebrionidae Zopherini, Elenophorini, Leptodini, Stenosini und Lachnogyini

aus der paldarktischen Fauna - Wien. ent. Zeit., Wien, 35 (5-7): 129-171.

SOLDATI F. & SOLDATI L., 1998 - Liste systématique remise a jour des Coléoptè-

res Tenebrionidae de la faune de France - Bull. Soc. linn. Bordeaux, 26 (4):

141-154.

SOLDATI F. & SOLDATI L., 2002 - Les Dichillus corso-sardes: taxonomie et géoné-

mie (Coleoptera, Tenebrionidae, Stenosini) - Bull. Soc. ent. Fr., Paris, 107 (2):

165-168.

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428 pp.

OSSERVAZIONI SU DICHILLUS CORSICUS 627

RIASSUNTO

Vengono discussi i problemi nomenclatoriali riguardanti Dichillus (Dichillus)

corsicus (Solier, 1838) (= pumilus Solier, 1838) che viene ridescritto. Vengono inoltre

descritte tre nuove specie: Dichillus (Dichillus) tyrrhenicus n. sp. (Sardegna, Corsica

e Toscana), in passato citato da vari Autori come pumilus (nec Solier), D. (D.) tapi-

nomae n. sp. (Sardegna) e D. (D.) matoricensis n. sp. (Baleari, Maiorca).

ABSTRACT

Remarks on Dichillus corsicus and description of three new species from West

Mediterranean (Coleoptera Tenebrionidae).

The nomenclatural problems concerning Dichillus (Dichillus) corsicus (Solier,

1838) (= pumilus Solier, 1838) are discussed, and a redescription of the species is

given. Moreover three new species are described: Dichillus (Dichillus) tyrrhenicus n.

sp. from Sardinia, Corsica and Tuscany, previously often quoted by many Authors

as pumilus (nec Solier), D. (D.) tapinomae n. sp. from Sardinia, and D. (D.) maiori-

censis n. sp. from Majorca (Balearic Islands).

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’

629

GIULIO GARDINI *

RICERCHE ZOOLOGICHE DELLA

NAVE OCEANOGRAFICA “MINERVA” (C. N. R.)

SULLE ISOLE CIRCUMSARDE. XXXI.

PSEUDOSCORPTONES'**

INTRODUZIONE

Le prime notizie relative a Pseudoscorpioni delle piccole isole

circumsarde si trovano in due contributi di GesTRO (1904) e di

ELLINGSEN (1909), 1 quali citano rispettivamente due specie per l’Isola

di San Pietro e tre per l'Isola Asinara. Più recentemente CALLAINI

(1983a, b) indica la presenza di sei specie sull’ Isola di Sant'Antioco,

quindi GRAFITTI & Muceppa (1996) e GRAFITTI et al. (1996) citano

due specie cavernicole dell'Isola Tavolara. Lo stato delle conoscenze

a tutt'oggi è quindi di 12 specie citate e di sole 4 isole esplorate.

Grazie alla cortesia del dott. Roberto Poggi, Direttore del Museo

civico di Storia naturale “G. Doria” di Genova, ho potuto studiare

circa 1.320 esemplari di Pseudoscorpioni, quasi tutti da lui raccolti

su 44 isole circumsarde nel corso delle ricerche zoologiche della nave

oceanografica “Minerva” del C. N. R. effettuate negli anni compresi

tra il 1986 e il 1994.

Le 24 specie identificate (e le 3 non identificate) sono elencate

secondo l’ordine sistematico seguito da GARDINI (2000); l’elenco dei

reperti e delle isole segue il criterio geografico proposto da BACCETTI

et al. (1989): da NE (Arcipelago de La Maddalena) in senso orario

sino a NW (Is. Asinara).

Qualora non diversamente specificato, il materiale si intende

raccolto da R. Poggi e depositato presso il Museo civico di Storia

naturale “G. Doria",

* c/o Dip.Te.Ris., Universita degli Studi, Corso Europa 26, I-16132 Genova GE

** Pseudoscorpioni d’Italia XLI.

630 G. GARDINI

ELENCO DELLE SPECIE

1) Chthonius (Chthonius) ischnocheles (Hermann, 1804)

BEIER 1963a: 25; GARDINI 2004: 126.

Is. La Maddalena, Case Fangotto, 4.VII.1990, 1 ®; Is. Asinara,

Cala Atena WNILISSA LIS, 2.

Specie euro-anatolica-macaronesica nota di tutte le regioni d’Ita-

lia; euriecia e antropofila, nelle regioni centromeridionali e in Sarde-

gna è particolarmente frequente nelle zone fotiche di cavità naturali

e artificiali. Delle isole circumsarde era già citata di Sant Antioco

(CALLAINI 1983b).

2) Chthonius (Chthonius) tenuis L. Koch, 1873

BEIER 1963a: 31.

Is. Spargi, Cala Granara, 16.X.1989, 1 d; id., Valle di Cala

Canniccio, 13.XII.1993, vaglio Arbutus unedo L., 1 ®; Is. La Mad-

dalena, La Trinità, 9.1V.1986, C. Manicastri e R. Argano leg., 2 dd;

Is. Caprera, Fonte Teialone, 7.IV.1986, sub Quercus ilex L. , 1 ®;

Is. Santo Stefano, 12.XI.1986, 1 6; Is. delle Bisce, 11.XI.1986, 1

T (***); Is. delle Rocche, 11.X1.1986, 1 d; Is. Sant'Antioco, Cala

Lunga, 12.XII.1993, V. Cottarelli leg., 1 d; id., Sa Scrocca Manna,

12.XII.1993, 1 6.

Specie euro-maghrebina nota di tutte le regioni italiane. Dif-

fusa, ma sporadica, in tutta la Sardegna, ma non ancora citata di

alcuna isola circumsarda.

3) Chthonius (Ephippiochthonius) berninii Callaini, 1983

CALLAINI 1983a: 406.

Is. Serpentara, 9.V.1988, 1 9; Is. Il Toro, 10.V.1988, 1 ®.

*** 'T’ = tritoninfa, D = deutoninfa, P = protoninfa.

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 631

Specie endemica della Sardegna meridionale. Delle isole cir-

cumsarde era già nota di Sant'Antioco (Callaini 1983a).

4) Chthonius (Ephippiochthonius) elbanus Beier, 1963

BEIER 1963b: 151; GARDINI 1993: 81.

Is. La Maddalena, Case Fangotto, 14.X.1989, 2 29; Is. Figarolo,

28.VII.1986, al vaglio sub Pistacia sp., 1 2; Is. Tavolara, versante

NW, 16.X.1989, 1 2; Is. Molara, 9.1V.1986, sub Quercus ilex L., 1 9.

Specie W-mediterranea a gravitazione tirrenica nota delle

seguenti regioni italiane: Friuli-Venezia Giulia, Veneto, Emilia-

Romagna, Lombardia, Piemonte, Liguria, Toscana e Arcipelago

Toscano, Lazio, Sicilia e Sardegna centrosettentrionale (GARDINI

2000, 2004). Non citata sinora di isole circumsarde.

5) Chthonius (Ephippiochthonius) sp. prope elbanus Beier, 1963

Is. Li Nibani 1, 11.XI.1986, 1 ®; Is. delle Rocche, 11.XI.1986,

al vaglio, 1 d; Is. Soffi, 10.IV.1986, N. Baccetti leg., 1 d.

Popolazioni conspecifiche affini a C. elbanus per la presenza

di un dente subterminale al dito mobile dei cheliceri, per la forma

della mano dei palpi e per il modello di dentatura delle dita degli

stessi: le dimensioni sono tuttavia nettamente minori (pinze dei

palpi 0.46-0.48 mm). Lo stato tassonomico di queste popolazioni

sarà formalizzato in una revisione delle specie italiane del subgen.

Ephippiochthonius Beier, 1930.

6) Chthonius (Ephippiochthonius) gibbus Beier, 1953

GARDINI 1980: 268;1993: 82.

Is. La Maddalena, Spiaggia dei Monti della Rena, 8.IX.1987,

sub Pistacia lentiscus L., 1 £; Is. San Pietro, pendici Monte Guar-

dia dei Mori, 10.VI.1989, 3 dd; Is. Asinara, dintorni Tumbarino,

13.X.1989, 1 d.

632 G. GARDINI

Specie W-mediterraneo-macaronesica nota, per l’Italia, di Ligu-

ria, Calabria, Sicilia, Isole Eolie e Sardegna (GARDINI 2000). Non

ancora citata di isole circumsarde.

7) Chthonius (Ephippiochthonius) tetrachelatus (Preyssler,

1790)

BEIER 1963a: 57.

Is. Molara, 13.IX.1987, sub Salix sp., 1 ®; Is. Serpentara,

30.VII.1986, sub Pistacia lentiscus L., 2 66, 1 ®; id., 7.VII.1990, 1

2: Is. San Pietro, Spiaggia Vinagra, 10.V.1988, V. Cottarelli leg., 1

?:; id., Stagno Cala Vinagra, 13.V.1988, 1 2.

Specie ad ampia distribuzione euro-mediterranea-macaronesica,

importata in U. S. A. e in Australia. Nota di quasi tutte le regioni

italiane, isole maggiori comprese (GARDINI 2000). Non ancora citata

di isole circumsarde.

8) Spelyngochthonius sardous Beier, 1955

GARDINI 1994: 6.

Is. Tavolara, Inghiottitoio della Mandria 189 Sa/SS, 90 m,

20.111.1994, M. Mucedda leg., 1 £; id., Grotta dei Trolls 2402 Sa/

SS, 150 m, 19.IX.1998, A. Gillono leg., 1 T (tutti in coll. Gardini).

Specie troglobia già citata dell’Inghiottitoio della Mandria

(GRAFITTI & MucEDDA 1996, sub Pseudoscorpiones; GRAFITTI et al.

1996, sub Spelyngochthonius n. sp.), attribuita a S. sardous da GAR-

DINI (in stampa).

9) Neobisium (Neobisium) incertum J. C. Chamberlin, 1930

BEIER 1963a: 97.

Is. Budelli, pendici Monte Budello, 14.X1II.1993, 1 9; Is. Spargi,

Cala’ Granate 0 X98 720 VI*' id. ~ Cala Canniécio, V3UX 11993,

sotto Pistacia lentiscus L. in spiaggia, 1 d; id., Valle di Cala Cannic-

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 633

cio, 13.XII.1993, vaglio Arbutus unedo L., 2 63, 2 2; Is. La Mad-

dalena, Case Fangotto, 14.X.1989, 4 TT; id., I Pozzoni, 14.X.1989,

vaglio lentisco, 1 T; id., Case Fangotto, 4.VII.1990, 2 ®9; Is.

Caprera, Stagnali, 7.1V.1986, sub Myrtus communis L., 1 &; id.,

Invaso Ferracciolo, 8.IX.1987, 2 TT; Isolotto Roma, 10.XI.1986,

L. Tonon, 1 2; Is. delle Bisce, 11.XI.1986, 1 gd; Is. Li Nibani 1,

11.X1.1986) ? TPs ds? Camere WP T11 Nd98 6; fd = AE? | TDs Ts.

Tavolara, 16.X.1989, 1 T; id., 9.X1.1986, 1 d, 2 29; Is. Molara,

10.XI.1986, al vaglio sub Pistacia sp., 2 dd; id., 13.IX.1987, sub

Salix sp., 2 TT; Is. Sant'Antioco, Cala Lunga, 12.V1.1989, 3 TT;

id., Cussorgia, 12.VI.1989, 2 2%; id., Stagno de Cirdu, 12.VI.1989,

1 T; id., Cala Lunga, 13.VI.1989, sotto lentisco, 1 9,2 TT; id., Sa

Scrocca Manna, 13.VI.1989, 1 ®; Is. San Pietro, Stagno della Viva-

gna, 10.V.1988, al vaglio, 1 9, 1 T; id., Canale del Geniale, dintorni

Carloforte, 11.XII.1993, 2 dd; Is. Asinara, Cala Arena, 12.X.1989,

3 66, 1 T; id., vaglio lentisco, 1 d, 1 2, 1 T; id., Monte Scomu-

nica, 12.X.1989, lecceta, 3 TTT, 1 D, 1 P; id., dintorni Tumbarino,

13.X.1989, 2 dd, 3 29; id., sotto Euphorbia dendroides L.,1 6,1 ®;

id., Cala Scombro di dentro, 13.X.1989, 1 2.

Specie endemica sarda citata sinora di due sole località: Asuni

(non Sorgono: cfr. GARDINI 2000) e Monte Sette Fratelli.

10) Roncus abditus (J. C. Chamberlin, 1930)

GARDINI & RIZZERIO 1985: 66.

Is, Razzoli, .13.X11986,.2 06.5 9° id Cala della Noce.

10.VII.1990, 1 6; id., a S del Faro, 14.XII.1993, 1 6, 4 99; Is.

Budelli, pendici Monte Budello, 14.X11.1993, 5 dd, 8 29, 1 T; Is.

La Presa, 16.X.1989, sotto lentisco, 1 2, 1 T; Isola Spargi, Cala

Granara, 16.X.1989, 11 dé, 6 29, 3 TT; id., Valle di Cala Cannic-

cio, 13.XII.1993, vaglio Arbutus unedo L., 6 66, 7 99,2 TT (3 3d,

3 92 coll. Gardini); Isola La Maddalena, Spiaggia dei Monti della

Rena, 8.IX.1987, sub Pistacia lentiscus L., 2 dd, 2 22; id., Case

Fangotto, 14.X.1989, 4 dd, 3 2%; id., Guardia Vecchia, 14.X.1989,

1 9; id., Cala Bassa Trinità, 14.X.1989, 1 gd, 2 ®9; Is. Caprera,

Fonte Teialone, 7.1V.1986, sub Quercus ilex L., 1 6, 1 9; id., Sta-

gnali, 7.1V.1986, sub Myrtus communis L.,:2 36,3 99,2 TT; id.,

12.XI.1986, M. Bologna leg., 4 66, 1 2, 6 TT; id., 13.XI.1986, R.

634 G. GARDINI

Argano leg., 1 ®; id., Forte Settentrionale, 15.X.1989, 1 6, 2 99;

id., pendici Monte Teialone, 15.X.1989, lecceta, 11 dd, 9 99, 2 TT;

id., Forte Settentrionale, 5.VII.1990, sotto corbezzolo, 1 6, 2 99; Is.

Santo Stefano, 12.XI.1986, 2 66; id., Forte San Giorgio, 17.VI.1989,

sotto mirto, 1 d, 1 2; Isolotto Roma, 12.XI.1986, 1 9, 1 T; Is. delle

Bisce, 11.XI.1986, 7 dd, 1 9; Is. Li Nibani 1, 11.XI.1986, 10 dd 1

2,2 TT, 1 D; Is. Li Nibani 3, 11.XI.1986, vaglio sub Pistacia sp.,

4 dé, 1 9; id. 3, 11.X1.1986, 1 3; Is. delle Rocche, 11.XI.1986, al

vaglio, 2 dé, 2 99; Is. Mortorio, 11.1V.1986, 1 D; id., 11.IV.1986,

sub Pistacia lentiscus L., 2 66, 3 TT, 1 D; id. , 3.VII.1987, 1 3; Is.

Camere W, 11.IV.1986, 2 od; id., 11.IV.1986, al vaglio sub Pista-

cia lentiscus L., 1 6, 1 ®; Is. Camere E, 11.IV.1986, al vaglio sub

Pistacia lentiscus L., 1 9; Is. Soffi, 10.IV.1986, N. Baccetti leg.,

1 3; Is. Figarolo, 28.VII.1986, al vaglio sub Pistacia sp., 1 6; Is.

Pavolatay (8.0V01986) 032 desc coll: Gardini); idz,..9X11986,) 23

Sd, 21 22 (5 34,5 22 coll. Gardini); id., zona bassa dietro l’abitato,

9.XI.1986, M. Zapparoli leg., 1 d; id., versante NW, 16.X.1989,

3 TT; id., 16.X.1989, 1 6, 3 99; Is. Molara, 9.IV.1986, sub Acer

sp.,1 ®; id., 10.XI.1986, al vaglio sub Pistacia sp., 10 66, 9 99; Is.

Serpentara, 9.V.1988, 1 6; id., 7.VII.1990, 1 6, 1 T, 2 DD; Is.

Asinara, Cala Arena, 12.X.1989, vaglio lentisco, 10 dd, 9 92 (coll.

Gardini); id., Monte Scomunica, 12.X.1989, lecceta, 10 dd, 7 99,

4 Ver ed dintorni “Dumbarino, 13.%.1989, 5 dd, 6 92 1 T: id.

dintorni Tumbarino, 13.X.1989, sotto Euphorbia dendroides L., 1 4;

id., Cala Scombro di dentro, 13.X.1989, 1 6, 1 9.

