Locomotion and posture in Lagothrix lagotricha

This long-term study of woolly monkey (Lagothrix) locomotor and postural behaviour employs methods identical to those used during a previous study of the locomotion and posture of two species of Ateles, allowing a detailed comparison between the two genera, which are strong competitors in extensive parts of the Amazon basin and northern Andes. As in Ateles, Lagothrix locomotion can be divided into five patterns, based on limb usage: quadrupedal walking and running, suspensory locomotion, climbing, bipedalism (very rare in wild woolly monkeys) and leaping. Lagothrix differs from Ateles primarily in its greater reliance on quadrupedal locomotion during both travel and feeding and on its de-emphasis of the use of suspensory locomotion as compared to Ateles, while the use of climbing and leaping is roughly equal in the two genera. Lagothrix exhibits more generalised (primitive) locomotive behaviour in accordance with its morphology, in comparison to the more specialised Ateles. The generic differences reflect differences in habitat use and particularly foraging ecology.     

Phylogenetic inferences of Atelinae (Platyrrhini) based on multi-directional chromosome painting in Brachyteles arachnoides, Ateles paniscus paniscus and Ateles b. marginatus

We performed multi-directional chromosome painting in a comparative cytogenetic study of the three Atelinae species Brachyteles arachnoides, Ateles paniscus paniscus and Ateles belzebuth marginatus, in order to reconstruct phylogenetic relationships within this Platyrrhini subfamily. Comparative chromosome maps between these species were established by multi-color fluorescence in situ hybridization (FISH) employing human, Saguinus oedipus and Lagothrix lagothricha chromosome-specific probes. The three species included in this study and four previously analyzed species from all four Atelinae genera were subjected to a phylogenetic analysis on the basis of a data matrix comprised of 82 discrete chromosome characters. The results confirmed that Atelinae represent a monophyletic clade with a putative ancestral karyotype of 2n = 62 chromosomes. Phylogenetic analysis revealed an evolutionary branching sequence [Alouatta [Brachyteles [Lagothrix and Ateles]]] in Atelinae and [Ateles belzebuth marginatus [Ateles paniscus paniscus [Ateles belzebuth hybridus and Ateles geoffroyi]]] in genus Ateles. The chromosomal data support a re-evaluation of the taxonomic status of Ateles b. hybridus.     

RFLP analysis of mtDNA from six platyrrhine genera: phylogenetic inferences

This study investigates the phylogenetic relationships of 10 species of platyrrhine primates using RFLP analysis of mtDNA. Three restriction enzymes were used to determine the restriction site haplotypes for a total of 276 individuals. Phylogenetic analysis using maximum parsimony was employed to construct phylogenetic trees. We found close phylogenetic relationships between Alouatta, Lagothrix and Ateles. We also found a close relationship between Cebus and Aotus, with Saimiri clustering with the atelines. Haplotype diversity was found in four of the species studied, in Cebus albifrons, Saimiri sciureus, Lagothrix lagotricha and Ateles fusciceps. These data provide additional information concerning the phylogenetic relationships between these platyrrhine genera and species.     

Delimitation of major lineages within Cuscuta subgenus Grammica (Convolvulaceae) using plastid and nuclear DNA sequences

Subgenus Grammica, the largest and most diverse group in the parasitic genus Cuscuta, includes 130 species distributed primarily throughout the New World, with Mexico as its center of diversity. To circumscribe the subgenus and assess the relationships among its major lineages, we conducted the first phylogenetic study of Grammica using plastid trnL-F and nrITS sequences from a wide taxonomic sampling covering its morphological, physiological, and geographical diversity. With the exception of one species belonging elsewhere, the subgenus was found to be monophyletic. The results further indicate the presence of 15 well-supported major clades within Grammica. Some of those lineages correspond partially to earlier taxonomic treatments, but the majority of groups are identified in this study for the first time. The backbone relationships among major clades, however, remain weakly supported or unresolved in some cases. The phylogenetic results indicate that the fruit dehiscence character is homoplastic, thus compromising its value as a major taxonomic and evolutionary feature. While several striking cases of long-distance dispersal are inferred, vicariance emerges as the most dominant biogeographical pattern for Cuscuta. Species placed within one of the clades with a predominantly South American distribution are hypothesized to have substantially altered plastid genomes.     

