morón y ramírez-ponce, 2012. mesoamerican genera of anomalini

8/12/2019 Morn y Ramrez-Ponce, 2012. Mesoamerican Genera of Anomalini

1/18

ABSTRACT

Shiny chafers of the tribe Anomalini arerepresented in Mesoamerica by 14 genera and240 species, but revisions, keys to species and

biological information are scarce, despite the factthat a number of species have been cited as pestsof diverse cultures, mainly as part of the whitegrub guilds. Past studies on the taxonomy of thesespecies offer many problems, because the generawere not adequately defined, the species usuallyoffer character variability, genitalia were rarelydescribed, and many authors do not agree with thestatus of the names. After phylogenetic studieswith a generic sample of Anomalini from theworld, Ramrez-Ponce and Morn proposed therevalidation of the genera Pachystethus Blanchardand Paranomala Casey, as well as the redefinitionof the genera Anomala Samouelle and Callistethus Blanchard, because the representatives of theselineages have unique combinations of morphologicalcharacters and some synapomorphies are supporting

adequately the nodes in the strict concensustree obtained during such analysis. After the publication of the cited work, new evidences werefound suggesting that other groups of species needto be separate in distinct genera from Paranomalawhile other taxa are part of the Paranomala clade.

Mesoamerican genera of Anomalini (Coleoptera:Melolonthidae: Rutelinae): A brief review

In the present review we provide brief nomenclaturalhistory for each genus and synopsis of the

phylogenetic hypothesis that support a new proposal for the classification of the generarepresented in Mesoamerica. Diagnosis andcomments on the distribution of the 14 genera are

provided. Color photographs, illustrations ofdiagnostic characters and a key to genera are alsoincluded.

KEYWORDS: shiny chafers, taxonomy,distribution, Mexico, Central America, importance,white grubs

INTRODUCTION

Members of the tribe Anomalini are a nearlycosmopolitan scarab group, widely diverse, buteasy to recognize because of its nine segmentedantennae, elytra with membranous border andtransverse labrum. We had world wide records fornear 2000 species and 34 - 44 genera [1, 2] buttaking account of recent studies on Americanfauna, these numbers have increased to 53 genera,including new taxa and changes in thenomenclature and tribal classification [3, 4, 5].

Aside from its ecological importance derivedfrom the great specific richness, wide distributionand diversity of wild food preferences, larvae ofmany species of Anomalini are included asmembers of the white grub complex, sometimesas debris consumers that aid to increase thefertility of soils used for agriculture or, on the

1Red de Biodiversidad y Sistemtica, Instituto de Ecologa A. C. Apartado Postal 63, Xalapa, Veracruz,91000, 2Posgrado en Ciencias Biolgicas, Instituto de Ecologa, UNAM, Ciudad Universitaria,D. F. 04510, Mxico

Miguel Angel Morn 1,* and Andrs Ramrez-Ponce 1,2

*Corresponding author: Dr. Miguel Angel Morn,Instituto de Ecologa, A.C., Carretera antigua a Coatepec351, El Haya, Xalapa, Veracruz, 91070, [email protected]

T r e n d s i n

Entomology

Vol. 8, 2012


2/18

98 Miguel Angel Morn & Andrs Ramrez-Ponce

taxonomy of the group [10] tacitly applied bymost of the authors [11, 12, 13]. During the past200 years the advancement on the general

classification of Anomalini is scarce [14], and theacceptance of the generic names were accordingto the criteria of each author (Table 1).

Historically, the first works dealing withAmerican species of Anomalini were written inaccordance with the species groups previouslyknown in the Old World. So that, some subgenera,divisions or sections was small parts of groupslargely represented in Eurasia [15, 16]. Otherworks were focused on the species found in awell-limited geographical region [13, 17]. Recently,a number of descriptions of new American genera

and species of Anomalini were published,including some brief reviews; proposals foradjustments, and for a new classification wassupported based on comparative morphology and

phylogenetic analysis [3, 4, 5, 8, 9].

Arrow said that one problem with the classificationssupported on species from some geographicalregion is that many characters are not useful whenapplied to other faunal assemblages or to groupslarger or shorter than those originally studied [18].Some of the characters used in this kind of studieswere frequently used with much flexibility bydistinct authors who offered speculations on theaffinities of the genera and species. Proposals

based on integral comparative morphologicalevidences may be true, but others supported byadaptative characters may result in convergencesor homoplasies included in artificial groups [4, 12].

Phylogenetical affinities for the AmericanAnomalini are not easy to find, because the groupis formed by some mixture of elements withdifferent biogeographical origin. Apparently, agood number of taxa in the New World are

closely related with taxa from the Old World,inclusively some Mesoamerican genera mayrepresent relicts with Palearctic or Paleotropicalorigins. Some authors argued in favor of theabove cited hypothesis, as an example: accordingto Lacordaire and Potts, Strigoderma is theAmerican representative of Asiatic and AfricanPopillia , because they share a good number ofmorphological characters and also present a stronggradation in some of these structures [10, 11]. Onthe other hand, Bates argued that Phylloperthatolucana Bates is much distinct from the

other hand, are cited as pest for the root system ofmany cultivate plants [6, 7].

Notwithstanding, the taxonomic knowledge of thegroup is limited. Most of the collections lackcorrect specific determination for most part ofthe specimens. Other problems including vaguegeneric limits, distinct criteria for use ofmorphological characters and diverse drafts on thesystematics of the genera that supported the listsof species represented in each geographicalregion. Really, diverse facts do not aid to obtaintaxonomical consensus. Many species show widevariation in color patterns, body size, andtegumentary sculpture. A large number of specieswere described with one of few specimens,

lacking information on the specific variation,mouth parts, genital structures and precisedistribution. Original descriptions are much brief,including uninformative details. Some type seriesare formed by representatives of two or morespecies [8, 4, 9].

One of the main problems is the heterogeneouscomposition of the hyperdiverse genus Anomala Samouelle. During two centuries many newspecies from Old and New Worlds were includedin such genus, using vague criteria and delayingdeep comparative studies on external morphology,with the result of numerous characters repeated indistinct taxa, or species included in the genus bythe absence of some other characters [10]. Widediscordances in the generic limits of Anomala arbitrarily applied to a great number of speciesoffer a large list of names without true relationsor supposed paraphyly [4]. In recent years thecomparative study of all described species isnearly impossible, so that we need to userepresentative samples for the preliminary

phylogenetic analysis.

Definitions of other less diverse American generaof Anomalini with recent taxonomic reviews as ofStrigoderma (40 species) and Epectinaspis (ninespecies) show some phyletic problems. Strigoderma includes some species with intermediate charactersin reference to Epectinaspis , and the phylogeneticevidence shows that the latter genus is

paraphyletic [5]. Strongly supported limits foreach genus are not easy to obtain. According toour experience, intermediate or transitional forms

between taxa are very frequent in Anomalini.These allowed arbitrary rules for the alpha


3/18

Mesoamerican genera of Anomalini shiny chafers 99

T a

b l e 1

. S y n o p s

i s o f

t h e

h i s t o r y o f

t h e c l a s s i

f i c a t

i o n o f

t h e

A m e r

i c a n g e n e r a o f

A n o m a l

i n i .