Specie citata dell’Italia centro-meridionale, Sicilia, Pantelleria,

Corsica e Sardegna. È probabile che le popolazioni estranee al com-

plesso corso-sardo debbano riferirsi ad altre specie (GARDINI 2000).

La citazione di Obisium lubricum per Is. Asinara (ELLINGSEN 1909)

è da riferirsi quasi certamente a R. abditus.

11) Roncus caralitanus Gardini, 1981

GARDINI 1981: 129.

Is. dei Cavoli, 30.VII.1986, 1 d; Is. Il Toro, 10.V.1988, 8 dé,

32 Gods, La Vaccas 10.V.1988, 1.9. Is. Sant'Antioco, Sa Screcca

Manna, 12.XII.1993, 3 dd, 3 29, 2 TT; Is. San Pietro, Pendici

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 635

Monte Guardia dei Mori, 27.VI.1987, 1 ®; id., NW de Le Colonne,

11.VI.1989, lentisco, 1 d, 1 @ (coll. Gardini); id., La Caletta,

11.XII.1993, sotto lentisco, 1 2; id., Canale del Geniale, dintorni Car-

loforte, 11.XII.1993, 5 66, 6 29,9 TT; Is. dei Ratti, 13.V.1988, 1 2.

Specie endemica della Sardegna meridionale nota sinora con

certezza solo della località tipica: Quartu Sant'Elena presso Cagliari

(GARDINI 2000). La citazione di Obtsium lubricum per VIs. San Pietro:

Carloforte (GESTRO 1904) è da riferirsi quasi certamente a R. caralitanus.

12) Roncus sp. prope turritanus Gardini, 1982

Is. Tavolara, Grotta dei Fiori d’Arancio 191 Sa/SS, 190 m,

27.VI.1993, G. Dore leg., 1 2, 1 T; id., 3.V.1998, J. De Waele leg.,

16,1 ®; id., Grotta dei Trolls 2402 Sa/SS, 150 m, 19.IX.1998, A.

Gillono leg., 1 6 (tutti in coll. Gardini)

Popolazioni cavernicole dell’Is. Tavolara, già citate della Grotta

dei Fiori d’Arancio (GRaFITTI & Muceppa 1996; GRAFITTI et al.

1996, sub Roncus n. sp.), con stato tassonomico da definire nell’am-

bito del complesso riferito a Roncus turritanus, comprendente popo-

lazioni ipogee delle aree carsiche di Sassari, Sedini, Ittiri e Mores,

nella Sardegna nordoccidentale (GARDINI 2000).

13) Microcreagrina hispanica (Ellingsen, 1910)

BEIER 1963a: 223 (sub M. maroccana)

Specie mediterraneo-macaronesica nota, per l’Italia, solo della

Sicilia meridionale e della Sardegna sudoccidentale; è l’unico rap-

presentante non cavernicolo italiano della famiglia Syarinidae. Citata

dell’ Isola Sant'Antioco (CALLAINI 1983b).

14) Garypus beauvoisi (Savigny & Audouin, 1826)

BEIER 1963a: 239.

Is. Santa Maria, 6.VIII.1986, R. Argano leg., 1 9; Is. Soffi,

3.VII.1987, R: Argano leg., 1 T; Is: der Cavoli, $0;Vi1.1986, R.

636 G. GARDINI

Argano leg., 1 D, 1 P; Is. Sant'Antioco, Capo Sperone, 31.V.1991, G.

Gardini e R. Rizzerio leg., 2 dd, 1 £, 1 T (coll. Gardini); Is. Mal di

Ventre, 3.VIII.1986, R. Argano leg., 1 T, 3 DD; id., 4.VIII.1986, C.

Manicastri leg., 1 6; Is. Piana dell’ Asinara, 4.VIII.1986, R. Argano

leg., 1 T, 1 P; Is. Asinara, Cala Arena, 1.VII.1987, R. Argano leg.,

16,1 ® (coll. Gardini); id., salicornieto, 23.V.2005, P. Marcia leg.,

1 (coll. Gardini):

Specie alofila mediterraneo-macaronesica, vicariante ecologica

(di coste sabbiose) di G. levantinus. Citata di pochissime località cen-

tromediterranee, senza dubbio per difetto di ricerche specializzate:

Corsica (Bonifacio), Sardegna (Is. Asinara) e Isole Maltesi (GAR-

DINI 2000). La conosco, per l’Italia, di Toscana: Parco dell’ Uccellina

(Grosseto), 20-21.V.1997, A. Zanetti leg., spiaggia, 1 6, 1 9, 2 juv.;

Sicilia: Isola Grande dello Stagnone (Trapani), 6.V.1991, R. Argano

e C. Manicastri leg., 1 6, 1 T; Sardegna: Alghero (Sassari), Capo

Caccia, 21.V.1982, G. Maceli leg., 1 D; Villaputzu (Cagliari), Porto

Corallo, 2.VII.1986, C. Torti leg., 2 DD, 17 PP; Pula (Cagliari),

Chia, spiaggia della Maddalena, 15.VIII.1997, M. Pisano leg., 1 D,

1 P (tutti in coll. Gardini). Nuova per la Toscana e per la Sicilia.

15) Garypus levantinus Navas, 1925

BEIER 1963a: 238; GARDINI 1975: 390.

Is. Razzoli, 30.VI.1987, R. Argano leg., 1 é (coll. Gardini);

Is. Serpentara, 30.VII.1986, R. Argano leg., 1 2, 4 TT; Isolotto

il Corno, 3.VIII.1986, 1 palpo; Isolotto Il Catalano, 3.VIII.1986, 1

3,2 99; Is. Asinara, Cala Sgombro, 13.X.1989, M. Cobolli leg., 1

2 (coll. Gardini).

Specie alofila N-mediterranea, vicariante ecologica (di coste

rocciose) di G. beauvoisi, citata per l’Italia di Liguria, Toscana e

Arcipelago Toscano, Isole Eolie, Lampedusa e Sardegna (GARDINI

2000). Di Sardegna era nota solo di Dorgali: Cala Gonone (CAL-

LAINI 1983b); la conosco anche di Buggerru (Oristano), Capo Pecora,

30.III.1991, G. Gardini e R. Rizzerio leg., 1 6, 2 TT, 1 D; Santa

Caterina di Pittinuri (Oristano), 26.III.1993, P. Leo leg., scogliera,

1 ®; Gairo Marina (Nuoro), 15.V.1980, G. Gardini leg., 1 2 (tutti

in coll. Gardini).

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 637

16) Geogarypus nigrimanus (Simon, 1879)

BEIER 1963a: 243.

Is. Razzoli, 30.VI.1987, al vaglio sub Pistacia lentiscus L., 3

dd, 1 2; Is. Budelli, c/o Spiaggia Rosa, 10.VII.1990, 2 9°; id.,

pendici Monte Budello, 14.XII.1993, 1 ¢; Is. La Presa, 16.X.1989,

sotto lentisco, 1 2; Is. Spargi, Cala Granara, 6.IX.1987, 1 6; Is. La

Maddalena, Spiaggia dei Monti della Rena, 8.IX.1987, sub Pistacia

lentiscus L., 1 36; id., Cala Bassa Trinità, 14.X.1989, 1 2; id., Case

Fangotto, 14.X.1989, 1 gd; id., Guardia Vecchia, 14.X.1989, 1 6; id.,

Case Fangotto, 4.VII.1990, 2 dd, 1 ®; id., I Pozzoni, 14.X.1989,

vaglio lentisco, 1 6; Is. Caprera, Forte Settentrionale, 15.X.1989,

1 ®; id., pendici Monte Teialone, 15.X.1989, lecceta, 2 dd, 1 9;

Is. Li Nibani 1, 11.XI.1986, 1 d; Is. Li Nibani 2, 11.X1.1986, C.

Manicastri leg., 1 ¢; Is. Li Nibani 3, 11.XI.1986, vaglio sub Pistacia

sp., 2 £®; Is. delle Rocche, 11.XI.1986, al vaglio, 2 29°; Is. Poveri

II, 11.1V.1986, 1 6; Is. Mortorio, 11.IV.1986, sub Cistus monspe-

liensis L., 1 2; Isola Camere W, 11.IV.1986, al vaglio sub Pistacia

lentiscus L.,1 6,2 9; id., 3.VII.1987, 1 4, 1 2; Isola Camere E,

11.1V.1986, sub Euphorbia dendroides L.,1 3, 2 9%; id., 3.VII.1987,

P's? Ts Soft VII 1987 Ts oe Ao AMan)

vaglio sub Pistacia sp., 1 2; Is. Tavolara, 9.XI.1986, M. Bologna

e R. Poggi leg., 2 9%; id., 8.VI.1989, al vaglio, 1 d; id., versante

NW, 16.X.1989, 1 ®; id., Fornaci, 19.V.1994, 1 ®; Is. Molara,

8.VI.1989, vaglio sotto lentisco, 2 29; Is. Molarotto, 10.IV.1986, 1

Pe ad. ot? PS Pos ae dite TO alias ARE

lotto Varaglioni, 30.VII.1986, al vaglio sub Pistacia sp., 1 &; Is.

Serpentara, 30.VIT.1986, sub Pistacia lentiscus TU. 1./6,-1. 9: id,

TNTIL1990, 1. 97 Issa eno, TO Val oases. 4 conic. sayy ares cup

Ecballium elaterium (L.), 2 2; id., 14.VI.1989, sub Ecballium elate-

riumd{L.), 8 od, 7 9, 5'TIRGS0,3 0011 Gardini. lady acca.

26.VI.1987, 1 3; id., 26.VI.1987, vaglio sub Malvacea, 3 dd, 1 9, 1

T; id., 14.VI.1989, 1 9; Is. Sant'Antioco, Cala Lunga, 11.11.1984,

L. Fancello e P. Leo leg., 1 2 (coll. Gardini); id., Stagno de Cirdu,

12,V1.1989, 2 o¢..2 P°-.id., Sa.,Sorocca IMamna, 19.VI.198% Là,

292% Is. San Pietro, La Caletta NARO, 1c. 2 fed. pen

dici Monte Guardia dei Mori, 8.VII.1990, 1 6; Is. Mal di Ventre,

2 VIII.1986,' to 2 ed TT aida 28, V1 OST Ld td 5 NT1080:02

29; Is. dei Porri, 29.VI.1987, sub Atriplex sp., 1 6, 1 ®; Is. Piana

638 G. GARDINI

dell’ Asinara, 4.VIII.1986, 1 9; Is. Asinara, Cala Reale, 9.VII.1990,

Drag

Specie mediterraneo-macaronesica, xerofila, propria della gariga

e della macchia mediterranea bassa. Nota per l’Italia di Liguria, Arci-

pelago Toscano, Isole Pontine, Puglia, Calabria, Sicilia, Isole Eolie e

Sardegna (GARDINI 2000). La citazione di G. minor (L. Koch, 1873)

per l’Is. Sant'Antioco (CALLAINI 1983b) è a mio avviso da riferire a

G. nigrimanus: le due specie sono comunque da revisionare.

17) Olpium pallipes (Lucas, 1849)

HEURTAULT 1979: 925.

Is. dei Porri, 29.VI.1987, 1 6.

Specie psammoalofila mediterraneo-macaronesica nota, per

l’Italia, di Liguria (probabilmente estinta), Toscana e Arcipelago

Toscano, Campania, Puglia, Basilicata, Sicilia, Is. Linosa e Sardegna

(GARDINI 2000). Citata da GESTRO (1904) dell’Is. di San Pietro.

18) Calocheiridius olivieri (Simon, 1879)

HEURTAULT 1982: 218.

Is. Razzoli, 13.XI.1986, 1 6, 2 99; Is. Spargi, Cala Granara,

6.VIII.1986, al vaglio, 2 dd, 1 2, 1.T; Is. La Maddalena, I Poz-

zoni, 14.X.1989, vaglio lentisco, 1 d; Is. Caprera, Forte Settentrio-

nale, 15.X.1989, 1 2; Is. delle Bisce, 11.XI.1986, 1 6, 1 ®; Is. Li

Nibani 1, 11.XI.1986, 1 3; Is. delle Rocche, 11.XI.1986, al vaglio, 1

6,2 reo is. vtortorio, PI V.19386) sub" Pisracia lentiscus Li 1 SUIS

Camete I PNTETOS7NI ‘08 Sor, 8VITI987r83! 8, 1399 1s;

Figarolo, 28.VII.1986, al vaglio sub Pistacia sp., 2 66, 1 £; Isolotto

Verde, 2.VII.1987, 1 6, 1 9; Is. Tavolara, settore SW, 9.XI.1986, 1

3, 1 2; id’, 8.VI.1989, al vaglio, 1 dt £; Is. Molara, 8.VI.1989,

vaglio sotto lentisco, 2 dd, 2 29; Is. Molarotto, 12.IX.1987, 15 dd,

16 22 (5 dd, 5 2° coll. Gardini); Is. Ogliastra, 6.VII.1990, 1 6;

Isolotto Varaglioni, 30.VII.1986, al vaglio sub Pistacia sp.,1 3; Is.

Serpentara, 30.VII.1986, sub Pistacia lentiscus L., 5 66, 6 22, 1 T;

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 639

id., 25.VI.1987, sub Pistacia lentiscus L., 1 3; id., 3.V.1988, M. Mei

leg., 1 3; id., 9.V.1988, 2 9%; id., 9.VI.1989, Pistacia lentiscus L.,

2 dd, 4 29; Is. dei Cavoli, 30.VII.1986, 1 3; id., 9.VI.1989, sotto

Pistacia sp., 1 3, 1 2; Is. La Vacca, 31.VII.1986, sub Pistacia sp.,

Fri DEAL O8Tie RIePoglie ti Edicola Gace Pros Ge

26.VI.1987, vaglio sub Malvacea, 2 dd, 1 2, 3 TT; id., 10.V.1988, 1

3,2 22; Is. Sant'Antioco, Stagno de Cirdu, 12.VI.1989, 6 dd, 6 29;

id., Triga, 12.VI.1989, sub Pistacia sp., 1 6; Is. San Pietro, pendici

Monte Guardia dei Mori, 27.VI.1987, 1 £; id., Bacino Acquedotto,

12.V:1988,, 2) OO: lid). babCalettay: te V1I.1989 1.6, 1 12:3 Isolotto

Il Catalano, 3.VIII.1986, 1 ®; Is. Piana di Alghero, 15.VI.1989,

1 ®; Is. Asinara, Diga Ruda, 15.V.1988, 1 6; \id., Campu Perdu,

T50V.t0 33 uns |

Specie xerofila nota sinora della Francia meridionale e della Sar-

degna. Olpium olivieri Simon, 1879, descritta su esemplari dell’Île

de Porquerolles (Iles d’Hyéres, Var), venne ridescritta e trasferita

nel genere Calocheiridius Beier & Turk, 1952 da HEURTAULT (1982).

Il genere Calocheiridius, istituito per C. mavromoustakisi Beier &

Turk, 1952 (Cipro), comprende 31 specie nell’area afrotropicale-

indo-mediterranea e, di queste, solo due erano note, sino al 1982,

del Mediterraneo: C. mavromoustakisi, segnalata anche di Creta,

Italia peninsulare e Sicilia e C. libanoticus Beier, 1955, descritta

del Libano e citata dell’Italia peninsulare, Sicilia, Sardegna, Malta,

Grecia, Turchia, Israele e Azerbaijan (HARVEY 1991; GARDINI 2000).

Come già sottolineato in altra sede (GARDINI 2000), le conoscenze

tassonomiche di questo gruppo di specie sono del tutto insufficienti

ed è assai probabile che le citazioni relative a popolazioni del Medi-

terraneo centro-occidentale, sinora attribuite a C. mavromoustakisi e

a C. libanoticus, debbano essere attribuite a C. oliviert. Non citata in

precedenza di isole circumsarde.

19) Amblyolpium dollfusi Simon, 1898

LAZZERONI 1970: 106; ScHAWALLER 1981: 46.

Is. Razzoli, Cala della Noce, 10.VII.1990, 1 ®, 1T; Is. La

Presa, 16.X.1989, sotto lentisco, 1 2, 1 D; Is. Mortorio, 3.VII.1987,

1 2; Is. Tavolara, versante NW, 29.VII.1986, 1 d; id., 8.VI.1989,

640 G. GARDINI

al vaglio, 7 66, 7 99, 1 T; id., settore SW, 19.V.1994, sotto len-

tisco, 6 ad., 1 T (3 ad. coll. Gardini); Is. Molara, 28.VII.1986, 1

3, 1 D; id., 8.VI.1989, vaglio sotto lentisco, 1 T; Isolotto Vara-

glioni, 30.VII.1986, al vaglio sub Pistacia sp., 1 d; Is. dei Cavoli,

9.VI.1989, sotto Pistacia sp., 1 2; Is. La Vacca, 31.VII.1986, sub

Pistaca spy VIS PO 7 145 PEVISB4 Ae “hh? Aids TON: 1988051

6; Is. Mal di Ventre, 3.VIII.1986, 1 6, 1 9; id., 28.VI.1987, 16,1

2: id., 15.VI.1989, 14 dd, 6 22 (4 dé, 1 £ coll. Gardini); Is. Piana

di Alghero, 29.VI.1987, 1 ®; Is. Piana dell’Asinara, 4.VIII.1986, 2

dona PRI TT,

Specie tirrenica, xerofila della gariga e della macchia mediterra-

nea bassa, nota di Francia meridionale, Arcipelago Toscano, Cala-

bria, Sardegna, Corsica (GARDINI 2000) e Sicilia: env. Caltabellotta

(Agrigento), 850 m, 22.VIII.1993, V. Mahnert leg., tamisage Quer-

cus tlex L., 1 es. (coll. Gardini). Non ancora citata di isole circum-

sarde e nuova per la Sicilia.