Social Behavior,Reproductive Strategies,and Population Genetic Structure of <Emphasis Type="Italic">Lagothrix poeppigii</Emphasis>

For species of primates in which females emigrate, we would expect males within groups to be related to one another. Kin selection theory suggests that these males should associate preferentially with one another, be more affiliative and cooperative with one another than females are, and compete less overtly with one another over reproductive opportunities than males in female philopatric taxa do. Precisely these patterns of social behavior characterize well-studied populations of 2 of the 3 atelin primate genera: spider monkeys (Ateles) and muriquis (Brachyteles). For the third atelin genus, Lagothrix, patterns of intragroup social behavior have been less well-documented. We studied the social and reproductive behavior of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in Ecuador during a one-year observational study and subsequently used molecular techniques to investigate population genetic structure and dispersal patterns for this taxon. Among adult male woolly monkeys, both affiliative and agonistic interactions were rare, and males were seldom in close proximity to one another. Relationships among male woolly monkeys are best characterized as tolerant, especially in the context of mating wherein direct competition among males was minimal despite the fact that females mated with multiple males. Relationships among females were likewise generally tolerant but nonaffiliative, though females often directed harassment towards copulating pairs. Affiliative interactions that did occur among woolly monkeys tended to be directed either between the sexes—primarily from female to male—or from younger towards older males, and the proximity partners of females tended to be members of the opposite sex. These results suggest that bonds between the sexes may be more important than same-sex social relationships and that direct female-female competition is an important feature of woolly monkey reproductive biology. Our genetic results indicate that, as in other atelins, dispersal by females is common, but some male dispersal likely occurs as well. In some but not all groups we studied, nonjuvenile males within social groups were more closely related to one another on average than females were, which is consistent with greater male than female philopatry. However, differences in these patterns among our study groups may reflect local variation in dispersal behavior.     

The natural place to begin: The ethnoprimatology of the Waorani

Ethnoprimatology is an important and growing discipline, studying the diverse relationships between humans and primates. However there is a danger that too great a focus on primates as important to humans may obscure the importance of other animal groups to local people. The Waorani of Amazonian Ecuador were described by Sponsel [Sponsel (1997) New World Primates: Ecology, evolution and behavior. New York: Aldine de Gruyter. p 143–165] as the “natural place” for ethnoprimatology, because of their close relationship to primates, including primates forming a substantial part of their diet. Therefore they are an ideal group in which to examine contemporary perceptions of primates in comparison to other types of animal. We examine how Waorani living in Yasuní National Park name and categorize primates and other common mammals. Although there is some evidence that the Waorani consider primates a unique group, the non‐primate kinkajou and olingo are also included as part of the group “monkeys,” and no evidence was found that primates were more important than other mammals to Waorani culture. Instead, a small number of key species, in particular the woolly monkey (Lagothrix poeppigii) and white‐lipped peccary (Tayassu pecari), were found to be both important in the diet and highly culturally salient. These results have implications for both ethnoprimatologists and those working with local communities towards broader conservation goals. Firstly, researchers should ensure that they and local communities are referring to the same animals when they use broad terms such as “monkey,” and secondly the results caution ethnoprimatologists against imposing western taxonomic groups on indigenous peoples, rather than allowing them to define themselves which species are important. Am. J. Primatol. 75:1117–1128, 2013. © 2013 The Authors. American Journal of Primatology Published by Wiley Periodicals, Inc.     

Double invasion of Tertiary island South America by ancestral New World monkeys?

Comparative Determinant Analysis was used to characterize antigenic determinants from 69 functionally diverse serum proteins for the phylogenetic study of anthropoid cladogenesis, of which 32 homologues with 66 antigenic determinants were compared between Lagothrix lagotricha and Cebus albifrons. Nine epitopes (one each of complement factors Cls, C3, C6, C7, C9, factor B, retinol-binding protein, β₂-glycoprotein III, and histidine rich α₂-glycoprotein) defined a robust clade uniting Lagothrix lagotricha with the Old World primates Macaca, Papio and Homo to the exclusion of Cebus albifrons. This tree indicates platyrrhine paraphyly or demonstrates an accelerated antigen substitution rate of Cebus when compared with Lagothrix. Narrow phylogenetic sampling does not permit the final resolution of this problem but strongly different evolutionary rates are considered unlikely because ancestral serum protein determinants, which are plesiomorphic for Primates as an order, are equally conserved in both Cebus and Lagothrix. If platyrrhine paraphyly is correct, our molecular-immunological clock estimates a divergence date of Cebus of 52.5 × 10⁶ years before present, i.e. 6.5 × 10⁶ years before the ateline clade diverged (possibly at 46.0 × 10⁶ years BP) from catarrhine ancestors. A scenario of platyrrhine paraphyly implies the necessity for double invasion of the isolated Tertiary South American island continent by primates across open ocean, and suggests a considerably earlier phylogenetic emergence of platyrrhines than is proposed by the oldest simian fossils from the Neotropics.     