A u t

h o r s r e

f e r e n c e s

G e n u s n a m e

[ 1 5 , 1 9 ]

[ 1 6 ]

[ 1 1 ]

[ 1 2 ]

[ 1 3 ]

[ 1 ]

[ 2 , 2

0 ]

[ 3 ]

[ 5 ]

[ 9 ]

P r e s e n

t w o r k

A n o m a l a

G

G

G

G

G

G

G

G

G

G

V a l

i d g e n u s

O W *

E u c h

l o r a

S G

S G

V a l

i d s u

b g e n u s

? O W

S p i l o t a

S G

S G

G

S G

X o c

h i c o

t l i a m n

N W

S t r i g o

d e r m a

G

G

G

G

G

G

G

G

G

G

V a l

i d g e n u s

N W

P o p i

l l i a

G

G

G

G

G

G

G

V a l

i d g e n u s

O W * *

C a l

l i s t e t h u s

G

G

G

G

G

G

G

V a l

i d g e n u s

O - N

W

P a c h y s

t e t h u s

G

G

S G

G

V a l

i d g e n u s

N W

P h y l

l o p e r t

h a

G

G

G

G

G

V a l

i d g e n u s

O - N

W

E p e c

t i n a s p i s

G

G

G

G

G

G

G

G

G

V a l

i d g e n u s

N W

C a l

l i r h i n u s

G

G

G

G

G

G

G

G

G

V a l

i d g e n u s

N W

D i l o p h o c

h i l a

G

G

G

G

G

V a l

i d g e n u s

N W

A l a m o n a

G

S y n

S t r i g o

d e r m a

A n o m a l e p

t a

G

S y n

S t r i g o

d e r m a

L a m o a n a

G

S y n

S t r i g o

d e r m a

R h o m

b o n a

l i a

G

S y n

S t r i g o

d e r m a

S t r i g o

d e r m e l

l a

G

S y n

S t r i g o

d e r m a

A n o m a l a c r a

G

G

G

S y n

P a r a n o m a l a

P a r a n o m a l a

S G

G

V a l

i d g e n u s

N W

O l i g a n o m a l a

S G

S y n

A n o m a l a

A n o m a l o p u s

S G

S y n

A n o m a l a

R u g o p e r t

h a

G

G

V a l

i d g e n u s

N W

M a z a h u a p e r t

h a

G

G

G

V a l

i d g e n u s

N W

N a y a r

i t a

G

G

G

V a l

i d g e n u s

N W

Y a a x

k u m u k

i a

G

G

G

V a l

i d g e n u s

N W

C h e l i l a b

i a

G

G

G

S u b g e n u s

P a r a n o m a l a

L e p t o h o p

l i a

G

G

S u b g e n u s

P a r a n o m a l a

A n o m a l o r

h i n a

G

G

V a l

i d g e n u s

N W

B a l a n o g o n i a

G

G

V a l

i d g e n u s

N W

A b b r e v i a t

i o n s : G - g e n u s , m

n - m a n u s c r

i p t n a m e ,

N W

- N e w

W o r

l d , O

W -

O l d W o r

l d , S

G - s u b g e n u s .

* A . o

r i e n

t a l i s i n t r o d u c e

d i n N W

, * * P . j

a p o n

i c a

i n t r o d u c e

d i n N W

.


4/18


are the main extensive references for the shinychafers of America [1, 2, 11, 12, 13, 15, 16, 20].Comparative study of the classifications proposed

by each of the above cited authors is not easy(Table 1), because each author used distincttaxonomic criteria, and the purposes of each workfrequently were different.

Burmeister

For the first time, a classification for the tribe as part of other groups of Rutelids was proposed by Burmeister. In the book of 1844, a detailedassembly for each one of the six genera thenrecognized is included [15]. The genus Anomala Samouelle was formed with nine subgenera and

74 species. Subgenus Anomala s.str. was formedwith 25 species separated into three groups, ninespecies from Java, China, Japan and Europe, and16 species from America. Subgenus Spilota Dejean was formed by three species from Asiaand eight new world species. The genusStrigoderma Dejean only included six Americanspecies. Main characters used for the genus rankclassification are the form of the clypeus and

protibia, form of pronotal basal margin, punctuationof pronotum and structure of mesosternum; for thesubgenus rank the characters were shape of tarsalclaws, form of mentum, pronotal, and elytralmargin, and prominence of mesosternum. In the

book of 1855, a new species from Brazil isdescribed as A. chromicolor , and is included inthe subgenus Euchlora Macleay with other threespecies from Asia [19].

Blanchard

As part of the catalogus of the beetles deposited inthe collection of the Museum of Natural Historyin Paris, this author listed 14 genera of Anomalinifor the world, seven of which were described asnew. Callistethus Blanchard is represented inAmerica as well as in Asia. Epectinaspis Blanchardand Pachystethus Blanchard are restricted toMesoamerica. Callirhinus is endemic of westernMexico. The genus Anomala is divided indivisions some of them without name andothers with the names used by Burmeister. Hisdivision VII includes only one Mexican species A.rhizotrogoides ; divisions VIII and IX are formed

by American taxa, 29 and 11 species respectively.The classification of the tribe is more complex

European species of such genus, but is clearlyrelated to it [12]. More than a century laterP. tolucana was transferred to the monotypic new

genus Mazahuapertha , and proposed as relictelement with Eurasiatic affinities [3].

Motivated to enface this problem, Ramrez-Ponceand Morn carried an exercise of phylogeneticinference based on one sample of diverse generaof American, European and Asiatic Anomalini,obtaining evidence to support the separation of thespecies of Anomala from the Old and New Worldas distinct genera, Anomala and Paranomala respectively [9]. These results corroborate theopinions of Ohaus and Casey about somemorphological differences between both groups ofspecies [1, 13]. The new generic definitions forParanomala and Anomala also aid to separateother genera with taxonomic problems, asCallistethus and Pachystethus , offering newdiagnostic characters useful for updating theclassification of the tribe, and some tools for theadvancement in the study on the phylogeny ofAnomalini and related groups.

According to the above cited ideas and consciousof that we need to obtain a complex classificatoryscheme supported with good phylogenetic data,

including both American and Old World lineages,the present work represent an updated synthesison the history of classification and the systematicsof the American Anomalini, previous to theformal discussion of the phylogenetic analysisand the new proposal for the supraspecificclassification of the genus Paranomala Caseycurrently in progress. Such study will includecomments on the affinities between diverselineages, supported by phylogenetic evidencesobtained by means of the analysis of a number ofrepresentative samples, larger than that in

preceding studies conducted by us. Also, we madedetailed comparative studies of many morphologicalcharacters used in old works, rejected by some

past authors, that reveal useful values at differenttaxonomic ranks of the Anomalini applied to ourresults.

