Ant Diversity Research Articles

Ant Diversity Declines with Increasing Elevation along the Udzungwa Mountains, Tanzania

Biodiversity patterns along elevational gradients are generally characterised by monotonic decreases or mid-elevational peaks in species richness, while elevational zones may be characterised by distinct assemblages, or higher zones may be subsets of lowland assemblages. Elevational gradients in diversity have been less studied in the Afrotropical region. This study documents ant diversity patterns in three forest types associated with the tropical mountains of Udzungwa; we hypothesise that: (1) ant diversity and activity will show a monotonic decrease from mid-elevation with increasing elevation and (2) that forests associated with different elevations will have a distinct ant assemblage. Pitfall traps were deployed at three targeted elevations (650–800, 800–1400, and 1400–1500 m a.s.l.). Ant species richness declined with increasing elevation from 650 m a.s.l. and formed three elevational assemblages with lower elevation forests having almost twice as many species as sub-montane forests and three times as many as that of the montane forests. In contrast, overall ant activity peaked at 800–1400 m a.s.l. The ant assemblages associated with the lower elevation forest were very distinct, while assemblages associated with the sub-montane and montane forests shared species. Our study reveals valuable and relevant information for biodiversity monitoring and conservation planning as the species associated with each forest type may be used as indicator species for assessing biodiversity responses to climate change and anthropogenic activities on these mountains.

Diversity and Seasonal Occurrence of Native and Nonnative Ants (Hymenoptera: Formicidae) in Long-Term Experimental Chinese Privet (Lamiales: Oleaceae) Plots in Georgia, USA

Diversity and Seasonal Occurrence of Native and Nonnative Ants (Hymenoptera: Formicidae) in Long-Term Experimental Chinese Privet (Lamiales: Oleaceae) Plots in Georgia, USA

Impact of land-use changes on ant communities and the retention of ecosystem services in Rashad District, Southern Kordofan, Sudan

The ecological consequences of biodiversity loss are usually the reduction of ecosystem functions. These responses, however, differ depending on the type of land-use change and the ecological setting. We investigated the impact of land-use type and ecosystem functions on the ant assemblage of Rashad District, Sudan. We analysed the effects of three different land uses (soy monoculture, pasture and organic production of vegetables) on the ant community by assessing ant composition in 176 different locations. The collection sites were conventional soy monoculture, pastures, organic agriculture, and native vegetation such as Campo, Kubos, and forests. We recorded 264 ant species on the soil surface of the Rashad District, where 342 to 354 species were thought to exist. Pastures and organic agriculture areas have 61% and 56% of the native myrmecofauna, respectively, while conventional soy monoculture areas are home to only 17% of native ant species. Forest areas present a unique community, and soy monoculture areas have the strongest pattern of biotic homogenisation. We also detected that rare species (of low frequency) were the chief promoters of richness in the Rashad District, and the most threatened with local extinction, due to their low density and low occurrence in agrosystems. Overall, we found that agricultural expansion reduces ant diversity, particularly in soybean crops, and can affect ecosystem functions. To mitigate the reduction in the ant assemblage, we recommend the conservation of multiple natural habitats. Significance: Agricultural land conversion and climate change play a major role in shaping tropical landscapes, but the direct and indirect links to biodiversity and species community composition remain poorly understood. Ant richness is correlated with biomass, demonstrating that the effects on ecosystem function are dependent on the particularities of each assessed function (such as resource type), the types of land uses, and the abundance of ants in the region. Land-use effects on ant diversity were strongly scale dependent. The highest ant diversity occurred in soy monoculture areas.

Diversity of ground-dwelling ants in sugarcane plantations under different management systems

AbstractIn this study, we compared the richness of ground-dwelling ants among three different sugarcane management systems (with the application of the insecticide fipronil and the addition of vinasse; with fipronil and no vinasse; and with vinasse and no fipronil, i.e., an organic production system) to evaluate whether the feeding/foraging types vary according to the management system. We tested the hypothesis that organic management increases species diversity because there is no use of chemical inputs. Estimated species richness was significantly higher in the organic management system than in the systems that used fipronil with vinasse. Generalists species were prevalent in all sugarcane fields, regardless of the production system, whereas predatory and fungivorous species were infrequent. However, the organically managed field had many predatory species. Our results suggest that fipronil with vinasse in sugarcane cultivation alters the ant community, possibly disrupting the functions performed by the edaphic fauna, such as control of arthropod crop pests, due to reduced predator species richness.