Fruit Choice by Woolly Monkeys in Tinigua National Park,Colombia

I combined morphological fruit data, phenological, and demographic information for 128 plant species, with nutritional information for 78 species, to assess feeding preferences of woolly monkeys (Lagothrix lagothricha lugens) in Tinigua National Park, Colombia. I used multiple and simple regression analyses to predict fruit feeding time and one index of fruit preference from 20 independent variables presumed to affect fruit choice. The results of the independent regressions indicated a trend to spend more time consuming good tasting fruits from abundant plants that produce large quantities of fruits and produce large crops. Among these variables, abundant fruit production and the astringency index are the most important variables explaining feeding times when other variables are statistically controlled in a multiple regression analysis. Furthermore, the preference index showed that woolly monkeys prefer fruits from large trees that produce in periods of fruit scarcity, with high pulp/seed ratios, low lipid contents, and clumped spatial distributions. The results suggest that woolly monkeys are primarily generalized, opportunistic frugivores that avoid some fruits with secondary compounds. Their feeding behavior is determined by many factors, but the predictive power of each one is always low (<15%). No multiple regression model explained >69% of the variation in feeding time. I suggest that a detailed quantification of secondary compounds might increase the predictive power of the models to explain fruit choice by frugivorous primates.     

Yellow-tailed woolly monkey (Lagothrix flavicauda) proximal spacing and forest strata use in La Esperanza,Peru

Inter-individual spacing of primates and their use of forest strata depend on temporal and spatial changes in the environment and on predator avoidance, group demography, and social conditions. Greater proximity also increases the chances of agonistic and affinitive contact between individuals. I collected behavioral data for three groups of yellow-tailed woolly monkeys (Lagothrix flavicauda) by instantaneous sampling of focal animals for 15 months in La Esperanza, northeastern Peru. By use of combined data for all groups I examined the effects of season, activity, and age/sex class on nearest-neighbor distances and forest strata use. Small differences were observed for nearest-neighbor distances, forest strata use by different age/sex classes, and activity. Adult males had the lowest contact index scores. Contact index scores were low for juvenile females, for which nearest-neighbor distances were largest. Very little aggressive behavior was observed. Focal animals preferred upper levels of the forest with little difference in height for different activities. Lagothrix flavicauda have very cohesive groups with little seasonal or activity-dependent difference between nearest-neighbor distances or proximity. These results suggest that this species has less variable social organization and greater group cohesion than other Atelini. However, more studies are needed on other populations of L. flavicauda to better determine the species’ social organization. Studies are also required to determine the extent to which dispersal times and kinship affect proximity, nearest-neighbor distances, and aggression.     

Phylogeny of Philippine slender skinks (Scincidae: Brachymeles) reveals underestimated species diversity, complex biogeographical relationships, and cryptic patterns of lineage diversification

The spectacular, virtually endemic radiation of Philippine semi-fossorial skinks of the genus Brachymeles represent one of the few radiations of scincid lizards to possess both fully limbed and limbless species. And yet, nothing is known of the phylogenetic relationships of this exceptional group. Morphologically similar body plans have made it difficult to assess species-level diversity, and the genus has long been recognized as one of the more modest radiations of southeast Asian lizards. However, recent large-scale survey efforts have resulted in the discovery of numerous new species, and taxonomic studies indicate that the diversity within the genus Brachymeles is grossly underestimated. In this study we provide the first robust estimate of phylogenetic relationships within the genus Brachymeles using a multi-locus dataset and nearly complete taxonomic sampling. We provide statistical tests of monophyly for all polytypic species and two widespread limb-reduced species and our results indicate wholesale deviations from past summaries and taxonomic evaluations of the genus. With few exceptions, we are able to reject the monophyly of all polytypic and widespread species, thereby validating the need for large-scale taxonomic revisions. Our results reveal that the limbless, monotypic, genus Davewakeum is nested within Brachymeles. Mapping of body form on our preferred phylogenetic tree suggests that limb-reduction and digit loss has occurred on multiple occasions in the history of the genus. A Bayesian reconstruction of ancestral areas indicates strong statistical support for a minimum of five major dispersal events that have given rise to a major component of the observed species diversity on separate Pleistocene aggregate island platforms of the archipelago.     