20) Hysterochelifer tuberculatus (Lucas, 1849)

BEIER 1963a: 285.

Is. Razzoli, Cala della Noce, 10.VII.1990, 2 TT; Is. Budelli,

dintorni Spiaggia Rosa, 10.VII.1990, 1 2, 1 T; id., pendici Monte

Budello, 14.XII.1993, 1 T; Is. La Presa, 13.X1.1986, 2 TT, 1 D;

id., 16.X.1989, sotto lentisco, 1 D; Is. Barrettini, 5.VIII.1986, al

vaglio sub Lavatera sp., 1 T; Is. Spargi, Cala Granara, 6.VIII.1986,

al’ vaslio, 1° 614) Gala’ Granara, 6.1X.1987, 179, 1 T;-1d.,° Cala

Granara, 16.X.1989, 2 66, 1 9, 1 T, 2 DD; Is. La Maddalena,

Spiaggia dei Monti della Rena, 8.IX.1987, sub Pistacia lentiscus L.,

Ge, Mere La? DD dl es id Spam 8 VIT9S9 2 oor id.:

Guardia Vecchia, 14.X.1989, 1 ®; id., I Pozzoni, 14.X.1989, vaglio

lentisco, 1 2; id., Cala Bassa Trinità, 14.X.1989, 1 2; id., Case Fan-

gotto, 4.VII.1990, 2 dg, 1 2, 3 TT; Is. Caprera, Fonte Teialone,

7.IV.1986, sub Quercus ilex L., 1 D; id., 12.XI.1986, M. Bologna

leg., 1 9; id., pendici Monte Teialone, 15.X.1989, lecceta, 2 dd, 1

2, 1D; id., Fosso di Stefano, 5.VII.1990, 1 6, 1 9, 1 P; id., Forte

Settentrionale, 5.VII.1990, sotto corbezzolo, 3 2%; Is. Santo Ste-

fano, 12.X1.1986, 1 6; id., 6.IX.1987, 1 T; id., Forte San Giorgio,

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 641

17.VI.1989, sotto mirto, 1 d, 1 T; Is. delle Bisce, 11.X1.1986, 1 T;

Is. Soffi, 3.VIH.198%; 1 6, 1990: RD. Die RE Lcarolo,

28.VII.1986f alvaslio sub Psa i TE td. VOT 1986:

229, ID; ide dO. X1.1986e vaslio-sub-Pistactausp., Woy “EF? id’)

14 TK..1987,°2. SSD 7Ase Tavolarayl SaNelosor D105 1 PPh eT eel

settore SW, 9.XI.1986, al vaglio, 6 9; id., 13.IX.1987, vaglio sub

Ficus carica L., 1 T; id., 8.VI.1989, al vaglio, 2 66, 4 29, 5 TT, 1

D; id., versante NW, 16.X.1989, 2 DD; id., settore SW, 19.V.1994,

sotto lentisco, 2 29, 1 T; id., Fornaci, 19.V.1994, 1 4,1 %, 1 D;

Is. Molara, 9.I1V.1986, sub Quercus ilex L., 1 6; id., 9.I1V.1986, sub

Acer. sp., 1.6; id., 28.V11.1986, 1d: 1d., 8.VI.1989,, vaglio sotto

lentisco, 2 dd, 2 99, 5 TT; Is. Serpentara, 20.VII.1986, M. Mei

leg., al vaglio, 1 T; id., 30.VII.1986, sub Pistacia lentiscus L., 2 66,

2 29, 1 T,2 DD; ids, 28.VL1987,; sub. Pisigera dentscus IL 6.04,

2, 22, 1 Py id., 8.V.1988y al vaglio d'entisconi id alt 130 RD

id., 9. VI.1989. Pistacia lentisciis Me. Baie, RT id,

7.VII.1990, 4 63,1 2,5 TT; Is. dei Cavoli, 30.VII.1986, al vaglio,

3 dé; Is. La Vacca, 31.VII.1986, sotto sasso con nido di Pheidole

ballidula Nyl., 1.1); td. 26,V11987, 1 SS id 26,V 1.1987, vacho

sub Malvacea, 1 6, 1 T, 1 D; Is. Sant'Antioco, Triga, 12.VI.1989,

sub Pistacia sp., 1 £, 1 D; id., Stagno de Cirdu, 12.VI.1989, 3 66,

2 292,4 TT, 3 DD: id., Sa Scrocca Manta IBN e

Is. San Pietro, pendici Monte Guardia dei Mori, 27.VI.1987, 1 3,

1 2; id., Monte Guardia dei Mori, 12.V.1988, 1 d, 1 ®; id., pen-

dici Monte Guardia dei Mori, 10.VI.1989, 3 DD, 2 PP; id., Cala

Vinagra, 10.VI.1989, 3 dg, 3 99, 1 T, 5 DD; id., Monte Guardia

dei Mori, 11.VI.1989, al vaglio, 5 99, 2 TT, 3 DD; id., NW de

Le Colonne, 11.VI.1989, lentisco, 1 ®; id., pendici Monte Guar-

dia dei Mori, 8.VII.1990, 3 TT; Is. Mal di Ventre, 3.VIII.1986, al

vaglia, 2 (86; 2 "PT IDE id 28. V LAOS Ficliedy ia Podi

29/VI.1987; sub. Atriplex isputlad plo PRCLTENLID i QPP iis. Briana

dell’Asinara, 4.VIII.1986, 1 6, 1 2, 1 T, 1 D, 1 P; Is. Asinara,

dintorni Tumbarino, 13.X1989) 2° dd.ol ®52 DD; idijndintorni

Tumbarino, 13.X.1989, sotto Euphorbia dendroides L., 1 ®.

Specie mediterranea nota di quasi tutte le regioni d’Italia (GAR-

DINI 2004) comune, in quelle centro-meridionali, in formazioni

vegetazionali della macchia mediterranea. Già nota dell’Is. Asinara

(ELLINGSEN 1909) e dell’Is. Sant'Antioco (CALLAINI 1983b).

642 G. GARDINI

21) Dactylochelifer latreillii (Leach, 1817)

BEIER 1963a: 291; MAHNERT 1977: 68.

Is. Budelli, Stagno, 10.VII.1990, 1 D; Is. Spargi, Cala Gra-

nara, 6.VIII.1986, al vaglio, 1 T; Is. Sant'Antioco, Cala Lunga,

12.VI.1989, 1 2, 1 juv.; Is. Asinara, Cala Arena, 1.VII.1987, 4 dd,

4 99; id., Cala Arena, 12.X.1989, 1 6, 2 99, 1 T; id., Fornelli,

9.VII.1990, 1 9.

Specie euro-mediterraneo-macaronesica nota di buona parte

delle regioni d’Italia (GARDINI 2000). Non ancora citata di isole cir-

cumsarde.

22) Rhacochelifer corcyrensis (Beier, 1930)

BEIER 1963a: 296; MAHNERT 1977: 69.

Is. Tavolara, 9.XI.1986, 1 d.

Specie mediterranea centro-orientale citata, per l’Italia, del

Veneto e di Puglia (GARDINI 2000). Nuova per la Sardegna.

23) Rhacochelifer maculatus (L. Koch, 1873)

BEIER 1963a: 294; HEURTAULT 1980: 161.

Is. Budelli, dintorni Spiaggia Rosa, 10.VII.1990, 1 gd; Is. La

Maddalena, Cala Bassa Trinità, 14.X.1989, 1 6; id., Case Fangotto,

4.VII.1990, 1 2, 1 T; Is. Serpentara, 9.V.1988, 1 3, 1 T; id.,

SAAS: aPistaord lentiscis0.] 2 166, IT De ide 7 VE 990) 62

dd; Is. Il Toro, 31.VII.1986, V. Cottarelli leg., 1 9; Is. Sant'An-

tioco, Triga, 12.VI.1989, sub Pistacia sp., 1 2; Is. San Pietro, pen-

dici Monte Guardia dei Mori, 27.VI.1987, 1 6.

Specie mediterraneo-macaronesica nota, per l’Italia, di Veneto,

Lombardia, Liguria, Toscana e Arcipelago Toscano, Marche, Lazio,

Abruzzo, Puglia, Calabria, Sicilia e Sardegna (GARDINI 2000). Non

ancora citata di isole circumsarde.

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 643

24) Rhacochelifer sp. prope maculatus (L. Koch, 1873)

Is. Caprera, Forte Settentrionale, 15.X.1989, 1 T; Is. Tavolara,

9.XI.1986, 1 T; Is. Il Toro, 10.V.1988, 2 es.; id., 14.VI.1989, sotto

Ecballium elaterium L., 3 29°, 2 TT; Is. La Vacca, 31.VII.1986, sub

Pistacia sp.,1:9;.1d::26.VI.1987,,4 66,229 ITT(+16,1 £ coll. Mus:

Ginevra); id., 26.VI.1987, vaglio sub Malvacea, 2 dd; id., 14.VI.1989,

1 %,,3.TT; Is. Sant'Antioco; Stagno de Cirdu, 12.V1.1989, 1 :d.;. Is;

Mal di Ventre, 3.VIII.1986, 1 2, 1 T; Is. Foradada, 29.VI.1987, 1

6; Is. Asinara, alta Valle Rio Baddi Longa, 14.V.1988, lecceta, 1 d.

Popolazioni che differiscono da R. maculatus per la forma dei

palpi, nel complesso più tozzi, e per l’assenza di grossi granuli sulla

superficie mediale del femore e della patella degli stessi. Lo statu-

men convolutum dei 6d è tuttavia pressoché identico a quello di

R. maculatus, mentre la placca cribrosa delle 99 è un poco più

dilatata apicalmente rispetto a quella di R. maculatus. La simpatria

con R. maculatus sull’ Is. Il Toro e a Sant'Antioco avvalora la sup-

posizione che si tratti di una specie inedita, la cui descrizione è da

subordinarsi, in altra sede, ad una attenta revisione delle popolazioni

mediterranee di R. maculatus.

25) Pselaphochernes italicus Beier, 1966

BEIER 1966: 109.

Is. Caprera, Invaso Ferracciolo, 8.IX.1987, 1 2; Is. delle Bisce,

11.XI.1986, V. Cottarelli leg., 2 od; Is. Figarolo, 28.VII.1986, al

vaglio sub Pistacia sp., 2 dc.

Specie descritta di Campania: Ascea, Scavi di Velia (BEIER 1966)

e citata di Sardegna, dubitativamente, da CALLAINI (1983b). Prima

segnalazione per le isole circumsarde.

26) Pselaphochernes lacertosus (L. Koch, 1873)

BEIER 1963a: 254.

Ts: «Caprera; Invaso* Ferracciolo, BVITI9IR a Sd hai S\oL D:

Is. Poveri 1, 11.1V.1986, sub Lavatera sp., 4 64, 12 29,3 TT, 1

644 G. GARDINI

D, 1 P; Is. Tavolara, 9.XI.1986, M. Bologna e V. Cottarelli leg., 1

Os, tue Pils. wee Vacca 20019871, 1 25 id 10:V.1988, 1 9,

2 TT; Is. San Pietro, Cala Vinagra, 8.VIII.1986, C. Manicastri leg.,

CEREA EVITO 897 NE Baccettr lem ro TT? 1d,

pendici Monte Guardia dei Mori, 8.VII.1990, 4 66,5 22,1 D.

Specie mediterraneo-macaronesica nota, per l’Italia, di Veneto,

Liguria, Arcipelago Toscano, Lazio, Puglia, Sicilia, Isole Eolie e

Sardegna (GARDINI 2000). Non ancora citata di isole circumsarde.

27) Atemnus politus (Simon, 1878)

BEIER 1963a: 247.

Is. Figarolo, 28.VII.1986, al vaglio sub Pistacia sp., 1 2, 1 T;

Is. dei Cavoli, 9.VI.1989, sotto Pistacia sp., 2 99; Is. La Vacca,

31.VII.1986, sub Pistacia sp., 5 TT; id., 26.VI.1987, 9 od, 5 P9, 3

TT, 1 Dj id., 10.V.1988, 5 3d, 1 ®; id., 14.VI.1989, 3 dd, 4 99,

5 TT; Is. Sant'Antioco, Su Pruini, 11.V.1988, 2 99, 1 T; id., Cala

Lunga, 12.VI.1989, 4 dd, 1 ®; id., Cala Lunga, 13.VI.1989, sotto

lentisco, 3 dd, 1 £; Is. Mal di Ventre, 3.VIII.1986, 1 2, 1 T; id.,

oe OS ae Sd 15.1 b1989 4 °¢, 2 29: Is. Piana di

Alghero, 29.VI.1987, 2 36d; id., 29.VI.1987, sub Pistacia lentiscus L.,

6 22, 1 P; Is. dei Porri, 29.VI.1987, sub Atriplex sp., 2 64, 6 99,

oi te ds, Asinarca Cala Arena, 1.V111987,.2 4,2 292.1 Te ids

Cala Arena, 1.VII.1987, R. Argano leg., 2 6g, 1 2 (coll. Gardini);

id., Cala Arena, 9.IX.1987, 2 63, 1 ®; id., Cala Arena, 12.X.1989,

1 g; id., Cala Arena, 12.X.1989, vaglio lentisco, 1 6, 2 99, 3 DD;

id., Monte Scomunica, 12.X.1989, lecceta, 1 9, 3 TT; id., dintorni

Tumbarino, 13.X.1989, sotto Euphorbia dendroides L., 1 6, 1 ®,

1 T; id., Cala Reale, 9.VII.1990, 2 dd, 4 99, 3 TT; Is. Scombro,

191989: 366,1. 9-7“ DD.

Specie centroasiatico-mediterraneo-macaronesica già citata, per

l’Italia, di Liguria, Emilia-Romagna, Toscana, Lazio, Abruzzo,

Puglia e Isole Tremiti, Calabria, Sicilia e Isole Eolie, Is. Pantelleria,

Is. Linosa e Sardegna (GARDINI 2000). Già nota dell’Is. Sant’ An-

tioco (CALLAINI 1983b).

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 645

CONCLUSIONI

Tra le regioni d’Italia, intesa in senso geografico e quindi

incluse la Corsica e le Isole Maltesi, la Sardegna è quella di cui è

noto a tutt'oggi il maggior numero di specie di Pseudoscorpioni:

73 su 218, cioè il 33,5% dell'intero popolamento (GARDINI 2000,

aggiornato). Ben 28 (il 38%) sono endemiche dell’isola e, di queste,

20 sono cavernicole e le restanti 8 epigee; le altre 45 specie, di cui

2 sole troglofile, presentano nel complesso corotipi mediterranei,

W-mediterranei o tirrenici, poche (8) sono più ampiamente diffuse

nella regione W-paleartica.

Sulle isole circumsarde sono state rinvenute 27 specie (tab. I):

24 sono presenti anche in Sardegna e le tre non ancora note dell’isola

madre (Chthonius sp. prope elbanus, Rhacochelifer corcyrensis e R.

sp. prope maculatus) vi saranno molto probabilmente rinvenute in

futuro. L’Isola di Tavolara è l’unica che ospita popolazioni troglo-

bie attribuibili, o strettamente affini, a due specie cavernicole pre-

senti in Sardegna: Spelyngochthonius sardous e Roncus turritanus. Il

debole differenziamento morfologico rilevabile nei pochi esemplari

a disposizione non consente di trarre considerazioni tassonomiche

certe (cfr. GARDINI in stampa per S. sardous) e ritengo comunque sia

conseguenza dell'isolamento carsico, e non marittimo, dal momento

che Tavolara risulta effettivamente separata dalla Sardegna da non

più di 8000 anni (ANTONIOLI & TRAINITO 2005).

L’esame dei corotipi delle specie delle isole circumsarde (tab.

I), intesi nel senso di VIGNA TAGLIANTI et al. (1999), ripropone

quanto già constatato a proposito delle specie presenti in Sardegna:

quelli dominanti sono i corotipi mediterranei s. l. e i tirrenici (11

e 3) a cui seguono gli endemici sardi (5), gli euro-mediterranei (4),

1 turano-mediterranei (1) e i centroasiatico-mediterranei (1). Due

specie (Chthonius sp. prope elbanus e Rhacochelifer sp. prope macu-

latus) hanno corotipo non definibile.