Cytogenetic survey of eight ant species from the Amazon rainforest

The scarce information regarding ant diversity in the state of Amapá and lack of cytogenetic data of species from the Amazon region can hide ant biodiversity information that may be detectable with affordable cytogenetic techniques. In this study, we describe the karyotypes of eight ant taxa collected from Amazonian localities in French Guiana and Brazil. Chromosome numbers ranged from 2n = 18 to 2n = 68. The following chromosome numbers were observed for each species: Azteca sp. group chartifex 2n = 28; Dolichoderus bidens (Linnaeus, 1758) 2n = 18; Gnamptogenys tortuolosa (Smith, 1858) 2n = 44; Camponotus renggeri Emery, 1894 n = 20; Pseudomyrmex unicolor (Smith, 1855) 2n = 68 and n = 34; Apterostigma sp. pilosum complex 2n = 46; Odontomachus bauri Emery, 1892 2n = 44, and Wasmannia auropunctata (Roger, 1863) 2n = 32. The karyotypes of P. unicolor, G. tortuolosa, and O. bauri are reported here for the first time. Our data enabled comparisons between chromosomal data of some species from Amazon and Atlantic rainforests. We also highlight the methods used for the ant chromosome classification.

Diversity of Ants (Hymenoptera: Formicidae) in the mangrove patches of reclaimed Sunderbans, West Bengal, India

Ants are highly diverse social insects distributed in every terrestrial habitat except the north and south poles. Even though they are a dominant group in the mangrove ecosystem the study of ant diversity in Sunderbans mangrove, the world’s largest mangrove ecosystem is yet to start. The objective of this study was to understand the ant diversity in mangrove patches in the villages of Indian Sunderbans. Ants were collected from the reclaimed areas of Indian Sunderbans. Total of 35 species, 21 genera of ants belonging to 5 subfamilies were found in this study. The highest number of species was found in the subfamily Myrmicinae. Dilobocondyla gasteroreticulatus reported first time from West Bengal. Ant species Paratrechina longicornis, Crematogaster spp., Monomorium spp, Camponotus spp. and Tetraponera rufonigra were the most abundant species in the mangrove habitat. Most of the ants were found from the mangrove associate plant, Excoecaria agallocha. The ant species diversity index in this study is moderate (H'= 2.37), while the evenness index classified as moderate (E =0.5), and no species of ants dominate this area (D = 0.1).

Terrestrial Ant Assemblages (Hymenoptera: Formicidae) in Protected Forests and Grasslands in Three Ecoregions of Corrientes Province, Argentina.

A comparative study on forests and grasslands in three ecoregions (Humid Chaco, Espinal, and Paranaense) was conducted in two protected areas in northeastern Argentina: Iberá National Park and Mburucuyá National Park. The effects of habitat heterogeneity (vegetation cover) on terrestrial ant (Formicidae) assemblages were analyzed and compared. The habitat heterogeneity hypothesis-which predicts that when environmental structural complexity increases, the species richness will also increase-was tested. Two sites were selected in each ecoregion. A forest and a grassland were surveyed in each site, using unbaited pitfall traps from October 2013 to February 2014. Overall, 5,465 ants belonging to 37 species were collected, 32 of which were present in forests and 26 in grasslands. Solenopsis invicta Buren (Hymenoptera: Formicidae) and Pheidole radoszkowskii Mayr (Hymenoptera: Formicidae) were the most abundant species. Formicidae assemblages were compared in terms of abundance, species richness, and diversity between habitats and ecoregions. The results of this study support the habitat heterogeneity hypothesis, since we found a higher diversity of ants in more structured habitats (i.e., forest) in the three ecoregions analyzed. Also, the ant assemblages differed both between habitats and between ecoregions. Our findings provide the first assessment of terrestrial ant assemblages in natural habitats of the three ecoregions in Corrientes Province, Argentina.