Ranging behavior and foraging ecology of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in Yasuní National Park,Ecuador

In a year-long study, I investigated the ranging behavior of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in a terra firma rainforest in Yasuní National Park, Ecuador, and examined the relationship between ranging, diet, food availability, and food patch use for this population. In Yasuní the total home range sizes for two social groups were 124 and 108 ha, which are much smaller than has been reported previously for Lagothrix elsewhere in its geographic distribution. The mean yearly day range estimates for these same groups were 1,792 m and 1,878 m, which are well within the range of variation previously reported. Ranging behavior was not correlated with the current habitat-wide abundance of ripe fruit, which comprises 76.3% of the yearly diet for this population, but was associated with one measure of likely insect prey abundance and with the availability of immature fruits, a minimal part of the diet. Specifically, one study group moved significantly greater distances during months of high likely insect prey abundance and when immature fruits were abundant. The second study group also traveled farther when likely insect prey abundance was high and when immature fruits were abundant, although the latter relationship only approached significance. This group also devoted significantly more of its daily activity budget to travel during these times. These results indicate that variation in ripe fruit abundance makes no meaningful contribution to explaining variation in ranging behavior for this population of woolly monkeys. Instead, the results raise the possibility that some aspects of the ranging behavior of frugivorous primates may be related to the availability of alternative food sources, such as animal prey, or to monitoring the phenological status of important fruit trees, rather than simply reflecting the degree of intragroup feeding competition.     

Effects of Seed Dispersal by Three Ateline Monkey Species on Seed Germination at Tinigua National Park, Colombia

We examined the effect of seed ingestion by three ateline primates: woolly monkeys, Lagothrix lagothricha; spider monkeys, Ateles belzebuth; and, red howler, Alouatta seniculus on germination rates and latency periods of seeds of several plant species in Tinigua National Park, Colombia. We collected dispersed seeds from feces and control seeds from the parental trees and washed them for germination trials. For the majority of plants, dispersed seeds germinated as well or better than control seeds did. Although spider monkeys depend more heavily on fruits than the other monkey species do, they were not more efficient than howlers or woolly monkeys at improving germination rates. A considerable proportion of the seeds dispersed by howlers and woolly monkeys showed reduced latency periods to germination, but spider monkeys showed less effect on reducing germination time. This result may be related to longer gut retention times, but such a trend has not been observed in other primate species. We conclude that, like many other primates, ateline monkeys are effective seed dispersers in terms of their effects on the seeds they swallow because they rarely decrease their germination rates. We discuss problems that make interspecific comparisons difficult, such as inappropriate control seeds and differences associated with germination substrates, and we stress the importance of studying other components of seed dispersal effectiveness.     

Time Allocation Patterns of Lowland Woolly Monkeys (Lagothrix lagotricha poeppigii) in a Neotropical Terra Firma Forest

We investigated the time allocation decisions of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in a terra firma forest in eastern Ecuador where they occur sympatrically with 9 other primate species. Woolly monkeys spent considerable amounts of time searching for and attempting to procure animal prey—roughly as much time as they spent consuming plant material: ripe fruits, leaves, and flowers. The amount of time spent foraging for animal prey is positively related to the habitat-wide availability of ripe fruits (the predominant component of the woolly monkey diet), and negatively related to both ambient temperature and the abundance of potential prey items in the habitat. Time spent resting showed exactly the opposite pattern with respect to these ecological variables. These results suggest that woolly monkeys follow an energy-maximizing strategy of food acquisition during times of fruit abundance—focusing on animal foods and perhaps laying down fat reserves to utilize when ecological conditions worsen—and follow an energy-minimizing strategy when fruit resources are scarce. Such a strong and seasonal commitment to animal prey foraging is unique among the ateline primates and is not ubiquitous even among lowland woolly monkeys. We suggest that this foraging strategy, and the greater intragroup cohesion that characterizes some populations of Lagothrix, are both opportunistic responses to regional differences in habitat quality. Identifying and accounting for such intraspecific variation should be a goal of any analysis of comparative socioecology.     