Le presenze di Roncus caralitanus e Microcreagrina hispanica

nelle piccole isole sudoccidentali sottolineano la peculiarità del popo-

lamento sulcitano, mentre la presenza di Chthonius berninii sull’ Isola

Serpentara e su Il Toro conferma l’endemicità della specie nella

Sardegna meridionale.

Le altre 24 specie sono distribuite in tutti i settori in cui sono

ripartite le isole circumsarde ed è evidente, dall'esame della tab. I,

che le isole più estese, e quindi con più elevata diversità ambien-

G. GARDINI

646

- Pseudoscorpioni presenti nelle isole circumsarde. Le specie sono elencate in

Tab. I

ordine sistematico, le isole in ordine geografico da NE a NW. Sono indicati

inoltre: i sette gruppi di isole definiti con criterio geografico (prima riga), la

ie eestor det Fay IT O

Th, 2) 8 ono

2| 2

Eee

ISOLARE

see] TRE

cwanrra{ evo | | 1 | 1111 |

Cis

is Ses Ra

eal A} s Ala

= = ea

EREBBERE

È S =

E ù s

= & si mS

è Bek SI N oy RS S

È 3 x 8 rS =

S| î| $$ a) s] 8

“<3 = sD D N a0 me

nal nl al al sal ai a»

Sa] Ero OSS OS | OS aS

= S a < S S <

S ©) S S S S S

INI ENIESNEGNESNENEOS

SSA SS SP SS Pe eS |S] SS

PL | ky ky LS] &

Ci ONOR) DI

et IAA

i i OO i CI i se ca

re | On ZIoEna

mad eee Re

erat hs DE ct Pa ee

ol wel CSR

stf_sest) fe] [| TT {| |xlxe[ {I [| De] [xl [el | | [xl] 10

si] roc] >|] [x] [xl | [tx] [||| [e] [el [el | [el {LIL LS

er] ew aE [555 ee deci dei Sasa

SI CE EET

sarD| |

Spelyngochthonius sardous

Neobisium incertum

N° SPECIE/ISOLA —>

10 fo a TTVAR x

Roncus caralitanus

Olpium pallipes

Amblyolpium dollfusi

Hysterochelifer tuberculatus

Atemnus politus

Roncus sp. prope turritanus

Calochetridius olivieri

647

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE

in ciascuna isola (ultima riga), il corotipo di ogni specie (seconda colonna),

il numero di isole in cui ciascuna specie è presente (ultima colonna).

superficie di ciascuna isola in km? (terza riga), il numero di specie presente

Tr i ERRE RENDE RASSNZEENSE

TITO th Pek Ae abc, me SSS a) Blea Seles RAI BRE

a | nf eet FE at babel abet hel he

BIBUISY vUeIg |

Pel BP”

OJ9YS]Y euelg °

epepeio |

9IZUIA IP [PII °

648 G. GARDINI

tale, sono quelle che ospitano il maggior numero di specie. È il caso

delle isole Molara, Spargi, Tavolara, Caprera, La Maddalena, Asi-

nara, San Pietro e Sant'Antioco, con superficie compresa tra 3,4 e

109 km? e numero di specie compreso tra 7 e 12. Non sempre però

vi è correlazione diretta superficie/numero di specie: di Santo Ste-

fano (3,05 km?) ne sono note solo 3, mentre di La Vacca (0,09 km?)

e di Serpentara (0,29 km?) se ne conoscono ben 8. È probabile che

siano intervenute variabili nei tempi o nelle condizioni di raccolta, al

momento non valutabili, che hanno reso contraddittori alcuni risultati.

La distanza dall’isola madre sembra non avere influito sulla com-

posizione qualitativa e neppure quantitativa dei singoli popolamenti

microinsulari: basti per tutti ancora l’esempio di Santo Stefano, che

dista dalle coste sarde solo 1 km, mentre La Vacca ne dista 12, Ser-

pentara 3,2 e addirittura Il Toro, di cui sono note 5 specie, 17,1 km.

Sei specie (su 27) sono presenti in almeno 10 delle 44 isole cir-

cumsarde indagate (tab. I): Geogarypus nigrimanus (nota di 30 isole),

Calocheiridius oliviert (25), Hysterochelifer tuberculatus (22), Roncus

abditus (21), Neobisium incertum (13) e Amblyolpium dollfusi (11). Tre

di queste (R. abditus, N. incertum e H. tuberculatus) hanno in Sardegna

valenza ecologica relativamente ampia nell’ambito delle formazioni

vegetazionali della macchia mediterranea; le altre tre, più stenoecie,

sono proprie della macchia bassa e della gariga. È lecito supporre che

anche altre specie stenoecie siano diffuse in buona parte delle pic-

cole isole circumsarde, ove presenti gli ambienti idonei. È il caso di

Garypus levantinus, specie alofila e termofila (HEURTAULT & VANNIER

1990) delle coste rocciose, citata di sole 5 isole, senz'altro presente

in quasi tutte le altre ma reperibile solo a seguito di ricerche mirate.

Non sono state rinvenute specie ubiquiste indicatrici di ambienti

degradati ed eccessivamente antropizzati, con l'eccezione di Chtho-

nius ischnocheles, limitatamente antropofila, nota solo dell’ Asinara e

della Maddalena.

NOTA

A lavoro ultimato e già in composizione ho avuto modo di reperire 1 £ e 1D di

Paraliochthonius singularis (Menozzi, 1924) sull'Isola di Caprera, Cala Napoletana,

24.IV.2008, in scogliera sotto pietre presso il mare (coll. G. Gardini). Specie alofila

N-mediterranea-orientale nota per l’Italia di Liguria, Arcipelago Toscano, Campa-

nia, Isole Eolie e Sardegna (Cagliari e Dorgali: Cala Gonone) (GARDINI 1994, 2000).

Non ancora citata di isole circumsarde.

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 649

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di Orosei: nuovi dati e loro significato paleoambientale - http://clima.casaccia.

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BACCETTI B., COBOLLI SBORDONI M. & Pocci R., 1989 - Ricerche zoologiche della

nave oceanografica “Minerva” (C. N. R.) sulle isole circumsarde. I. Introdu-

zione - Annali Mus. civ. St. nat. “G. Doria”, Genova, 87: 127-136.

BrieR M., 1963a - Ordnung Pseudoscorpionidea (Afterskorpione) - Bestimmungs-

biicher zur Bodenfauna Europas, 1, Akademie-Verlag, Berlin, 313 pp.

BEIER M., 1963b - Pseudoskorpione aus dem Museum ,, Enrico Caffi“ in Bergamo -

Rend. Ist. lomb. Sc. Lett., Milano, 97 B: 147-156.

BEIER M., 1966 - Ein neuer Pselaphochernes (Pseudoscorp.) aus Sùditalien - Fragm.

ent., Roma, 4: 109-111.

CALLAINI G., 1983a - Notulae Chernetologicae. XI. Il sottogenere Ephippiochthonius

in Sardegna (Arachnida, Pseudoscorpionida, Chthoniidae) - Annali Mus. civ.

St. nat. “G. Doria”, Genova, 84: 401-423.

CALLAINI G., 1983b - Notulae Chernetologicae. XII. Nuovi reperti sugli Pseu-

doscorpioni della Sardegna - Lavori Soc. it. Biogeogr., Forlì, n.s., 8 (1980):

279-322.

ELLINGSEN E., 1909 - Contribution to the knowledge of the Pseudoscorpions from

the material belonging to the Museo Civico in Genova - Annali Mus. civ. St.

nat. Genova, 44: 205-220.

GARDINI G., 1975 - Pseudoscorpioni dell’Isola di Capraia (Arcipelago Toscano) -

Lavori Soc. it. Biogeogr., Forlì, n.s., 5 (1974): 385-396.

GARDINI G., 1980 - Identità di Chthonius tetrachelatus fuscimanus Simon, 1900 e

ridescrizione di C. (E.) nanus Beier, 1953 (Pseudoscorpionida Chthoniidae)

(Pseudoscorpioni d’Italia. XI) - Annali Mus. civ. St. nat. “G. Doria”, Genova,

88: 261-270.

GARDINI G., 1981 - Roncus caralitanus n.sp. della Sardegna meridionale (Pseudo-

scorpionida Neobisiidae) (Pseudoscorpioni d’Italia. XIII) - Boll. Soc. ent. it.,

Genova, 113: 129-135.

GARDINI G., 1993 - Chthonius (E.) poseidonis (Pseudoscorpionida, Chthoniidae)

nuova specie cavernicola della Nurra (Sardegna nord-occidentale) (Pseudoscor-

pioni d’Italia. XXVI) - Boll. Soc. sarda Sc. nat., Sassari, 29: 75-83.

GARDINI G., 1994 - I generi Paraliochthonius Beier, 1956 e Spelyngochthonius Beier,

1955 in Italia (Pseudoscorpionida, Chthoniidae) (Pseudoscorpioni d’Italia

XXIX) - Fragm. ent., Roma, 26: 1-10.

GARDINI G., 2000 - Catalogo degli Pseudoscorpioni d’Italia (Arachnida) (Pseudo-

scorpioni d’Italia XXXIV) - Fragm. ent., Roma, 32 (suppl.): 1-181.

GARDINI G., 2004 - Note faunistiche e tassonomiche sugli Pseudoscorpioni (Arach-

nida) epigei. (Pseudoscorpioni d’Italia XXXV) - In: Latella L. (ed.). Il Monte

Pastello. Mem. Mus. civ. St. nat. Verona - 2*. Serie, Monografie Naturalistiche,

15 4232129,

GARDINI G., in stampa - Pseudoscorpioni d’Italia XLII. Chthoniidae cavernicoli di

Sardegna (Pseudoscorpiones) - Mem. Soc. ent. it., Genova, 87

GARDINI G. & RIZZERIO R., 1985 - Materiali per una revisione del genere Roncus L.

650 TRA Ur ee G. GARDINI

Koch, 1873. I. Ridescrizione dei tipi di alcune specie italiane non cavernicole

(Pseudoscorpionida, Neobisiidae) - Fragm. ent., Roma, 18: 47-79.

Gestro R., 1904 - Una gita in Sardegna. Divagazioni biogeografiche - Boll. Soc.

geogr. it., Roma, (4) 5 (4): 1-39.

Grarirtti G. & Muceppa M., 1996 - Le grotte dell'Isola di Tavolara e la loro fauna

- Biogeographia, Siena, 18 (1995): 51-62.

Grafitti G., LACANA G., Loru R. & Muceppa M., 1996 - Trent'anni del Gruppo

Speleologico Sassarese - Boll. Gruppo Spel. Sass., Sassari, 16: 60-68.

Harvey M. S., 1991 - Catalogue of the Pseudoscorpionida - Manchester University

Press, Manchester & New York, 726 pp.

HeuRTAULT J., 1979 - Complément a la description de Olpium pallipes Lucas, 1845, type

de la famine Olpiidae (Arachnides, Pseudoscorpions) - Rev. suisse Zool., Genève,

86: 925-931,

HeurtauLT J., 1980 - Quelques remarques:sur les:espéces francgaises du genre Rha-

cochelifer Beier (Arachnides, Pseudoscorpions, Cheliferidae) - Bull. Mus. natn.

Hist. nat., Paris, Sect. A, (4) 2: 161-173,

HeurtauLT J., 1982 - Présence et signification dans la France méditerranéenne hide

espéces del genres Beterochelifer, Cheirochelifer et Calocheiridius (Arachnides,

Pseudoscorpions) - Atti Soc. tosc. Sc. nat., Mem., Pisa, (B) 88 (1981)(suppl.):

209-222.

HeurrauLT J. & VANNIER G., 1990 - Thermorésistance chez deux /Pseudoscorpions

(Garypidae), l’un du désert de Namibie, l’autre de la région de ‘Génes (Italie)

- Acta zool. fenn., Helsinki, 190: 165171. |

LAZZERONI G., 1970 - Ricerche sugli Pséudoscorpioni. V. L’Isola di Giannutri - Atti

Soc. tosc. Sc. nat., Mem., Pisa, (B) 76: 101-112.

MAHNERT V., 1977 - Uber einige ‘Atemnidae und Cheliferidae Griechenlands (Pseu-

doscorpiones) - Mitt. schwéiz. ent. Ges., Zirich, 50: 67-74.

SCHAWALLER W., 1981 - Pseudoskorpione von Korsika (Arachnida :Pseudoscorpioni-

dea) - Ent. basil., Basel, 6: 42-51.

Vigna TAGLIANTI A., Aupisio P. A., Bionpi M., BoLocna M. A., CARPANETO

G. M., De Biase A,, Fatrorini S., PIATTELLA E., SINDACO R., ‘VENCHI A.

& ZappParoLI M., 1999 - A proposal for chorotype classification of’ the Near

East fauna, in the framework of the Western Palearctic tegion--: Biogeographia,

Siena, 20: 31-59.

RIASSUNTO

Sono illustrati i risultati delle ricerche zoologiche della nave oceanografica

“Minerva” del C. N. R. effettuate sulle isole circumsarde tra il 1986 e il 1994. Su

44 isole sono state rinvenute 27 specie di Pseudoscorpioni, cioè il 37,5% dell’intero

popolamento sardo, che è costituito attualmente da 72 specie, di cui 18 cavernicole

e 28 endemiche. Delle specie presenti nelle isole circumsarde, 24 sono conosciute

anche dell’isola maggiore. Le specie a corologia mediterranea s. l. e tirrenica sono

14, 5 sono endemiche, 4 euro-mediterranee, 1 turano-mediterranea, 1 centroasia-

tico-mediterranea e 2 con corotipo non definibile, È analizzato il popolamento in

relazione alla superficie delle isole circumsarde e alla conseguente diversità ambien-

tale e si sottolinea il fatto che non sempre vi è correlazione diretta tra superficie e

PSEUDOSCORPIONI DELLE ISOLE CIRCUMSARDE 651

numero di specie; anche la distanza dall’isola madre non sembra avere influito sulla

composizione quantitativa e qualitativa dei singoli popolamenti microinsulari. Sono

elencate le specie più diffuse nelle isole circumsarde e si accenna alle loro esigenze

ecologiche. Garypus beauvoisi (Savigny & Audouin, 1826) è citata per la prima volta

di Toscana e di Sicilia, Amblyolpium dollfust Simon, 1898 è nuova per la Sicilia e

Rhacochelifer corcyrensis (Beier, 1930) per la Sardegna.

ABSTRACT

Zoological researches of the oceanographic ship “Minerva” (C. N. R.) on the

circumsardinian islands. XX XI. Pseudoscorpiones.

‘Twenty-seven species of Pseudoscorpions have been found in 44 small islands

around Sardinia, i.e. 37,5% of the whole Sardinian population, which consists of 72

species (18 are troglobitic and 28 endemic). Between them, twenty-four species also

occur in Sardinia.

A chorological analysis reveals that Mediterranean (‘Tyrrhenian included) species

dominate over the other patterns of distribution: i. e. 14 species are Mediterranean

s. 1 and Tyrrhenian, 5 endemic, 4 Europeo-Mediterranean, 1 Turano-Mediterra-

nean, 1 Centralasiatic-Mediterranean and 2 with indefinite chorotype. The number

of species of each island is not always according to the species/area relationship, and

the size of the area is not the only parameter to explain species abundance.

Garypus beauvoist (Savigny & Audouin, 1826) is firstly recorded from ‘Tuscany

and Sicily, Amblyolpium dollfusi Simon, 1898 from Sicily and Rhacochelifer corcyren-

sis (Beier, 1930) from Sardinia.

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653

KRISTOFER M. HELGEN*, TANYA LEARY**,

GIULIANO Dorta*** & GIOVANNI AMORI****

CATALOGUE OF MELANESIAN RODENTS IN THE

MUSEUM OF GENOVA

(MAMMALIA, RODENTIA)

INTRODUCTION

Rodent specimens from Melanesia (New Guinea and surround-

ing islands) have been stored at the Museo Civico di Storia Naturale

“Giacomo Doria” in Genova since the latter decades of the nine-

teenth century. The main body of this collection (206 samples) is

represented by 152 specimens collected by the Italian explorer and

collector Lamberto Loria (1855-1913) during his pioneering bio-

logical and anthropological survey efforts between 1889 and 1896

in south-eastern New Guinea (the modern day Milne Bay, Central

Provinces and National Capital District of Papua New Guinea).