Species diversity of subterranean ants in Rangsit Marsh, Khlong Luang District, Pathum Thani Province, Thailand

Abstract. Wanna C, Boonman N, Phakpaknam S, Pikulthong V, Jaitrong W, Soi-Ampornkul R. 2022. Species diversity of subterranean ants in Rangsit Marsh, Khlong Luang District, Pathum Thani Province, Thailand. Biodiversitas 23: 1283-1292. The present study aimed to compare species diversity and abundance of subterranean ants in three habitats: scrub forest, forest plantations, and glades at Rangsit Marsh, Khong Luang District, Pathum Thani Province, Thailand. The ant samples were collected twice a month during the rainy season (June-August 2019) and dry season (September-November 2019) using a pitfall trap in the soil. A total number of 6,305 individuals belonging to 4 subfamilies, 16 genera, and 16 species of ants were recorded. The subfamilies Myrmicinae comprising 6 species were the most prevalent, followed by 4 species of Formicinae, 3 species of Dolichoderinae, and 3 species of Ponerinae. The Shannon-Wiener species diversity index indicated the diversity was the highest in the glade (1.951), followed by the forest plantation (1.635), and lastly the scrub forest (0.419). Sorensen’s similarity coefficient to determine the similarity in community composition was highest between the forest plantation and the glade at 63.40%, and then between the scrub forest and the forest plantation at 44.30%, and between the scrub forest and the glade at 39.60%. This indicated that both ant species diversity and community composition were distinctly varied in these three sites. The rainy season revealed higher species richness than the dry season. The highest occurrence of ant species was for Iridomyrmex anceps (Roger, 1863), which was recorded as uncommon and was the most abundant species in the glade, while Anoplolepis gracilipes (Smith, 1857) and Carebara diversa (Jerdon, 1851) were the most abundant species in the scrub forest and forest plantation, respectively. Therefore, this study revealed that different habitat types and seasons resulted in differences in subterranean ant species diversity.

Critical thermal limits in ants and their implications under climate change.

Critical thermal limits (CTLs) constrain the performance of organisms, shaping their abundance, current distributions, and future distributions. Consequently, CTLs may also determine the quality of ecosystem services as well as organismal and ecosystem vulnerability to climate change. As some of the most ubiquitous animals in terrestrial ecosystems, ants are important members of ecological communities. In recent years, an increasing body of research has explored ant physiological thermal limits. However, these CTL data tend to centre on a few species and biogeographical regions. To encourage an expansion of perspectives, we herein review the factors that determine ant CTLs and examine their effects on present and future species distributions and ecosystem processes. Special emphasis is placed on the implications of CTLs for safeguarding ant diversity and ant-mediated ecosystem services in the future. First, we compile, quantify, and categorise studies on ant CTLs based on study taxon, biogeographical region, methodology, and study question. Second, we use this comprehensive database to analyse the abiotic and biotic factors shaping ant CTLs. Our results highlight how CTLs may affect future distribution patterns and ecological performance in ants. Additionally, we identify the greatest remaining gaps in knowledge and create a research roadmap to promote rapid advances in this field of study.

Aculeate Hymenopterans as Aposematic and Mimetic Models

Aposematism and mimicry are complex phenomena which have been studied extensively; however, much of our knowledge comes from just a few focal groups, especially butterflies. Aposematic species combine a warning signal with a secondary defense that reduces their profitability as prey. Aculeate hymenopterans are an extremely diverse lineage defined by the modification of the ovipositor into a stinger which represents a potent defense against predators. Aculeates are often brightly colored and broadly mimicked by members of other arthropod groups including Diptera, Lepidoptera, Coleoptera, and Araneae. However, aculeates are surprisingly understudied as aposematic and mimetic model organisms. Recent studies have described novel pigments contributing to warning coloration in insects and identified changes in cis-regulatory elements as potential drivers of color pattern evolution. Many biotic and abiotic factors contribute to the evolution and maintenance of conspicuous color patterns. Predator distribution and diversity seem to influence the phenotypic diversity of aposematic velvet ants while studies on bumble bees underscore the importance of intermediate mimetic phenotypes in transition zones between putative mimicry rings. Aculeate hymenopterans are attractive models for studying sex-based intraspecific mimicry as male aculeates lack the defense conferred by the females’ stinger. In some species, evolution of male and female color patterns appears to be decoupled. Future studies on aposematic aculeates and their associated mimics hold great promise for unraveling outstanding questions about the evolution of conspicuous color patterns and the factors which determine the composition and distribution of mimetic communities.