Brief analysis of the taxonomic history of thetribe anomalini

The publications of Burmeister, Blanchard,Lacordaire, Bates, Casey, Ohaus and Machatschke


5/18


Casey

The results of this author may represent the mostdetailed and contrasting classificatory system for

American shiny chafers. Such a system wasdeveloped using the personal collection of theauthor with specimens mainly obtained in NorthAmerican localities. Many new genera were

proposed in this work as well as three newsubgenera for Anomala . The subgenus Paranomala was the most diverse, some species were groupedin similar manner to that of Burmeisters sectionsor to division VII of Blanchard, but is not easy tocorrelate the classification of Casey with the oldauthors. Like Bates, Anomala rizotrogoides is

placed apart, as the type species of the new genus Anomalopus that also included A. tibialisSchaeffer. On the other hand, Spilota wasrecognized in the genus rank, divided into foursubgenera. All other genera of Anomalini

proposed previously were recognized as valid.The main advantage of the work of Casey is thatthe morphological criteria used by the division ofgenera, subgenera and sections, were described indetail. The characters used included general bodyshape and size, surface sculpture, color, details oftarsal claws, legs, mesosternum, mouth parts, aswell as geographical distribution [13].

Many authors after Casey said that suchclassification was much complex and excessive,including many over valuated importance ofcharacters included in the definition ofsupraspecific categories. As a result, many generaand subgenera were placed in synonymy.

Ohaus

Volume 20 of Coleopterorum Catalogus included3073 species of Rutelinae [21], an impressivedifference from previous 818 known species listedin previous works [22]. As part of such cataloguethe Anomalini were grouped in subgenera usingthe names previously assigned in the groups ofspecies of Burmeister. Near 700 species of Anomala were divided into four subgenera ( Aprosterna Hope, Anomala Samouelle, Euchlora MacLeayand Spilota Burmeister). Species in each subgenuswere subdivided according to geographicalregions. The American species were included inthree subgenera. Ohaus synonymized nearly allthe genera proposed by Casey ( Strigodermella ,

Alamona , Anomaleopta , Rhombonalia , etc.) and

because it includes a large number of species andmore morphological characters. Main charactersused for the genus rank are: the form of

tarsomeres and shape of tarsal claws, form oflabium and palpomeres, body shape and size,structures of mesosternum and details of elytralstriae. Proposals of Burmeister and Blanchardare much similar, because divisions VIII and IXcorrespond to the subgenera Anomala and Spilota .The differences are that Blanchard did notrecognize the subgenus Euchlora, and separate thedivision VII [16].

Lacordaire

A total of 13 genera of Anomalini from the world

were described in detail and placed in an old styletaxonomic key. No new classification is includedin his work, but useful morphological characterswere discussed and comments on the affinities ofeach genus were included. Main characters usedfor genus rank were exposition of mesepimeraand basal width of elytra. Genera from Oldand New World were mixed, and did not findstrong characters for the separation of Spilota ,Callistethus and Pachystethus from Anomala [11].

Bates

This big work dealt only with the species ofMexico and Central America, but increased muchknowledge on the Anomalini. The new genus

Dilophochila and the first American species ofPhyllopertha were described. Into the genus

Anomala 65 new species were described tocomplete 108 species recorded in the study area.Proposed classification is much distinct from the

preceding authors; the Mexican and CentralAmerican species were divided into three groupsaccording to the shape of tarsal claws. The firstgroup was divided into five sections attending tothe form and extension of the meso-metasternum.Sections II and III of Anomala proposed byBurmeister, and division VIII of Anomala in theclassification of Blanchard, were similar to thesections I and II in the book of Bates. The speciesincluded in the subgenus Spilota by Burmeisterand in the division IX of Blanchard, characterized

by the long mesosternal projection, are grouped inthe sections 4 and 5 of the first group of Anomala

proposed by Bates. Anomala rhizotrogoides remain isolated in a separate group supported bythe simple tarsal claws [12].


6/18


other groups of Rutelinae [18, 24]. Casey proposedthat complete pronotal margin is the primitivecondition. Genera that do not present such

character are Callistethus , Yaaxkumukia , Nayarita ,Pachystethus and Xochicotlia; incompletemargin is found in species of Strigoderma ; andmargin completely formed is always presentin Paranomala , Anomalorhina , Epectinaspis ,

Mazahuapertha , Callirhinus and Dilophochila .

2) Form and extension of the meso-metasternal process, or a wide intercoxal prominent space atthe middle of mesosternum, also has beenfrequently used as character for separation ofgenus or subgenus. The degree of development ofsuch structures is widely variable but in somegenera is easy to recognize some patterns, as inCallistethus and Yaaxkumukia the process isusually curved, much long and acutely pointed; inPachystethus , Callirhinus , Xochicotlia andStrigoderma is short, rounded and stout; in

Nayarita is long but narrowed from it base; in Epectinaspis is brief, weakly formed; meanwhilein Dilophochila , Anomalorhina and Paranomala is not developed.

3) Shape, position and extension of themesepimera is another frequently used character

all through the history of the group. In somegenera, as Strigoderma , is directed upwards; inother genera as Epectinaspis and Callirhinus iseasily visible from above.

Other characters has been used with lessfrequency by authors, some times weak changesnot confirmed in posterior works, such asmodifications in mouth parts [13], or in the formof tarsal claws [12], without other combinedcharacters, favored artificial grouping. Accordingto our phylogenetic analysis a great number ofnew characters have a strong phyletic mark, as themesothoracic wings, genitalia of two sexes, aswell as other meristic characters; the taxonomicvalue of some abandoned characters has beenreinterpreted, such as form and details in tarsalclaws and mouth parts.

All the new information used in diverse andvariable combinations allows to recognize fourmain lineages in the American fauna ofAnomalini, and gives us elements for proposinglimits in diagnosis of each genus (Table 2). Manycharacters are strongly correlated, as has been

do not accept the genus Phyllopertha as part ofthe fauna in America [1].

An interesting point is the inclusion of sevenspecies from tropical America as members of thesubgenus Aprosterna . Previously, Burmeister andBlanchard recorded the genus only for Asia. Onthe other hand, the species of the subgenusSpilota , previously cited only for America andAsia, were also registered for Australia andAfrica. But the main problem with this work isdealing with the absence of comments or data thatexplain the taxonomic changes, as is common inthe catalogues or checklists. So, that classificationneeds to be used with caution in the absence ofsupports.

Machatschke

In his works this author proposed synonymies for30 genera and subgenera described during the past100 years simplifying excessively the classificationof the Anomalini [23]. However, as the lastworldwide catalogue published was used asobligate reference of authority, but as is commonin other similar lists, due to the lack of details onthe criteria, do not have real taxonomic value.Some taxa were revalidated in recent studiesgiven precise diagnostic characters for theidentification and comparative studies, as withCallistethus and Pachystethus [9]. In hiscatalogues Machatschke listed Spilota as synonymof Callistethus but grouped the species not as

preceding authors did. Placed Dilophochila assynonym of Anomala , and listed Phyllopertha asvalid for New World fauna, including the newmonotypic genus Rugopertha for Honduras inCentral America. Into the genus Anomala was

proposed an elaborate scheme with 15 groups ofspecies, but without explicit criteria [2, 20].