Earlier specimens derive from the collections of the Italian natural-

ist-explorers Odoardo Beccari (1843-1920), 18 specimens from north-

western New Guinea, and Luigi Maria D'Albertis (1841-1901), 34

specimens from the Fly River in south-central and Mount Arfak in

the Vogelkop Peninsula of north-western New Guinea (D’ALBERTIS

1880; GesTRro 1928; VAN STEENIS KRUSEMAN 1950; CAPOCACCIA &

Pocci 1982). A single specimen was collected by the Dutch collector

A. A. Bruijn in 1875 (PETERS & Doria 1876) on the island of Salawati

off north-western New Guinea, and another by the British explorer

* Division of Mammals, National Museum of Natural History, MRC 108,

Room 390, Smithsonian Institution, Washington D.C., 20013-7012, USA

(helgenk@si.edu)

** Parks and Wildlife Division, Department of Environment and Climate Change,

P.O. Box 95, Parramatta, NSW 2124, Australia

*** Museo Civico di Storia Naturale “G. Doria”, Via Brigata Liguria 9, 16121

Genova, Italy

**** CNR — Institute of Ecosystem Studies, Via A. Borelli 50, 00161 Roma, Italy

654 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

C. M. Woodford in 1889 in the Solomon Islands (WOODFORD 1890).

Though modest with respect to the number of specimens that

it holds, the Museum of Genova represents one of the most impor-

tant collections of New Guinea mammals (PETERS & Doria 1876,

1881; THomas 1897; FLANNERY 1995a). The collection of Melane-

sian rodents is important both for its scientific and historical value.

It contains, for instance, the first specimens of three endemic New

Guinean murine genera to be deposited in a museum (Leptomys

Thomas, 1897, Paramelomys Rummler, 1936 and Pogonomelomys

Rummler, 1936); it also houses a good number of type specimens

(as identified below) as well as exemplars of several species very rare

in collections, such as the large endemic insular murines Uromys rex

(of Guadalcanal in the Solomon Archipelago) and Uromys siebersi

(of the Kai Archipelago of Indonesia).

Previous overviews of Melanesian rodent holdings in Genova

include the publications of PETERS (1874) and PETERS & DORIA (1876,

1881), who reviewed the Melanesian material gathered by Beccari,

D'Albertis and Bruijn (including new species of monotremes, mar-

supials, rodents and bats), and that of THomas (1897), who provided

an overview of mammal specimens later obtained by Loria. During

the twentieth century, a number of rodent taxonomists discussed

particular Melanesian specimens in the collection (RUMMLER 1938;

TATE 1951; DENNIS & MENZIES 1979; TavLOR et al. 1982; MEN-

zies 1989, 1990, 1996; Groves & FLANNERY 1994; FLANNERY 1995a)

and Genova specimens continue to influence important works in

New Guinea murine systematics today (MUSSER & CARLETON 2005;

HELGEN 2007; MUSSER et al. in press).

The purpose of this catalogue is to bring together in one place

a summary of information on these holdings and to bring up to date

the identifications of the museum’s material according to current

taxonomy and nomenclature (e.g. MussER & CARLETON 2005),

TYPE SPECIMENS

Amongst the Melanesian rodent holdings at the Museum of

Genova are a number of type specimens, including six holotypes,

one lectotype, one syntype, paralectotypes for five taxa, and four

paratypes (terms used according to INTERNATIONAL COMMISSION ON

ZOOLOGICAL NOMENCLATURE 1999), Descriptions of new taxa based

on Genova material were published by two sets of authors: Wilhelm

MELANESIAN RODENTS 655

Peters (curator of the Museum of Berlin), both alone and with Giacomo

Doria (Director of the Museum of Genova) as co-author, and Michael

Rogers Oldfield Thomas of the British Museum (Natural History)

in London (BMNH; today the Natural History Museum, London).

For several taxa named by Peters based on material collected by

Beccari, D'Albertis, and Bruijn, syntype series were split between

the Museum of Genova and the Museum fiir Naturkunde in Berlin

(ZMB). For those named by Oldfield Thomas based on material

collected by Loria, for all taxa except Leptomys elegans, the unique

example of which was retained in Genova, two designated syn-

types were in each case automatically divided between London and

Genova, an arrangement presumably both convenient to Thomas

and deferential to Genova. 'THomas (1897: 607) wrote:

Of each of the new species two “co-types’ [i.e. syntypes] have been specially selected

for description, one of these being permanently preserved in the Genoa and the

other in the British Museum. The remaining members of the series: would offcourse

be simple paratypes. Leptomys elegans is alone represented by a single example.

For most of these divided syntype series, a lectotype was selected

by subsequent revisers, either presumably or explicitly for conve-

nience on the part of future taxonomic workers (RUMMLER 1938;

TATE 1951; TAYLOR et al. 1982; MENZIES 1989; Musser & CARLE-

TON 2005). In all but a single case (that of Hydromys beccanii, see

below), the lectotype was selected at BMNH or ZMB rather than

at Genova. To the best of our knowledge, only one specimen at

Genova remains a syntype - the original “co-type” of Pogonomys

loriae Thomas, 1897, for which no lectotype has been chosen (the

other syntype remains at BMNH; see below).

Finally, we note that other (non-type) mammal specimens

secured by Loria have been traded amongst many major European

museums (including London, Dresden and Oslo among others), such

that the great majority but by no means all of the rodent specimens

collected during Loria’s efforts are preserved in Genova.

Holotypes

MSNG 32988, holotype of Leptomys elegans Thomas, 1897 (Figs 1, 2),

MSNG 3534, holotype of Mus beccarii Peters & Doria, 1881 (not Mus

beccarii Jentink, 1880) and Mus doriae Trouessart, 1897 (currently

regarded as synonyms of Rattus rattus (Linnaeus, 1758); see below),

656 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

MSNG 3491, holotype of Mus mollipilosus Peters & Doria, 1881

(now classified as Pogonomys mollipilosus (Peters & Doria, 1881); see

below) (Figs 3, 4).

MSNG 3460, holotype of Mus ringens Peters & Doria, 1881 (cur-

rently regarded as a synonym of Rattus leucopus (Gray, 1867); see

below).

MSNG 3677, holotype of Uromys bruijni Peters & Doria, 1876

(now classified as Pogonomelomys bruijnit (Peters & Doria, 1876); see

below) (Figs 5, 6).

MSNG 3248, holotype of Uromys validus Peters & Doria, 1881

(currently regarded as a synonym of Uromys caudimaculatus (Krefft,

1867); see below).

Lectotypes

MSNG 3637, lectotype of Hydromys beccari Peters, 1874 (currently

regarded as a synonym of Hydromys chrysogaster E. Geoffroy, 1804;

see below), selected by TATE (1951: 235) (Figs 7, 8).

Syntypes

MSNG 3492, syntype of Pogonomys loriae Thomas, 1897 (the other

syntype is apparently BMNH 97.8.7.47; see TATE 1951: 280).

Paralectotypes

MSNG 3501, paralectotype of Mus gestri Thomas, 1897 (currently

regarded as a synonym of Rattus sordidus (Gould, 1858); see below);

lectotype (BMNH 97.8.7.36) selected by RUMMLER (1938: 208); see

also TAYLOR et al. (1982: 269).

MSNG 3495 (b), paralectotype of Pogonomys lamia 'Thomas, 1897

(now classified as Chiruromys lamia (Thomas, 1897); see below); lec-

totype (BMNH 97.8.7.65) selected by RUMMLER (1938: 73).

MSNG 3600 (a), paralectotype of Uromys levipes Thomas, 1897

(now classified as Paramelomys levipes (Thomas, 1897); see below);

lectotype (BMNH 97.8.7.72) selected by RUMMLER (1938: 139); see

also MENZIES (1989) and Musser & CARLETON (2005).

MELANESIAN RODENTS 657

Fig. 1 - MSNG 32988, holotype of Leptomys elegans Thomas, 1897; dorsal and

ventral view of the skin.

Fig. 2 - MSNG 32988,

holotype of Leptomys elegans

Thomas, 1897; skull.

658 K.M, HELGEN, T. LEARY, G. DORIA & G, AMORI

Fig. 3 - MSNG 3491, holotype of Mus mollipilosus Peters & Doria, 1881 (now clas-

sified as Pogonomys mollipilosus (Peters & Doria, 1881)); dorsal and ventral

view of the skin,

Fig. 4 - MSNG 3491, holotype

of Mus mollipilosus Peters

dl & Doria, 1881 (now clas-

sified as Pogonomys mol-

lipilosus (Peters & Doria,

1881)); skull; only the

left part of the mandible

is present.

MELANESIAN RODENTS 659

Fig. 5 - MSNG 3677, holotype of Uromys bruijnii Peters & Doria, 1876 (now clas-

sified as Pogonomelomys bruijnii (Peters & Doria, 1876)); dorsal and ventral

view of the skin.

Fig, 6 - MSNG 3677, holotype of Uromys

bruijnit Peters & Doria, 1876 (now

on or

classified as Pogonomelomys bruijni

(Peters & Doria, 1876)); skull,

660 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

Fig. 7 - MSNG 3637, lectotype of Hydromys beccarit Peters, 1874 (currently

regarded as a synonym of Hydromys chrysogaster E. Geoffroy, 1804); dorsal

and ventral view of the skin.

Fig. 8 - MSNG _ 3637, lectotype. of

Hydromys beccarit Peters, 1874

(currently regarded as a syn-

onym of Hydromys chrysogaster

E. Geoffroy, 1804); skull.

MELANESIAN RODENTS 661

MSNG 3474 (a-d), four paralectotypes of Mus albertist Peters &

Doria, 1881 (currently regarded as a synonym of Mus musculus Lin-

naeus, 1758; see below); lectotype (ZMB 5423) selected by RÙM-

MLER (1938: 218).

MSNG 3257-57, paralectotype of Pogonomys lepidus "Thomas, 1897

(currently regarded as a synonym of Pogonomys macrourus (Milne-

Edwards, 1877); see below), lectotype (BMNH 97.8.7.51) selected

by TATE (1951: 279).

GAZETTEER

To maximize the usefulness of our catalogue, we have compiled

a gazetteer of collecting localities mentioned in the pages below (Fig.

9). Some localities, such as Port Moresby and Yule Island in south-

eastern Papua New Guinea, and Humboldt Bay (near Jayapura),

Salawati, and Sorong in West Papua refer to well known histori-

cal place names in the region and need no detailed explanation to

students of the region. Other place names refer to lesser known vil-

140°0'0"E 150°0'0"E 160°0'0"E

0 130 260 520 780 1,040

——__ ee Kilometers

0°0'0”

0°0'0"

3 Sorong PAS n°

a Ni ecs Hatam SIN

‘Ara N = tb

» we Fa poe <

Salawati ls. }< NG =r, TN

2 È To a LT È > a

4 Box ~

Via

\3pf \ x are, he nggnlia Bay

angle ba

zi lan } =

eter 1\ 7 }

me LE N (OS

JEllangowan Is >q_ | De

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<hr

10°0'0"S

ara Bara

~—

140°0'0"E 150°0'0"E 160°0'0"E

Fig. 9 - Map of collecting localities.

662 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

lages or collecting camps, and our understanding of the situation of

these localities is clarified and briefly discussed. For locality details

we have depended especially on information provided by ‘THOMAS

(1897), TATE (1951), LAURIE & HitL (1954), TAYLOR et al. (1982)

and Bonaccorso (1998).

INDONESIA

Maluku Province: Weri, Kai Bandan (Kai Besar), Kai Islands

(Beccari collecting locality) The Kai (or Kei) Islands are an oce-

anic archipelago in the south-eastern Moluccas, lying just off the

bathymetric contours of the Australo-Papuan continental shelf, near

the Aru Islands. We have been unable to locate this specific local-

ity (Weri) within the archipelago. BECCARI (1924: 239-241) wrote

that “Keibandan” is a village lying in a bay placed almost in the

middle of the western coast of the Great Kei (the island of Kai

Besar) and that, on August 1st [1873], he reached a place [probably

Weri] with a forest at about 7-8 miles from Keibandan by a ship; he

spent there some days and, on August 9th, he received a “Kedir”, a

rodent living underground in small holes [probably Hydromys bec-

cart], by indigenous collectors.

Maluku Province: Wokam (05°44’S, 134°11’E), Aru Islands, < 250

m above sea level

(Beccari collecting locality) Wokam is the northernmost of the

large islands in the Aru Archipelago, a land-bridge group of islands

between New Guinea and the Australian continent. The islands are

low-lying, with maximum elevation less than 250 m above sea level.

Papua Province: Humboldt Bay (02°30’S, 140°50’E), near sea level

(Beccari collecting locality) Humboldt Bay is the harbour associ-

ated with the port city of Jayapura, formerly Hollandia, the modern

capital of Papua Province.

West Irian Jaya Province: Hatam (01°14’S, 134°01’E), circa

1000-1500 m above sea level

(Beccari and D’Albertis collecting locality) Hatam is a village

on Mt. Arfak in the Arfak Mountains of the Vogelkop (“Bird’s

MELANESIAN RODENTS 663

Head”) Peninsula of northwestern New Guinea. According to WAL-

LACE (1880), Hatam is “a village on Mount Arfak, about 3500 feet

above the sea”. D’ALBERTIS (1879) referred to the locality as “Mt.

Hatam” and gave the maximum elevation of the mountain as 5000

feet. TAYLOR et al. (1982: 327) gave the elevation of Hatam as “1200

m” and its coordinates as “01°10’S, 133°40’E”.

West Irian Jaya Province: Salawati Island (01°05’S, 130°53’E), <

700 m above sea level

(Bruijn collecting locality) Salawati (= Salawatti or Salwatti) is

a medium sized (circa 1600 km?), low-lying (0-700 m) land-bridge

island in the Raja Ampat group off the northwestern tip of the

Vogelkop Peninsula (see D’ALBERTIS 1879).

West Irian Jaya Province: Sorong (00°53’S, 131°15’E), near sea level

(Beccari and D'Albertis collecting locality) The town of Sorong

is located on the north-western coast of the Vogelkop Peninsula (see

‘D’ALBERTIS 1879). This locality, often named or labelled “Sorong

Island” in older literature or on older specimens, may refer either to

the town of Sorong itself or to Pulau Doom, a small island off the

coast of Sorong which was an important historical centre of trading

in the region (T. Flannery, n litt.).

PAPUA NEW GUINEA

Central Province: Aroma (10°08’S, 147°59’E), near sea level

(Loria collecting locality) “... Aroma, still further east (10.8 S.

147.59 E.) [of Kapa Kapa and Cape Hood]” (THomas 1897: 607).

LAURIE & HILL (1954: 140) gave the same coordinates of the modern

settlement of Aroma (= Aroma Coast) (10°08’S, 147°59’E).

Central Province: Epa (08°44’S, 146°43°E)

(D’Albertis collecting locality) LAURIE & HILL (1954: 144) gave

‘the coordinates: of Epaas “08°44’S, 146°43’E.”

Central District: ‘Gerekanumu (09°31’S, 147°22’E), circa 0-200 m

above sea level

664 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

(Loria collecting locality) “Gerekanumu, on the southern slope

of the Astrolabe Range (09.31 S. 147.22 E.)” (THomas 1897: 607).

LAURIE & Hutt (1954: 145) and TAYLOR et al. (1982: 322) quoted

the same coordinates for this locality (as “Gerekanamu”). TAYLOR et

al. (1982: 322) gave the elevation at Gerekanamu as “200 m.” The

modern settlement of Gereka is situated at 09°30’S, 147°17°E, near

sea level.

Central Province: Haveri (09°25’S 147°35’E), circa 700 m above sea level

(Loria collecting locality) “Haveri (09.25 S. 147.35 E.) ... among

the mountains behind the Astrolabe Range, near Mt. Wori-Wori”

(THomas 1897: 607). TAYLOR et al. (1982: 321) provided these same

coordinates, but LAURIE & HILL (1954: 145) and later Bonaccorso

(1998: 431) associated this collecting locality with the coordinates

‘09°25’5S, 147°22' E”. TAYLOR et al. (1982: 321) gave the elevation

of Haveri as “750 m”, though most of Loria’s specimen labels quote

the elevation as “700 m”. Tags on the Genova specimens identify

this locality with the “Bartholomew Range”. “Mt. Wori-Wori”

is today referenced as “Vori Vori Bluff” (A. ALLISON, in litt.).

Central Province: Hula (10°06’S, 147°43’E), near sea level

(Loria collecting locality) “Hula, close to the mouth of the

[Kemp Welch] river on the promontory that ends in Cape Hood

(10.10 S. 147.44 E)” (THomas 1897: 607). Some specimen labels

give this locality as “Hula, Hood Point”. The modern settlement of

Hula is situated at 10°06°S, 147°43’E.

Central Province: Ighibirei (09°43’S, 147°45’E), elevation unknown

(Loria collecting locality) “Ighibirei, on the Kemp Welch river

some little way inland” (THOMAS 1897: 607); “just inland from the

mouth of the Wanigela River” (LauRIE & HILL 1954: 146). PUTHZ

(1971: 453) at first was unable to locate this locality, but later iden-

tified it as Boku (09°43’S, 147°45°E) on the basis of a handmade

map (PUTHZ 1982: 121). Bonaccorso (1998: 431) gave the coor-

dinates of “Ighibierei” as “10°02°S, 147°46’° E”. Only Chiruromys

lamia (Thomas, 1897), a mouse only known from montane forests

(recorded altitudes from 1200-2300 m; FLANNERY 1995a) is repre-

sented amongst Loria’s collections from Ighibirei at Genova—an

MELANESIAN RODENTS 665

indication that this collection site was probably situated within this

same altitudinal range.