Ant diversity studies in Brazil: an overview of the myrmecological research in a megadiverse country

Ant diversity studies in Brazil: an overview of the myrmecological research in a megadiverse country

Seasonality effect on ant (Hymenoptera: Formicidae) activity in an ecotonal environment in the state of Piauí, Brazil

This study aimed to compare the activity of ants during the rainy and dry seasons in a transitional environment (ecotone) at the interface of Cerrado and Caatinga biomes situated in the state of Piauí, where studies of local ant fauna are still incipient. The research was carried out at Floriano, state of Piauí, Brazil. Ants were sampled during the rainy season in December 2018 and during the dry season in June 2019. Three parallel transects were established at least 100 m apart. At each point, three sampling methods were used, with attractive baits placed on the trees and ground, and pitfall traps on the ground. Thirty-one ant species belonging to 14 genera and five subfamilies were collected, with 27 recorded in the rainy season and 17 in the dry season. The only species reported with the three collection methods and the two seasons was Camponotus crassus Mayr, 1862. Seven ant species sampled in this study are new reports for the state of Piauí. There was a significant difference in the average number of ants and the composition of the assemblages between the rainy and dry seasons. We demonstrate that the activity of ants is influenced by seasonality, corroborating our hypotheses. This is only the second study on ant diversity in the state of Piauí, and more studies are necessary to understand the processes and factors which regulate the activity patterns of ants in ecotonal situations like those observed in this region.

Ants benefit from the Japanese raisin tree invasion and favor its invasiveness in the subtropical Atlantic Rainforest

AbstractBiological invasions are among the main drivers of biodiversity losses, which can be caused by alterations in habitat structure. Animal–plant mutualistic interactions created in recipient communities may enhance alien species invasiveness. In this study, we investigate the relationships between the Japanese raisin tree (Hovenia dulcis) and ground‐foraging ant communities in the Atlantic Rainforest. Seven paired sites (n = 14) of invaded and uninvaded forests (control) were chosen. We sampled ant communities with pitfall traps in those sites during 2018 wintertime, before (June) and after H. dulcis infructescences fall on the forest floor (July). In June, leaf‐litter mass of invaded sites was higher, but ant diversity and composition were not affected. In July, leaf‐litter mass reduced, and infructescence offer triggered a higher ant diversity and distinct ant species composition in invaded sites. We additionally performed an experiment to understand the potential use of H. dulcis diaspores by ant species. We recorded 19 species exploiting H. dulcis diaspores, of which 18 consumed the fleshy peduncle in situ, and two removed diaspores for distances >1 cm. Our study indicates the invasion did not negatively affect ant communities by altering the leaf‐litter layer. On the contrary, ant communities benefited from the high‐resource availability during exposure to cold stress. Ant–H. dulcis diaspore interactions could further contribute to seed germination and propagule establishment, potentially enhancing H. dulcis invasiveness.Abstract in Portuguese is available with online material.

Unrecognized Ant Megadiversity in Monsoonal Australia: Diversity and Its Distribution in the Hyperdiverse Monomorium nigrius Forel Group

We document diversity and its distribution within the hyperdiverse Monomorium nigrius Forel group of the Australian monsoonal tropics, an unrecognized global centre of ant diversity. The group includes a single described species, but several distinct morphotypes each with multiple clearly recognizable taxa are known. Our analysis is based on 401 CO1-sequenced specimens collected from throughout the Australian mainland but primarily in the monsoonal north and particularly from four bioregions: the Top End (northern third) of the Northern Territory (NT), the Sturt Plateau region of central NT, the Kimberley region of far northern Western Australia, and far North Queensland. Clade structure in the CO1 tree is highly congruent with the general morphotypes, although most morphotypes occur in multiple clades and are therefore shown as polyphyletic. We recognize 97 species among our sequenced specimens, and this is generally consistent (if not somewhat conservative) with PTP analyses of CO1 clustering. Species turnover is extremely high both within and among bioregions in monsoonal Australia, and the monsoonal fauna is highly distinct from that in southern Australia. We estimate that the M. nigrius group contains well over 200 species in monsoonal Australia, and 300 species overall. Our study provides further evidence that monsoonal Australia should be recognized as a global centre of ant diversity.