Diagnostic characters in Anomalini

Despite the changes of criteria among distinctauthors cited above, it is possible to find a numberof common characters typical of the Anomalini,that aid to support the taxonomic ranks andto propose classificatory hypothesis. Mainmorphological characters used by various authorsare:

1) presence-absence and extension of the pronotal basal margin [13] of great taxonomic value also in


7/18


T a

b l e 2

. D i s t i n c t

i v e c h a r a c

t e r s i n

t h e

A n o m a l

i n i g e n e r a .

G e n e r a

B a s a l

m a r g i n

p r o n o t u m

A p i c a

l

m a r g i n

p r o n o t u m

M e s o s

t e r n a l

p r o j e c

t i o n

P a r a m e r e s

P a r a m e

r e s -

v e n t r a

l

p l a t e

M e s e p

i m e r o n

A p e x

e l i t r a l

s u t u r e

B a s a l

w i d t h

p r o n o t u m

S c u t e l

l u m

S e x u a l

d i m

o r p h

i s m

N a y a r

i t a

A b s e n

t

A b s e n

t

S m a l

l

P a r a

l l e l

F r e e

H i d d e n

R o u n d e d

B r o a d

T r i a n g u l a r

P r e s e n

t

X o c

h i c o

t l i a

A b s e n

t

C o m p l e t e

P r o m

i n e n

t ,

b r o a

d

P a r a

l l e l

P a r t i a l

l y

f u s e

d

S l i g h t l y

e x p o s e

d

S p i n e -

l i k e

B r o a d

T r i a n g u l a r

P r e s e n

t

C a l

l i r h i n u s

P r e s e n

t

C o m p l e t e

P r o m

i n e n

t ,

b r o a

d

P e r p e n

d i c u l a r

F r e e

E x p o s e

d

S p i n e -

l i k e

L i t t l e

n a r r o w

T r i a n g u l a r

P r e s e n

t

S t r i g o

d e r m a

P r e s e n

t

C o m p l e t e

N o t a b

l e ,

b r o a

d

S o m e

p e r p e n

d i c u l a r

F r e e

E x p o s e d ,

a s c e n d

i n g

S p i n e -

l i k e

N a r r o w

T r i a n g u l a r

P r e s e n

t

R u g o p e r t

h a

P r e s e n

t

C o m p l e t e

D e v e l o p e d

n . a .

n . a .

E x p o s e

d

n . a .

S l i g h t

n a r r o w

n . a .

n . a .

E p e c

t i n a s p i s

P r e s e n

t

C o m p l e t e

S l i g h t l y

b r o a

d

S o m e s l

i g h t l y

p e r p e n

d i c u l a r

F r e e

E x p o s e d

S p i n e -

l i k e

B r o a d

T r i a n g u l a r

P r e s e n

t

B a l a n o g o n i a

P r e s e n

t

C o m p l e t e

S l i g h t l y

b r o a

d

P e r p e n

d i c u l a r

F r e e

S l i g h t l y

e x p o s e

d

n . a .

B r o a d

n . a .

n . a .

P a c h y s

t e t h u s

A b s e n

t

I n c o m p l e t e

P r o m

i n e n

t ,

b r o a

d

P a r a

l l e l

F u s e

d

H i d d e n

S p i n e -

l i k e

B r o a d

T r i a n g u l a r

P r e s e n

t

C a l

l i s t e t h u s

A b s e n

t

I n c o m p l e t e ,

s o m e

c o m p l e t e

V e r y

l o n g ,

a c u t e

P a r a

l l e l

F u s e

d

H i d d e n

S p i n e -

l i k e

B r o a d

T r i a n g u l a r

S c a r c e

Y a a x

k u m u k

i a

A b s e n

t

I n c o m p l e t e

V e r y

l o n g ,

a c u t e

P a r a

l l e l

F r e e

H i d d e n

R o u n d e d

B r o a d

P a r a

b o l i c

S c a r c e

A n o m a l o r

h i n a

P r e s e n

t

C o m p l e t e

N a r r o w

P a r a

l l e l

F r e e

S l i g h t l y

e x p o s e

d

n . a .

B r o a d

P a r a

b o l i c

A b s e n t

D i l o p h o c

h i l a

P r e s e n

t

C o m p l e t e

A b s e n

t

P e r p e n

d i c u l a r

F r e e

H i d d e n

R o u n d e d

N a r r o w

P a r a

b o l i c

P r e s e n

t

M a z a h u a p e r t

h a

P r e s e n

t

C o m p l e t e

B r i e

f

P e r p e n

d i c u l a r

F r e e

S l i g h t l y

e x p o s e

d

R o u n d e d

N a r r o w

T r i a n g u l a r

P r e s e n

t

P a r a n o m a l a

P r e s e n

t

C o m p l e t e

A b s e n

t

P a r a

l l e l

F r e e

H i d d e n

R o u n d e d

B r o a d

P a r a

b o l i c

S c a r c e

D a t a n o

t a v a

i l a b l e

i n d i c a

t e d a s n . a .


8/18


Distribution. Restricted. Slopes of Tepetlilticvolcano, Santa Mara del Oro, Nayarit, and sierrade Mascota, Jalisco, Mexico.

Comments. Originally it was related withCallistethus and Anomala , but its position in therecent cladogram and some conspicuous charactersshared with Mimela and Anomala is clear thathave Palearctic affinities.

Genus Callistethus Blanchard, 1851

(Figures 2, 28, 32)

Type species: Mimela auronitens Hope, 1835: 114

Diagnosis. Body size big to medium; bodyconvex, weakly depressed, stout; six maxillary

teeth; pronotum without posterior margin, withincomplete anterior margin; pronotum usuallywith discrete fine punctuation; sutural apex ofelytron spiniform; epipleural fold wide; meso-metaesternum much projected reaching procoxae;dorsal coloration variable, but usually intensegreen. Male genitalia: genital capsule much curved,

parameres and ventral plate completely fused.

Distribution. Wide. Asia and America.

Comments. This genus is mainly found insoutheast of Asia and some authors do not accept

its representation in America, but we found phylogenetic evidences to support at least fewspecies of Central and northern South Americaas members of this genus [9]. Recent study ofthe type specimens of C. chrysanthe (Bates),C. phospora (Bates), C. specularis (Bates),C. xipostetha (Bates), C. pseudolepida Morn and

Nogueira, and C. tlapanecus Ramrez-Ponce andMorn confirm the existence of Callistethus intropical or subtropical areas of the New World [25].

Genus Yaaxkumukia Morn and Nogueira, 1998

(Figures 3, 29)Type species: Yaaxkumukia ephemera Morn and

Nogueira, 1998: 27

Diagnosis. Body size medium; body muchconvex, stout; six maxillary rounded teeth;

pronotum without posterior margin and withincomplete anterior margin; pronotum withdiscrete, much fine punctures; sutural apex ofelytron rounded; epipleural fold narrowed; meso-metasternum clearly projected until procoxae;males with last pair of respiratory stigma placed

observed by previous authors, facts that facilitatethe assembly of packages of character states torecognize groups with different taxonomic rank

(as the ascendant mesepimera associate with theexpansion of the epipleura [11] or the ascendantmesepimera related with the protuberatmesosternum [13]).