Central Province: Inawi (08°36’S, 146°33’E), near sea level

(Loria collecting locality) “Inawi, on the S. Giuseppe river,

which runs into Hall Sound” (THomas 1897: 607). Laurie & HILL

(1954: 140) gave the coordinates for “Inawi” (= Inawa) as “08°31’S,

146°35’E”. The modern settlement of Inawi (also spelled Inawae) is

situated at 08°36’S, 146°33’E.

Central Province: Irupara (10°05’S, 147°43’E), near sea level

(Loria collecting locality) “... Irupara ... close to the mouth of

the [Kemp Welch] river on the promontory that ends in Cape Hood

(10.10 S. 147.44 E)” (THomas 1897: 607); “near Kamali” (LAURIE

& Hitt 1954: 146). The modern settlement of Irupara is situated at

10°05°S, 147°43°E.

Central Province: Kamali (10°03’S, 147°45’E), near sea level

(Loria collecting locality) “Kamali ... close to the mouth of the

[Kemp Welch] river on the promontory that ends in Cape Hood

(10.10 S. 147.44 E)” (THomas 1897: 607). Bonaccorso (1998: 432)

gave the same coordinates quoted for Cape Hood (i.e. 10°10’S,

147°44’E) as the position of Kamali. Some specimen labels give this

locality as “Hood Bay, Kamali”. LAURIE & HiLL (1954: 147) gave

the coordinates of Kamali as “10°02’S, 147°45’E”, and the modern

settlement of Kamali is situated at 10°03’S, 147°45’E.

Central Province: Kapa Kapa (09°50’S, 147°30’E), near sea level

(Loria collecting locality) “Kapa Kapa (9.50 S. 147.30 E.) ...

a little further east [of Cape Hood] ... on the sea coast” (THOMAS

1897: 607); at the “mouth of the Kemp Welch River” (Tate 1951:

420). LAURIE & HiLL (1954: 147) quoted these same coordinates for

Kapa Kapa (i.e. 09°50’S, 147°30° E). Bonaccorso (1998: 432) gave

the coordinates for Kapa Kapa as “09°48°S, 147°30’ E”.

Central Province: Moroka (09°25’S, 147°41’E), circa 1300 m above

sea level

(Loria collecting locality) “Moroka (09.25 S. 147.41 E.), among

the mountains behind the Astrolabe Range, near Mt. Wori-Wori”

666 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

(THOMAS 1897: 607); at the “headwaters of the Musgrave River”

(TATE 1940: 2). LAURIE & HILL (1954: 151) gave the coordinates of

this locality as “09°24’S, 147°32’E”. Loria’s specimen labels quote

the elevation of Moroka as “1300 m.”. ‘Tags on the Genova specimens

identify this locality with the “Bartholomew Range”. “Mt. Wori-

Wori” is today referenced as “Vori Vori Bluff” (A. ALLISON, in litt.).

Central Province: Yule Island (08°49’S, 146°32’E), near sea level

(D’Albertis collecting locality) Yule is a small, low-lying land

bridge island near the modern settlement of Poukama. Between

Yule and the mainland is Hall Sound, long an important harbour in

Papua New Guinea.

Milne Bay Province: Bara Bara (10°19’S, 150°41’E), near sea level

(Loria collecting locality) “... one locality, Bara Bara, is oppo-

site Killerton Island in Milne Bay at the extreme S.E. corner of

New Guinea” (THomas 1897: 607). Bonaccorso (1998: 428) gave

the coordinates for this locality (erroneously, in our assessment) as

09°07°S, 149°19°E. Laurie & Hic (1954: 140) gave coordinates

for the “Bara Bara River” on mainland New Guinea as 10°19’S,

150°41’E, and Killerton Island is located at 10°21’S, 150°40’E. The

modern settlement of Barabara is now located on Normanby Island

in the D’Entrecasteaux Archipelago and is situated at 10°08’S,

151°07'E. As a result, MCDOWELL (1975) erroneously associated this

collecting locality with Normanby Island.

National Capital District: Port Moresby (09°28’S, 147°08’E)

Port Moresby is the capital and largest city in the nation of

Papua New Guinea.

Two specimens are labelled “Port Moresby, D’Albertis” but

L. M. D’Albertis never visited this locality; these specimens were

instead probably collected by L. Loria.

Western Province: Fly River (southwestern region)

(D’Albertis collecting locality) D’Albertis made three expe-

ditions up the Fly River in 1875, 1876 and 1877, and collected

extensively along the river (particularly birds and plants), although

MELANESIAN RODENTS 667

exact localities are unclear. On his first journey he reached as far

as Ellangowan Island (7°49’S 141°41’E). On his second expedition

up the Fly River he claimed to have reached 05°30’S before being

forced back by low waters and lack of supplies, but Sir William

MacGregor who later explored the Fly, doubted that he had passed

06°11’S (GIBBNEY 1972). His final expedition was beset with prob-

lems, and he did not reach as far as he had on his second expedition.

Western Province: Katau (09°00’S, 143°00’E)

(D’Albertis collecting locality) Katau lies in the south-eastern

TransFly area to the west of the Fly River, and D’Albertis visited

this locality on each of his three expeditions up the Fly River: “at

anchor here in the little river Kataw, in sight of the houses of Moatta

(= Mawatta), which are 800 or 1000 yards from us” (D’ALBERTIS

1880: 162). He had an extended stay there (August 7° to Novem-

ber 2"? 1876) due to bad weather on his second expedition and in

this period probably the majority of his specimens were collected.

He also purchased animals from people from surrounding villages,

including the village of Matzingare. This river is also known as the

Binaturi River (LOVERIDGE 1948).

SOLOMON ISLANDS

Guadalcanal Province: “Guadalcanar” (= Guadalcanal), altitude

unknown

(Woodford collecting locality) One specimen of Uromys rex at

Genova was collected in Guadalcanal by Woodford; the specimen is

labelled “Guadalcanar” which is the archaic name of Guadalcanal.

Other native murids collected by Woodford on Guadalcanal were all

taken at Aola (09°32’S, 160°29’E), on the north-eastern coast of the

island (THOMAS 1888).

FORMAT

‘The species are listed in alphabetical order according to genus

and species. For each record we give the following information:

Catalogue number of the Museum of Genova (MSNG); for

668 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

numerous series the number is followed by a letter, a number or

a roman number in brackets, or a number after a line. In some

cases, specimens from other museums are discussed, under the fol-

lowing museum abbreviations: BMNH (Natural History Museum,

London); ZMB (Museum ftir Naturkunde, Humboldt Universitat,

Berlin).

Locality of collection: country, province, and geographical local-

ity and, when clearly indicated on the original label, altitude above

sea level. Refer to the gazetteer (above) for further details on some

collection localities.

Details of collection: the collecting date is given in arabic numer-

als (day), roman numerals (month) and arabic numerals (year), for

instance “3.VI.1873”. The collector is indicated with the first letter

of the first (Christian) name (and sometimes of the middle name)

and the entire surname, e.g. “L. Loria” or “L. M. D'Albertis”.

Nature of specimen: number of specimens, sex and stage of

development are given: 6 = male, 2 = female, juv. = young. Type

specimens have been identified above and in the accounts that

follow. Specimens are preserved as one of the following prepara-

tions: skin; skull; skin and skull; or alcohol (preserved in ethanol);

for some specimens preserved in alcohol the skull has been prepared

to better identify the species. We note that many of the skins in the

collection were prepared in 1937 from carcasses stored in fluid, long

after their date of collection.

Order RODENTIA Bowdich, 1821

Family MuRIDAE Illiger, 1811

Subfamily MuURINAE Illiger, 1811

Chiruromys forbesi Thomas, 1888

MSNG 3258 (a-c). Papua New Guinea, Milne Bay Province, Bara

Bara, I-11.1890, L. Loria; 1 3 (b), 1 d juv. (a), 1 2 (c), skin and skull.

MSNG 3521. Papua New Guinea, Milne Bay Province, Bara Bara,

I-II1.1890, L. Loria; 1 d juv., skin and skull.

MELANESIAN RODENTS 669

MSNG 3521 (a-d). Papua New Guinea, Milne Bay Province, Bara

Bara, I-I1I.1890, L. Loria; 2 dd (b and d), 2 22 (a and c), skin and skull.

MSNG 52759. Papua New Guinea, Milne Bay Province, Bara Bara,

I-II.1890, L. Loria; 1 9, alcohol and skull.

Remarks. The type series of Pogonomys forbesi vulturnus

Thomas, 1920 (a synonym of Chiruromys forbest) are specimens

from this same series, collected by Loria at Bara Bara in 1890, but

sent to Oldfield Thomas in London (see ‘THOMAS 1920).

Chiruromys lamia (Thomas, 1897)

MSNG 3495 (a). Papua New Guinea, Central Province, Ighibirei,

1890, L. Loria; 1 d, skin and skull, paratype of Pogonomys lamia

Thomas, 1897.

MSNG 3495 (b). Papua New Guinea, Central Province, Ighibirei,

1890, L. Loria; 1 ®, skin and skull, paralectotype of Pogonomys

lamia Thomas, 1897.

Remarks. THomas (1897) originally described this spe-

cies based on seven specimens collected by Loria from Ighibirei

(designated as the type locality) and Haveri. Of the five specimens

from Ighibirei, Thomas identified two as “cotypes” (= syntypes—

apparently the specimens now designated as BMNH 97.8.7.65 and

MSNG 3495 (b)). We accept RUMMLER’S (1938: 73) reference to the

BMNH specimen as “the type” (i.e. “der Typus”) as a lectotype

designation for lamia. MSNG 3495 (b) is thus a paralectotype.

Chiruromys vates (Thomas, 1908)

MSNG 54203. Papua New Guinea, Central Province, Bar-

tholomew Range, Haveri, VIII-X1I.1893, L. Loria; 1 9, skin and skull,

paratype of Pogonomys lamia "Thomas, 1897. Ex MSNG 3495 (a).

Remarks. This specimen formed part of Thomas’ original

hypodigm of Chiruromys lamia (see THomas 1897). However, we now

670 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

identify it instead as Chiruromys vates (Thomas, 1908), on account

of its richer coloration, smaller molars, shorter incisive foramina,

and less expanded zygomata relative to the type (and other) speci-

mens of C. lamia (cf. THomas 1908). Further, C. lamia is a montane

mouse (firmly identified specimens are known only from elevations

at and above 1200 m; FLANNERY 1995a), while C. vates is widely dis-

tributed in the lowlands of southern and southeastern New Guinea

(sea level to 1500 m; FLANNERY 1995a). Haveri, the site where this

specimen is collected, is apparently situated at approximately 700

m above sea level (see Gazetteer, above). Unfortunately, the first

and second authors made only cursory examination of this specimen

when visiting Genova, so we recommend that this identification be

revisited by future examiners.

Hydromys chrysogaster E. Geoffroy, 1804

MSNG 3637. Indonesia, Maluku Province, Kai Islands, Kai Bandan,

Weri, without collecting date, O. Beccari; 1 3, skin and skull, lec-

totype of Hydromys beccarit Peters, 1874. TATE (1951) gave errone-

ously the number 3627 for this specimen (Figs 7, 8).

MSNG 3663. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 £ juv., skin and skull.

Remarks. PETERS (1874) based his description of Hydromys

beccarii on two syntypes, one deposited in Genova (MSNG 3637),

the other in Berlin (ZMB 4805). TATE (1951: 191) acknowledged

the two syntypes before referring to the Genova specimen as the

“type” of beccarti (p. 235; though he gave the number as MSNG

3627), and we accept this as a lectotype designation.

Though Hydromys beccarit Peters, 1874 is usually regarded as a

synonym of the large and widespread species Hydromy® chrysogaster,

patterns of geographic variation in H. chrysogaster remain little stud-

ied (TATE 1951; HELGEN 2005), and as currently recognized, this

widespread species is likely to represent a complex of related species.

As the oldest name applied to a Melanesian population of Hydromys,

further study may show that the epithet beccari is the most appro-

priate name for a distinct species endemic to the Melanesian region.

MELANESIAN RODENTS 671

Leptomys elegans Thomas, 1897

MSNG 32988. Papua New Guinea, Central Province, Astrolabe

Range, without collecting date, L. Loria; 1 9, skin and skull, holo-

type of Leptomys elegans ‘Thomas, 1897 (Figs 1, 2).

Remarks. MUSSER et al. (in press) provided a comprehensive

overview of the taxonomy and biology of the genus Leptomys, the type

species of which is L. elegans. The original details of collection for the hol-

otype of elegans are not recorded. According to MUSSER et al. (in press):

The tag of the holotype bears no definite locality other than “British N.G.”,

and no exact provenance was published in the original description of L. elegans,

although ‘Thomas (1897: 607) wrote that Loria’s localities “are mostly between the

Owen Stanley Range and the sea, in or near the watershed of the Kemp Welch

river”, an area that would include the Astrolabe Range and adjacent Sogeri Plateau.

TATE (1951: 223) observed that “Loria collected in the Astrolabe Range behind Port

Moresby”, and we regard specimens from the Astrolabe Range (available at AMNH

and ZMB) as topotypical.

Melomys leucogaster (Jentink, 1908)

MSNG 3609 (a and b). Papua New Guinea, Central Province, Hood

Bay, Kamali, III.1891, L. Loria; 1 d (b), 1 2 (a), skin and skull.

MSNG 3616. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 dé, skin and skull.

MSNG 54204. Papua New Guinea, Central Province, Yule Island,

IV.1875, L. M. D‘Albertis; 1 2, skin and skull. Ex MSNG 3200 (2).

MSNG 54207. Papua New Guinea, Central Province, Yule Island,

1875, L. M. D‘Albertis; 1 6, skin and skull. Ex MSNG 3240-17.

Melomys lutillus (Thomas, 1913)

MSNG 3601. Papua New Guinea, Central Province, Astrolabe

Range, Gerekanumu, IJ.1893, L. Loria; 1 d juv., skin and skull.

MSNG 43956. Papua New Guinea, without collecting date, L. M.

D'Albertis; 1 specimen, alcohol.

672 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

MSNG 54206 (a and b). Papua New Guinea, Central Province,

Epa, IV.1875, L. M. D’Albertis; 1 dé (a) ex MSNG 3240-14, 1 9

(b) ex MSNG 3240-16, skin and skull.

MSNG 54212. Papua New Guinea, National Capital District, Port

Moresby, without collecting date, [L. M. D'Albertis in the label but

probably L. Loria; see Gazetteer, above]; 1 dé, skin and skull. Ex

MSNG 3507 (1).

Melomys rufescens (Alston, 1877)

MSNG 3240- (13, 15 and 18). Papua New Guinea, Western Prov-

ince, Fly River, 1875, L. M. D’Albertis; 2 66 (13 and 18), 1 2 (15),

skin and skull.

MSNG 3507 (2). Papua New Guinea, National Capital District, Port

Moresby, without collecting date, [L. M. D'Albertis in the label but

probably L. Loria; see Gazetteer, above]; 1 d, skin and skull.

MSNG 42139 (I and II). Papua New Guinea, Western Province,

Fly River, 1875, L. M. D’Albertis; 1 £ (1), 1 juv. (II), alcohol and

skull.

MSNG 42139 (III and IV). Papua New Guinea, Central Province,

Epa, 1875, L. M. D'Albertis; 2 juv., alcohol and skull.

MSNG 42139 (V). Papua New Guinea, without collecting date, L.

M. D'Albertis; 1 juv., alcohol and skull.

MSNG 54202. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 d juv., skin and skull.

Ex MSNG 3669.

MSNG 54205. Indonesia, West Irian Jaya Province, Mount Arfak,

Hatam, 1875, L. M. D'Albertis; 1 ®, skin and skull. Ex MSNG

3240-12.

MSNG 54209 (a and b). Papua New Guinea, Central Province,

Kapa Kapa, VILISS9, L. Loria; 1 6 (b), 1 & (a), skin. and. skull.

Exe MSNG;: 3250.

MSNG 54214. Papua New Guinea, Central Province, Bartholomew

MELANESIAN RODENTS 673

Range, Haveri, VIII-XI.1893, L. Loria; 1 ®, skin and skull. Ex

MSNG 3616 (2).

Mus musculus Linnaeus, 1758

MSNG 3474 (a-d). Indonesia, West Irian Jaya Province, Sorong,

without collecting date, L. M. D’Albertis; 1 6 juv. (d), 2 99 (a

and b), 1 2 juv. (d), skin and skull, paralectotypes of Mus albertisi

Peters & Doria, 1881.