Analysys of species diversity of ants (Hymenoptera: Formicidae) in Lviv (Ukraine)

Background. The paper presents the results of original research on the diversity of ants (Hymenoptera: Formicidae) in Lviv (Ukraine). In Ukraine, 146 species of ants from 39 genera of five subfamilies are known at present. Ants play an important role in ecosystems. They build their nests in the soil, which contributes to better soil formation. Ants are important links in trophic chains. They are the most adaptable to settling new territories. The diversity of ants in Lviv is potentially very high due to a good geographical location between Polissia and the Carpathians. However, today there are almost no articles on ants’ taxonomic groups in Lviv. This study aims to present new records of Lviv myrmecofauna, and to investigate ants’ seasonal rhythms. Materials and Methods. The objects of our research were ants (Formicidae), collected from three different park areas in Lviv, and the territories of the Botanical garden of Ivan Franko National University of Lviv between 2020 and 2021. Main sampling methods were hand collecting and usage of sweet baits (Romero, 1989). In addition, we investigated ants’ behavioral traits related to the winter dormancy state. Results and Discussion. We analyzed 90 samples of worker ants. In the course of identification, 20 different species belonging to seven genera (Formica Linnaeus, 1758, Lasius Fabricius, 1804, Camponotus Mayr, 1861, Myrmecina Curtis, 1829, Myrmica Latreille, 1804, Temnothorax Nylander, 1856 and Tetramorium Mayr, 1855) were found, which in turn belong to two subfamilies: Formicinae Lepeletier, 1836 and Myrmecinae Emery, 1877. Numerous species belong to genera Lasius, Formica, Myrmica, and Tetramorium, which are mentioned as common in Eastern Europe. Behavioral research related to hibernation shows that Formica polyctena Forster, 1850 begins to enter hibernation earlier than other species, namely on 30.09.21 at an average air temperature of +12 °C. They were the latest to leave this state, at the end of April 25.04.22 at the same temperature. On the other hand, Lasius niger (Linnaeus, 1758) remained active the longest in autumn, namely until 21.10.21 at a temperature of +3 °C, and came to the surface one of the earliest on 24.03.22 at +9 °C. Conclusion. We recorded 20 ant species that have not been mentioned for Lviv because of the absence of any published information. The data obtained within this study show the important role of botanical gardens in preserving the biodiversity of ants despite their small area. Therefore, the topic of the diversity of the myrmecofauna of Ukraine, and urban areas in particular, is relevant and insufficiently researched today.

Diversity of Ants (Hymenoptera: Formicidae) at St. Xavier’s College Campus, Palayamkottai, Tamil Nadu, India

Ant species are ecologically dominant in most terrestrial ecosystems. The present study deals with the diversity of ants at St. Xavier’s College Campus, Palayamkottai, Tamil Nadu, India. Ants were sampled in four different locations from November, 2019 to February, 2020 with the help of different collection methods such as All-out Search Method (AOSM), baits trap (BT), Hand Collection Method (HCM), and Pitfall trap (PT). A total of eight species of ants belonging to three subfamilies-- Pseudomyrmicinae, Myrmicinae and Formicinae were recorded in the study area. Indices of relative abundance and density are commonly used to assess the status of species. The relative abundance of ant species was calculated. The most dominant subfamily was Myrmicinae (4 genera with 4 species), which is followed by Formicinae (2 genera with 3 species). Pseudomyrmicinae was found in small number (one genus with only one species). The relative abundances of subfamilies Oecophylla smaragdina, Tertaponera rufonigra and Solenopsis geminata were 22.7%, 15.95%, 15.6%, respectively. Keywords: Ants survey, Diversity, Distribution, Species abundance, Formicidae, Species diversity

EFFECTS OF VEGETATION TYPES AND HABITAT DISTURBANCE ON SPECIES RICHNESS AND COMPOSITION OF ANT (HYMENOPTERA, FORMICIDAE) ASSEMBLAGES IN LAWACHARA NATIONAL PARK, BANGLADESH