Other useful characters for supraspecific diagnosisare: the sexual dimorphism (expressed asenlargement of fore legs and antennae, expansionof epipleura); form of fore tarsal claws; form ofthe distal part of elytra; shape of the apex ofelytral suture; basal width of pronotum; form ofthe scutellum; exposition of prepygidium; lengthof metatarsomeres; and shape of clypeus. Withinmale genital capsule main useful supraspecificcharacters are: position of the parameres inreference to basal piece; tubular or flattened formof the parameres; and degree of fusion of theventral plate with parameres.

Comments on each American genus ofAnomalini

The following information is based on the originaldescriptions and in some regional brief reviews[3, 4, 5, 9]. The genus Popillia Dejean is

represented in America only by one introducedspecies, and the genus Phyllopertha Stephens wasconsidered as incertae sedis because the typespecimen was not found and no specimen similarto the description is available at present; bothgenera were commented in such reference and arenot included in this section [4].

Genus Nayarita Morn and Nogueira, 1998

(Figures 1, 13, 26, 30)

Type species: Nayarita viridinota Morn and Nogueira, 1998: 16.

Diagnosis. Body size big; body weakly convex,depressed, stout; clypeus with anterior borderwidened; five maxillary teeth; pronotum withoutanterior or posterior margins; pronotum with deepand dense punctuation; sutural apex of elytronrounded; epipleural fold much wide; meso-metaesternum projected; simple claws on middleand hind tarsi; dorsal coloration contrasted goldenyellow with metallic green. Male genitalia:

parameres distally curved; ventral plate long, bilobed, separated from parameres.


9/18


of species in America. Most interesting aspectsdeal with the new position of Leptohoplia Saylorand Chelilabia Morn and Nogueira as subgenera

of Paranomala , and additions to renew the groupof species capito sensu Machatschke [2] as thenew subgenus Bucaphallanus Ramrez-Ponce andMorn [41].

Genus Pachystethus Blanchard, 1851

(Figures 5, 25)

Type species: Popillia vidua Newman, 1838 bymonotypy.

Diagnosis. Body size small, body stout,depressed; six maxillary teeth; pronotum without

posterior margin and anterior margin incomplete; pronotal surface with moderate punctuation;sutural apex of elytron spiniform; epipleural foldwide; meso-metasternum wide with projection;contrasting coloration or completely black. Malegenitalia: parameres separated medially; ventral

plate completely fused with the parameres.

Distribution. Limited. East and southeast ofMexico to Guatemala.

Comments. This genus was revalidated byRamrez-Ponce and Morn [9], and the recentrevision included three new species [27]. Accordingto the phylogenetic analysis Pachystethus sharecharacters with Callistethus , Strigoderma ,

Balanogonia , Epectinaspis, Popillia and Callirhinus , as the increased and wide meso-metasternal projection, depressed body, elytranarrowed towards the apex, protarsi and protibiaewidened, and sutural apex of each elytronspiniform.

Genus Xochicotlia Ramrez-Ponce and Morn(unpublished)

(Figures 6, 16, 27)Type species Spilota micans Burmeister, 1844:269

Diagnosis. Body size small to moderate; bodystout, depressed; fourth antennomere muchelongate, five times longer than fifth antennomere;

posterior width of pronotum briefly shorter than basal width of elytra; mesepimera weakly exposeddorsally; elytra clearly reduced in width towardsthe apex, flattened, with deep striae; sutural apexof elytron spiniform; meso-metasternal space

on conspicuous tubercles; coloration dorsallyintense green, easy to lost by degradation,ventrally with submetallic shine. Male genitalia:

parameres distally expanded and curved, alignedwith basal piece; ventral plate short, separate from parameres. Distribution. Limited. Southeast ofMexico (Chiapas), Guatemala and Honduras.

Comments. After the description of this genus, thestudy of large series of specimens and thedescription of two new species, point out theimpact of deforestation in the mountains ofCentral America on the populations of theselimitedly distributed shiny chafers [26]. GenusYaaxkumukia is closely related to the species ofCallistethus ; one of the shared characters is thetuberculiform last pair of respiratory stigma, butin the former is bigger than in the latter, mainly inmales.

Genus Paranomala Casey 1915

(Figures 4, 10, 14, 19, 24, 31)

Type species: Melolontha marginata Fabricius,1792: 164

Diagnosis. Body size small to large, ovate orelongate, convex; fronto-clypeal suture complete;

pronotum with anterior and posterior borderscomplete; epipleural fold narrowed; sutural apexof elytron rounded; meso-metasternum withintercoxal space narrow or moderately wide butwithout projection; mesepimera not exposeddorsally. Coloration diverse, dark to shiny. Malegenitalia: parameres tubiform, aligned with basal

piece.

Distribution. Wide. America (southern Canada to Northern Argentina), Africa, India and southeasternAsia.

Comments. This genus was validated with supporton phylogenetic inferences to accommodate mostof the American species previously included in

Anomala Samoulle [9]. Such classificationreflects part of the scheme of Casey [13], whoexplained the differences between European andAmerican species proposed the name Paranomala for a number of species in the New World.Current unpublished results obtained by Ramrez-Ponce and Morn with phylogenetic analysis of81 taxa show topologies for well supported cladesthat will be ranked as four subgenera and 12 groups


10/18


Comments. Some of the species presently groupedin this new genus were associated with the oldname Spilota Dejean. Burmeister used Spilota assubgenus including Asiatic and American species[15, 19]. In the same usage Blanchard and Caseyonly listed American species, but Ohaus alsoadded species from Australia and Africa [16, 13, 1].

wide and prominent; prepygidium exposed in part.Male genitalia: parameres nearly cylindrical,longer than half of the length of tectum, partiallyfused with basal piece.

Distribution. Limited. Central and southernMexico, with some records in Panama and

Nicaragua.

Figures 1-12. Dorsal view of some American genera of Anomalini. 1. Nayarita Morn and Nogueira, 2. Callistethus Blanchard, 3. Yaaxkumukia Morn and Nogueira, 4. Paranomala Casey, 5. Pachystethus Blanchard, 6. Xochicotlia

Ramrez-Ponce and Morn, 7. Callirhinus Blanchard, 8. Epectinaspis Blanchard, 9. Strigoderma Burmeister,10. Paranomala ( Bucaphallanus ) Ramrez-Ponce and Morn, 11. Mazahuapertha Morn and Nogueira,12. Dilophochila Bates.


11/18


offer good support for such clade, but is better notto use the name Spilota in this genus-group

because the name of Dejean is in homonymy withSpilota Billberg, 1820 (Coccinelidae) and withSpilota Huebner, 1822 (Lepidoptera). Xochicotlia

Machatschke synonimized Spilota under Anomala , but Frey take it again for a largenumber of American species [2, 20, 28]. Thecombination of characters is different from othergroups of Anomalini, and phylogenetic analysis

Figures 13-33. Diagnostic characters of genera of American Anomalini. 13-18. Dorsal view of pronotum and elytrain: 13. Nayarita , 14. Paranomala , 15. Dilophochila , 16. Xochicotlia , 17. Callirhinus , 18. Strigoderma :19-22. Dorsal view of tibia and tarsus of left fore leg in: 19. Paranomala , 20. Mazahuapertha , 21. Epectinaspis ,22. Dilophochila . 23-25. Lateral view of male genital capsule in: 23. Callirhinus , 24. Paranomala , 25. Pachystethus .26-29. Lateral view of meso-metaesternal projection in: 26. Nayarita , 27. Xochicotlia , 28. Callistethus ,29. Yaaxkumukia . 30-33. Dorsal detail of the apex of left elytron in: 30. Nayarita , 31. Paranomala . 32. Callistethus y 33. Strigoderma .