MSNG 3669. Papua New Guinea, without collecting date, L. Loria;

1, skin and skull.

Remarks. Peters & Doria’s (1881) name Mus albertisi is a

synonym of Mus musculus Linnaeus, 1758; the name was applied to

a series of commensal house mice collected by D’Albertis at Sorong,

including four of the specimens listed above. RUMMLER (1938: 218)

identified a syntype in Berlin (ZMB 5423) as the “type” of albertisz,

which we accept as a lectotype designation. The four former syn-

types MSNG 3474 (a-d) are thus paralectotypes.

Paramelomys levipes (Thomas, 1897)

MSNG 3600 (a). Papua New Guinea, Central Province, Bartholomew

Range, Haveri, 700 m above sea level, VIII-XI.1893, L. Loria; 1

6, skin and skull, paralectotype of Uromys levipes Thomas, 1897.

MSNG 3600 (b). Papua New Guinea, Central Province, Astrolabe

Range, Gerekanumu, II.1893, L. Loria; 1 d, skin and skull, para-

type of Uromys levipes Thomas, 1897.

Remarks. THomas’ (1897) description of Uromys levipes

was based on these two “co-types” (= syntypes) collected by Loria

at Haveri, apparently BMNH 97.8.7.72 (received by London from

Genova) and MSNG 3600 (a). MENZIES (1989) formally designated

BMNH 97.8.7.72 as the lectotype of levipes, but we judge that RUMMLER

(1938) had already fixed a lectotype by his indication that ‘Thomas’

674 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

BMNH syntype should be considered “the type” (see also MUSSER

& CARLETON 2005: 1432). MSNG 3600 (a) is thus a paralectotype.

Paramelomys lorentzii (Jentink, 1908)

MSNG 3200 (1). Papua New Guinea, Western Province, Fly River,

IV.1875, L. M. D'Albertis; 1 d, skin and skull.

Pogonomelomys bruijnii (Peters & Doria, 1876)

MSNG 3677. Indonesia, West Irian Jaya Province, Salawati Island,

1875, A. A. Bruyn; 1 6, skin and skull, holotype of Uromys bruijnii

Peters & Doria, 1876 (Figs 5, 6).

Remarks. Two species are now recognized within the

genus Pogonomelomys (MUSSER & CARLETON 2005), but the true

diversity represented among museum collections is much greater

(HELGEN & APLIN, in litt.). The holotype of Pogonomelomys brutjnit

from Salawati remains morphologically unique. FLANNERY (1995a:

312) referred a series of Pogonomelomys specimens collected on the

mainland of the Vogelkop Peninsula (from the localities Djitman

and Aijinjo, deposited in the Naturalis Museum, Leiden) to bruzjnii.

‘This may be a correct attribution, but we note that these specimens

are considerably smaller than the holotype of bruijnii in cranial and

dental size (e.g. crown length of molar row 8.1 mm in the holotype

versus 6.9-7.3 mm in the series of three adults from the Vogelkop).

Only subfossil material from the Vogelkop Peninsula reported by

APLIN et al. (1999: 379) matches the holotype of bruzjnii in size; the

Leiden series better corresponds to a taxon that APLIN et al. (1999:

379-380) identify as “Pogonomelomys sp.” A full review of the genus

based on all available specimens, modern and subfossil, is needed.

Pogonomys loriae Thomas, 1897

MSNG 3259. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, 700 m above sea level, VIII-XI.1893, L. Loria; 1

2. skin and skull.

MELANESIAN RODENTS 675

MSNG 3492. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, 700 m above sea level, VIII-X1I.1893, L. Loria; 1 &,

skin and skull, syntype of Pogonomys loriae Thomas, 1897.

MSNG 3492 (a and b). Papua New Guinea, Central Province, Bar-

tholomew Range, Haveri, 700 m above sea level, VIII-XI.1893, L.

Loria; 1 6 (a) paratype, 1 d juv. (b), skin and skull.

MSNG 3493 (a). Papua New Guinea, Central Province, Astrolabe

Range, Gerekanumu, II.1893, L. Loria; 1 2, skin and skull.

MSNG 34292. Papua New Guinea, Central Province, Bartholomew

Range, Moroka, 1300 m above sea level, VII-VIII.1893, L. Loria;

1 2, skin and skull.

MSNG 54210. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 dé, skin and skull. Ex

MSNG 3257-66.

Remarks. ‘THomas’ (1897) two original “co-types” of

Pogonomys loriae are apparently MSNG 3492 and BMNH 97.8.7.47

(see TATE 1951: 280). As far as we are aware, no lectotype has been

designated.

Pogonomys macrourus (Milne-Edwards, 1877)

MSNG 3257- (50-71; except 66, see Pogonomys loriae MSNG 54210).

Papua New Guinea, Central Province, Bartholomew Range, Haveri,

VIII-XI.1893, L. Loria; 1 d (57) paralectotype of Pogonomys lepi-

dus Thomas, 1897, 13 do (51, 56, 59, 60, 61, 62, 63, 64, 67, 68,

69, 70 and 71), 7 99 (50, 52, 53, 54, 55, 58 and 65), skin and skull.

MSNG 3670. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, 700 m above sea level, VIII-XI.1893, L. Loria; 1 9

juv., skin and skull.

MSNG 34293-2. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 gd, skin and skull.

MSNG 34294-1. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 d, skin and skull.

676 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

MSNG 34295 (a-c). Papua New Guinea, Central Province, Bartho-

lomew Range, Haveri, VIII-XI.1893, L. Loria; 3 29°, skin and skull.

MSNG 34296. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 £ juv., skin and skull.

MSNG 42217 (I-IV). Papua New Guinea, Central Province, Bar-

tholomew Range, Haveri, 700 m above sea level, VIII-XI.1893, L.

Loria; 2 dé (I and II), 2 2 (III and IV), alcohol and skull.

MSNG 54211. Papua New Guinea, Central Province, Astrolabe

Range, Gerekanumu, 1I.1893, L. Loria; 1 9, skin and skull. Ex

MSNG 3493 (b).

Remarks. ‘THOMAS (1897) based his description of Pogono-

mys lepidus on two syntypes, apparently MSNG 3257-57 and BMNH

97.8.7.51. We accept TaTe’s (1951: 279) reference to the BMNH

specimen as the “type” of lepidus as a lectotype designation. MSNG

3257-57 is thus a paralectotype.

Pogonomys mollipilosus (Peters & Doria, 1881)

MSNG 3491. Papua New Guinea, Western Province, Katau, with-

out collecting date, L. M. D’Albertis; 1 é, skin and skull, holotype

of Mus mollipilosus Peters & Doria, 1881 (Figs 3, 4).

Remarks. Few specimens of Pogonomys have been collected

in the TransFly region of southern New Guinea, and the taxonomic

status of Pogonomys mollipilosus has never been satisfactorily resolved.

DENNIS & MENZIES (1979: 322) observed that “the possibility that

mollipilosus is an Australian species occurring in New Guinea only

in the lower Fly River region cannot be ignored”, and Australian

samples of Pogonomys have been explicitly discussed under the name

Pogonomys mollipilosus (WINTER & WHITFORD 1995), thus assumed

to represent a distinct species occurring in Australia and the Trans-

Fly. Musser & CARLETON (1993, 2005) believed that the holotype of

mollipilosus to be a specimen of the widespread lowland New Guinea

taxon Pogonomys macrourus and synonymized it accordingly, but the

last revisionary treatment of Pogonomys that involved firsthand com-

MELANESIAN RODENTS 677

parisons of the holotype of mollipilosus was the review published by

DENNIS & MENZIES (1979).

Our own examination of the young holotype (HELGEN 2007:

747) indicates that it is a larger-toothed rat compared to samples of

P. macrourus, probably does not belong within the synonymy of that

taxon as currently recognized (MUSSER & CARLETON 2005), and pos-

sibly shows a closer morphological resemblance to the larger-bodied

species P. loriae. "Taxonomic boundaries within the genus Pogonomys

remain very poorly understood and must be evaluated in the context

of morphometric and genetic examinations of a sufficient sample

drawing from type material and the very large number of speci-

mens and sampling localities now represented in world museums.

Rattus exulans (Peale, 1848)

MSNG 3151. Papua New Guinea, Central Province, Kapa Kapa,

VII,1891, L. Loria, 1 è and 1.9. skin and shall,

MSNG 3167. Papua New Guinea, Milne Bay Province, Bara Bara,

I-IT.1890> L. Loria; 2 2S. skim and skull:

MSNG 3249- (6-11). Indonesia, Papua Province, Humboldt Bay,

XII.1875, O. Beccari; 3 dé (7, 9 and 11) ), 3 22 (6, 8 and 10), skin

and skull.

MSNG 3250 (a-d). Indonesia, West Irian Jaya Province, Mount

Arfak, Hatam, VI.1875, O. Beccari; 1 d (c), 3 22 (a, b and d), skin

and skull.

MSNG 3255. Papua New Guinea, Central Province, Aroma Bay,

IX.1889, L. Loria; 17 juv., alcohol.

MSNG 3255- (20-49; except 22, 28, 29). Papua New Guinea, Cen-

tral Province, Aroma Bay, IX.1889, L. Loria; 13 dd (21, 23, 24, 25,

27, 34, 35, 38, 44, 46, 47, 48 and 49), 14 99 (20, 26, 30, 31, 32, 33,

36, 37, 39, 40, 41, 42, 43, 45), skin and skull.

MSNG 3255- (001-008). Papua New Guinea, Central Province,

Aroma Bay, IX.1889, L. Loria; 5 dd (001, 002, 005, 006 and 007),

3 22 (003, 004 and 008), skin and skull.

678 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

MSNG 3256 (c and d). Papua New Guinea, Central Baines Kapa

Kapa, VII.1891, L. Loria; 1 d (d), 1 9 (c), skin and skull.

MSNG 3505. Papua New Guinea, Central Province, Mekeo Dis-

trict, Inawi, IX.1892, L. Loria; 1 ¢, skin and skull.

MSNG 3508. Papua New Guinea, Central Province, Hula, Hood

Point, VIII.1889, L. Loria; 1 2, skin and skull.

MSNG 3509. Papua New Guinea, Central Province, Irupara,

VIII.1890, L. Loria; 1 6, skin and skull.

MSNG 3518 (a-e). Papua New Guinea, Central Province, Aroma Bay,

IX.1889, L. Loria; 1 d juv. (d), 4 92 (a, b, c and e), skin and skull.

MSNG 5043. Papua New Guinea, Milne Bay Province, Bara Bara,

ICRZIS90,. IL, Losa 14? sti

MSNG 38829 (a-d). Papua New Guinea, Central Province, Aroma

Bay, IX.1889, L. Loria; 1 d (a), 1 £ (b), 2 specimens (c and d),

skin and skull.

MSNG 52753. Papua New Guinea, Milne Bay Province, Bara Bara,

I-I1.1890, L. Loria; 1 3d, alcohol and skull.

MSNG 52755. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, XI.1893, L. Loria; 1 2, alcohol and skull.

Rattus leucopus (Gray, 1867)

MSNG 3219. Papua New Guinea, Western Province, Katau, with-

out collecting date, L. M. D’Albertis; 1 d juv., skin and skull.

MSNG 3244 (a-e). Indonesia, Maluku Province, Aru Islands,

Wokam, 29.II-3.VI.1873, O. Beccari; 2 dé (b and c), 2 22 (a and

d), skin and skull; 1 juv. (e), alcohol.

MSNG 3460. Papua New Guinea, Western Province, Fly River,

without collecting date, L. M. D’Albertis; 1 2, skin and skull, holo-

type of Mus ringens Peters & Doria, 1881.

MSNG 3506. Papua New Guinea, Central Province, Astrolabe

Range, Gerekanumu, II.1893, L. Loria; 1 d, skin and skull.

MELANESIAN RODENTS 679

MSNG 52752. Papua New Guinea, Central Province, Kapa Kapa,

VI-VII.1891, L. Loria; 1 é juv.; alcohol and skull.

MSNG 54201. Papua New Guinea, Western Province, Katau, with-

out collecting date, L. M. D'Albertis; 3 juv., alcohol. Ex MSNG

S219.

MSNG 54208. Papua New Guinea, Fly River, 1875, L. M.

D’Albertis; 1 d juv., skin and skull. Ex MSNG 3240-19.

Remarks. As far as we are aware, FLANNERY (1995b) pro-

vided the first published record of a native Rattus species in the

Aru Islands when he reported R. leucopus from Aru based on two

specimens deposited at the Australian Museum in Sydney. Flannery

and earlier reviewers, such as TAYLOR et al. (1982), apparently over-

looked Beccari’s series of Rattus from the Aru Islands in Genova,

which we refer to Rattus leucopus.

According to FLANNERY (1995b), the Aru specimens referred to

R. leucopus in the Australian Museum have four pairs of mammae,

but Rattus leucopus from mainland New Guinea and Australia have

only three pairs (TAYLOR et al. 1982; TAYLoR & HORNER 1973), as

does the only one of the Genova Aru specimens in which the for-

mula can be determined, MSNG 3244 (d). Aru specimens (both in

Sydney and Genova) warrant further examination to establish their

taxonomic status relative to other populations identified as R. leuco-

pus (MusseR & CARLETON 2005).

Rattus mordax (Thomas, 1904)

MSNG 3494. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, 700 m above sea level, VIII-XI.1893, L. Loria; 1

6, skin and skull.

Rattus rattus (Linnaeus, 1758)

MSNG 3534. Indonesia, West Irian Jaya Province, Sorong, 1875,

O. Beccari; 1 9, skin and skull, holotype of Mus beccarii Peters &

Doria, 1881 and Mus doriae 'Trouessart, 1897.

680 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

MSNG 3535. Indonesia, West Irian Jaya Province, Sorong, without

collecting date, L. M. D’Albertis; 1 d, skin and skull.

MSNG 3536. Indonesia, West Irian Jaya Province, Sorong, without

collecting date, L. M. D'Albertis; 1 2, skin and skull.

MSNG 3537. Papua New Guinea, Western Province, Fly River,

without collecting date, L. M. D’Abertis; 1 £ juv., skin and skull.

MSNG 3538. Indonesia, West Irian Jaya Province, Sorong, 1872,

L. M. D'Albertis; 1 juv., alcohol.

MSNG 54194. Indonesia, West Irian Jaya Province, Sorong, with-

out collecting date, L. M. D'Albertis; 1 juv., alcohol.

Remarks. PETERS & Doria (1881) based the name Mus bec-

cart on MSNG 3534, a commensal house rat collected at Sorong.

Noting that this name was preoccupied by Mus beccari Jentink,

1880 (now Margaretamys beccarit; see MusseR & CARLETON 2005),

TROUESSART (1897: 472) introduced the name Mus dorzae to replace it.

Rattus sordidus (Gould, 1858)

MSNG 3260 (a and b). Papua New Guinea, Central Province, Kapa

Kapa, NENTEAS9ITI Loria 1 o (a) 19), skin and skull.

MSNG 3501. Papua New Guinea, Central Province, Kapa Kapa,

V-VI.1891, L. Loria; 1 gd, skin and skull [MSNG 3501 (a)], para-

lectotype of Mus gestri Thomas, 1897.

MSNG 3501 (b - e). Papua New Guinea, Central Province, Kapa

Kapa, V-VII.1891,..L. Loria; 1 Glcp 2a (brand 1d); skin and

skull; 1 6 (e), skated:

Remarks. THomas’ (1897) description of Mus gestri was

based on these two “co-types” (= syntypes) collected by Loria at Kapa

Kapa, apparently MSNG 3501 and BMNH 97.8.7.72. TAYLOR et al.

(1982: 269) formally designated BMNH 97.8.7.72 as the lectotype of

levipes, but we judge that RUMMLER (1938: 208) had already fixed a

lectotype by his indication that Thomas’ BMNH syntype should be

considered the “type” (“der Typus”). MSNG 3501 is a paralectotype.

MELANESIAN RODENTS 681

Rattus verecundus (Thomas, 1904)

MSNG 3517. Papua New Guinea, Central Province, Bartholomew Range,

Moroka, 1300 m above sea level, X.1893, L. Loria; 1 2, skin and skull.

Uromys caudimaculatus (Krefft, 1867)

MSNG 3248. Papua New Guinea, Western Province, Katau, with-

out collecting date, L. M. D'Albertis; 1 specimen, skin, holotype of

Uromys validus Peters & Doria, 1881.

MSNG 3605 (a). Papua New Guinea, Central Province, Kapa Kapa,

VII.1891, L. Loria; 1 d, skin and skull.

MSNG 3605 (b). Papua New Guinea, Central Province, Aroma

Bay, IX.1889, L. Loria; 1 specimen, skin and skull.

Remarks. ‘The holotype of Uromys validus is represented at

Genova only by a skin, and ‘TATE (1951) suspected that the accom-

panying skull was lost. The first author recently discovered the type

skull at BMNH, misplaced in a drawer with type specimens of other

Uromys. We advocate the return of the skull to Genova to reunite

the type material of Uromys validus.