Atelinae phylogenetic relationships: The trichotomy revived?

This research examines phylogenetic relationships between members of the Atelinae subfamily (Alouatta, Ateles, Brachyteles, and Lagothrix), based on analysis of three genetic regions. Two loci, cytochrome c oxidase subunit II (COII) and the hypervariable I portion of the control region, are part of the mitochondrial genome. The other is a single-copy nuclear gene, Aldolase A Intron V. Analysis of these genetic regions provides support for tribe Alouattini containing the Alouatta species, while tribe Atelini contains the other three genera. However, these three genetic regions produce conflicting results for relationships among tribe Atelini members. Previous genetic studies supported grouping Brachyteles with Lagothrix, leaving Ateles in a separate subclade. The present data sets vary based on the genetic region analyzed and method of analysis suggesting all possible cladistic relationships. These results are more consistent with investigations of morphology and behavior among these primates. The primary cause of discrepancy between this study and previous genetic studies is postulated to reside in increased sampling in the present study of genetic variation among members of the Atelinae, specifically Ateles. The present study utilized samples of Ateles from all postulated species for this genetically variable primate, while previous studies used only one or two species of Ateles. This paper demonstrates that shifting relationships are produced when different species of Ateles are used to reconstruct phylogenies. This research concludes that a trichotomy should still be supported between members of tribe Atelini until further analyses, which include additional Atelinae haplotypes are conducted.     

Revival of the genus Tropicoperdix Blyth 1859 (Phasianidae,Aves) using multilocus sequence data

Although Tropicoperdix has been considered to be either a full genus or a species complex within the Phasianid genus Arborophila (hill partridges), there is long‐standing uncertainty regarding the degree of difference that warrants generic separation, including reported anatomical cranial differences. In addition, the intra‐generic taxonomy remains under dispute. Most studies hypothesize that Tropicoperdix comprises three species, while others postulate from one to four species. However, no molecular study has been performed to clarify the systematic and taxonomic uncertainties surrounding Tropicoperdix. In the present study, we performed a series of molecular phylogenetic analyses of Tropicoperdix and Arborophila taxa based on two mitochondrial genes and five nuclear introns. All the results are consistent with the finding that Tropicoperdix and Arborophila are phylogenetically distinct and distant genera, although the precise phylogenetic position of Tropicoperdix remains undetermined. Retrospective examination of external characteristics also supports the generic separation, as well as providing evidence of remarkable multiple character convergence. We propose that Tropicoperdix comprises at least two full species based on mitochondrial data obtained from museum specimens by using a next‐generation sequencing method. © 2015 The Linnean Society of London     

A taxonomic revision of Paradoris sea slugs (Mollusca, Gastropoda, Nudibranchia, Doridina)

Paradoris , traditionally a generic taxon of discodorid sea slugs, is revised for the first time. One hundred and thirty specimens were examined, including all types and most of the nontype specimens available. New records for Paradoris are provided: South Africa, Tanzania, Seychelles, Western Australia, Thailand, Philippines, shallow waters of New Caledonia, southern Japan, and Hawaii. The individual variation of all taxonomic characters is thoroughly evaluated. Prior to the present study, Paradoris included 15 nominal species: 12 valid names and three synonyms. Three additional names, Discodoris erythraeensis , D. lora , and D. cavernae , are re-allocated to Paradoris , based on phylogenetic analysis. A phylogenetic diagnosis is provided for Paradoris : in particular, two new synapomorphies are described. Eight species names are regarded as valid: P. araneosa , P. dubia , P. erythraeensis , P. indecora , P. liturata , P. lopezi , P. mulciber , and P. tsurugensis . However, most of these species are poorly known, i.e. from very few specimens, and their taxonomic status might change when more individuals are available. P. lora is regarded as a nomen dubium . Six new synonymies are proposed, and explained by the fact that: (1) species names were created for one or a few specimens, without considering individual variation; (2) authors have not worked within a phylogenetic framework and have created new species names without considering all the existing species names already available within Paradoris . Three new morphospecies are described, but not formally named because their taxonomic status is still uncertain for several reasons (e.g. lack of knowledge of individual variation for some critical features).  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 147 , 125–238.     