12/18


Comments. Paucar-Cabrera reviewed the taxonomyof the genus and made phylogenetic analysis, atthe side detected paraphylic topology that support

the related genus Balanogonia [5]. Both generaand Strigoderma share a number of characters,such as the form of pronotum, extension ofmesepimera and elytral sculpture, but is easy toseparate with the diagnostic characters cited above.

Genus Strigoderma Burmeister, 1844

(Figures 9, 18, 33)

Type species: Strigoderma sulcipennisBurmeister, 1844: 316

Diagnosis. Body size medium; body narrowed,

with dorsal surface weakly flattened; sixmaxillary teeth; pronotum usually with completeanterior and posterior margins; posterior width of

pronotum shorter than basal width of elytra;mesepimera usually ascendant and prominent;sutural apex of elytron spiniform; epipleural foldwide; meso-metaesternal area wide, protuberant;coloration variable. Male genital: parameres andventral plate not fused.

Distribution. Wide. From United States to Argentina.

Comments. Historically, the taxonomic limits of

this genus have been unclear. Casey accept theutility of the typical body shape, but Potts remarksthat such character in some atypical species,induced to grouping these into erroneous groups,inclusive from the Old World [13, 10]. Badermade the alpha taxonomic revision of the genus,and also found evidences about the group not

being a natural genus, and is in need of phylogeneticanalysis [30]. At first hand, some species appearto be best situated in the genus Paranomala , butother species may form a new genus.

Genus Rugopertha Machatschke, 1957Type species: Phyllopertha sericeomicans

Nonfriend, 1891: 233 by monotypy.

Diagnosis. Body size small; body elongated;mesepimera exposed; mesosternal process brieflydeveloped; coloration bright green to cooperygreen.

Distribution. Restricted. Central Honduras.

Comments. Blanchard described Phylloperthavillosella and P. moreletiana from Mexico [16],

share characters with Pachystethus , Strigoderma and Callistethus .

Genus Callirhinus Blanchard, 1851

(Figures 7, 17, 23)

Type species: Callirhinus metallescens Blanchard,1851: 176

Diagnosis. Body size medium; body stout,depressed; clypeus narrowed and distally curvedas laminar projection; six maxillary teeth;

pronotum with anterior and posterior marginscomplete; epipleural fold widened; sutural apex ofelytron spiniform; meso-metasternal space wideand prominent; coloration diverse, contrasting

dark and brilliant. Male genitalia: parameresdepressed, plate-like, setose and perpendicular to basal piece; ventral plate long, separated from parameres.

Distribution. Limited. Western Mexico.

Comments. Wide variation in the color patterns ofC. metallescens , and one interesting feeding habiton sugarcane leaves were described and illustrated[29]. Machatschke placed Callirhinus close toother species of Anisoplina [2]. The possiblerelictual condition of the genus in western Mexico

was hypothesized [14], but the results of the phylogenetic analysis conduced by do not offergood support for the relation of Callirhinus with

Anisoplia . According with our recent studiesCallirhinus is part of the second lineage, relatedwith Strigoderma, Pachystethus and Xochicotlia .

Genus Epectinaspis Blanchard, 1851

(Figures 8, 21)

Type species: Anomala (Phyllopertha ) mexicana Burmeister, 1844: 241

Diagnosis. Body size medium; body stout, weaklydepressed; six maxillary teeth; clypeus of male briefly elevated; pronotum with anterior and posterior margins complete; anterior angles of pronotum acute, prominent; sutural apex ofelytron spiniform; epipleural fold wide; meso-metasternal area without projection; mesepimera

briefly seen from above. Male genitalia: parameres and ventral plate separate.

Distribution. Limited. Southern Mexico to northernVenezuela.


13/18

teeth; fifth antennomere with angulate keel; pronotum with both margins complete; pronotal basal width as the width of the base of elytra;

mesepimera hidden; sutural apex of elytronrounded; epipleural fold narrow; meso-metasternal area narrowed, briefly projected;

protibial spur absent; coloration contrasting, darkmetallic green on head, pronotum, and scutellum,and dark yellow on elytra. Male genitalia:

parameres plate-like, perpendicular to basal piece;ventral plate elongated, not fused with parameres.

Distribution. Restricted. Only known by typeseries from Toluca, state of Mexico, Mexico.

Comments. Original description detailed a

number of characters of Phyllopertha tolucana not shared by species of Phyllopertha in the OldWorld, that support their proposal for the genus

Mazahuapertha [3]. Our recent comparativestudies offer shared characters of Mazahuapertha with Epectinaspis , Strigoderma, Rugopertha ,Callirhinus and Balanogonia, as exposedmesepimera, narrowed base of pronotum, andmeso-metasternal process briefly developed.Other characters are shared with Strigoderma ,Callirhinus , Balanogonia and Dilophochila , suchas depressed, plate-like parameres, perpendicular

to basal piece, covered with short setae, as well asthe free, elongated ventral plate.

Genus Dilophochila Bates, 1888

(Figures 12, 15, 22)

Type species: Dilopochila bolacoides Bates 1888:261

Diagnosis. Body size small; body elongated,moderately convex; clypeus bilobated; threemaxillary teeth; pronotum with both marginscomplete; sutural apex of elytron rounded;

epipleural fold narrow, discrete; meso-metasternalarea small, not produced; mesepimera hiddenunder humeral calla. Male genitalia: parameres

plate-like, perpendicular to basal piece; ventral plate elongated, not fused with parameres.

Distribution. Limited. Southern Mexico (Chiapas,Oaxaca, Veracruz), Guatemala and Honduras.

Comments. This genus was listed as uniquemember of the subtribe Dilophochilina [31].Later, the genus-name was placed as synonym of

Anomala without listing D. bolacoides in any

and Bates argued that the genus Phyllopertha was badly defined, and placed such species as membersof Anomala and Epectinaspis , respectively, but

with reluctance described one new species inPhyllopertha (P. tolucana ) [12]. Some years later Nonfried described P. sericeomicans , thatMachatschke change to his newly proposed genus

Rugopertha [2]. Such species looks likeStrigoderma by the elongate body and theexposition of mesepimera, and particularlyresemble species of the group costulipennis by thedorsal surface finely rugo-punctate [4].

Genus Balanogonia Paucar-Cabrera, 2003

Type species: Epectinaspis freudei Frey, 1968:14

Diagnosis. Body size small; body slightlyelongated; six maxillary teeth; pronotum withcomplete anterior and posterior margins; anteriorangles of pronotum obtuse, not prominent;epipleural fold poorly developed; meso-metaesternal area small, not produced; base ofmesepimera briefly exposed near base of eachelytron. Male genitalia: parameres setose,

perpendicular to basal piece; ventral plate notfused with parameres.