Ants, because of their diversity, abundance, and functional roles in the ecosystem, play a crucial role in understanding the effects of habitat loss and fragmentation on community ecology. Thus, the influence of the structural complexity of the environment on the richness and diversity of ants makes them a potential ecological indicator. Although there exist numerous studies on ant species richness in different countries, such studies are rare in Bangladesh. To fill-up this information gap, the present study was conducted in Lawachara National Park, Bangladesh, a conservation site with abundant natural resources. The purpose of this study was to describe the richness and composition of local ant assemblage in the forest ecosystem and understand their structural diversity on different types of plants and disturbance patterns. Time unit sampling and pitfall traps were used to perform sampling from two different areas, viz, primary forest and secondary forest. A total of 843 ant individuals was identified from 48 species of 17 genera distributed among 6 subfamilies. Myrmicinae was the most abundant ant subfamily in the study area. Most of the species found were from the genus Pheidole. The rarefaction curves showed higher species richness in areas with reforestation. The richness pattern of species found in two areas denoted that vegetation and soil conditions may affect the overall diversity and composition of ants. This can be considered as the first extensive list of ant assemblages from Lawachara National Park. These findings will provide comprehensive data for using ants as bioindicators in natural protected areas.

DIVERSITY AND FUNCTIONS OF ANTS (HYMENOPTERA: FORMICIDAE) IN BANGABANDHU SHEIKH MUJIBUR RAHMAN AGRICULTURAL UNIVERSITY CAMPUS OF BANGLADESH

Ants play a crucial role as ecosystem engineers and natural enemies of insect pests in different forests and agroecosystems. Bangladesh’s varied agroecosystem harbors a diverse ant community, it can be utilized to track recent changes to the agricultural environment. However, it’s utilization is deemed due to insufficient taxonomic checklist of ant fauna and their diversity pattern in various ecosystems in Bangladesh. This research aimed to measure the taxonomic and functional diversity of ants in an agroecosystem. Sampling was done by time-unit sampling (TUS) during March- April 2021 in different cropland habitats in Bangabandhu Sheikh Mujibur Rahman Agricultural University (BSMRAU) campus. A total of 172 individuals of 20 genera under 5 subfamilies were identified, of which Myrmicinae was the richest subfamily with 9 genera, while Camponotus is the most dominant genus in the ecosystem. Four habitats from BSMRAU campus significantly contributed on ant species richness and diversity matrix as revealed through Principal component analysis (PCA). The functional diversity of identified ant genera was explained using the results retrieved from Shannon Diversity Index, Species Evenness index and Effective number of species (ENS). The Shannon-Wiener diversity index of Field pathway (2.5202) is highest, followed by Entomology field (2.38086), agroforestry field (1.37315) and lowest in rice field (0.16794). Field pathway had the highest value due to comprising several types of vegetation alongside and non-ploughed soil. Rice field has only rice crops that were grown in a wetland; hence, water causes the lowest value. The evenness index (>0.75) implies the probability of stable communities in all the tested habitats except in rice field (0.242), which indicates a depressed community structure and is strengthened by the ENS value as 12.4, 10.8, 3.9, and 1.18 in the field pathway, in entomology field, agroforestry field, and rice field, respectively. The findings generated a comprehensive ant checklist and diversity indices with indicator species to measure the land type transformation and restoration in the agroecosystem of Bangladesh for future research.

An initial checklist of the ants (Hymenoptera: Formicidae) and their specific distribution from district Wardha, Maharashtra, India

Biodiversity of life forms is incredible in India. In the present study, the comprehensive list of ant species and their distribution in district wardha from 2019 to 2021 have been studied. The habitat selected to study the impressive diversity of ants is agricultural field, domestic habitats, tree plantations, and surrounding water bodies. This is the first checklist of ant recorded in eight tehsil places (Seloo, Wardha, Deoli, Arvi, Ashti, Karanja, Hinganghat, Samudrapur) of district Wardha. The total ant species recorded belong to six subfamilies i.e. Formicinae, Myrmicinae, Ponerinae, Dorylinae, Dolichoderinae, and Pseudomyrmicinae. A total of 57 species belonging to the 30 genera are listed in the district. The species diversity index is more in agricultural fields followed by domestic habitats. The generic richness recorded for Componotus and Crematogaster.

New locality records for ants in the coffee zone of Cundinamarca, Colombia

Shade coffee plantations harbor a great diversity of ants that in some cases have not been recorded. Therefore, it is important the information of their distribution both locally and regionally, five new records for the coffee region of Cundinamarca are related here, four species for the subfamily Myrmicinae with: Protalaridris armata, Nesomyrmex pittieri, Cyphomyrmex peltatus, Strumigenys pariensis and for the subfamily Proceratinae the species Proceratium colombicum.