Footfall patterns, stride length and speed of vertical climbing in spider monkeys (Ateles fusciceps robustus) and woolly monkeys (Lagothrix lagotricha)

Vertical climbing is central to theories surrounding the locomotor specialisations of large primates. In this paper, we present spatiotemporal gait parameters obtained from video recordings of captive spider monkeys (Ateles fusciceps robustus) and woolly monkeys (Lagothrix lagotricha) in semi-natural enclosures, with the aim of discovering the influence of body weight and differences in general locomotor behaviour on vertical climbing kinematics on various substrates. Results show that there are only few differences between gait parameters of climbing on thin trees, vertical and oblique ropes, while climbing on large-diameter trees differs considerably, reflecting the higher costs of locomotion on the latter. At the same speed, Ateles takes longer strides and the support phase takes a smaller percentage of cycle duration than in Lagothrix. Footfall patterns are more diverse in Ateles and include a higher proportion of ipsilateral limb coupling. Compared to other primates, the gait characteristics of vertical climbing of atelines most closely resemble those of African apes.     

Primate Abundance in an Unhunted Region of the Northern Peruvian Amazon and the Influence of Seismic Oil Exploration

The western Amazon, a relatively remote and unstudied region, is experiencing unprecedented levels of oil and gas exploration. Despite the widespread use of seismic reflection technology for oil and gas exploration, no studies have investigated the response of primate populations to this disturbance in the Amazon. We conducted distance sampling along transects in pristine, unhunted lowland rain forest inside a large oil concession (Block 39) in the northern Peruvian Amazon with ongoing 2D seismic exploration. We aimed to investigate seismic exploration effects on local primate abundance, with a particular focus on the region’s most endangered primates, lowland woolly monkeys (Lagothrix poeppigii) and white-bellied spider monkeys (Ateles belzebuth). We sampled transects before the arrival of exploration crews (control) and during the seismic operation (disturbance) and compared primate counts using mixed-effect models. We confirmed the presence of nine species of primates along sampled transects. Abundance of primate groups (species pooled) was not different between the control and disturbance period, yet abundance of primate individuals was significantly lower in the disturbance period. Although we encountered groups of Ateles belzebuth equally often during the control and disturbance periods, overall counts of individuals of this species were lower during the disturbance period owing to lower subgroup sizes. This suggests this species may have responded, at least temporarily, with localized spatial avoidance, and that it may be particularly sensitive to human disturbance regardless of hunting pressure. Our overall density estimate for Lagothrix poeppigii (30.6/km²) approaches the highest reported for the species. Although the relatively temporary nature of 2D seismic exploration may limit its effect on primate species, our data indicate some species may respond more negatively than others.     

Review on the use of sexually dimorphic characters in the taxonomy of Diabroticites (Galerucinae,Luperini, Diabroticina)

Sexual dimorphism occurs frequently in Chrysomelidae Latreille, 1802 and is particularly variable in subfamily Galerucinae Latreille, 1802. This diversity has been early noted by authors a potential source of taxonomic characters. The section Diabroticites (Luperini Gistel, 1848) is one of the largest assemblies of chrysomelid genera with currently 823 valid species in 17 genera (12 based on dimorphic characteristics), being most diverse in the neotropical region. Apart from a revision work on the type specimens for the section, there are no general taxonomic studies for this group. The occurrence of sexually dimorphic characteristics in the section Diabroticites is revised and their practical taxonomic relevance evaluated. A total of 240 species was studied (145 species with males available), representing 15 out of the 17 genera included in Diabroticites. The analysis of characters was based on the study of specimens in south-american collections, literature and the aid of photos in online databases. Sexual dimorphism occurred in most species analyzed. Dimorphic features were divided in general (i. e., occur in higher taxa) and special characters (those that support the definition of species and genera). Special dimorphism was observed in every tagma, and most modifications occur in antennae. Characters used as diagnostic of genera often do not correspond to the modifications present in species included in them. Many modifications were considered by earlier authors as a single character, probably due to vague definitions. Most generic definitions are, therefore, inaccurate. The study of morphology and the homology assessment of characters are needed to increase understanding of the genera in Diabroticites.