Distribution. Restricted. Western Mexico (unique

precise record is from Colima volcano, Jalisco,Mexico).

Comments. The two species presently describedagree with the combination of charactersoriginally proposed, as the setose parameres are

perpendicular to basal piece [5]. This characteris also found in species of Strigoderma ,

Mazahuapertha , Callirhinus , some species ofParanomala and other Old World taxa such asPopillia and Anomala . A new phylogeneticanalysis including representatives of all American

genera of Anomalini is necessary to obtain a better hypothesis on the relationships of Balanogonia, Epectinaspis, Strigoderma andParanomala .

Genus Mazahuapertha Morn y Nogueira, 1999

(Figures 11, 20)

Type species: Phyllopertha tolucana Bates, 1888:216

Diagnosis. Body size small; body form stout,shortened; maxillary lacinia reduced to three



14/18


Middle and hind tarsal claws simple. WesternMxico (Nayarit) . Nayarita Morn and Nogueira

1 Clypeus with anterior border narrow (near aquarter of the length of clypeus in dorsalview). Maxilla with 2, 3, 4, 6 teeth. Anterior

pronotal margin present. Middle third oflateral border of each elytron some timesthickened in females. Only hind tarsal clawssimple. .. 2

2 Posterior margin of pronotum absent. .... 3

2 Posterior margin of pronotum present. ................. 6

3 Meso-metasternal projection long, with apexacute and/or curved, reaching posterior sideof procoxae. Body size medium to large (14 -22 mm). .................. 4

3 Meso-metasternal projection short, prominentand widely rounded, reaching at apex ofmesocoxae. Body size small (10 - 13 mm) ............. 5

4 Dorsal coloration bright emerald green. Lastabdominal spiracle tuberculiform. Malegenital capsule with parameres distallywidened, ventrally not fused one with theother or with ventral plate. .... Yaaxkumukia Mornand Nogueira

4 Dorsal coloration reddish brown, yellowish,green or bluish, with metallic or semimetallicluster. Last abdominal spiracle swollen orannular. Male genital capsule with parameresnot widened distally, ventrally fused betweenit and with ventral plate. Mexico, CentralAmerica and Colombia. ..Callistethus Blanchard

5 Male protarsi notably shortened and widened.Punctures on elytral striae small, simple.Parameres completely fused between it andwith ventral plate. Coloration variable, bright

black, reddish brown, dark brown, orangeyellow or bright yellow without metallicluster. Central and southern Mexico. ..... Pachystethus Blanchard

5 Male protarsi briefly shortened. Punctures onelytral striae large, umbiliform or annulated.Parameres nearly fused to ventral plate but notfused one with the other. Coloration greenish

group of species [20]. Posteriorly the type specieswas redescribed, revalidating the genus-name anddescribing five new species from Mexico,

Guatemala and Honduras, including a key for thesix species in the genus [3, 32]. As was discussed,no much characters are shared between

Dilophochila and other American genera, as pronotal base narrower than the elytral base (also present in Strigoderma and Mazahuapertha ),notably protibial widening (also in Strigoderma ,

Epectinaspis and Pachystethus ), and depressed, plate-like parameres perpendicular to basal piece (shared with Callirhinus , Strigoderma , Balanogonia and Mazahuapertha ) [3].

Genus Anomalorhina Jameson, Paucar-Cabreraand Sols, 2003

Type species: Anomala turrialbana Ohaus, 1928:393

Diagnosis. Body size small; body ovate elongated;clypeus notably upturned; frons of male with

basal tubercles; frontoclypeal suture erased atmiddle; six maxillary teeth; pronotum withcomplete anterior and posterior margins; pronotumof male with shallow depression on anteriormiddle; epipleural fold narrow, discrete; meso-metasternum with narrow process; base ofmesepimera briefly exposed near humeral calla.Male genitalia: parameres aligned with basal piece.

Distribution. Limited. Costa Rica and Nicaragua.

Comments. Ohaus originally described only onefemale. The peculiar form of body, labrum, hindtibiae and coloration, offer an isolated position forthis species between the Anomalini, but theaddition of male dimorphic characters completethe diagnosis for another genus, much distinctfrom other American shiny chafers. Without adetailed comparative study of the morphology of

A. turrialbana is not easy to hypothesized aboutits relationships within the Anomalini, but at firsthand look more related to the lineage of Paranomala .

Key to the genera of Anomalini from America. Modified from [3, 4, 5, 9].

1 Clypeus with anterior border much wide (nearhalf the length of clypeus in dorsal view).Maxilla with 5 teeth. Anterior pronotal marginabsent. Middle third of lateral border of eachelytron notably thickened in both sexes.


15/18

yellow or yellowish brown with intensemetallic luster. Central and southern Mexico,Panam and Nicaragua.

... Xochicotlia Ramrez-Ponce and Morn(unpublished)

6 Clypeus with anterior border bilobed. Apex of protibia ventrally with mesial proyection.Maxillary lacinia with 4 teeth. Protarsal innerclaw narrow, nearly as long as distal

protarsomere. Southern Mexico, Guatemalaand Honduras. . Dilophochila Bates

6 Clypeus with anterior border straight, roundedo projected. Apex of protibia ventrally withoutmesial projection.Maxillary lacinia with 2, 3

or 6 teeth. Protarsal inner claw wide, notablyshorter than distal protarsomere. ..... 7

7 Dorsal part of mesepimere hidden under elytralhumerus. Sexual dimorphism scarce.Southeastern Canada to Northern Argentina... Paranomala Casey

7 Dorsal part of mesepimere partially seen fromabove, not covered by elytral humerus. Sexualdimorphism notably in protibiae, protarsi andtarsal claws, elevation of clypeus and length ofantennal club. .. 8

8 Fronto-clypeal suture incomplete at middle.Lateral borders of clypeus clearly elevatedover the base of ocular canthus. Male withantero-mesial pronotal depresin. Female withV shaped posterior pronotal border. CostaRica and Nicaragua. ..... Anomalorhina Jameson,Paucar-Cabrera and Sols

8 Fronto-clypeal suture complete. Lateral borders of clypeus nearly flat or scarcely,elevated over the base of ocular canthus.Males and females with completely convex

pronotum, and posterior pronotal borderstraight or curved. ........................ 9

9 Anterior border of clypeus elongate, plate-like,narrowed and curved towards apex. CentralWestern Mexico. ...... Callirhinus Blanchard

9 Form of clypeal border variable, but notnarrowed to the apex. .... 10

10 Dorsal surface of pronotum and elytra finelyrugo-punctate. Elytra without striae or

longitudinal keels. Honduras. ...................................... Rugopertha Machatschke

10 Dorsal surface of pronotum and elytra with

variable punctuation, but not finely rugo- punctate. Elytral surface with punctate striae,with longitudinal keels or nearly smooth.....................................................................................................................11

11 Protibial spur absent. Fifth antennomere withnotably angled ventral keel. Toluca, Mxico... Mazahuapertha Morn and Nogueira

11 Protibial spur present. Fifth antennomerenearly cylindrical, without notably keel. 12

12 Mesosternum with wide intercoxal area.

Meso-metasternal projection more or lessnotably. Anterior border of clypeus semi-trapezoidal, nearly as the level of clypealdisk. USA. to Argentina. .... Strigoderma Burmeister