Though Uromys validus Peters & Doria, 1881 is usually regarded

as a synonym of the large and widespread species Uromys caudi-

maculatus, patterns of geographic variation in Melanesian Uromys

remain poorly understood (Musser & CARLETON 2005). As cur-

rently recognized, “Uromys caudimaculatus” is likely to represent a

complex of related species. As the oldest name applied to a Melane-

sian population of Uromys, further study may show that the epithet

validus is the most appropriate name for a distinct species endemic

to the Melanesian region.

Uromys nero Thomas, 1913

MSNG 54213. Papua New Guinea, Central Province, Bartholomew

Range, Haveri, VIII-XI.1893, L. Loria; 1 d, skin and skull. Ex

MSNG 3605 (c).

682 K.M. HELGEN, T. LEARY, G. DORIA & G. AMORI

Remarks. Following the revision presented by TATE (1951),

many large scansorial rat species from New Guinea and north-east-

ern Australia have been lumped together under the name “Uromys

caudimaculatus” (e.g. FLANNERY 1995a, 1995b). These include speci-

mens from populations that HELGEN (2007) refers to a number of

distinct species, including U. nero and U. scaphax, in advance of a

formal revision (MUSSER & CARLETON 2005). We tentatively associ-

ate this specimen with the name Uromys nero 'Thomas, 1913, an

identification that awaits detailed exposition in a future review of

the caudimaculatus species-complex (HELGEN 2007).

Uromys rex (Thomas, 1888)

MSNG 3459. Solomon Islands, Guadalcanal Province, Guadalcanal,

6.V.1889, C. M. Woodford; 1 6, skin and skull; bought by Gerrard.

Remarks. Only a handful of specimens of Uromys rex are

preserved in world museums (see GROVES & FLANNERY 1994). The

holotype and three paratypes of Uromys rex, all collected at Aola by

Woodford, are preserved at BMNH (THOMAS 1888; TATE 1951). It

is unclear whether MSNG 3459 was collected at the same time or

place as the type series (i.e. with the date listed above an indication

of its receipt by the museum, rather than the date of collection) or

whether it was truly secured later than Thomas’ original series.

Uromys siebersi Thomas, 1923

MSNG 3245. Papua New Guinea, Kai Islands, without collecting

date, O. Beccari; 1 dé, skin and skull.

Remarks. The Uromys of the Kai Archipelago is U. siebersz,

a distinctive species endemic to these islands (MUSSER & CARLETON

2005), formerly included within the synonymy of U. caudimacu-

latus (see above). The species was last collected in 1922 (THoMmas

1923) and very few specimens of U. sieberst are preserved in world

museums. Apart from the specimen at Genova, there are only three

others—the holotype at BMNH (THomas 1923), a paratype in the

MELANESIAN RODENTS 683

Museum Zoologicum Bogoriense in Cibinong, Indonesia (formerly

the Buitenzorg Museum), and a skull in Berlin (ZMB 5682).

ACKNOWLEDGEMENTS

We wish to thank Roberto Poggi, Director of the Museo Civico

di Storia Naturale “G. Doria” of Genova for his kind support.

The first author’s visit to Genova was supported by funding

from the U.S. National Science Foundation and the Smithsonian

Institution, and he thanks Don Wilson for ongoing assistance, Allen

Allison for advice and support at the Bernice P. Bishop Museum in

Honolulu, Tim Flannery and Guy Musser for helpful discussion.

The second author’s visit to Genova was partly supported by

funding from the Postgraduate Research Support Scheme of the

University of Sydney, and she thanks Chris Dickman for ongo-

ing assistance. The second author would also like to thank Robert

Sheaffe for allowing her to take numerous extended leaves of absence

from her substantive position with Parks and Wildlife Division to

pursue her passion for New Guinea rodents.

Thanks to Maria Bruna Invernici (Museo Civico di Storia Natu-

rale “G. Doria” of Genova), Alessia Lantieri and Cristina Camattari

for technical support, and to Marino Superina and Luciano Ber-

lingieri (Centro Video, Comune di Genova) for the digital photos.

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ABSTRACT

Catalogue of Melanesian Rodents (Mammalia: Rodentia) in the Museum of

Genova.

The Museum of Genova holds an historically important collection of 206 rodent

specimens from Melanesia (the large tropical island of New Guinea and adjacent

smaller islands), collected during the late nineteenth century. The collection houses

a number of Melanesian rodent type specimens, including six holotypes, one lecto-

type, one syntype, paralectotypes for five taxa, and four paratypes. Also to be found

in the collection are exemplars of several species very rare in collections, such as the

large endemic insular murines Uromys rex (of Guadalcanal in the Solomon Archi-

pelago) and Uromys stebersi (of the Kai Archipelago of Indonesia).

The Authors present revised identifications of all Melanesian rodents repre-

sented in the collection, discuss the localities where these specimens were obtained,

and provide photographic illustrations of selected type material.

RIASSUNTO

Catalogo dei Roditori (Mammalia: Rodentia) della Melanesia nel Museo di

Genova.

Nel Museo di Genova è conservata un’importante collezione di 206 Roditori

della Melanesia (comprendente la grande isola tropicale della Nuova Guinea e le

più piccole isole adiacenti) raccolti nel XIX secolo. Nella collezione sono presenti

anche esemplari tipici: sei olotipi, un lectotipo, un sintipo, paralectotipi di cinque

specie e quattro paratipi. Vi sono anche rappresentanti di specie molto rare nelle

collezioni museali come i grossi murini, endemiti insulari, Uromys rex (di Guadal-

canal nell’Arcipelago delle Solomon) e Uromys siebersi (dell’ Arcipelago delle Kai

dell’ Indonesia).

Vengono riportate le determinazioni aggiornate di tutti gli esemplari, sono

esaminate le località di raccolta e vengono fornite le fotografie di alcuni esemplari

tipici.

GRUPPI E FORME NUOVE DESCRITTI NEL PRESENTE

VOLUME

INSECTA

Coleoptera

Cala bad ave

Pag.

Carabus (Apotomopterus) mecynodes mitis Cavazzuti, n. ssp. . STI

Carabus (Apotomopterus) patroclus xinshibanus Cavazzuti, n. ssp. 575

Carabus (Archaeocarabus) lacertosus Cavazzuti, n. sp. ........... 576

Carabus (Coptolabrus) ishizukat huberanus Cavazzuti, n. ssp. . Sot

Carabus (Cupreocarabus) laevithorax arduus Cavazzuti, n. ssp. 584

Carabus (Cupreocarabus) lixianensis pseudomorettoi Cavazzuti, n. ssp. 585

Carabus (Lamprostus) seroulikbin Cavazzuti, n. sp. ................ 588

Carabus (Piocarabus) vladimirskyi tianzhuiensis Cavazzuti, nomen

novum pro tianzhuicus Cavazzuti, 2006, nec Imura, 1996 . BAD

Carabus (Rhigocarabus) impavidus Cavazzuti, n. sp. .............. 581

Cychrus brugget lianghensis Cavazzuti, n. ssp. «1.0.0... cece cece eee 596

Cychrus toledanot -Cavazzani WI spe DB Sess Se aes 593

Cychrus toledanot maerkangensis Cavazzuti, n. ssp. ............... 595

Psephenidae

Eetopria tranensis Lee &liCh Mocha) ok aan eal

Macroeubria shimadgi Lea:&dlach n spagna: 235

Schinostethus (Sundodrupeus) sumatrensis Lee & Jach, n. sp. . 237

Schinostethus (Sundodrupeus) transversus Lee & Jach, n. sp. . 239

Monotomidae

tadolentibsusipnazi Balestrieri 204

Cer ty-homa dave

Pag.

Pres Apri et LIO ZA

Discolomidae

-phanocephalus HGS Al, We Spread 205

Cephalephanusechhakkackal ,Miehspra lite he Lad Abstain hs 214

Pavajaiie Wavra Pal) n sp. rotaia Ni 207

RavamaschomazaiaktbPal, isp, ia 240.

Cioe Amine ha ciare

Oly peaspis Gordon & (Cane paris, n. Se. ida 278

Diazonema boothi Gordon & Canepari, n. sp. ....................... 267

Diazonema eccentrica Gordon & Canepari, n. sp................... 269

Diazonema murilloi Gordon & Canepari, n. sp. .................... 268

Hyperaspis abertha Gordon & Canepari, n. sp. .................... 365

Hyperaspis aemulata Gordon & Canepari, n. sp. ................... 367

Hyperaspis apicaspis Gordon & Canepari, n. sp. .................. 342

Hyperaspis arida Gordon & Canepari, n. sp..................000..... 389

Hyperaspis atra Gordon Sj Canepariy span iitoreondi). 350

Hyperaspis ayacucho Gordon & Canepari, n. sp. .................. 3/1

Hwyperaspis bisignata Gordon & Canepari; Ne-Spre-}. carbon. 313

Hivperasprs brethest’ GordonsXvCanepari i a Spe), sai dew rdnotien.. 325

Hyperaspis camargoi Gordon & Canepari, n. sp. .................. 345

Hyperaspis campbelli Gordon & Canepari, n. sp.................... 331

Elwperaspis ehapinGordon"& Cancpari ni spt seta 307

Elyperaspis:chocor Gordon. & Canepari, ni spit i) 348

Hyperaspis circumclusa Gordon & Canepari, n. sp. ................ 366

Hyperaspis colombiensis Gordon & Canepari, n. sp. .............. Boz

Hyperaspis cracentis Gordon & Canepari, n. sp. .................... 308

Pivperaspis dispar Gordon & Canepari, n. Sp. si... 368

Hyperaspis dissidens Gordon & Canepari, n. sp. .................... 333

Hyperaspis helveola Gordon & Canepari, n. sp. .................... 302

Hvyperaspis herrerat Gordon & Canepari, n. sp. .................... SZ

Pag.

Hyperaspis howdeni Gordon & Canepari, n. sp...................... 347

Hyperaspis tmitatrix Gordon & Canéeparin Sp, liti 305

Hyperaspis istmina Gordon & Canepari, n. sp....................... 363

Hyperaspis joannae Gordon & Canepari, n. sp. ..................... 339

Hyperaspis laterirmacula Gordon & Canepari, n. sp. .............. 346

Hyperaspis latitibia Gordon & Canepari, n. sp...................... 27

Hyperaspis lindae Gordon, & Ganepariia. spa, LIE 357

Hyperaspis mariposa Gordon & Canepari, n. sp. .................. 354

Hwvperaspis mimica Gordon & Canepari, isp. iena 344

Hyperaspis octonotata Gordon & Canepari, n. sp. ................ DIRI

Hyperaspis orthivora Gordon & Canepari, n. sp. .................. 322

ITyperaspis prolata Gordone& Canepari, a spin eee ee 394

Hyperaspis pseudodonzeli Gordon & Canepari, n. sp. ........... 320

Hyperaspis pseudopavida Gordon & Canepari, n. sp. ............ 362

Hyperaspis rosariensis Gordon & Canepari, n. sp. ................ 327

Hyperaspis satipoensis Gordon & Canepari, n. sp. ................. 364

Hyperaspis stiladesma Gordon & Canepari, n. sp. .................. 328

Hyperaspis simlaensis Gordon & Canepari, n. sp. ................. 355

Hyperaspis tayronensis Gordon & Canepari, n. sp. ............... 319

Hyperaspis uninotata Gordon & Canepari, n. sp. ................. 335

Hyperaspis vredenburgi Gordon & Canepari, n. sp. ............... 374

Hyperaspis zonula Gordon & Ganepanieràrsp. sh .aitoes.2ees 329

Peruaspis Gordon-& Canepagi, n acta RA 282

Peruaspis hypocrita Gordon & Canepari, n. sp... ui 284

Peruaspis papravehu Gordon & Canepa) spl 283

Prognataspis Gordon & £'anepagiemni pene ani 280

Prognataspis surreptiva Gordon & Canepari, n. sp. ............... 281

Tenuisvalvae caucaensis Gordon & Canepari, n. sp. .............. 409

Tenuisvalvae gnoma Gordon & Canepari, 1. Sp rg 412

Tenuisvalvae parenthesis Gordon & Canepari, n. sp. ............ 408

Tenebrionidae

Dichillus (Dichillus) maionese Leo: Dip era 621

Dichillus (Dichillus) tapinonae, Leo. AM iSelect Rk a 618

Dichillus (Dichilus)-tvavbemons IE I sica 610

Inopeplidae

TRO DEDE SIAE RE DISEASE SI RE Me iii pie

LORIS AAA BA ASI ici a. bade ew ei

Hymenoptera

Formicidae

Basiceros onorei Baroni Urbani & de Andrade, n. sp. ...........

Basiceros papuanum de, Andrade tiny spi List. iii

Strumigenys aduncomala de Andrade, n. sp. ............... 20.

Strumigenys aequinoctialis de Andrade, n. sp.........................

perumicenys buddhasia:de Andrade Ne, SP) locati notara

Strumigenys caniophanoides de Andrade, n. sp.......................

pirunncenys hindu desAndrade;ctv SP) selicrlapssidgarari

Strumigenys longimala de Andrade, n. sp. ..........cccceceeeeeeeee eee

Strumigenys nageli Baroni Urbani & de Andrade, n. sp. .......

Strumigenys onorei Baroni Urbani & de Andrade, n. sp. .......

7 otrumicenys pilosulasde Andrade, ny sp. lis). Wade. dI

| Strumigenys poinari Baroni Urbani & de Andrade, n. sp. ..

Strumigenys veddha de Andrade, n. sp................

Pag.

197

133

137

150

165

159

153

156

147

162

168

173

141

INDICE

Bapano D. - Contributo alla conoscenza dei Neurotteri

della Liguria (Insecta, Neuropterida). (16.1.2008) .......

BAGHINO L. - Monitoraggio della migrazione post-riprodut-

tiva dei rapaci diurni nella Zona di Protezione Speciale

IT 1331578 “Beigua-Turchino” (Aves, Accipitriformes

et Falcontiornes 101.2060815 tha

BARONI URBANI C. & DE ANDRADE M. L. - The ant tribe

Dacetini: limits and constituent genera, with descrip-

tions of new species (Hymenoptera, Formicidae). (28.

a, ROOT ee OI TRONI

BATTISTON R. & FONTANA P. - Ricerche zoologiche della

nave oceanografica “Minerva” (C.N.R.) sulle isole cir-

cumsarde. XXX. Embiidina (Insecta). (4.XII.2007)....

CavazzuTI P. - Nuovi Carabus L. e Cychrus F. di Cina e

Iran (Coleoptera, Carabidae). Descrizioni e note siste-

matiche. (9.05. 2008) le, ta ee

GARDINI G. - Ricerche zoologiche della nave oceanografica

“Minerva” (C.N.R.) sulle isole circumsarde. XXXI.

Pseudoscorpiones (16) V, 2008 )e a eee

GORDON R. D. & CANEPARI C. - South American Cocci-

nellidae (Coleoptera), Part XI: a systematic revision of

Hyperaspidini (Hyperaspidinae). (8.1.2008) .................

HeLGEN K. M., Leary T., Doria G. & AMORI G. - Cata-

logue of Melanesian rodents in the Museum of Genova

(Mammalia, Rodentia), CRI

Lee C.-F & JAcH M. - New species and new records of

Psephenidae from Asia (Coleoptera). (10.XII.2007).....

Leo P. - Osservazioni su Dichillus corsicus e descrizione

di tre nuove specie del Mediterraneo occidentale (Co-

leoptera, Tenebrionidae). (28.IV.2008).........................

Pagg.

535-548

513-534

1-191

223-227

573-601

629-651

245-512

653-686

229-244

603-627

Pagg.

Par T. K. - On a collection of Heteromera and Cucujoidea

of the Museo Civico di Storia Naturale “G. Doria”,

Genova (Coleoptera, Polyphaga). (4.XII.2007)............ 193-221

PSOMADAKIS P. N., BoTraro M., Doria G., GARIBALDI F,

GIUSTINO S. & VaccHI M. - Notes on the Regalecus

glesne occurring in the Gulf of Genova and in Liguro-

Provencal waters (NW Mediterranean) (Pisces, Lam-

priditormes, Regalecidae). (7.11. 2008)... nre 549-571

REGISTRATO AL TRIBUNALE DI GENOVA AL N. 74 IN DATA 17 giugno 1949

DOTT. ROBERTO POGGI - DIRETTORE RESPONSABILE

Finito di stampare nel mese di maggio 2008

PRINTED IN ITALY

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BINAGHI G., 1950 - Coleotteri d’Italia. Vita, ambienti, utilità, mezzi di lotta - Ed. Briano,

Genova, 210 pp., 104 figg. [nel caso di un’opera monografica]

Capra F.,, 1987 - Italian and European Dermaptera of the museums of Florence and

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