12 Mesosternum with narrow intercoxal area.Meso-metasternal projection weak or absent.Anterior border of clypeus rounded orsubquadrate, notably or moderately elevated...... 13

13 Pronotum with posterior border rounded,

anterior angles rounded, not reaching the posterior half of each eye. Anterior border ofclypeus moderately elevate. Hind wings withanterior region to RA 3+4 without setae.Southwestern Mxico. ...... Balanogonia Paucar-Cabrera

13 Pronotum with posterior border sinuate,anterior angles acute, projected, coveringmore than 1/3 of posterior part of each eye.Anterior border of clypeus notably elevate.Hind Kings with anterior region to RA 3+4setose. Southern Mexico to northernVenezuela. ... Epectinaspis Blanchard

Synopsis on the phylogeny of AmericanAnomalini

Using morphological characters of nearly all thegenera of Anomalini known to the Americas and arepresentative sample of many groups of speciesof Paranomala , we conduct two series of

phylogenetic analysis. The first series was madewith the aid of traditional heuristic search in



16/18


17/18

3. Morn, M. A. and Nogueira, G. 1998, FoliaEntomol. Mex., 103, 15.

4. Jameson, M. L., Paucar-Cabrera, A. andSolis, A. 2003, Ann. Entomol. Soc. Am.,96(4), 415.

5. Paucar-Cabrera, A. 2003, Coleopts. Soc.Monogr., 2, 10.

6. Potter, D. A. and Held, D. W. 2002, Annu.Rev. Entomol., 47, 175.

7. Morn, M. A. 2010, Plagas del suelo, L. A.Rodrguez y M. A. Morn (Eds.), Mundi-Prensa, Mxico, 41.

8. Morn, M. A. and Nogueira, G. 2002, FoliaEntomol. Mex., 41(1), 31.

9. Ramrez-Ponce, A. and Morn, M. A. 2009,Rev. Mex. Biodivers., 80, 357.

10. Potts, R. W. L. 1974, Pan-Pac. Entomol.,50, 148.

11. Lacordaire, T. 1856, Histoire Naturelle desInsectes, Librairie Enciclopedique de Roret.,Paris.

12. Bates, H. W. 1888, Biologia Centrali-Americana, Insecta Coleoptera, II (2),Taylor and Francis, London.

13. Casey T. L. 1915, Memoirs on theColeoptera, 6, 304.

14. Morn, M. A., Ratcliffe, B. C. and Deloya,C. 1997, Atlas escarabajos de Mxico,CONABIO, Mxico.

15. Burmeister, H. C. C. 1844, Handbuch derEntomologie, 4(1), Berlin.

16. Blanchard, E. 1851, Catalogue collectionentomologique, Coleopteres, I(1), Gidee andBaudry, Paris.

17. Potts, R. W. L. 1977, Pan-Pac. Entomol.,53, 34.

18. Arrow, G. J. 1917, The fauna of BritishIndia, Coleoptera, Lamellicornia, II, Taylorand Francis, London.

19. Burmeister, H. C. C. 1855, Handbuch derEntomologie, 4(2), Berlin.

20. Machatschke, J. W. 1957, GeneraInsectorum, 199-B, Ed. Mercurius, Anvers,Belgique.

21. Kohlmann, B. and Morn, M. A. 2003, ActaZool. Mex. (n.s.), 90, 175.

22. Gemminger, M. and Harold, E. 1869,Catalogus coleopterorum, 4, Monachii.

23. Mic, E. 2001, Los escarabeidos antfilosde la pennsula ibrica, UniversidadAlicante, Espaa.

molecular evidence and precise ecologicalinformation will be also important to complete thehypothesis of classification. We expect that in

future new tests the approximations will be progressively refined and better supported than present proposals.

Exposed results also indicate significant advancesin the description and definition of supraspecifictaxa that aid to recognize particular problems.Some of these problems have been included inrecent papers and others are nearly ready for

publication. Other questions require more detailedstudies, more specimens to review using freshcriteria and open mind. In our experience oldideas also will be useful with renewed points ofview. Our results include a large number ofcharacters never used before, as well as manycharacters continuously used since years back andother forgotten by some authors. Some old ideasabout the structure of classification of shinychafers were refreshed, gaining good support tothe actual hypothesis.

One of the main problems dealing with the studyof these beetles is in the collections. Nearly all themuseums and institutions have large number ofspecimens mounted in the cabinets, but most

part are poorly studied, not determined or provisionally identified. This fact reflects thereduced number of taxonomists interested in thisabundant and diverse, but complex group.Unfortunately during our studies we had at handvery few correctly identified specimens fromsoutheastern Asia or sub-Saharan Africa to makecomparative descriptions, useful to confirm some

problems with the genera Anomala, Paranomala and Callistethus .

Our main current objectives deal with the

supraspecific classification of the large genusParanomala in the Neotropical realm, includingthe definition, revision and phylogeny of eachsubgenus, the groups of species and the descriptionof many new species.

REFERENCES

1. Ohaus, F. 1918, Coleopterorum Catalogus,20(66), 124.

2. Machatschke, J. W. 1972, ColeopterorumCatalogus Supplementa, 66(2), 363.



18/18

33. Goloboff, P. A. 1999, NONA Ver., 2,Tucuman, Argentina.

34. Goloboff, P. A, Farris, J. S. and Nixon, K.

C. 2008, Cladistics, 24, 774.35. Goloboff, P. A. 1993, Cladistics, 9, 83.36. Hillis, D. M. 1998, Syst. Biol., 47, 3.37. Poe, S. 1998, Syst. Biol., 47, 18.38. Pollock, D. D., Zwickl, D. J., McGuire,

J. A. and Hillis, D. M. 2002, Syst. Biol.,51(4), 664.

39. Zwickl, D. J. and Hillis, D. M. 2002, Syst.Biol., 51, 588.

40. Pol, D. and Escapa, H. I. 2009, Cladistics,25, 515.

41. Ramrez-Ponce, A. and Morn, M. A. 2012,Ann. Entomol. Soc. Am., 105, 781.

24. Jameson, M. L. 1997, Bull. Univ. NebraskaState Mus., 14, 84.

25. Ramrez-Ponce, A. and Morn, M. A. 2012,

Rev. Bras. Entomol., 56, 142.26. Mic, E., Gmez, B. and Galante, E. 2006,Ann. Entomol. Soc. Am., 99, 1.

27. Ramrez-Ponce, A. and Morn, M. A. 2012,Zootaxa, 3394, 1.

28. Frey, G. 1968, Ent. Arb. Mus. Frey, 19, 281.29. Morn, M. A. and Hernndez-Rodrguez, S.

1996, G. it. Ent., 8, 105.30. Bader, A. M. 1992, T. Am. Entomol. Soc.,

118(2), 269.31. Blackwelder, R. 1944, US Nat. Mus. Bull.,

185, 243.

32. Morn, M. A. and Howden, H. F. 2001,Coleopts. Bull., 55(1), 51.


morón y ramírez-ponce, 2012. mesoamerican genera of anomalini

Documents